929 lines
147 KiB
XML
929 lines
147 KiB
XML
<document id="52CF9F9FDA48B5692267472FDEAB7424" ID-CLB-Dataset="276833" ID-DOI="10.11646/zootaxa.5209.3.7" ID-GBIF-Dataset="f6643467-b4da-45f8-8896-94409dc6d74d" ID-ISSN="1175-5326" ID-Zenodo-Dep="7329737" ID-ZooBank="0576E9E6-22F1-4A77-ACC8-439662DF53FC" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1668669530257" checkinUser="plazi" docAuthor="Wörheide, Gert, Kaltenbacher, Emilie, Cowan, Zara-Louise & Haszprunar, Gerhard" docDate="2022" docId="817B87971B48FF90AEAE5E1DFE68F8C6" docLanguage="en" docName="zootaxa.5209.3.7.pdf" docOrigin="Zootaxa 5209 (3)" docStyle="DocumentStyle:647186512141C8FC8976D5BCC54AEB7D.9:Zootaxa.2013-.journal_article" docStyleId="647186512141C8FC8976D5BCC54AEB7D" docStyleName="Zootaxa.2013-.journal_article" docStyleVersion="9" docTitle="Acanthaster benziei Wörheide & Kaltenbacher & Cowan & Haszprunar 2022, sp. nov." docType="treatment" docVersion="6" lastPageNumber="390" masterDocId="7D42FFEF1B4CFF9BAE395A6BFFAFFFB0" masterDocTitle="A new species of crown-of-thorns sea star, Acanthaster benziei sp. nov. (Valvatida Acanthasteridae), from the Red Sea" masterLastPageNumber="393" masterPageNumber="379" pageNumber="383" updateTime="1699414052955" updateUser="ExternalLinkService" zenodo-license-document="CLOSED">
|
||
<mods:mods id="B322A3CD0F1C8AB745535ABA5CA2E6A6" xmlns:mods="http://www.loc.gov/mods/v3">
|
||
<mods:titleInfo id="E9FECDD509B86034063312D19F433766">
|
||
<mods:title id="2A9ECAFB20B2EDCE513BC850FB913853">A new species of crown-of-thorns sea star, Acanthaster benziei sp. nov. (Valvatida Acanthasteridae), from the Red Sea</mods:title>
|
||
</mods:titleInfo>
|
||
<mods:name id="20B080A66D1EFF26DFC30438127BA748" type="personal">
|
||
<mods:role id="79A3B2BE8614E41B277847380B2AB768">
|
||
<mods:roleTerm id="BBA91F5B972DEE64010D4DE20C4AA1F9">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="628F0FD138FC0ACE00D0794E7A724DE8">Wörheide, Gert</mods:namePart>
|
||
<mods:affiliation id="4110888882E3F01BA5BE811D5621FCA2">Department of Earth and Environmental Sciences Palaeontology and Geobiology, Ludwig-Maximilians-Universität München, Munich, Germany. & GeoBio-Center, Ludwig-Maximilians-Universität München, Munich, Germany. & SNSB-Bavarian State Collection of Palaeontology and Geology, Munich, Germany.</mods:affiliation>
|
||
</mods:name>
|
||
<mods:name id="B34A521BA5CBF09AB5FF7921300871F3" type="personal">
|
||
<mods:role id="45680FC1FF9236F559EC82168E873BFF">
|
||
<mods:roleTerm id="0FB677ECFF4D19CAC8F632314B9F97A2">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="2FAF8DA2629E3D09A625DF7C66B8ECB4">Kaltenbacher, Emilie</mods:namePart>
|
||
<mods:affiliation id="73BA6A0C4CC26B417BF87B3802010510">Department of Earth and Environmental Sciences Palaeontology and Geobiology, Ludwig-Maximilians-Universität München, Munich, Germany.</mods:affiliation>
|
||
</mods:name>
|
||
<mods:name id="85A3BF2678126C3621E19ABBC749F04D" type="personal">
|
||
<mods:role id="9E02BDA24B2AA17C37A79F90BC93FD94">
|
||
<mods:roleTerm id="112C715AEAE5DCF84D62ADC5A4CE53F8">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="D451F8EC7AD7B875C0ED44A96561793B">Cowan, Zara-Louise</mods:namePart>
|
||
<mods:affiliation id="6FE5365C9E11B0B1D3EADB738D8885F2">Department of Earth and Environmental Sciences Palaeontology and Geobiology, Ludwig-Maximilians-Universität München, Munich, Germany.</mods:affiliation>
|
||
</mods:name>
|
||
<mods:name id="B281B7CCF23E698FFFC8D88D08A8DD65" type="personal">
|
||
<mods:role id="DDCDA0B4AF6CCC60175F97773913FEA5">
|
||
<mods:roleTerm id="F5274D27CF1524605805F68534EC6BA9">Author</mods:roleTerm>
|
||
</mods:role>
|
||
<mods:namePart id="4C1C42001D3CF8D5265112E8A31BB280">Haszprunar, Gerhard</mods:namePart>
|
||
<mods:affiliation id="70D9BE1D5EAA2784D4CF6EC7C28A5BDB">GeoBio-Center, Ludwig-Maximilians-Universität München, Munich, Germany. & SNSB-Zoological State Collections, Munich, Germany.</mods:affiliation>
|
||
</mods:name>
|
||
<mods:typeOfResource id="153BE9EF479DAD537D63BA587A12E17C">text</mods:typeOfResource>
|
||
<mods:relatedItem id="A716340F56869E56A82DDE3BEBD3D654" type="host">
|
||
<mods:titleInfo id="DEDEE298C69C09A33515B289EAEA1959">
|
||
<mods:title id="A9D4427EB654E89FBF620358DEAA4364">Zootaxa</mods:title>
|
||
</mods:titleInfo>
|
||
<mods:part id="7B49271E4E24C4A77CF920AE12DB7F91">
|
||
<mods:date id="3EB59DBB2C0671436D2F3D1030A4A501">2022</mods:date>
|
||
<mods:detail id="DAACD1AC98A6145285454B43A01DB7F8" type="pubDate">
|
||
<mods:number id="72DB1F691341B8D06AFBE0D2DEAFEFA2">2022-11-17</mods:number>
|
||
</mods:detail>
|
||
<mods:detail id="5DB33DC29186D42137D0E07B6ABAFD17" type="volume">
|
||
<mods:number id="106035DC69117914B1DD549EC13B9751">5209</mods:number>
|
||
</mods:detail>
|
||
<mods:detail id="1A2D9BDB384F6EFE6168EF3C0E513F38" type="issue">
|
||
<mods:number id="A67A1B404CDC069AEC8C1A339DE8C5F4">3</mods:number>
|
||
</mods:detail>
|
||
<mods:extent id="E2E0F363F5AD519D2C9C5FB7CDEA5F26" unit="page">
|
||
<mods:start id="1536719DE79B5520832F38483FB6EFC4">379</mods:start>
|
||
<mods:end id="237ACFFF5B20EA4E0EDBF88C4AC7D9A2">393</mods:end>
|
||
</mods:extent>
|
||
</mods:part>
|
||
</mods:relatedItem>
|
||
<mods:classification id="CDE420AAD2544402821BAEE23EBEBAFF">journal article</mods:classification>
|
||
<mods:identifier id="E1A8172D86817B3B88FC3D89D741788B" type="CLB-Dataset">276833</mods:identifier>
|
||
<mods:identifier id="391D927754DE1794DC66E3B760676FC7" type="DOI">10.11646/zootaxa.5209.3.7</mods:identifier>
|
||
<mods:identifier id="7457C33BA24244245C32B4F86E9BDEB7" type="GBIF-Dataset">f6643467-b4da-45f8-8896-94409dc6d74d</mods:identifier>
|
||
<mods:identifier id="F373ABE915E44477CB6A33AAB927653A" type="ISSN">1175-5326</mods:identifier>
|
||
<mods:identifier id="C21080873B74C886145560829AE8038A" type="Zenodo-Dep">7329737</mods:identifier>
|
||
<mods:identifier id="D1C21E0D152CF1DA4C2B1656F8C420BC" type="ZooBank">0576E9E6-22F1-4A77-ACC8-439662DF53FC</mods:identifier>
|
||
</mods:mods>
|
||
<treatment id="817B87971B48FF90AEAE5E1DFE68F8C6" ID-DOI="http://doi.org/10.5281/zenodo.7334240" ID-GBIF-Taxon="217323666" ID-Zenodo-Dep="7334240" LSID="urn:lsid:plazi:treatment:817B87971B48FF90AEAE5E1DFE68F8C6" httpUri="http://treatment.plazi.org/id/817B87971B48FF90AEAE5E1DFE68F8C6" lastPageId="11" lastPageNumber="390" pageId="4" pageNumber="383">
|
||
<subSubSection id="41C8650A1B48FF9FAEAE5E1DFBB0FB62" pageId="4" pageNumber="383" type="nomenclature">
|
||
<paragraph id="096D36811B48FF9FAEAE5E1DFCEFFB20" blockId="4.[151,832,1142,1169]" box="[151,832,1142,1169]" pageId="4" pageNumber="383">
|
||
<heading id="522581ED1B48FF9FAEAE5E1DFCEFFB20" bold="true" box="[151,832,1142,1169]" fontSize="11" level="1" pageId="4" pageNumber="383" reason="1">
|
||
<emphasis id="3BA6EA931B48FF9FAEAE5E1DFCEFFB20" bold="true" box="[151,832,1142,1169]" pageId="4" pageNumber="383">
|
||
<taxonomicName id="CED24D021B48FF9FAEAE5E1DFE29FB21" ID-CoL="e9966371-fd1a-42c0-ac85-cdf0d7b0f138" authority="Wörheide & Kaltenbacher & Cowan & Haszprunar, 2022" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[151,390,1142,1169]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="4" pageNumber="383" phylum="Echinodermata" rank="species" species="benziei" status="sp. nov.">
|
||
<emphasis id="3BA6EA931B48FF9FAEAE5E1DFE29FB21" bold="true" box="[151,390,1142,1169]" italics="true" pageId="4" pageNumber="383">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="209557E81B48FF9FAFB45E1DFE44FB20" box="[397,491,1142,1168]" pageId="4" pageNumber="383" rank="species">sp. nov.</taxonomicNameLabel>
|
||
Wörheide & Kaltenbacher
|
||
</emphasis>
|
||
</heading>
|
||
</paragraph>
|
||
<paragraph id="096D36811B48FF9FAEAE5ED0FBB0FB62" blockId="4.[151,1055,1210,1235]" box="[151,1055,1210,1235]" pageId="4" pageNumber="383">
|
||
Zoobank LSID:
|
||
<uri id="7D433A831B48FF9FAF045ED1FBB0FB62" box="[317,1055,1210,1234]" pageId="4" pageNumber="383">
|
||
urn:lsid:zoobank.org:act:
|
||
<uuid id="7D740C541B48FF9FAC0C5ED1FBB0FB62" box="[565,1055,1210,1234]" pageId="4" pageNumber="383">4C462EF3-39AF-4767-96DF-C3B8CC5D9388</uuid>
|
||
</uri>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="41C8650A1B48FF9FAEAE5F6AFA23FAAB" box="[151,1420,1281,1307]" pageId="4" pageNumber="383" type="description">
|
||
<paragraph id="096D36811B48FF9FAEAE5F6AFA23FAAB" blockId="4.[151,1437,1281,2027]" box="[151,1420,1281,1307]" pageId="4" pageNumber="383">
|
||
<emphasis id="3BA6EA931B48FF9FAEAE5F6AFE96FAAB" bold="true" box="[151,313,1281,1307]" pageId="4" pageNumber="383">Formal name.</emphasis>
|
||
<taxonomicName id="CED24D021B48FF9FAF065F6AFC9CFAAB" authority="Worheide & Kaltenbacher" authorityName="Worheide & Kaltenbacher" box="[319,819,1281,1307]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="4" pageNumber="383" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B48FF9FAF065F6AFDBEFAAB" box="[319,529,1281,1307]" italics="true" pageId="4" pageNumber="383">Acanthaster benziei</emphasis>
|
||
Wörheide & Kaltenbacher
|
||
</taxonomicName>
|
||
in Wörheide, Kaltenbacher, Cowan & Haszprunar 2022
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="41C8650A1B48FF9FAEFE5F4EFD39FA1B" pageId="4" pageNumber="383" type="etymology">
|
||
<paragraph id="096D36811B48FF9FAEFE5F4EFD39FA1B" blockId="4.[151,1437,1281,2027]" pageId="4" pageNumber="383">
|
||
<emphasis id="3BA6EA931B48FF9FAEFE5F4EFEE1FA8F" bold="true" box="[199,334,1317,1343]" pageId="4" pageNumber="383">Etymology.</emphasis>
|
||
The species name pays tribute to Professor John Benzie, who has decisively promoted research on CoTS, with numerous publications and his own collection. He was among the first scientists to genetically analyse
|
||
<taxonomicName id="CED24D021B48FF9FAEAE5F06FEFBFA37" box="[151,340,1389,1415]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="4" pageNumber="383" phylum="Echinodermata" rank="species" species="undetermined">
|
||
<emphasis id="3BA6EA931B48FF9FAEAE5F06FEB1FA37" box="[151,286,1389,1415]" italics="true" pageId="4" pageNumber="383">Acanthaster</emphasis>
|
||
spp.
|
||
</taxonomicName>
|
||
and his collection was the basis of the work of
|
||
<bibRefCitation id="6D434B701B48FF9FAD555F06FBE4FA37" author="Vogler, C. & Benzie, J. & Lessios, H. & Barber, P. & Worheide, G." box="[876,1099,1389,1415]" pageId="4" pageNumber="383" pagination="696 - 699" refId="ref9248" refString="Vogler, C., Benzie, J., Lessios, H., Barber, P. & Worheide, G. (2008) A threat to coral reefs multiplied? Four species of crownof-thorns starfish. Biology Letters, 4 (6), 696 - 699. https: // doi. org / 10.1098 / rsbl. 2008.0454" type="journal article" year="2008">
|
||
Vogler
|
||
<emphasis id="3BA6EA931B48FF9FAD875F05FC58FA37" box="[958,1015,1389,1415]" italics="true" pageId="4" pageNumber="383">et al.</emphasis>
|
||
(2008)
|
||
</bibRefCitation>
|
||
, which represents a milestone in the species identification of these sea stars.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="41C8650A1B48FF9FAEFE5FDEFDDAF85B" pageId="4" pageNumber="383" type="materials_examined">
|
||
<paragraph id="096D36811B48FF9FAEFE5FDEFB2AF98B" blockId="4.[151,1437,1281,2027]" pageId="4" pageNumber="383">
|
||
<materialsCitation id="B9BA3CDC1B48FF9FAEFE5FDEFB2AF98B" ID-GBIF-Occurrence="4435992302" country="Saudi Arabia" county="Farasan" elevation="10" latitude="18.841667" location="Sara Campana" longLatPrecision="1" longitude="40.781387" municipality="Miskah" pageId="4" pageNumber="383" specimenCount="10" typeStatus="holotype">
|
||
<emphasis id="3BA6EA931B48FF9FAEFE5FDEFE94FA7F" bold="true" box="[199,315,1461,1487]" pageId="4" pageNumber="383">
|
||
<typeStatus id="D66988231B48FF9FAEFE5FDEFE99FA7F" box="[199,310,1461,1487]" pageId="4" pageNumber="383" type="holotype">Holotype</typeStatus>
|
||
.
|
||
</emphasis>
|
||
SNSB-BSPG.GW.4202, adult individual (
|
||
<figureCitation id="91E92A041B48FF9FAD225FDEFCD9FA7F" box="[795,886,1461,1487]" captionStart="FIGURE 2" captionStartId="5.[152,255,1323,1347]" captionTargetBox="[151,1436,181,1300]" captionTargetId="figure-19@5.[151,1436,181,1300]" captionTargetPageId="5" captionText="FIGURE 2. Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm." figureDoi="http://doi.org/10.5281/zenodo.7329743" httpUri="https://zenodo.org/record/7329743/files/figure.png" pageId="4" pageNumber="383">Fig. 2A</figureCitation>
|
||
), collected in 2017 by
|
||
<location id="0C0D605A1B48FF9FAA405FDEFAB2FA7F" LSID="urn:lsid:plazi:treatment:817B87971B48FF90AEAE5E1DFE68F8C6:0C0D605A1B48FF9FAA405FDEFAB2FA7F" box="[1145,1309,1461,1487]" country="Saudi Arabia" county="Farasan" latitude="18.841667" longLatPrecision="1" longitude="40.781387" municipality="Miskah" name="Sara Campana" pageId="4" pageNumber="383">Sara Campana</location>
|
||
and
|
||
<location id="0C0D605A1B48FF9FAB6C5FDEFF7CFA43" LSID="urn:lsid:plazi:treatment:817B87971B48FF90AEAE5E1DFE68F8C6:0C0D605A1B48FF9FAB6C5FDEFF7CFA43" country="Saudi Arabia" county="Farasan" latitude="18.841667" longLatPrecision="1" longitude="40.781387" municipality="Miskah" name="Oliver Voigt" pageId="4" pageNumber="383">Oliver Voigt</location>
|
||
at
|
||
<collectingMunicipality id="E909ACFB1B48FF9FAEC15FB2FEE0FA43" box="[248,335,1497,1523]" pageId="4" pageNumber="383">Miskah</collectingMunicipality>
|
||
,
|
||
<collectingCounty id="E00C4E0D1B48FF9FAF655FB2FE19FA43" box="[348,438,1497,1523]" pageId="4" pageNumber="383">Farasan</collectingCounty>
|
||
,
|
||
<collectingCountry id="71C576111B48FF9FAFFB5FB2FDFAFA43" box="[450,597,1497,1523]" name="Saudi Arabia" pageId="4" pageNumber="383">Saudi Arabia</collectingCountry>
|
||
(
|
||
<geoCoordinate id="6CE650461B48FF9FAC5C5FB2FD55FA43" box="[613,762,1497,1523]" degrees="18.84166667" direction="north" orientation="latitude" pageId="4" pageNumber="383" precision="1" value="18.841667">18.84166667</geoCoordinate>
|
||
/
|
||
<geoCoordinate id="6CE650461B48FF9FAD2B5FB2FC08FA43" box="[786,935,1497,1523]" degrees="40.78138889" direction="east" orientation="longitude" pageId="4" pageNumber="383" precision="1" value="40.781387">40.78138889</geoCoordinate>
|
||
) in a water depth of
|
||
<specimenCount id="1FD4FD081B48FF9FAAAE5FB2FB77FA43" box="[1175,1240,1497,1523]" pageId="4" pageNumber="383" type="generic">
|
||
<quantity id="CE2A9B641B48FF9FAAAE5FB2FB7BFA43" box="[1175,1236,1497,1523]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="4" pageNumber="383" unit="m" value="10.0">
|
||
<elevation id="82FFD1B21B48FF9FAAAE5FB2FB7BFA43" box="[1175,1236,1497,1523]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="4" pageNumber="383" unit="m" value="10.0">10 m</elevation>
|
||
</quantity>
|
||
.
|
||
</specimenCount>
|
||
The sea star was narcotized with menthol, fixed in 4% formaldehyde, and preserved in 70% EtOH. Some tube feet for DNA analyses were preserved in 95% EtOH and are stored at SNSB-BSPG together with the specimens.
|
||
</materialsCitation>
|
||
</paragraph>
|
||
<paragraph id="096D36811B48FF9FAEFE5C2EFD74F97B" blockId="4.[151,1437,1281,2027]" pageId="4" pageNumber="383">
|
||
<materialsCitation id="B9BA3CDC1B48FF9FAEFE5C2EFD78F97B" ID-GBIF-Occurrence="4435992303" country="Saudi Arabia" elevation="10" latitude="20.12456" location="Coast" longLatPrecision="1" longitude="40.258747" municipality="AlLith" pageId="4" pageNumber="383" specimenCount="1" typeStatus="paratype">
|
||
<emphasis id="3BA6EA931B48FF9FAEFE5C2EFEE9F9EF" bold="true" box="[199,326,1605,1631]" pageId="4" pageNumber="383">
|
||
<typeStatus id="D66988231B48FF9FAEFE5C2EFEEEF9EF" box="[199,321,1605,1631]" pageId="4" pageNumber="383" type="paratype">Paratypes</typeStatus>
|
||
.
|
||
</emphasis>
|
||
SNSB-BSPG.GW.4081, adult individual (
|
||
<figureCitation id="91E92A041B48FF9FAD015C2EFC39F9EF" box="[824,918,1605,1631]" captionStart="FIGURE 2" captionStartId="5.[152,255,1323,1347]" captionTargetBox="[151,1436,181,1300]" captionTargetId="figure-19@5.[151,1436,181,1300]" captionTargetPageId="5" captionText="FIGURE 2. Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm." figureDoi="http://doi.org/10.5281/zenodo.7329743" httpUri="https://zenodo.org/record/7329743/files/figure.png" pageId="4" pageNumber="383">Fig. 2B</figureCitation>
|
||
), collected from
|
||
<location id="0C0D605A1B48FF9FAA5D5C2EFB0BF9EF" LSID="urn:lsid:plazi:treatment:817B87971B48FF90AEAE5E1DFE68F8C6:0C0D605A1B48FF9FAA5D5C2EFB0BF9EF" box="[1124,1188,1605,1631]" country="Saudi Arabia" latitude="20.12456" longLatPrecision="1" longitude="40.258747" municipality="AlLith" name="Coast" pageId="4" pageNumber="383">Coast</location>
|
||
guard reef, near
|
||
<collectingMunicipality id="E909ACFB1B48FF9FAB4E5C2EFF67F933" pageId="4" pageNumber="383">AlLith</collectingMunicipality>
|
||
,
|
||
<collectingCountry id="71C576111B48FF9FAEE15C02FEC3F933" box="[216,364,1641,1667]" name="Saudi Arabia" pageId="4" pageNumber="383">Saudi Arabia</collectingCountry>
|
||
(
|
||
<geoCoordinate id="6CE650461B48FF9FAF465C02FE57F934" box="[383,504,1641,1668]" degrees="20.124560" direction="north" orientation="latitude" pageId="4" pageNumber="383" precision="1" value="20.12456">20.124560</geoCoordinate>
|
||
/
|
||
<geoCoordinate id="6CE650461B48FF9FAC2C5C02FD21F934" box="[533,654,1641,1668]" degrees="40.258746" direction="east" orientation="longitude" pageId="4" pageNumber="383" precision="1" value="40.258747">40.258746</geoCoordinate>
|
||
) in a water depth of <
|
||
<quantity id="CE2A9B641B48FF9FADA25C02FC76F933" box="[923,985,1641,1667]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="4" pageNumber="383" unit="m" value="10.0">
|
||
<elevation id="82FFD1B21B48FF9FADA25C02FC76F933" box="[923,985,1641,1667]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="4" pageNumber="383" unit="m" value="10.0">10 m</elevation>
|
||
</quantity>
|
||
, narcotized with menthol, fixed in 4% formaldehyde, and preserved in 70% EtOH. Some tube feet for DNA analyses were preserved in 95% EtOH and are stored at SNSB-BSPG together with the specimens
|
||
</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="096D36811B48FF9FAEFE5CBEFD8AF8EB" blockId="4.[151,1437,1281,2027]" pageId="4" pageNumber="383">
|
||
<materialsCitation id="B9BA3CDC1B48FF9FAEFE5CBEFD8EF8EB" ID-GBIF-Occurrence="4435992301" country="Saudi Arabia" elevation="10" latitude="19.094444" location="Mubarak" longLatPrecision="1" longitude="40.379166" municipality="Farasan Banks" pageId="4" pageNumber="383" specimenCount="1">
|
||
SNSB-BSPG.GW.4168, adult individual (
|
||
<figureCitation id="91E92A041B48FF9FAC935CBEFCA7F95F" box="[682,776,1749,1775]" captionStart="FIGURE 2" captionStartId="5.[152,255,1323,1347]" captionTargetBox="[151,1436,181,1300]" captionTargetId="figure-19@5.[151,1436,181,1300]" captionTargetPageId="5" captionText="FIGURE 2. Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm." figureDoi="http://doi.org/10.5281/zenodo.7329743" httpUri="https://zenodo.org/record/7329743/files/figure.png" pageId="4" pageNumber="383">Fig. 2C</figureCitation>
|
||
), collected from
|
||
<location id="0C0D605A1B48FF9FADEA5CBEFB94F95F" LSID="urn:lsid:plazi:treatment:817B87971B48FF90AEAE5E1DFE68F8C6:0C0D605A1B48FF9FADEA5CBEFB94F95F" box="[979,1083,1749,1775]" country="Saudi Arabia" latitude="19.094444" longLatPrecision="1" longitude="40.379166" municipality="Farasan Banks" name="Mubarak" pageId="4" pageNumber="383">Mubarak</location>
|
||
,
|
||
<collectingMunicipality id="E909ACFB1B48FF9FAA725CBEFB59F95F" box="[1099,1270,1749,1775]" pageId="4" pageNumber="383">Farasan Banks</collectingMunicipality>
|
||
,
|
||
<collectingCountry id="71C576111B48FF9FAB3F5CBEFA33F95F" box="[1286,1436,1749,1775]" name="Saudi Arabia" pageId="4" pageNumber="383">Saudi Arabia</collectingCountry>
|
||
(
|
||
<geoCoordinate id="6CE650461B48FF9FAEA65C92FE9BF8A3" box="[159,308,1785,1811]" degrees="19.09444444" direction="north" orientation="latitude" pageId="4" pageNumber="383" precision="1" value="19.094444">19.09444444</geoCoordinate>
|
||
/
|
||
<geoCoordinate id="6CE650461B48FF9FAF735C92FE70F8A3" box="[330,479,1785,1811]" degrees="40.37916667" direction="east" orientation="longitude" pageId="4" pageNumber="383" precision="1" value="40.379166">40.37916667</geoCoordinate>
|
||
) in a water depth of <
|
||
<quantity id="CE2A9B641B48FF9FACE05C92FCBBF8A3" box="[729,788,1785,1811]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="4" pageNumber="383" unit="m" value="10.0">
|
||
<elevation id="82FFD1B21B48FF9FACE05C92FCBBF8A3" box="[729,788,1785,1811]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="4" pageNumber="383" unit="m" value="10.0">10 m</elevation>
|
||
</quantity>
|
||
, narcotized with menthol, fixed in 4% formaldehyde, and preserved in 70% EtOH. Some tube feet for DNA analyses were preserved in 95% EtOH and are stored at SNSBBSPG together with the specimens
|
||
</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="096D36811B48FF9FAEFE5D0EFDDAF85B" blockId="4.[151,1437,1281,2027]" pageId="4" pageNumber="383">
|
||
<materialsCitation id="B9BA3CDC1B48FF9FAEFE5D0EFDDFF85B" ID-GBIF-Occurrence="4435992304" country="Saudi Arabia" elevation="10" latitude="19.127777" location="Tidhkar Island" longLatPrecision="1" longitude="40.669445" municipality="Farasan Banks" pageId="4" pageNumber="383" specimenCount="1">
|
||
SNSB-BSPG.GW.4266, juvenile individual (
|
||
<figureCitation id="91E92A041B48FF9FACF15D0EFC8AF8CF" box="[712,805,1893,1919]" captionStart="FIGURE 2" captionStartId="5.[152,255,1323,1347]" captionTargetBox="[151,1436,181,1300]" captionTargetId="figure-19@5.[151,1436,181,1300]" captionTargetPageId="5" captionText="FIGURE 2. Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm." figureDoi="http://doi.org/10.5281/zenodo.7329743" httpUri="https://zenodo.org/record/7329743/files/figure.png" pageId="4" pageNumber="383">Fig. 2D</figureCitation>
|
||
), collected from
|
||
<location id="0C0D605A1B48FF9FADD55D0EFB3AF8CF" LSID="urn:lsid:plazi:treatment:817B87971B48FF90AEAE5E1DFE68F8C6:0C0D605A1B48FF9FADD55D0EFB3AF8CF" box="[1004,1173,1893,1919]" country="Saudi Arabia" latitude="19.127777" longLatPrecision="1" longitude="40.669445" municipality="Farasan Banks" name="Tidhkar Island" pageId="4" pageNumber="383">Tidhkar Island</location>
|
||
,
|
||
<collectingMunicipality id="E909ACFB1B48FF9FAA9D5D0EFAE1F8CF" box="[1188,1358,1893,1919]" pageId="4" pageNumber="383">Farasan Banks</collectingMunicipality>
|
||
,
|
||
<collectingCountry id="71C576111B48FF9FAB655D0EFF4CF813" name="Saudi Arabia" pageId="4" pageNumber="383">Saudi Arabia</collectingCountry>
|
||
(
|
||
<geoCoordinate id="6CE650461B48FF9FAEC85DE2FE29F813" box="[241,390,1929,1955]" degrees="19.12777778" direction="north" orientation="latitude" pageId="4" pageNumber="383" precision="1" value="19.127777">19.12777778</geoCoordinate>
|
||
/
|
||
<geoCoordinate id="6CE650461B48FF9FAFA35DE2FD8EF813" box="[410,545,1929,1955]" degrees="40.6694444" direction="east" orientation="longitude" pageId="4" pageNumber="383" precision="1" value="40.669445">40.6694444</geoCoordinate>
|
||
) in a water depth of <
|
||
<quantity id="CE2A9B641B48FF9FAD2A5DE2FCE3F813" box="[787,844,1929,1955]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="4" pageNumber="383" unit="m" value="10.0">
|
||
<elevation id="82FFD1B21B48FF9FAD2A5DE2FCE3F813" box="[787,844,1929,1955]" metricMagnitude="1" metricUnit="m" metricValue="1.0" pageId="4" pageNumber="383" unit="m" value="10.0">10 m</elevation>
|
||
</quantity>
|
||
, narcotized with menthol, fixed in 4% formaldehyde, and preserved in 70% EtOH. Some tube feet for DNA analyses were preserved in 95% EtOH and are stored at SNSB-BSPG together with the specimens
|
||
</materialsCitation>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<caption id="5DAD66091B49FF9EAEA15F40FC07FAD7" ID-DOI="http://doi.org/10.5281/zenodo.7329743" ID-Zenodo-Dep="7329743" httpUri="https://zenodo.org/record/7329743/files/figure.png" pageId="5" pageNumber="384" startId="5.[152,255,1323,1347]" targetBox="[151,1436,181,1300]" targetPageId="5">
|
||
<paragraph id="096D36811B49FF9EAEA15F40FC07FAD7" blockId="5.[151,1437,1323,1384]" pageId="5" pageNumber="384">
|
||
<emphasis id="3BA6EA931B49FF9EAEA15F40FEB6FAF4" bold="true" box="[152,281,1323,1348]" pageId="5" pageNumber="384">FIGURE 2.</emphasis>
|
||
Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm.
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection id="41C8650A1B49FF93AEFE5FD1FC5BF84B" lastPageId="8" lastPageNumber="387" pageId="5" pageNumber="384" type="diagnosis">
|
||
<paragraph id="096D36811B49FF9EAEFE5FD1FDC0F944" blockId="5.[151,1437,1466,1924]" pageId="5" pageNumber="384">
|
||
<emphasis id="3BA6EA931B49FF9EAEFE5FD1FEEDFA64" bold="true" box="[199,322,1466,1492]" pageId="5" pageNumber="384">Diagnosis.</emphasis>
|
||
DNA barcoding analysis of partial COI sequences reasserted
|
||
<taxonomicName id="CED24D021B49FF9EAA1F5FD1FAA6FA64" box="[1062,1289,1466,1492]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B49FF9EAA1F5FD1FAA6FA64" box="[1062,1289,1466,1492]" italics="true" pageId="5" pageNumber="384">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
as a deeply divergent clade (
|
||
<figureCitation id="91E92A041B49FF9EAF6D5FB5FE37FA48" box="[340,408,1502,1528]" captionStart="FIGURE 3" captionStartId="6.[152,255,1199,1223]" captionTargetBox="[395,1195,188,1170]" captionTargetId="figure-35@6.[374,1212,181,1175]" captionTargetPageId="6" captionText="FIGURE 3. A: Indo-Pacific ‘Acanthaster planci’ species complex, COI ML tree based on the haplotype alignment of Vogler et al. (2008), supplemented with five samples from Israel (see text for details) extracted from full mitochondrial genome sequences from Yuasa et al. (2021) and four sequences of the type series of Acanthaster benziei sp. nov. (GW4xxx, highlighted in bold). The tree was rooted with Acanthaster brevispinus (accession number AB231476), showing the deep divergence among, and little diversity within, species/geographic clades. ML bootstrap values are above branches, bootstrap values of the NJ clustering of haplotypes below branches. B: Geographic distribution of COI-barcoded clades and of type localities of names (Figure 1 from Haszprunar et al. 2017): red—Red Sea (RS) species; blue—Southern Indian Ocean (SIO) species (A. mauritiensis); yellow—Northern Indian Ocean (NIO) species (A. planci); green—Pacific Ocean (PO) species (A. cf. solaris). Location of type localities of nominal Acanthaster species: asterisk—A. planci; cross—A. echinites; triangle—A. solaris, square—A. mauritiensis; circle—A. ellisii pseudoplanci; “?” - the type locality of A. ellisii was not specified: in South American waters of the East Pacific." figureDoi="http://doi.org/10.5281/zenodo.7329745" httpUri="https://zenodo.org/record/7329745/files/figure.png" pageId="5" pageNumber="384">Fig. 3</figureCitation>
|
||
, see also
|
||
<bibRefCitation id="6D434B701B49FF9EAC3F5FB5FD4AFA48" author="Vogler, C. & Benzie, J. & Lessios, H. & Barber, P. & Worheide, G." box="[518,741,1502,1528]" pageId="5" pageNumber="384" pagination="696 - 699" refId="ref9248" refString="Vogler, C., Benzie, J., Lessios, H., Barber, P. & Worheide, G. (2008) A threat to coral reefs multiplied? Four species of crownof-thorns starfish. Biology Letters, 4 (6), 696 - 699. https: // doi. org / 10.1098 / rsbl. 2008.0454" type="journal article" year="2008">
|
||
Vogler
|
||
<emphasis id="3BA6EA931B49FF9EAC615FB4FD3EFA48" box="[600,657,1502,1528]" italics="true" pageId="5" pageNumber="384">et al.</emphasis>
|
||
[2008]
|
||
</bibRefCitation>
|
||
) with distinct geographic distribution (Red Sea).
|
||
<taxonomicName id="CED24D021B49FF9EAB2C5FB5FF49F9AC" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B49FF9EAB2C5FB5FF49F9AC" italics="true" pageId="5" pageNumber="384">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
possesses diagnostic mutations in its partial mitochondrial COI gene sequenced here that are unique for the Red Sea and not shared with any other species of the species’ complex, i.e.,
|
||
<taxonomicName id="CED24D021B49FF9EAA3A5C4DFBC5F9F0" box="[1027,1130,1574,1600]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B49FF9EAA3A5C4DFBC5F9F0" box="[1027,1130,1574,1600]" italics="true" pageId="5" pageNumber="384">A. planci</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="CED24D021B49FF9EAA405C4DFA8BF9F0" box="[1145,1316,1574,1600]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B49FF9EAA405C4DFA8BF9F0" box="[1145,1316,1574,1600]" italics="true" pageId="5" pageNumber="384">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="CED24D021B49FF9EAB665C4DFF48F9D4" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" isUncertain="true" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="solaris">
|
||
<emphasis id="3BA6EA931B49FF9EAB665C4DFAD8F9F0" box="[1375,1399,1574,1600]" italics="true" pageId="5" pageNumber="384">A.</emphasis>
|
||
cf.
|
||
<emphasis id="3BA6EA931B49FF9EAEAE5C21FF48F9D4" box="[151,231,1610,1636]" italics="true" pageId="5" pageNumber="384">solaris</emphasis>
|
||
</taxonomicName>
|
||
<emphasis id="3BA6EA931B49FF9EAEDE5C21FF44F9D4" box="[231,235,1610,1636]" italics="true" pageId="5" pageNumber="384">.</emphasis>
|
||
Specifically, mutations in the following positions are diagnostic for
|
||
<taxonomicName id="CED24D021B49FF9EAA395C21FBDFF9D4" box="[1024,1136,1610,1636]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B49FF9EAA395C21FBDFF9D4" box="[1024,1136,1610,1636]" italics="true" pageId="5" pageNumber="384">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
(the position refers to the position in the COI gene extracted from the mitochondrial genome of a specimen from
|
||
<collectingCountry id="71C576111B49FF9EAA485C05FB00F938" box="[1137,1199,1646,1672]" name="Israel" pageId="5" pageNumber="384">Israel</collectingCountry>
|
||
[GenBank accession number
|
||
<accessionNumber id="1681AB621B49FF9EAECD5CF9FEDDF91D" box="[244,370,1682,1709]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/LC566218" pageId="5" pageNumber="384">LC566218</accessionNumber>
|
||
]; the first nucleotide is the one in
|
||
<taxonomicName id="CED24D021B49FF9EACD25CF9FCF6F91C" box="[747,857,1682,1708]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B49FF9EACD25CF9FCF6F91C" box="[747,857,1682,1708]" italics="true" pageId="5" pageNumber="384">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
, the second one in the other three species): 150 (T / C), 426 (G / A), 495 (T / C), 504 (G / A), 555 (T / C), 585 (G / A), 588 (T / C), 612 (G / A), 711 (T / C), 714 (C / T). All these are silent third-codon mutations.
|
||
</paragraph>
|
||
<paragraph id="096D36811B49FF9EAEFE5C95FB09F834" blockId="5.[151,1437,1466,1924]" pageId="5" pageNumber="384">
|
||
The following diagnostic morphological characters differentiate
|
||
<taxonomicName id="CED24D021B49FF9EADB55C95FC57F8A8" box="[908,1016,1790,1816]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B49FF9EADB55C95FC57F8A8" box="[908,1016,1790,1816]" italics="true" pageId="5" pageNumber="384">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
from its congeners, and were assessed from the
|
||
<typeStatus id="D66988231B49FF9EAEC65D49FE80F88C" box="[255,303,1826,1852]" pageId="5" pageNumber="384">type</typeStatus>
|
||
series at size measured (see
|
||
<tableCitation id="4450033A1B49FF9EAC4A5D49FD67F88C" box="[627,712,1826,1852]" captionStart="TABLE 3" captionStartId="7.[152,245,151,175]" captionTargetPageId="7" captionText="TABLE 3: Basic measurements of type specimens, including number of arms and madreporites, mean length of arms (R) and radius of disk (r). Note that GW4266 is a juvenile specimen." httpUri="http://table.plazi.org/id/5DAD66091B4BFF9CAEA15AFCFCE5FF63" pageId="5" pageNumber="384" tableUuid="5DAD66091B4BFF9CAEA15AFCFCE5FF63">Table 3</tableCitation>
|
||
): fanned spine tips in primary and latero-oral spines; a wider tip or tapering shape in circumoral spines; and rhombus-shaped oral pedicellariae. Additionally,
|
||
<taxonomicName id="CED24D021B49FF9EAA8E5D2DFA89F8D0" box="[1207,1318,1862,1888]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B49FF9EAA8E5D2DFA89F8D0" box="[1207,1318,1862,1888]" italics="true" pageId="5" pageNumber="384">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
has fewer arms than its congeners (up to
|
||
<quantity id="CE2A9B641B49FF9EAFD65D01FD87F834" box="[495,552,1898,1924]" metricMagnitude="-1" metricUnit="m" metricValue="3.556" pageId="5" pageNumber="384" unit="in" value="14.0">14 in</quantity>
|
||
<taxonomicName id="CED24D021B49FF9EAC165D01FD32F834" box="[559,669,1898,1924]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B49FF9EAC165D01FD32F834" box="[559,669,1898,1924]" italics="true" pageId="5" pageNumber="384">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
vs. up to
|
||
<quantity id="CE2A9B641B49FF9EAD325D01FCEBF834" box="[779,836,1898,1924]" metricMagnitude="-1" metricUnit="m" metricValue="5.842" pageId="5" pageNumber="384" unit="in" value="23.0">23 in</quantity>
|
||
<taxonomicName id="CED24D021B49FF9EAD725D01FC76F834" box="[843,985,1898,1924]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" isUncertain="true" kingdom="Animalia" order="Valvatida" pageId="5" pageNumber="384" phylum="Echinodermata" rank="species" species="solaris">
|
||
<emphasis id="3BA6EA931B49FF9EAD725D01FCCCF834" box="[843,867,1898,1924]" italics="true" pageId="5" pageNumber="384">A.</emphasis>
|
||
cf.
|
||
<emphasis id="3BA6EA931B49FF9EADB45D01FC76F834" box="[909,985,1898,1924]" italics="true" pageId="5" pageNumber="384">solaris</emphasis>
|
||
</taxonomicName>
|
||
from the Pacific).
|
||
</paragraph>
|
||
<caption id="5DAD66091B4AFF9DAEA15EC4FBD7F99F" ID-DOI="http://doi.org/10.5281/zenodo.7329745" ID-Zenodo-Dep="7329745" httpUri="https://zenodo.org/record/7329745/files/figure.png" pageId="6" pageNumber="385" startId="6.[152,255,1199,1223]" targetBox="[395,1195,188,1170]" targetPageId="6">
|
||
<paragraph id="096D36811B4AFF9DAEA15EC4FE63FACB" blockId="6.[151,1437,1199,1584]" pageId="6" pageNumber="385">
|
||
<emphasis id="3BA6EA931B4AFF9DAEA15EC4FEBDFB77" bold="true" box="[152,274,1199,1223]" pageId="6" pageNumber="385">FIGURE 3</emphasis>
|
||
. A: Indo-Pacific ‘
|
||
<taxonomicName id="CED24D021B4AFF9DAFFE5EC4FD29FB77" box="[455,646,1199,1223]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B4AFF9DAFFE5EC4FD29FB77" box="[455,646,1199,1223]" italics="true" pageId="6" pageNumber="385">Acanthaster planci</emphasis>
|
||
</taxonomicName>
|
||
’ species complex, COI ML tree based on the haplotype alignment of
|
||
<bibRefCitation id="6D434B701B4AFF9DAB785EC4FF53FB5B" author="Vogler, C. & Benzie, J. & Lessios, H. & Barber, P. & Worheide, G." pageId="6" pageNumber="385" pagination="696 - 699" refId="ref9248" refString="Vogler, C., Benzie, J., Lessios, H., Barber, P. & Worheide, G. (2008) A threat to coral reefs multiplied? Four species of crownof-thorns starfish. Biology Letters, 4 (6), 696 - 699. https: // doi. org / 10.1098 / rsbl. 2008.0454" type="journal article" year="2008">
|
||
Vogler
|
||
<emphasis id="3BA6EA931B4AFF9DABB35EDBFF1DFB5B" italics="true" pageId="6" pageNumber="385">et al.</emphasis>
|
||
(2008)
|
||
</bibRefCitation>
|
||
, supplemented with five samples from Israel (see text for details) extracted from full mitochondrial genome sequences from
|
||
<bibRefCitation id="6D434B701B4AFF9DAEF75E93FE3FFABF" author="Yuasa, H. & Kajitani, R. & Nakamura, Y. & Takahashi, K. & Okuno, M. & Kobayashi, F. & Shinoda, T. & Toyoda, A. & Suzuki, Y. & Thongtham, N. & Forsman, Z. & Bronstein, O. & Seveso, D. & Montalbetti, E. & Taquet, C. & Eyal, G. & Yasuda, N. & Itoh, T." box="[206,400,1271,1295]" pageId="6" pageNumber="385" pagination="4" refId="ref9464" refString="Yuasa, H., Kajitani, R., Nakamura, Y., Takahashi, K., Okuno, M., Kobayashi, F., Shinoda, T., Toyoda, A., Suzuki, Y., Thongtham, N., Forsman, Z., Bronstein, O., Seveso, D., Montalbetti, E., Taquet, C., Eyal, G., Yasuda, N. & Itoh, T. (2021) Elucidation of the speciation history of three sister species of crown-of-thorns starfish (Acanthaster spp.) based on genomic analysis. DNA Research, 28 (4), dsab 012. https: // doi. org / 10.1093 / dnares / dsab 012" type="journal article" year="2021">
|
||
Yuasa
|
||
<emphasis id="3BA6EA931B4AFF9DAF2B5E93FEE9FABF" box="[274,326,1271,1295]" italics="true" pageId="6" pageNumber="385">et al.</emphasis>
|
||
(2021)
|
||
</bibRefCitation>
|
||
and four sequences of the type series of
|
||
<taxonomicName id="CED24D021B4AFF9DAD0A5E9CFC53FABF" authority="Wörheide & Kaltenbacher & Cowan & Haszprunar, 2022" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[819,1020,1271,1295]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="benziei" status="sp. nov.">
|
||
<emphasis id="3BA6EA931B4AFF9DAD0A5E9CFC53FABF" box="[819,1020,1271,1295]" italics="true" pageId="6" pageNumber="385">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="209557E81B4AFF9DAA3A5E93FBFCFABF" box="[1027,1107,1272,1295]" pageId="6" pageNumber="385" rank="species">sp. nov.</taxonomicNameLabel>
|
||
(GW4xxx, highlighted in bold). The tree was rooted with
|
||
<taxonomicName id="CED24D021B4AFF9DAF9D5F70FD35FA83" authorityName="Fisher" authorityYear="1917" box="[420,666,1307,1331]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="brevispinus">
|
||
<emphasis id="3BA6EA931B4AFF9DAF9D5F70FD35FA83" box="[420,666,1307,1331]" italics="true" pageId="6" pageNumber="385">Acanthaster brevispinus</emphasis>
|
||
</taxonomicName>
|
||
(accession number
|
||
<accessionNumber id="1681AB621B4AFF9DAD505F70FC73FA83" box="[873,988,1307,1331]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/AB231476" pageId="6" pageNumber="385">AB231476</accessionNumber>
|
||
), showing the deep divergence among, and little diversity within, species/geographic clades. ML bootstrap values are above branches, bootstrap values of the NJ clustering of haplotypes below branches.
|
||
</paragraph>
|
||
<paragraph id="096D36811B4AFF9DAEAE5FE3FBD7F99F" blockId="6.[151,1437,1199,1584]" pageId="6" pageNumber="385">
|
||
B: Geographic distribution of COI-barcoded clades and of type localities of names (Figure 1 from
|
||
<bibRefCitation id="6D434B701B4AFF9DAAA65FE3FA3DFA2F" author="Haszprunar, G. & Vogler, C. & Worheide, G." box="[1183,1426,1415,1439]" pageId="6" pageNumber="385" pagination="22" refId="ref8118" refString="Haszprunar, G., Vogler, C. & Worheide, G. (2017) Persistent gaps of knowledge for naming and distinguishing multiple species of crown-of-thorns-seastar in the Acanthaster planci species complex. Diversity, 9 (2), 22. https: // doi. org / 10.3390 / d 9020022" type="journal article" year="2017">
|
||
Haszprunar
|
||
<emphasis id="3BA6EA931B4AFF9DAB245FE3FAFDFA2F" box="[1309,1362,1415,1439]" italics="true" pageId="6" pageNumber="385">et al.</emphasis>
|
||
2017
|
||
</bibRefCitation>
|
||
): red—Red Sea (RS) species; blue—Southern Indian Ocean (SIO) species (
|
||
<taxonomicName id="CED24D021B4AFF9DADA25FC0FB98FA73" box="[923,1079,1451,1475]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B4AFF9DADA25FC0FB98FA73" box="[923,1079,1451,1475]" italics="true" pageId="6" pageNumber="385">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
); yellow—Northern Indian Ocean (NIO) species (
|
||
<taxonomicName id="CED24D021B4AFF9DAF085FA4FE24FA57" box="[305,395,1487,1511]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B4AFF9DAF085FA4FE24FA57" box="[305,395,1487,1511]" italics="true" pageId="6" pageNumber="385">A. planci</emphasis>
|
||
</taxonomicName>
|
||
); green—Pacific Ocean (PO) species (
|
||
<taxonomicName id="CED24D021B4AFF9DAD285FA4FC3FFA57" box="[785,912,1487,1511]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" isUncertain="true" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="solaris">
|
||
<emphasis id="3BA6EA931B4AFF9DAD285FA4FC88FA56" box="[785,807,1487,1510]" italics="true" pageId="6" pageNumber="385">A.</emphasis>
|
||
cf.
|
||
<emphasis id="3BA6EA931B4AFF9DAD725FA4FC3FFA57" box="[843,912,1487,1511]" italics="true" pageId="6" pageNumber="385">solaris</emphasis>
|
||
</taxonomicName>
|
||
). Location of type localities of nominal
|
||
<taxonomicName id="CED24D021B4AFF9DAB1B5FA4FF4BF9BB" pageId="6" pageNumber="385">
|
||
<emphasis id="3BA6EA931B4AFF9DAB1B5FA4FA33FA57" box="[1314,1436,1487,1511]" italics="true" pageId="6" pageNumber="385">Acanthaster</emphasis>
|
||
species
|
||
</taxonomicName>
|
||
: asterisk—
|
||
<taxonomicName id="CED24D021B4AFF9DAF6D5F98FE00F9BB" box="[340,431,1523,1547]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B4AFF9DAF6D5F98FE00F9BB" box="[340,431,1523,1547]" italics="true" pageId="6" pageNumber="385">A. planci</emphasis>
|
||
</taxonomicName>
|
||
; cross—
|
||
<taxonomicName id="CED24D021B4AFF9DAC335F98FD2FF9BB" baseAuthorityName="Ellis & Solander" baseAuthorityYear="1786" box="[522,640,1523,1547]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="echinites">
|
||
<emphasis id="3BA6EA931B4AFF9DAC335F98FD2FF9BB" box="[522,640,1523,1547]" italics="true" pageId="6" pageNumber="385">A. echinites</emphasis>
|
||
</taxonomicName>
|
||
; triangle—
|
||
<taxonomicName id="CED24D021B4AFF9DACCA5F98FCFBF9BB" box="[755,852,1523,1547]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="solaris">
|
||
<emphasis id="3BA6EA931B4AFF9DACCA5F98FCFBF9BB" box="[755,852,1523,1547]" italics="true" pageId="6" pageNumber="385">A. solaris</emphasis>
|
||
</taxonomicName>
|
||
, square—
|
||
<taxonomicName id="CED24D021B4AFF9DAD855F98FBFAF9BB" box="[956,1109,1523,1547]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B4AFF9DAD855F98FBFAF9BB" box="[956,1109,1523,1547]" italics="true" pageId="6" pageNumber="385">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
; circle—
|
||
<taxonomicName id="CED24D021B4AFF9DAA8D5F98FA3AF9BB" authorityName="Caso" authorityYear="1962" box="[1204,1429,1523,1547]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="subSpecies" species="ellisii" subSpecies="pseudoplanci">
|
||
<emphasis id="3BA6EA931B4AFF9DAA8D5F98FA3AF9BB" box="[1204,1429,1523,1547]" italics="true" pageId="6" pageNumber="385">A. ellisii pseudoplanci</emphasis>
|
||
</taxonomicName>
|
||
; “?” - the type locality of
|
||
<taxonomicName id="CED24D021B4AFF9DAFAA5C7CFE48F99F" baseAuthorityName="Gray" baseAuthorityYear="1840" box="[403,487,1559,1583]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="6" pageNumber="385" phylum="Echinodermata" rank="species" species="ellisii">
|
||
<emphasis id="3BA6EA931B4AFF9DAFAA5C7CFE48F99F" box="[403,487,1559,1583]" italics="true" pageId="6" pageNumber="385">A. ellisii</emphasis>
|
||
</taxonomicName>
|
||
was not specified: in South American waters of the East Pacific.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="096D36811B4AFF9DAEFE5CEAFCA8F8B8" blockId="6.[151,1437,1665,2016]" pageId="6" pageNumber="385">
|
||
<emphasis id="3BA6EA931B4AFF9DAEFE5CEAFDACF92B" bold="true" box="[199,515,1665,1691]" pageId="6" pageNumber="385">Morphological description</emphasis>
|
||
. Applies to the
|
||
<typeStatus id="D66988231B4AFF9DAC885CEAFCB9F92B" box="[689,790,1665,1691]" pageId="6" pageNumber="385" type="holotype">holotype</typeStatus>
|
||
, except otherwise noted. Large sea star with a convex disk and 13 arms (number of arms across type series 11-14,
|
||
<tableCitation id="4450033A1B4AFF9DAD295CCEFCCAF90F" box="[784,869,1701,1727]" captionStart="TABLE 3" captionStartId="7.[152,245,151,175]" captionTargetPageId="7" captionText="TABLE 3: Basic measurements of type specimens, including number of arms and madreporites, mean length of arms (R) and radius of disk (r). Note that GW4266 is a juvenile specimen." httpUri="http://table.plazi.org/id/5DAD66091B4BFF9CAEA15AFCFCE5FF63" pageId="6" pageNumber="385" tableUuid="5DAD66091B4BFF9CAEA15AFCFCE5FF63">Table 3</tableCitation>
|
||
), which have a subcylindrical cross section. Each arm tapers to an acute point and the arms are slightly variable in length. The mean disk radius (r) measures
|
||
<quantity id="CE2A9B641B4AFF9DAB7E5CA2FA37F954" box="[1351,1432,1737,1764]" metricMagnitude="-2" metricUnit="m" metricValue="5.8" pageId="6" pageNumber="385" unit="mm" value="58.0">58 mm</quantity>
|
||
, the mean length of rays (R) is
|
||
<quantity id="CE2A9B641B4AFF9DAFD25C86FD95F8B8" box="[491,570,1773,1800]" metricMagnitude="-2" metricUnit="m" metricValue="9.1" pageId="6" pageNumber="385" unit="mm" value="91.0">91 mm</quantity>
|
||
(R/r ratio = 1.57).
|
||
</paragraph>
|
||
<paragraph id="096D36811B4AFF9DAEFE5D7AFB69F86F" blockId="6.[151,1437,1665,2016]" pageId="6" pageNumber="385">
|
||
There are two rows of ambulacral tube feet (approximately
|
||
<quantity id="CE2A9B641B4AFF9DAD425D7AFC74F89C" box="[891,987,1809,1836]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" metricValueMax="3.0" metricValueMin="1.0" pageId="6" pageNumber="385" unit="mm" value="2.0" valueMax="3.0" valueMin="1.0">1–3 mm</quantity>
|
||
in diameter, with flattened tips and no sucker) in the ambulacral groove in the midline of the oral side of each arm. The stereom on both the oral and aboral side consists of a mesh of ossicles, concealed by soft tissue and a large number of spines and pedicellariae, both of which are sheathed in tissue, typically labyrinthic or elongated trabeculated. The aboral disc surface has many papulae with no clear arrangement. The tissue here is relatively soft and compressible. The anus – in the centre of the disc – has no papulae, is harder, and is spaciously encircled by six madreporites on the disk.
|
||
</paragraph>
|
||
<caption id="5DAD66091B4BFF9CAEA15AFCFCE5FF63" ID-Table-UUID="5DAD66091B4BFF9CAEA15AFCFCE5FF63" httpUri="http://table.plazi.org/id/5DAD66091B4BFF9CAEA15AFCFCE5FF63" pageId="7" pageNumber="386" startId="7.[152,245,151,175]" targetBox="[179,1325,227,626]" targetIsTable="true" targetPageId="7">
|
||
<paragraph id="096D36811B4BFF9CAEA15AFCFCE5FF63" blockId="7.[151,1436,151,212]" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAEA15AFCFEBFFF1F" bold="true" box="[152,272,151,175]" pageId="7" pageNumber="386">TABLE 3:</emphasis>
|
||
Basic measurements of type specimens, including number of arms and madreporites, mean length of arms (R) and radius of disk (r). Note that GW4266 is a juvenile specimen.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="096D36811B4BFF9CAC455A88FB74FDE6" pageId="7" pageNumber="386">
|
||
<table id="7BD2C4211B4B0064AE8A5A88FA82FDC2" box="[179,1325,227,626]" gridcols="5" gridrows="6" pageId="7" pageNumber="386">
|
||
<tr id="B7E234C31B4B0064AE8A5A88FA82FF4B" box="[179,1325,227,251]" gridrow="0" pageId="7" pageNumber="386" rowspan-0="1">
|
||
<th id="F4335DBF1B4B0064AC455A88FD74FF4B" box="[636,731,227,251]" gridcol="1" gridrow="0" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAC455A88FD74FF4B" bold="true" box="[636,731,227,251]" pageId="7" pageNumber="386">GW4202</emphasis>
|
||
</th>
|
||
<th id="F4335DBF1B4B0064AD695A88FC1FFF4B" box="[848,944,227,251]" gridcol="2" gridrow="0" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAD695A88FC1FFF4B" bold="true" box="[848,944,227,251]" pageId="7" pageNumber="386">GW4081</emphasis>
|
||
</th>
|
||
<th id="F4335DBF1B4B0064AA285A88FBDEFF4B" box="[1041,1137,227,251]" gridcol="3" gridrow="0" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAA285A88FBDEFF4B" bold="true" box="[1041,1137,227,251]" pageId="7" pageNumber="386">GW4168</emphasis>
|
||
</th>
|
||
<th id="F4335DBF1B4B0064AAF75A88FA82FF4B" box="[1230,1325,227,251]" gridcol="4" gridrow="0" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAAF75A88FA82FF4B" bold="true" box="[1230,1325,227,251]" pageId="7" pageNumber="386">GW4266</emphasis>
|
||
</th>
|
||
</tr>
|
||
<tr id="B7E234C31B4B0064AE8A5B74FA82FE88" box="[179,1325,287,312]" gridrow="1" pageId="7" pageNumber="386">
|
||
<th id="F4335DBF1B4B0064AE8A5B74FE1CFE88" box="[179,435,287,312]" gridcol="0" gridrow="1" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAE8A5B74FEC7FE87" bold="true" box="[179,360,287,311]" pageId="7" pageNumber="386">Number of Arms</emphasis>
|
||
</th>
|
||
<td id="F4335DBF1B4B0064AC455B74FD74FE88" box="[636,731,287,312]" gridcol="1" gridrow="1" pageId="7" pageNumber="386">13</td>
|
||
<td id="F4335DBF1B4B0064AD695B74FC1FFE88" box="[848,944,287,312]" gridcol="2" gridrow="1" pageId="7" pageNumber="386">11</td>
|
||
<td id="F4335DBF1B4B0064AA285B74FBDEFE88" box="[1041,1137,287,312]" gridcol="3" gridrow="1" pageId="7" pageNumber="386">14</td>
|
||
<td id="F4335DBF1B4B0064AAF75B74FA82FE88" box="[1230,1325,287,312]" gridcol="4" gridrow="1" pageId="7" pageNumber="386">11</td>
|
||
</tr>
|
||
<tr id="B7E234C31B4B0064AE8A5B34FA82FEC8" box="[179,1325,351,376]" gridrow="2" pageId="7" pageNumber="386">
|
||
<th id="F4335DBF1B4B0064AE8A5B34FE1CFEC8" box="[179,435,351,376]" gridcol="0" gridrow="2" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAE8A5B34FE1CFEC7" bold="true" box="[179,435,351,375]" pageId="7" pageNumber="386">Mean R (length of arm)</emphasis>
|
||
</th>
|
||
<td id="F4335DBF1B4B0064AC455B34FD74FEC8" box="[636,731,351,376]" gridcol="1" gridrow="2" pageId="7" pageNumber="386">91 mm</td>
|
||
<td id="F4335DBF1B4B0064AD695B34FC1FFEC8" box="[848,944,351,376]" gridcol="2" gridrow="2" pageId="7" pageNumber="386">106 mm</td>
|
||
<td id="F4335DBF1B4B0064AA285B34FBDEFEC8" box="[1041,1137,351,376]" gridcol="3" gridrow="2" pageId="7" pageNumber="386">125 mm</td>
|
||
<td id="F4335DBF1B4B0064AAF75B34FA82FEC8" box="[1230,1325,351,376]" gridcol="4" gridrow="2" pageId="7" pageNumber="386">50 mm</td>
|
||
</tr>
|
||
<tr id="B7E234C31B4B0064AE8A5BF4FA82FE64" box="[179,1325,415,468]" gridrow="3" pageId="7" pageNumber="386">
|
||
<th id="F4335DBF1B4B0064AE8A5BF4FE1CFE64" box="[179,435,415,468]" gridcol="0" gridrow="3" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAE8A5BF4FEFBFE64" bold="true" pageId="7" pageNumber="386">Mean r (radius of disc)</emphasis>
|
||
</th>
|
||
<td id="F4335DBF1B4B0064AC455BF4FD74FE64" box="[636,731,415,468]" gridcol="1" gridrow="3" pageId="7" pageNumber="386">58 mm</td>
|
||
<td id="F4335DBF1B4B0064AD695BF4FC1FFE64" box="[848,944,415,468]" gridcol="2" gridrow="3" pageId="7" pageNumber="386">51.5 mm</td>
|
||
<td id="F4335DBF1B4B0064AA285BF4FBDEFE64" box="[1041,1137,415,468]" gridcol="3" gridrow="3" pageId="7" pageNumber="386">65 mm</td>
|
||
<td id="F4335DBF1B4B0064AAF75BF4FA82FE64" box="[1230,1325,415,468]" gridcol="4" gridrow="3" pageId="7" pageNumber="386">28 mm</td>
|
||
</tr>
|
||
<tr id="B7E234C31B4B0064AE8A5B96FA82FDA6" box="[179,1325,509,534]" gridrow="4" pageId="7" pageNumber="386">
|
||
<th id="F4335DBF1B4B0064AE8A5B96FE1CFDA6" box="[179,435,509,534]" gridcol="0" gridrow="4" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAE8A5B96FEBDFDA5" bold="true" box="[179,274,509,533]" pageId="7" pageNumber="386">R:r ratio</emphasis>
|
||
</th>
|
||
<td id="F4335DBF1B4B0064AC455B96FD74FDA6" box="[636,731,509,534]" gridcol="1" gridrow="4" pageId="7" pageNumber="386">1.57</td>
|
||
<td id="F4335DBF1B4B0064AD695B96FC1FFDA6" box="[848,944,509,534]" gridcol="2" gridrow="4" pageId="7" pageNumber="386">2.06</td>
|
||
<td id="F4335DBF1B4B0064AA285B96FBDEFDA6" box="[1041,1137,509,534]" gridcol="3" gridrow="4" pageId="7" pageNumber="386">1.92</td>
|
||
<td id="F4335DBF1B4B0064AAF75B96FA82FDA6" box="[1230,1325,509,534]" gridcol="4" gridrow="4" pageId="7" pageNumber="386">1.79</td>
|
||
</tr>
|
||
<tr id="B7E234C31B4B0064AE8A5856FA82FDC2" box="[179,1325,573,626]" gridrow="5" pageId="7" pageNumber="386">
|
||
<th id="F4335DBF1B4B0064AE8A5856FE1CFDC2" box="[179,435,573,626]" gridcol="0" gridrow="5" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAE8A5856FECEFDC2" bold="true" box="[179,353,573,626]" pageId="7" pageNumber="386">Number of Madreporites</emphasis>
|
||
</th>
|
||
<td id="F4335DBF1B4B0064AC455856FD74FDC2" box="[636,731,573,626]" gridcol="1" gridrow="5" pageId="7" pageNumber="386">6</td>
|
||
<td id="F4335DBF1B4B0064AD695856FC1FFDC2" box="[848,944,573,626]" gridcol="2" gridrow="5" pageId="7" pageNumber="386">4</td>
|
||
<td id="F4335DBF1B4B0064AA285856FBDEFDC2" box="[1041,1137,573,626]" gridcol="3" gridrow="5" pageId="7" pageNumber="386">4</td>
|
||
<td id="F4335DBF1B4B0064AAF75856FA82FDC2" box="[1230,1325,573,626]" gridcol="4" gridrow="5" pageId="7" pageNumber="386">4</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
<paragraph id="096D36811B4BFF9CAEFE58B4FED3FB7D" blockId="7.[151,1437,735,2022]" pageId="7" pageNumber="386">
|
||
Six spine
|
||
<typeStatus id="D66988231B4BFF9CAF7B588BFED2FD4A" box="[322,381,736,762]" pageId="7" pageNumber="386">types</typeStatus>
|
||
are distinguished: primary and secondary spines on the aboral side, and subambulacral, circumoral, oral, and latero-oral spines on the oral side (see
|
||
<figureCitation id="91E92A041B4BFF9CAD7A5968FC2AFCAD" box="[835,901,771,797]" captionStart="FIGURE 1" captionStartId="3.[152,255,1515,1539]" captionTargetBox="[271,1317,666,1489]" captionTargetId="figure-208@3.[270,1318,666,1489]" captionTargetPageId="3" captionText="FIGURE 1. Illustrations of pedicellariae and spines (A–C). A: Subambulacral spines, top: bundle with two equally long spines each, bottom: bundle with four unequal long spines; B: Aboral pedicellariae with two valves; C: Close up of part of one arm of paratype GW4081, showing three different spine types indicated by arrows: latero-oral spines (dark blue), oral spines (yellow), subambulacral spines (turquoise). The actinal pedicellariae (red) are associated with the first row of oral spines." figureDoi="http://doi.org/10.5281/zenodo.7329741" httpUri="https://zenodo.org/record/7329741/files/figure.png" pageId="7" pageNumber="386">Fig. 1</figureCitation>
|
||
for definition). The primary and the secondary spines, that cover the aboral surface, differ in size and supporting ossicle (the pedicle), which is shorter and supported by a secondary ossicle in the secondary spines (
|
||
<bibRefCitation id="6D434B701B4BFF9CAC855920FCD5FCD5" author="Motokawa, T." box="[700,890,843,869]" pageId="7" pageNumber="386" pagination="247 - 253" refId="ref8475" refString="Motokawa, T. (1986) Morphology of spines and spine joint in the crown-of-thorns starfish Acanthaster planci (Echinodermata, Asteroida). Zoomorphology, 106 (4), 247 - 253. https: // doi. org / 10.1007 / BF 00312046" type="journal article" year="1986">Motokawa 1986</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="6D434B701B4BFF9CADBE5920FB85FCD5" author="Walbran, P." box="[903,1066,843,869]" pageId="7" pageNumber="386" pagination="1 - 45" refId="ref9389" refString="Walbran, P. (1987) An atlas of the skeletal components of the crown-of-thorns starfish (Acanthaster planci (L )). Technical Memorandum GBRMPA-TM- 11, 1 - 45 (including 13 plates). Available from: http: // www. gbrmpa. gov. au / __ data / assets / pdf _ file / 0005 / 9752 / gbrmpa-tm 11. pdf (accessed 13 March 2022)" type="book chapter" year="1987">Walbran 1987</bibRefCitation>
|
||
). Subambulacral spines are very short and occur next to the ambulacral groove or furrow. Long latero-oral spines intercross with those of the adjacent arm, while the oral spines are shorter, with a blunt tip, positioned in one or two rows next to the subambulacral spines. Most circumoral spines are longer than the oral spines and are located in a single row at the mouth opening. One group of circumoral spines is part of two adjoining rays. Spines on the oral side may have a bend in the lower quarter of the shaft, are more irregular than the aboral spines, and either lack or have a less dominant pedicle. The oral and circumoral row of spines is continuous throughout all of the arms. They fringe the ambulacral spines from the tip of one arm to the mouth opening, turning to the next arm, where they also fringe the ambulacral spines, remaining symmetrical on both sides of the ambulacral groove. Variations and intermediate forms of one or more of the spine tip shapes within one individual are possible, however, the pointed tip is common in any sea star studied from the
|
||
<typeStatus id="D66988231B4BFF9CAEC75EDFFE81FB7E" box="[254,302,1204,1230]" pageId="7" pageNumber="386">type</typeStatus>
|
||
series.
|
||
</paragraph>
|
||
<paragraph id="096D36811B4BFF9CAEFE5EBCFDDEFA31" blockId="7.[151,1437,735,2022]" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAEFE5EBCFED4FB41" bold="true" box="[199,379,1239,1265]" pageId="7" pageNumber="386">Primary spines</emphasis>
|
||
(
|
||
<figureCitation id="91E92A041B4BFF9CAFB35EBCFE42FB41" box="[394,493,1239,1265]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="7" pageNumber="386">Figs. 4A</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B4BFF9CAFC15EBCFDA8FB41" box="[504,519,1239,1265]" captionStart="FIGURE 6" captionStartId="10.[200,303,1266,1290]" captionTargetBox="[479,1103,191,1235]" captionTargetId="figure-35@10.[472,1115,181,1242]" captionTargetPageId="10" captionText="FIGURE 6. Typical primary spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329751" httpUri="https://zenodo.org/record/7329751/files/figure.png" pageId="7" pageNumber="386">6</figureCitation>
|
||
) are the longest spines on the aboral surface, ranging between approximately
|
||
<quantity id="CE2A9B641B4BFF9CAB5D5EBCFF6CFAA6" metricMagnitude="-2" metricUnit="m" metricValue="1.95" metricValueMax="3.3" metricValueMin="0.6" pageId="7" pageNumber="386" unit="mm" value="19.5" valueMax="33.0" valueMin="6.0">6–33 mm</quantity>
|
||
in length. They are straight and slender, slightly shorter (approx.
|
||
<quantity id="CE2A9B641B4BFF9CAD8E5E90FB8AFAA5" box="[951,1061,1275,1301]" metricMagnitude="-2" metricUnit="m" metricValue="1.65" metricValueMax="2.7" metricValueMin="0.6" pageId="7" pageNumber="386" unit="mm" value="16.5" valueMax="27.0" valueMin="6.0">6–27 mm</quantity>
|
||
) on the disc and longer (approx.
|
||
<quantity id="CE2A9B641B4BFF9CAEAE5F74FEBDFA89" box="[151,274,1311,1337]" metricMagnitude="-2" metricUnit="m" metricValue="3.0" metricValueMax="3.3" metricValueMin="2.7" pageId="7" pageNumber="386" unit="mm" value="30.0" valueMax="33.0" valueMin="27.0">27–33 mm</quantity>
|
||
) on the arms, consist of one to two parts and are supported by a basal/primary ossicle (= pedicle). The shape of spine tips is variable, but most common is a fanned spine tip with several small furrows (
|
||
<figureCitation id="91E92A041B4BFF9CAB395F28FAF3FAED" box="[1280,1372,1347,1373]" captionStart="FIGURE 6" captionStartId="10.[200,303,1266,1290]" captionTargetBox="[479,1103,191,1235]" captionTargetId="figure-35@10.[472,1115,181,1242]" captionTargetPageId="10" captionText="FIGURE 6. Typical primary spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329751" httpUri="https://zenodo.org/record/7329751/files/figure.png" pageId="7" pageNumber="386">Fig. 6A</figureCitation>
|
||
). The spines can be granulated in the upper half.
|
||
</paragraph>
|
||
<paragraph id="096D36811B4BFF9CAEFE5FE0FAA1FA5D" blockId="7.[151,1437,735,2022]" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAEFE5FE0FE38FA15" bold="true" box="[199,407,1419,1445]" pageId="7" pageNumber="386">Secondary spines</emphasis>
|
||
(
|
||
<figureCitation id="91E92A041B4BFF9CAF915FE0FE50FA15" box="[424,511,1419,1445]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="7" pageNumber="386">Fig. 4B</figureCitation>
|
||
) are found mainly on the disc, but also on the arms. They are less numerous and shorter than the primary spines, ranging between approximately
|
||
<quantity id="CE2A9B641B4BFF9CADB35FC4FC57FA7A" box="[906,1016,1455,1482]" metricMagnitude="-3" metricUnit="m" metricValue="9.5" metricValueMax="11.0" metricValueMin="8.0" pageId="7" pageNumber="386" unit="mm" value="9.5" valueMax="11.0" valueMin="8.0">8–11 mm</quantity>
|
||
in length. Secondary spines always consist of one part, but otherwise reflect the appearance of primary spines, also regarding granulation.
|
||
</paragraph>
|
||
<paragraph id="096D36811B4BFF9CAEFE5F9CFDEDF912" blockId="7.[151,1437,735,2022]" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAEFE5F9CFE0EF9A1" bold="true" box="[199,417,1527,1553]" pageId="7" pageNumber="386">Latero-oral spines</emphasis>
|
||
(
|
||
<figureCitation id="91E92A041B4BFF9CAF965F9CFDBEF9A1" box="[431,529,1527,1553]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="7" pageNumber="386">Figs. 4C</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B4BFF9CAC255F9CFD84F9A1" box="[540,555,1527,1553]" captionStart="FIGURE 7" captionStartId="11.[152,255,1418,1442]" captionTargetBox="[479,1108,204,1389]" captionTargetId="figure-19@11.[472,1115,199,1394]" captionTargetPageId="11" captionText="FIGURE 7. Typical latero-oral spines, showing species-specific variation among (A) Acanthaster benziei sp. nov., (B) A. planci, (C) A. mauritiensis, and (D) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329753" httpUri="https://zenodo.org/record/7329753/files/figure.png" pageId="7" pageNumber="386">7</figureCitation>
|
||
) are similar in size to the primary spines, ranging between approximately
|
||
<quantity id="CE2A9B641B4BFF9CAB5D5F9CFF6CF986" metricMagnitude="-2" metricUnit="m" metricValue="1.2" metricValueMax="2.0" metricValueMin="0.4" pageId="7" pageNumber="386" unit="mm" value="12.0" valueMax="20.0" valueMin="4.0">4–20 mm</quantity>
|
||
in length. They are located on marginal ossicles, forming no or short pedicles, compared to primary spines. Spine tips have similar shapes to primary spines: pointed to fanned or flat with a slight furrow in the middle, rounded tip with small furrows, mostly broader than tips of aboral spines, can be asymmetrical, can widen or taper upwards. Granulation may be present.
|
||
</paragraph>
|
||
<paragraph id="096D36811B4BFF9CAEFE5CC0FC91F872" blockId="7.[151,1437,735,2022]" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAEFE5CC0FE08F975" bold="true" box="[199,423,1707,1733]" pageId="7" pageNumber="386">Circumoral spines</emphasis>
|
||
(
|
||
<figureCitation id="91E92A041B4BFF9CAF825CC0FD8CF975" box="[443,547,1707,1733]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="7" pageNumber="386">Figs. 4D</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B4BFF9CAC0A5CC0FDEEF975" box="[563,577,1707,1733]" captionStart="FIGURE 8" captionStartId="12.[152,255,936,960]" captionTargetBox="[482,1106,192,868]" captionTargetId="figure-35@12.[472,1114,181,877]" captionTargetPageId="12" captionText="FIGURE 8. Circumoral spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329755" httpUri="https://zenodo.org/record/7329755/files/figure.png" pageId="7" pageNumber="386">8</figureCitation>
|
||
) form a single row surrounding the mouth, and range in length between approximately 9 and
|
||
<quantity id="CE2A9B641B4BFF9CAFB15CA4FE75F95A" box="[392,474,1743,1770]" metricMagnitude="-2" metricUnit="m" metricValue="1.1" pageId="7" pageNumber="386" unit="mm" value="11.0">11 mm</quantity>
|
||
. Groups of 8–12 spines are associated with two large oral ossicles and every ray has two oral ossicles (occurring symmetrically, one on each side of a ray), each with the same number of spines. One group of circumoral spines is part of two adjoining rays. Within each group, the spine that is closest to the mouth is the terminal spine. There are always two terminal spines, which are the longest, and the spines become shorter towards the adambulacral ossicles. All spines are wider and mostly flattened towards the tip, with the flat side facing either the ambulacral furrow or, if terminal, the mouth opening. Spine tips are mostly flat and may have furrows or be smooth; a pointed tip is rare. Granulation may be present.
|
||
</paragraph>
|
||
<paragraph id="096D36811B4BFF93AEFE5DA0FE9CFF49" blockId="7.[151,1437,735,2022]" lastBlockId="8.[151,1437,151,1041]" lastPageId="8" lastPageNumber="387" pageId="7" pageNumber="386">
|
||
<emphasis id="3BA6EA931B4BFF9CAEFE5DA0FEE0F855" bold="true" box="[199,335,1995,2021]" pageId="7" pageNumber="386">Oral spines</emphasis>
|
||
(
|
||
<figureCitation id="91E92A041B4BFF9CAF665DA0FE1CF855" box="[351,435,1995,2021]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="7" pageNumber="386">Fig. 4E</figureCitation>
|
||
) are very abundant, occurring in one or two rows on the oral-intermediate ossicles and in one row on the adambulacral ossicles, with a flat side facing the ambulacral furrow. They are similar to circumoral spines, but smaller, in the range of
|
||
<quantity id="CE2A9B641B44FF93AC2A5AD0FDC1FF65" box="[531,622,187,213]" metricMagnitude="-3" metricUnit="m" metricValue="5.5" metricValueMax="7.0" metricValueMin="4.0" pageId="8" pageNumber="387" unit="mm" value="5.5" valueMax="7.0" valueMin="4.0">4–7 mm</quantity>
|
||
length, and with deeper furrows. The upper outline can show a depression in the middle.
|
||
</paragraph>
|
||
<paragraph id="096D36811B44FF93AEFE5B69FC92FE61" blockId="8.[151,1437,151,1041]" pageId="8" pageNumber="387">
|
||
<emphasis id="3BA6EA931B44FF93AEFE5B69FE61FEAC" bold="true" box="[199,462,258,284]" pageId="8" pageNumber="387">Subambulacral spines</emphasis>
|
||
(
|
||
<figureCitation id="91E92A041B44FF93AFE75B68FD9EFEAD" box="[478,561,259,285]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="8" pageNumber="387">Fig. 4F</figureCitation>
|
||
) fringe the margin of the ambulacral grooves. They are the most abundant and smallest, ranging between
|
||
<quantity id="CE2A9B641B44FF93AFF95B4CFD85FEF1" box="[448,554,295,321]" metricMagnitude="-3" metricUnit="m" metricValue="6.5" metricValueMax="11.0" metricValueMin="2.0" pageId="8" pageNumber="387" unit="mm" value="6.5" valueMax="11.0" valueMin="2.0">2–11 mm</quantity>
|
||
in length, becoming shorter towards the tip of the arm. Three to four spines are grouped in an adambulacral comb on one ossicle, which is connected by tissue at the base; the outer spines within the grouped spines are always the smallest. Each group of spines is associated with one tube foot. Most spine tips are pointed; however, larger spines can have a flattened tip with slight furrows. The shaft of the spines can be bulbous on one side, increasing the width at the middle of the spines.
|
||
</paragraph>
|
||
<paragraph id="096D36811B44FF93AEFE5BB0FDC7FDA9" blockId="8.[151,1437,151,1041]" pageId="8" pageNumber="387">
|
||
All
|
||
<emphasis id="3BA6EA931B44FF93AECB5BB1FE25FE44" bold="true" box="[242,394,474,500]" pageId="8" pageNumber="387">pedicellariae</emphasis>
|
||
, aboral and oral, are straight, bivalved, and alveolar, positioned over a small cavity, or cupule in the underlying ossicle (see
|
||
<bibRefCitation id="6D434B701B44FF93AFDC5B94FDF2FDA9" author="Gale, A. S." box="[485,605,511,537]" pageId="8" pageNumber="387" pagination="1 - 112" refId="ref7872" refString="Gale, A. S. (2011) The phylogeny of post-Palaeozoic Asteroidea (Echinodermata, Neoasteroidea). Special Papers in Palaeontology, 85, 1 - 112. ISBN 978144435029 - 6" type="journal article" year="2011">Gale 2011</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="096D36811B44FF93AEFE5849FEB1FD41" blockId="8.[151,1437,151,1041]" pageId="8" pageNumber="387">
|
||
<emphasis id="3BA6EA931B44FF93AEFE5849FE14FD8C" bold="true" box="[199,443,546,572]" pageId="8" pageNumber="387">Aboral pedicellariae</emphasis>
|
||
(
|
||
<figureCitation id="91E92A041B44FF93AFF55848FD8AFD8D" box="[460,549,547,573]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="8" pageNumber="387">Fig. 4G</figureCitation>
|
||
) are located among the primary spines, secondary spines and papulae, and are mostly very frequent, giving the aboral side of the sea stars a hairy appearance. They are very common on the disc, but less frequent on the arms; however, abundance differs between individuals. They consist of two equally sized valves, ranging between approximately
|
||
<quantity id="CE2A9B641B44FF93AC6158E4FD1AFD19" box="[600,693,655,681]" metricMagnitude="-3" metricUnit="m" metricValue="2.5" metricValueMax="3.0" metricValueMin="2.0" pageId="8" pageNumber="387" unit="mm" value="2.5" valueMax="3.0" valueMin="2.0">2–3 mm</quantity>
|
||
in length, are long and slender, and are nearly symmetrical with a tapering tip. The outer rim of the valves has a fine, tooth-like structure that is uniform but can be more prominent on one side.
|
||
</paragraph>
|
||
<paragraph id="096D36811B44FF93AEFE5891FD03FBA1" blockId="8.[151,1437,151,1041]" pageId="8" pageNumber="387">
|
||
<emphasis id="3BA6EA931B44FF93AEFE5891FE0FFCA4" bold="true" box="[199,416,762,788]" pageId="8" pageNumber="387">Oral pedicellariae</emphasis>
|
||
(
|
||
<figureCitation id="91E92A041B44FF93AF8B5890FDB7FCA5" box="[434,536,763,789]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="8" pageNumber="387">Figs. 4H</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B44FF93AC1C5890FD9CFCA5" box="[549,563,763,789]" captionStart="FIGURE 8" captionStartId="12.[152,255,936,960]" captionTargetBox="[482,1106,192,868]" captionTargetId="figure-35@12.[472,1114,181,877]" captionTargetPageId="12" captionText="FIGURE 8. Circumoral spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329755" httpUri="https://zenodo.org/record/7329755/files/figure.png" pageId="8" pageNumber="387">8</figureCitation>
|
||
) occur at two locations: most commonly next to oral spines or the group of subambulacral spines on the adambulacral ossicle; or more rarely, between the oral spines on the oral ossicles, or exceptionally found on marginal ossicles. The valves are mostly of the same length, ranging between approximately
|
||
<quantity id="CE2A9B641B44FF93AEAE590DFEA7FC31" box="[151,264,870,897]" metricMagnitude="-3" metricUnit="m" metricValue="1.75" metricValueMax="2.0" metricValueMin="1.5" pageId="8" pageNumber="387" unit="mm" value="1.75" valueMax="2.0" valueMin="1.5">1.5–2 mm</quantity>
|
||
in length. The overall shape is very variable, however there are two main shapes: i) smaller and irregularly formed with a hook-shaped tip, wide from the side, slender from the front, rounded shaft under the “hook” which may have small, asymmetrically-arranged teeth; or ii) flatter, largely resembling the shape of a rhombus due to a widening in the middle part and a pointed tip, with teeth occurring asymmetrically on the outer rim and occasionally on the inner surface, if the area is large enough.
|
||
</paragraph>
|
||
<caption id="5DAD66091B44FF93AEA15D60FB54F8DB" ID-DOI="http://doi.org/10.5281/zenodo.7329747" ID-Zenodo-Dep="7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="8" pageNumber="387" startId="8.[152,255,1803,1827]" targetBox="[151,1436,1065,1779]" targetPageId="8">
|
||
<paragraph id="096D36811B44FF93AEA15D60FB54F8DB" blockId="8.[151,1437,1802,1899]" pageId="8" pageNumber="387">
|
||
<emphasis id="3BA6EA931B44FF93AEA15D60FCB3F893" bold="true" box="[152,796,1802,1827]" pageId="8" pageNumber="387">
|
||
FIGURE 4.
|
||
<taxonomicName id="CED24D021B44FF93AF275D61FE43F892" authority="Wörheide & Kaltenbacher & Cowan & Haszprunar, 2022" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[286,492,1802,1826]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="8" pageNumber="387" phylum="Echinodermata" rank="species" species="benziei" status="sp. nov.">
|
||
<emphasis id="3BA6EA931B44FF93AF275D61FE43F892" bold="true" box="[286,492,1802,1826]" italics="true" pageId="8" pageNumber="387">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="209557E81B44FF93AFCB5D60FDEDF893" box="[498,578,1803,1827]" pageId="8" pageNumber="387" rank="species">sp. nov.</taxonomicNameLabel>
|
||
(holotype GW4202)
|
||
</emphasis>
|
||
(A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of
|
||
<taxonomicName id="CED24D021B44FF93AF935D44FDDCF8F7" authority="Wörheide & Kaltenbacher & Cowan & Haszprunar, 2022" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[426,627,1839,1863]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="8" pageNumber="387" phylum="Echinodermata" rank="species" species="benziei" status="sp. nov.">
|
||
<emphasis id="3BA6EA931B44FF93AF935D44FDDCF8F7" box="[426,627,1839,1863]" italics="true" pageId="8" pageNumber="387">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="209557E81B44FF93AC435D44FD63F8F6" box="[634,716,1839,1862]" pageId="8" pageNumber="387" rank="species">sp. nov.</taxonomicNameLabel>
|
||
: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="096D36811B44FF93AEFE5DF2FC5BF84B" blockId="8.[151,1436,1945,2044]" pageId="8" pageNumber="387">
|
||
<emphasis id="3BA6EA931B44FF93AEFE5DF2FE8FF803" bold="true" box="[199,288,1945,1971]" pageId="8" pageNumber="387">Colour.</emphasis>
|
||
The colouration of live adult specimens is grey-green to grey-purple, with aboral spines that can be orange to reddish in colour (
|
||
<figureCitation id="91E92A041B44FF93AFF65DD6FDBDF868" box="[463,530,1981,2008]" captionStart="FIGURE 5" captionStartId="9.[152,255,1202,1226]" captionTargetBox="[151,1436,201,1178]" captionTargetId="figure-19@9.[151,1436,201,1178]" captionTargetPageId="9" captionText="FIGURE 5. Typical colouration of Acanthaster benziei sp. nov. (A) GW4081 (Paratype, hiding during the day under a crevice), Al-Lith, Saudi Arabia, (photo credit: Oliver Voigt), (B–D) Thuwal Reefs, Saudi Arabia (photo credit: Gert Wörheide). Approximate diameter of specimens is 25–30 cm." figureDoi="http://doi.org/10.5281/zenodo.7329749" httpUri="https://zenodo.org/record/7329749/files/figure.png" pageId="8" pageNumber="387">Fig. 5</figureCitation>
|
||
). Reddish papulae on the aboral surface may also give a bulls-eye appearance due to the formation of two darker rings (
|
||
<bibRefCitation id="6D434B701B44FF93AC0E5D8AFCE1F84B" author="Birkeland, C. & Lucas, J. S." box="[567,846,2017,2044]" pageId="8" pageNumber="387" refId="ref7443" refString="Birkeland, C. & Lucas, J. S. (1990) Acanthaster planci: MaJor Management Problem of Coral Reefs. CRC Press, Boca Raton, Florida, 262 pp." type="book" year="1990">Birkeland & Lucas 1990</bibRefCitation>
|
||
) (
|
||
<figureCitation id="91E92A041B44FF93AD5C5D8AFC47F84B" box="[869,1000,2017,2044]" captionStart="FIGURE 5" captionStartId="9.[152,255,1202,1226]" captionTargetBox="[151,1436,201,1178]" captionTargetId="figure-19@9.[151,1436,201,1178]" captionTargetPageId="9" captionText="FIGURE 5. Typical colouration of Acanthaster benziei sp. nov. (A) GW4081 (Paratype, hiding during the day under a crevice), Al-Lith, Saudi Arabia, (photo credit: Oliver Voigt), (B–D) Thuwal Reefs, Saudi Arabia (photo credit: Gert Wörheide). Approximate diameter of specimens is 25–30 cm." figureDoi="http://doi.org/10.5281/zenodo.7329749" httpUri="https://zenodo.org/record/7329749/files/figure.png" pageId="8" pageNumber="387">Fig. 5 B, D</figureCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<caption id="5DAD66091B45FF92AEA15ED9FD3FFAA2" ID-DOI="http://doi.org/10.5281/zenodo.7329749" ID-Zenodo-Dep="7329749" httpUri="https://zenodo.org/record/7329749/files/figure.png" pageId="9" pageNumber="388" startId="9.[152,255,1202,1226]" targetBox="[151,1436,201,1178]" targetPageId="9">
|
||
<paragraph id="096D36811B45FF92AEA15ED9FD3FFAA2" blockId="9.[151,1437,1202,1298]" pageId="9" pageNumber="388">
|
||
<emphasis id="3BA6EA931B45FF92AEA15ED9FEB8FB7A" bold="true" box="[152,279,1202,1226]" pageId="9" pageNumber="388">FIGURE 5</emphasis>
|
||
. Typical colouration of
|
||
<taxonomicName id="CED24D021B45FF92AC255ED9FD45FB7A" authority="Wörheide & Kaltenbacher & Cowan & Haszprunar, 2022" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[540,746,1202,1226]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="benziei" status="sp. nov.">
|
||
<emphasis id="3BA6EA931B45FF92AC255ED9FD45FB7A" box="[540,746,1202,1226]" italics="true" pageId="9" pageNumber="388">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="209557E81B45FF92ACCD5ED8FCE7FB7A" box="[756,840,1203,1226]" pageId="9" pageNumber="388" rank="species">sp. nov.</taxonomicNameLabel>
|
||
(A) GW4081 (Paratype, hiding during the day under a crevice), Al-Lith, Saudi Arabia, (photo credit: Oliver Voigt), (B–D) Thuwal Reefs, Saudi Arabia (photo credit: Gert Wörheide). Approximate diameter of specimens is 25–30 cm.
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection id="41C8650A1B45FF92AEFE5F0FFE8FFA76" pageId="9" pageNumber="388" type="distribution">
|
||
<paragraph id="096D36811B45FF92AEFE5F0FFE8FFA76" blockId="9.[151,1437,1380,1946]" pageId="9" pageNumber="388">
|
||
<emphasis id="3BA6EA931B45FF92AEFE5F0FFE55FACE" bold="true" box="[199,506,1380,1406]" pageId="9" pageNumber="388">Distribution and habitat.</emphasis>
|
||
So far known
|
||
<taxonomicName id="CED24D021B45FF92AC9E5F0FFC24FACE" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[679,907,1380,1406]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B45FF92AC9E5F0FFC24FACE" box="[679,907,1380,1406]" italics="true" pageId="9" pageNumber="388">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
is restricted to the Red Sea, where it inhabits coral reefs, predominantly the outer reef surfaces where it mostly hides in crevices during the day and feeds nocturnally.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="41C8650A1B45FF90AEFE5FBBFE68F8C6" lastPageId="11" lastPageNumber="390" pageId="9" pageNumber="388" type="diagnosis">
|
||
<paragraph id="096D36811B45FF92AEFE5FBBFCDCF92E" blockId="9.[151,1437,1380,1946]" pageId="9" pageNumber="388">
|
||
<emphasis id="3BA6EA931B45FF92AEFE5FBBFE7BFA5A" bold="true" box="[199,468,1488,1514]" pageId="9" pageNumber="388">Differential diagnosis.</emphasis>
|
||
<taxonomicName id="CED24D021B45FF92AFE25FBBFD11FA5A" authority="Wörheide & Kaltenbacher & Cowan & Haszprunar, 2022" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[475,702,1488,1514]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="benziei" status="sp. nov.">
|
||
<emphasis id="3BA6EA931B45FF92AFE25FBBFD11FA5A" box="[475,702,1488,1514]" italics="true" pageId="9" pageNumber="388">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="209557E81B45FF92ACFF5FBAFCB0FA5B" box="[710,799,1489,1515]" pageId="9" pageNumber="388" rank="species">sp. nov.</taxonomicNameLabel>
|
||
can clearly be distinguished by diagnostic mutations in the partial COI gene sequences analysed, all material examined fall within the deeply divergent monophyletic group of CoTS from the Red Sea (see
|
||
<figureCitation id="91E92A041B45FF92AC6E5C73FD32F982" box="[599,669,1560,1586]" captionStart="FIGURE 3" captionStartId="6.[152,255,1199,1223]" captionTargetBox="[395,1195,188,1170]" captionTargetId="figure-35@6.[374,1212,181,1175]" captionTargetPageId="6" captionText="FIGURE 3. A: Indo-Pacific ‘Acanthaster planci’ species complex, COI ML tree based on the haplotype alignment of Vogler et al. (2008), supplemented with five samples from Israel (see text for details) extracted from full mitochondrial genome sequences from Yuasa et al. (2021) and four sequences of the type series of Acanthaster benziei sp. nov. (GW4xxx, highlighted in bold). The tree was rooted with Acanthaster brevispinus (accession number AB231476), showing the deep divergence among, and little diversity within, species/geographic clades. ML bootstrap values are above branches, bootstrap values of the NJ clustering of haplotypes below branches. B: Geographic distribution of COI-barcoded clades and of type localities of names (Figure 1 from Haszprunar et al. 2017): red—Red Sea (RS) species; blue—Southern Indian Ocean (SIO) species (A. mauritiensis); yellow—Northern Indian Ocean (NIO) species (A. planci); green—Pacific Ocean (PO) species (A. cf. solaris). Location of type localities of nominal Acanthaster species: asterisk—A. planci; cross—A. echinites; triangle—A. solaris, square—A. mauritiensis; circle—A. ellisii pseudoplanci; “?” - the type locality of A. ellisii was not specified: in South American waters of the East Pacific." figureDoi="http://doi.org/10.5281/zenodo.7329745" httpUri="https://zenodo.org/record/7329745/files/figure.png" pageId="9" pageNumber="388">Fig. 3</figureCitation>
|
||
). There is full agreement with initial results that proposed species distinction of the Red Sea clade based on COI data alone (
|
||
<bibRefCitation id="6D434B701B45FF92AD025C57FBBFF9E6" author="Vogler, C. & Benzie, J. & Lessios, H. & Barber, P. & Worheide, G." box="[827,1040,1596,1622]" pageId="9" pageNumber="388" pagination="696 - 699" refId="ref9248" refString="Vogler, C., Benzie, J., Lessios, H., Barber, P. & Worheide, G. (2008) A threat to coral reefs multiplied? Four species of crownof-thorns starfish. Biology Letters, 4 (6), 696 - 699. https: // doi. org / 10.1098 / rsbl. 2008.0454" type="journal article" year="2008">
|
||
Vogler
|
||
<emphasis id="3BA6EA931B45FF92ADA85C56FC64F9E6" box="[913,971,1596,1622]" italics="true" pageId="9" pageNumber="388">et al.</emphasis>
|
||
2008
|
||
</bibRefCitation>
|
||
), recently corroborated by nuclear genome analyses (
|
||
<bibRefCitation id="6D434B701B45FF92AF485C0BFDE5F9CA" author="Yuasa, H. & Kajitani, R. & Nakamura, Y. & Takahashi, K. & Okuno, M. & Kobayashi, F. & Shinoda, T. & Toyoda, A. & Suzuki, Y. & Thongtham, N. & Forsman, Z. & Bronstein, O. & Seveso, D. & Montalbetti, E. & Taquet, C. & Eyal, G. & Yasuda, N. & Itoh, T." box="[369,586,1632,1658]" pageId="9" pageNumber="388" pagination="4" refId="ref9464" refString="Yuasa, H., Kajitani, R., Nakamura, Y., Takahashi, K., Okuno, M., Kobayashi, F., Shinoda, T., Toyoda, A., Suzuki, Y., Thongtham, N., Forsman, Z., Bronstein, O., Seveso, D., Montalbetti, E., Taquet, C., Eyal, G., Yasuda, N. & Itoh, T. (2021) Elucidation of the speciation history of three sister species of crown-of-thorns starfish (Acanthaster spp.) based on genomic analysis. DNA Research, 28 (4), dsab 012. https: // doi. org / 10.1093 / dnares / dsab 012" type="journal article" year="2021">
|
||
Yuasa
|
||
<emphasis id="3BA6EA931B45FF92AFFA5C0AFDAFF9CA" box="[451,512,1632,1658]" italics="true" pageId="9" pageNumber="388">et al.</emphasis>
|
||
2021
|
||
</bibRefCitation>
|
||
). The molecular-based species distinction of
|
||
<taxonomicName id="CED24D021B45FF92AA5E5C0AFB73F9CA" authority="Wörheide & Kaltenbacher & Cowan & Haszprunar, 2022" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[1127,1244,1632,1659]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="benziei" status="sp. nov.">
|
||
<emphasis id="3BA6EA931B45FF92AA5E5C0AFB73F9CA" box="[1127,1244,1632,1659]" italics="true" pageId="9" pageNumber="388">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="209557E81B45FF92AAD15C0AFAEAF9CB" box="[1256,1349,1633,1659]" pageId="9" pageNumber="388" rank="species">sp. nov.</taxonomicNameLabel>
|
||
is also substantially supported by diagnostic morphological characters.
|
||
</paragraph>
|
||
<paragraph id="096D36811B45FF92AEFE5CC3FE97F82A" blockId="9.[151,1437,1380,1946]" pageId="9" pageNumber="388">
|
||
<taxonomicName id="CED24D021B45FF92AEFE5CC3FE05F972" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[199,426,1704,1730]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B45FF92AEFE5CC3FE05F972" box="[199,426,1704,1730]" italics="true" pageId="9" pageNumber="388">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
has fewer arms than congeneric sea stars of comparable size from the other geographic regions/species. Our reported range between 11 and 14 arms in the
|
||
<typeStatus id="D66988231B45FF92ADA65CA6FC7FF957" box="[927,976,1741,1767]" pageId="9" pageNumber="388">type</typeStatus>
|
||
series (
|
||
<tableCitation id="4450033A1B45FF92AA105CA7FBDDF956" box="[1065,1138,1740,1766]" captionStart="TABLE 3" captionStartId="7.[152,245,151,175]" captionTargetPageId="7" captionText="TABLE 3: Basic measurements of type specimens, including number of arms and madreporites, mean length of arms (R) and radius of disk (r). Note that GW4266 is a juvenile specimen." httpUri="http://table.plazi.org/id/5DAD66091B4BFF9CAEA15AFCFCE5FF63" pageId="9" pageNumber="388" tableUuid="5DAD66091B4BFF9CAEA15AFCFCE5FF63">Tab. 3</tableCitation>
|
||
) is consistent with values previously reported for other individuals from the Red Sea (mean of 13 arms [
|
||
<bibRefCitation id="6D434B701B45FF92AA005C9BFA3EF8BA" author="Campbell, A. C. & Ormond, R. F. G." box="[1081,1425,1776,1802]" pageId="9" pageNumber="388" pagination="246 - 251" refId="ref7634" refString="Campbell, A. C. & Ormond, R. F. G. (1970) The threat of the " crown-of-thorns " starfish (Acanthaster planci) to coral reefs in the Indo-Pacific area: observations on a normal population in the Red Sea. Biological Conservation, 2 (4), 246 - 251. https: // doi. org / 10.1016 / 0006 - 3207 (70) 90004 - 2" type="journal article" year="1970">Campbell and Ormond 1970</bibRefCitation>
|
||
]; maximum of 13–14 arms [
|
||
<bibRefCitation id="6D434B701B45FF92AFFF5D7EFD79F89E" author="Haszprunar, G. & Vogler, C. & Worheide, G." box="[454,726,1812,1839]" pageId="9" pageNumber="388" pagination="22" refId="ref8118" refString="Haszprunar, G., Vogler, C. & Worheide, G. (2017) Persistent gaps of knowledge for naming and distinguishing multiple species of crown-of-thorns-seastar in the Acanthaster planci species complex. Diversity, 9 (2), 22. https: // doi. org / 10.3390 / d 9020022" type="journal article" year="2017">
|
||
Haszprunar
|
||
<emphasis id="3BA6EA931B45FF92AC6C5D7EFD20F89E" box="[597,655,1812,1838]" italics="true" pageId="9" pageNumber="388">et al.</emphasis>
|
||
2017
|
||
</bibRefCitation>
|
||
]). By contrast, the number of arms reported for sea stars from
|
||
<collectingCountry id="71C576111B45FF92AEAE5D53FF7DF8E2" box="[151,210,1848,1874]" name="India" pageId="9" pageNumber="388">India</collectingCountry>
|
||
(=
|
||
<taxonomicName id="CED24D021B45FF92AEC25D52FECDF8E2" box="[251,354,1848,1875]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B45FF92AEC25D52FECDF8E2" box="[251,354,1848,1875]" italics="true" pageId="9" pageNumber="388">A. planci</emphasis>
|
||
</taxonomicName>
|
||
) was 15 (Linnaeus 1758) and for
|
||
<taxonomicName id="CED24D021B45FF92ACD35D52FC37F8E2" box="[746,920,1849,1875]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B45FF92ACD35D52FC37F8E2" box="[746,920,1849,1875]" italics="true" pageId="9" pageNumber="388">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
13–16 (
|
||
<bibRefCitation id="6D434B701B45FF92ADCF5D53FB09F8E3" author="de Loriol, P." box="[1014,1190,1848,1875]" pageId="9" pageNumber="388" refId="ref8364" refString="de Loriol, P. (1885) Catalogue raisonne des Echinodermes recueillis par M. V. de Robillard a l'ile Maurice. Memoires de la societe de physique et d'histoire naturelle de Geneve, 29 (1 re Partie), No 4, 84 pp., pls. VII - XXII. [in French] https: // www. biodiversitylibrary. org / page / 36249315 (accessed 24 October 2022)" type="book" year="1885">de Loriol 1885</bibRefCitation>
|
||
), with a maximum of 23 arms reported for
|
||
<taxonomicName id="CED24D021B45FF92AFBE5D36FE42F8C6" box="[391,493,1884,1911]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B45FF92AFBE5D36FE42F8C6" box="[391,493,1884,1911]" italics="true" pageId="9" pageNumber="388">A. planci</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicName id="CED24D021B45FF92AFC55D36FD06F8C6" box="[508,681,1885,1911]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B45FF92AFC55D36FD06F8C6" box="[508,681,1885,1911]" italics="true" pageId="9" pageNumber="388">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
and the Pacific species
|
||
<taxonomicName id="CED24D021B45FF92AD805D36FAC7F8C6" authority="(Haszprunar et al. 2017)" baseAuthorityName="Haszprunar" baseAuthorityYear="2017" box="[953,1384,1884,1911]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" isUncertain="true" kingdom="Animalia" order="Valvatida" pageId="9" pageNumber="388" phylum="Echinodermata" rank="species" species="solaris">
|
||
<emphasis id="3BA6EA931B45FF92AD805D36FC7EF8C7" box="[953,977,1885,1911]" italics="true" pageId="9" pageNumber="388">A.</emphasis>
|
||
cf.
|
||
<emphasis id="3BA6EA931B45FF92ADC55D37FBE5F8C6" box="[1020,1098,1884,1910]" italics="true" pageId="9" pageNumber="388">solaris</emphasis>
|
||
(
|
||
<bibRefCitation id="6D434B701B45FF92AA605D37FACFF8C6" author="Haszprunar, G. & Vogler, C. & Worheide, G." box="[1113,1376,1884,1910]" pageId="9" pageNumber="388" pagination="22" refId="ref8118" refString="Haszprunar, G., Vogler, C. & Worheide, G. (2017) Persistent gaps of knowledge for naming and distinguishing multiple species of crown-of-thorns-seastar in the Acanthaster planci species complex. Diversity, 9 (2), 22. https: // doi. org / 10.3390 / d 9020022" type="journal article" year="2017">
|
||
Haszprunar
|
||
<emphasis id="3BA6EA931B45FF92AADC5D36FAB1F8C6" box="[1253,1310,1884,1910]" italics="true" pageId="9" pageNumber="388">et al.</emphasis>
|
||
2017
|
||
</bibRefCitation>
|
||
)
|
||
</taxonomicName>
|
||
(see Supp.
|
||
<tableCitation id="4450033A1B45FF92AED95DEBFE89F82A" box="[224,294,1920,1946]" captionStart="TABLE 1" captionStartId="1.[152,245,1448,1472]" captionTargetBox="[163,1354,1560,1951]" captionTargetPageId="1" captionText="TABLE 1. The type series investigated, including sampling location, details about preservation, and voucher location. BSPG refers to the ‘Bavarian State Collection for Palaeontology and Geology’ of the Bavarian Natural History Collections" httpUri="http://table.plazi.org/id/5DAD66091B4DFF9AAEA15FC3FB03F9B9" pageId="9" pageNumber="388" tableUuid="5DAD66091B4DFF9AAEA15FC3FB03F9B9">Tab. 1</tableCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="5DAD66091B46FF91AEF15E99FD23FA9D" ID-DOI="http://doi.org/10.5281/zenodo.7329751" ID-Zenodo-Dep="7329751" httpUri="https://zenodo.org/record/7329751/files/figure.png" pageId="10" pageNumber="389" startId="10.[200,303,1266,1290]" targetBox="[479,1103,191,1235]" targetPageId="10">
|
||
<paragraph id="096D36811B46FF91AEF15E99FD23FA9D" blockId="10.[151,1437,1266,1326]" pageId="10" pageNumber="389">
|
||
<emphasis id="3BA6EA931B46FF91AEF15E99FEEDFABA" bold="true" box="[200,322,1266,1290]" pageId="10" pageNumber="389">FIGURE 6</emphasis>
|
||
. Typical primary spines, showing species-specific variation among (A, B)
|
||
<taxonomicName id="CED24D021B46FF91AA0F5E99FB50FABA" authority="Wörheide & Kaltenbacher & Cowan & Haszprunar, 2022" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[1078,1279,1266,1290]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="benziei" status="sp. nov.">
|
||
<emphasis id="3BA6EA931B46FF91AA0F5E99FB50FABA" box="[1078,1279,1266,1290]" italics="true" pageId="10" pageNumber="389">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="209557E81B46FF91AB3D5E99FAFAFAB9" box="[1284,1365,1266,1289]" pageId="10" pageNumber="389" rank="species">sp. nov.</taxonomicNameLabel>
|
||
, (C)
|
||
<taxonomicName id="CED24D021B46FF91ABBF5E99FF79FA9E" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B46FF91ABBF5E99FF79FA9E" italics="true" pageId="10" pageNumber="389">A. planci</emphasis>
|
||
</taxonomicName>
|
||
, (D)
|
||
<taxonomicName id="CED24D021B46FF91AF345F7DFE09FA9E" box="[269,422,1302,1326]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B46FF91AF345F7DFE09FA9E" box="[269,422,1302,1326]" italics="true" pageId="10" pageNumber="389">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
, and (E)
|
||
<taxonomicName id="CED24D021B46FF91AC3D5F7DFD29FA9E" box="[516,646,1302,1326]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" isUncertain="true" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="solaris">
|
||
<emphasis id="3BA6EA931B46FF91AC3D5F7DFDBCFA9D" box="[516,531,1302,1325]" italics="true" pageId="10" pageNumber="389">A</emphasis>
|
||
. cf.
|
||
<emphasis id="3BA6EA931B46FF91AC785F7DFD29FA9E" box="[577,646,1302,1326]" italics="true" pageId="10" pageNumber="389">solaris</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="096D36811B46FF91AEFE5F18FAF4F9AD" blockId="10.[151,1437,1395,2033]" pageId="10" pageNumber="389">
|
||
The spines and pedicellariae of
|
||
<taxonomicName id="CED24D021B46FF91AC0C5F18FD0AFA3D" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[565,677,1395,1421]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B46FF91AC0C5F18FD0AFA3D" box="[565,677,1395,1421]" italics="true" pageId="10" pageNumber="389">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
are more variable, and spines are narrower and thinner than in its congeners. The pointy spine
|
||
<typeStatus id="D66988231B46FF91AFE25FFCFDA4FA01" box="[475,523,1431,1457]" pageId="10" pageNumber="389">type</typeStatus>
|
||
was not found in such high abundance in specimens outside the Red Sea (
|
||
<figureCitation id="91E92A041B46FF91AB6B5FFCFA38FA01" box="[1362,1431,1431,1457]" captionStart="FIGURE 6" captionStartId="10.[200,303,1266,1290]" captionTargetBox="[479,1103,191,1235]" captionTargetId="figure-35@10.[472,1115,181,1242]" captionTargetPageId="10" captionText="FIGURE 6. Typical primary spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329751" httpUri="https://zenodo.org/record/7329751/files/figure.png" pageId="10" pageNumber="389">Fig. 6</figureCitation>
|
||
, Suppl.
|
||
<figureCitation id="91E92A041B46FF91AEDF5FD0FEF8FA65" box="[230,343,1467,1493]" captionStart-0="FIGURE 2" captionStart-1="FIGURE 3" captionStart-2="FIGURE 4" captionStartId-0="5.[152,255,1323,1347]" captionStartId-1="6.[152,255,1199,1223]" captionStartId-2="8.[152,255,1803,1827]" captionTargetBox-0="[151,1436,181,1300]" captionTargetBox-1="[395,1195,188,1170]" captionTargetBox-2="[151,1436,1065,1779]" captionTargetId-0="figure-19@5.[151,1436,181,1300]" captionTargetId-1="figure-35@6.[374,1212,181,1175]" captionTargetId-2="figure-448@8.[151,1436,1065,1779]" captionTargetPageId-0="5" captionTargetPageId-1="6" captionTargetPageId-2="8" captionText-0="FIGURE 2. Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm." captionText-1="FIGURE 3. A: Indo-Pacific ‘Acanthaster planci’ species complex, COI ML tree based on the haplotype alignment of Vogler et al. (2008), supplemented with five samples from Israel (see text for details) extracted from full mitochondrial genome sequences from Yuasa et al. (2021) and four sequences of the type series of Acanthaster benziei sp. nov. (GW4xxx, highlighted in bold). The tree was rooted with Acanthaster brevispinus (accession number AB231476), showing the deep divergence among, and little diversity within, species/geographic clades. ML bootstrap values are above branches, bootstrap values of the NJ clustering of haplotypes below branches. B: Geographic distribution of COI-barcoded clades and of type localities of names (Figure 1 from Haszprunar et al. 2017): red—Red Sea (RS) species; blue—Southern Indian Ocean (SIO) species (A. mauritiensis); yellow—Northern Indian Ocean (NIO) species (A. planci); green—Pacific Ocean (PO) species (A. cf. solaris). Location of type localities of nominal Acanthaster species: asterisk—A. planci; cross—A. echinites; triangle—A. solaris, square—A. mauritiensis; circle—A. ellisii pseudoplanci; “?” - the type locality of A. ellisii was not specified: in South American waters of the East Pacific." captionText-2="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi-0="http://doi.org/10.5281/zenodo.7329743" figureDoi-1="http://doi.org/10.5281/zenodo.7329745" figureDoi-2="http://doi.org/10.5281/zenodo.7329747" httpUri-0="https://zenodo.org/record/7329743/files/figure.png" httpUri-1="https://zenodo.org/record/7329745/files/figure.png" httpUri-2="https://zenodo.org/record/7329747/files/figure.png" pageId="10" pageNumber="389">Fig. S2–4</figureCitation>
|
||
). Unique for
|
||
<taxonomicName id="CED24D021B46FF91AFD25FD0FDF7FA65" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[491,600,1467,1493]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B46FF91AFD25FD0FDF7FA65" box="[491,600,1467,1493]" italics="true" pageId="10" pageNumber="389">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
are the fanned primary spines (
|
||
<figureCitation id="91E92A041B46FF91AD805FD0FBBEFA65" box="[953,1041,1467,1493]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="10" pageNumber="389">Fig. 4A</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B46FF91AA245FD0FBEFFA65" box="[1053,1088,1467,1493]" captionStart="FIGURE 6" captionStartId="10.[200,303,1266,1290]" captionTargetBox="[479,1103,191,1235]" captionTargetId="figure-35@10.[472,1115,181,1242]" captionTargetPageId="10" captionText="FIGURE 6. Typical primary spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329751" httpUri="https://zenodo.org/record/7329751/files/figure.png" pageId="10" pageNumber="389">6A</figureCitation>
|
||
) and the distally fanned laterooral spines, which can also be granulated (
|
||
<figureCitation id="91E92A041B46FF91AC405FB4FD7EFA49" box="[633,721,1503,1529]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="10" pageNumber="389">Fig. 4C</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B46FF91ACE75FB4FCACFA49" box="[734,771,1503,1529]" captionStart="FIGURE 7" captionStartId="11.[152,255,1418,1442]" captionTargetBox="[479,1108,204,1389]" captionTargetId="figure-19@11.[472,1115,199,1394]" captionTargetPageId="11" captionText="FIGURE 7. Typical latero-oral spines, showing species-specific variation among (A) Acanthaster benziei sp. nov., (B) A. planci, (C) A. mauritiensis, and (D) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329753" httpUri="https://zenodo.org/record/7329753/files/figure.png" pageId="10" pageNumber="389">7A</figureCitation>
|
||
). The granulation of the latero-oral spines is rare, and may also be found in
|
||
<emphasis id="3BA6EA931B46FF91AF6B5C68FDADF9AC" box="[338,514,1539,1565]" italics="true" pageId="10" pageNumber="389">
|
||
<taxonomicName id="CED24D021B46FF91AF6B5C68FE51F9AC" box="[338,510,1539,1565]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="mauritiensis">A. mauritiensis</taxonomicName>
|
||
,
|
||
</emphasis>
|
||
where the longest spines have an arrow-head tip (
|
||
<figureCitation id="91E92A041B46FF91AA0D5C68FB23F9AD" box="[1076,1164,1539,1565]" captionStart="FIGURE 7" captionStartId="11.[152,255,1418,1442]" captionTargetBox="[479,1108,204,1389]" captionTargetId="figure-19@11.[472,1115,199,1394]" captionTargetPageId="11" captionText="FIGURE 7. Typical latero-oral spines, showing species-specific variation among (A) Acanthaster benziei sp. nov., (B) A. planci, (C) A. mauritiensis, and (D) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329753" httpUri="https://zenodo.org/record/7329753/files/figure.png" pageId="10" pageNumber="389">Fig. 7C</figureCitation>
|
||
; Suppl.
|
||
<figureCitation id="91E92A041B46FF91AADE5C68FAE1F9AD" box="[1255,1358,1539,1565]" captionStart="FIGURE 3" captionStartId="6.[152,255,1199,1223]" captionTargetBox="[395,1195,188,1170]" captionTargetId="figure-35@6.[374,1212,181,1175]" captionTargetPageId="6" captionText="FIGURE 3. A: Indo-Pacific ‘Acanthaster planci’ species complex, COI ML tree based on the haplotype alignment of Vogler et al. (2008), supplemented with five samples from Israel (see text for details) extracted from full mitochondrial genome sequences from Yuasa et al. (2021) and four sequences of the type series of Acanthaster benziei sp. nov. (GW4xxx, highlighted in bold). The tree was rooted with Acanthaster brevispinus (accession number AB231476), showing the deep divergence among, and little diversity within, species/geographic clades. ML bootstrap values are above branches, bootstrap values of the NJ clustering of haplotypes below branches. B: Geographic distribution of COI-barcoded clades and of type localities of names (Figure 1 from Haszprunar et al. 2017): red—Red Sea (RS) species; blue—Southern Indian Ocean (SIO) species (A. mauritiensis); yellow—Northern Indian Ocean (NIO) species (A. planci); green—Pacific Ocean (PO) species (A. cf. solaris). Location of type localities of nominal Acanthaster species: asterisk—A. planci; cross—A. echinites; triangle—A. solaris, square—A. mauritiensis; circle—A. ellisii pseudoplanci; “?” - the type locality of A. ellisii was not specified: in South American waters of the East Pacific." figureDoi="http://doi.org/10.5281/zenodo.7329745" httpUri="https://zenodo.org/record/7329745/files/figure.png" pageId="10" pageNumber="389">Fig. S3C</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="096D36811B46FF91AEFE5C4CFC15F8A9" blockId="10.[151,1437,1395,2033]" pageId="10" pageNumber="389">
|
||
Compared to spines of its congeners, there are some key differences: primary spines of
|
||
<taxonomicName id="CED24D021B46FF91AA8E5C4CFAB1F9F1" box="[1207,1310,1575,1601]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B46FF91AA8E5C4CFAB1F9F1" box="[1207,1310,1575,1601]" italics="true" pageId="10" pageNumber="389">A. planci</emphasis>
|
||
</taxonomicName>
|
||
show only one tip-shape, which resembles an arrowhead (
|
||
<figureCitation id="91E92A041B46FF91ACFA5C20FC8FF9D5" box="[707,800,1611,1637]" captionStart="FIGURE 6" captionStartId="10.[200,303,1266,1290]" captionTargetBox="[479,1103,191,1235]" captionTargetId="figure-35@10.[472,1115,181,1242]" captionTargetPageId="10" captionText="FIGURE 6. Typical primary spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329751" httpUri="https://zenodo.org/record/7329751/files/figure.png" pageId="10" pageNumber="389">Fig. 6C</figureCitation>
|
||
; Suppl.
|
||
<figureCitation id="91E92A041B46FF91ADBA5C20FC40F9D5" box="[899,1007,1611,1637]" captionStart="FIGURE 2" captionStartId="5.[152,255,1323,1347]" captionTargetBox="[151,1436,181,1300]" captionTargetId="figure-19@5.[151,1436,181,1300]" captionTargetPageId="5" captionText="FIGURE 2. Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm." figureDoi="http://doi.org/10.5281/zenodo.7329743" httpUri="https://zenodo.org/record/7329743/files/figure.png" pageId="10" pageNumber="389">Fig. S2A</figureCitation>
|
||
) – this was not found in
|
||
<taxonomicName id="CED24D021B46FF91AB195C20FA3BF9D5" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[1312,1428,1611,1637]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B46FF91AB195C20FA3BF9D5" box="[1312,1428,1611,1637]" italics="true" pageId="10" pageNumber="389">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
; secondary spines are longer compared to primary spines in
|
||
<taxonomicName id="CED24D021B46FF91AD0E5C04FC08F939" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[823,935,1647,1673]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B46FF91AD0E5C04FC08F939" box="[823,935,1647,1673]" italics="true" pageId="10" pageNumber="389">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
than in
|
||
<taxonomicName id="CED24D021B46FF91AA385C04FB01F938" box="[1025,1198,1647,1673]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B46FF91AA385C04FB01F938" box="[1025,1198,1647,1673]" italics="true" pageId="10" pageNumber="389">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
(around half to three quarters the size of the primary spines),
|
||
<taxonomicName id="CED24D021B46FF91AC675CF8FD6BF91D" box="[606,708,1683,1709]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B46FF91AC675CF8FD6BF91D" box="[606,708,1683,1709]" italics="true" pageId="10" pageNumber="389">A. planci</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="CED24D021B46FF91ACC25CF8FC25F91D" box="[763,906,1683,1709]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" isUncertain="true" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="solaris">
|
||
<emphasis id="3BA6EA931B46FF91ACC25CF8FC25F91D" box="[763,906,1683,1709]" italics="true" pageId="10" pageNumber="389">A. cf. solaris</emphasis>
|
||
</taxonomicName>
|
||
(around one quarter to half the size of primary spines); and the second articulation of primary spines is only found in
|
||
<taxonomicName id="CED24D021B46FF91ADF15CDCFB96F961" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[968,1081,1719,1745]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B46FF91ADF15CDCFB96F961" box="[968,1081,1719,1745]" italics="true" pageId="10" pageNumber="389">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
. Primary spines in CoTS from the Red Sea are also considered to be less harmful than spines of other regions, which could be connected to the shape, and they seem to have less toxins (
|
||
<bibRefCitation id="6D434B701B46FF91AC4D5C94FC01F8A9" author="Campbell, A. C. & Ormond, R. F. G." box="[628,942,1791,1817]" pageId="10" pageNumber="389" pagination="246 - 251" refId="ref7634" refString="Campbell, A. C. & Ormond, R. F. G. (1970) The threat of the " crown-of-thorns " starfish (Acanthaster planci) to coral reefs in the Indo-Pacific area: observations on a normal population in the Red Sea. Biological Conservation, 2 (4), 246 - 251. https: // doi. org / 10.1016 / 0006 - 3207 (70) 90004 - 2" type="journal article" year="1970">Campbell & Ormond 1970</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="096D36811B46FF91AEFE5D48FBB0F841" blockId="10.[151,1437,1395,2033]" pageId="10" pageNumber="389">
|
||
Some shapes of the circumoral spines are unique for
|
||
<taxonomicName id="CED24D021B46FF91AD1E5D48FC37F88D" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[807,920,1827,1853]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B46FF91AD1E5D48FC37F88D" box="[807,920,1827,1853]" italics="true" pageId="10" pageNumber="389">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="91E92A041B46FF91AD915D48FBABF88D" box="[936,1028,1827,1853]" captionStart="FIGURE 8" captionStartId="12.[152,255,936,960]" captionTargetBox="[482,1106,192,868]" captionTargetId="figure-35@12.[472,1114,181,877]" captionTargetPageId="12" captionText="FIGURE 8. Circumoral spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329755" httpUri="https://zenodo.org/record/7329755/files/figure.png" pageId="10" pageNumber="389">Fig. 8A</figureCitation>
|
||
). While the common shape of these spines in the species complex is straight, flattened, and with a blunt tip (e.g., in
|
||
<taxonomicName id="CED24D021B46FF91AA795D2CFB06F8D1" box="[1088,1193,1863,1889]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B46FF91AA795D2CFB06F8D1" box="[1088,1193,1863,1889]" italics="true" pageId="10" pageNumber="389">A. planci</emphasis>
|
||
</taxonomicName>
|
||
[
|
||
<figureCitation id="91E92A041B46FF91AA835D2CFAB9F8D1" box="[1210,1302,1863,1889]" captionStart="FIGURE 8" captionStartId="12.[152,255,936,960]" captionTargetBox="[482,1106,192,868]" captionTargetId="figure-35@12.[472,1114,181,877]" captionTargetPageId="12" captionText="FIGURE 8. Circumoral spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329755" httpUri="https://zenodo.org/record/7329755/files/figure.png" pageId="10" pageNumber="389">Fig. 8C</figureCitation>
|
||
; Supp.
|
||
<figureCitation id="91E92A041B46FF91AB565D2CFF62F835" captionStart="FIGURE 2" captionStartId="5.[152,255,1323,1347]" captionTargetBox="[151,1436,181,1300]" captionTargetId="figure-19@5.[151,1436,181,1300]" captionTargetPageId="5" captionText="FIGURE 2. Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm." figureDoi="http://doi.org/10.5281/zenodo.7329743" httpUri="https://zenodo.org/record/7329743/files/figure.png" pageId="10" pageNumber="389">Fig. S2D</figureCitation>
|
||
] and
|
||
<taxonomicName id="CED24D021B46FF91AF345D00FE14F834" box="[269,443,1899,1925]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B46FF91AF345D00FE14F834" box="[269,443,1899,1925]" italics="true" pageId="10" pageNumber="389">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
[
|
||
<figureCitation id="91E92A041B46FF91AFF25D00FD87F835" box="[459,552,1899,1925]" captionStart="FIGURE 8" captionStartId="12.[152,255,936,960]" captionTargetBox="[482,1106,192,868]" captionTargetId="figure-35@12.[472,1114,181,877]" captionTargetPageId="12" captionText="FIGURE 8. Circumoral spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329755" httpUri="https://zenodo.org/record/7329755/files/figure.png" pageId="10" pageNumber="389">Fig. 8D</figureCitation>
|
||
; Supp.
|
||
<figureCitation id="91E92A041B46FF91AC465D00FD44F835" box="[639,747,1899,1925]" captionStart="FIGURE 3" captionStartId="6.[152,255,1199,1223]" captionTargetBox="[395,1195,188,1170]" captionTargetId="figure-35@6.[374,1212,181,1175]" captionTargetPageId="6" captionText="FIGURE 3. A: Indo-Pacific ‘Acanthaster planci’ species complex, COI ML tree based on the haplotype alignment of Vogler et al. (2008), supplemented with five samples from Israel (see text for details) extracted from full mitochondrial genome sequences from Yuasa et al. (2021) and four sequences of the type series of Acanthaster benziei sp. nov. (GW4xxx, highlighted in bold). The tree was rooted with Acanthaster brevispinus (accession number AB231476), showing the deep divergence among, and little diversity within, species/geographic clades. ML bootstrap values are above branches, bootstrap values of the NJ clustering of haplotypes below branches. B: Geographic distribution of COI-barcoded clades and of type localities of names (Figure 1 from Haszprunar et al. 2017): red—Red Sea (RS) species; blue—Southern Indian Ocean (SIO) species (A. mauritiensis); yellow—Northern Indian Ocean (NIO) species (A. planci); green—Pacific Ocean (PO) species (A. cf. solaris). Location of type localities of nominal Acanthaster species: asterisk—A. planci; cross—A. echinites; triangle—A. solaris, square—A. mauritiensis; circle—A. ellisii pseudoplanci; “?” - the type locality of A. ellisii was not specified: in South American waters of the East Pacific." figureDoi="http://doi.org/10.5281/zenodo.7329745" httpUri="https://zenodo.org/record/7329745/files/figure.png" pageId="10" pageNumber="389">Fig. S3D</figureCitation>
|
||
]), the spines of
|
||
<taxonomicName id="CED24D021B46FF91AD9C5D00FBB9F835" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[933,1046,1899,1925]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B46FF91AD9C5D00FBB9F835" box="[933,1046,1899,1925]" italics="true" pageId="10" pageNumber="389">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="91E92A041B46FF91AA1E5D00FB21F835" box="[1063,1166,1899,1925]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="10" pageNumber="389">Figs. 4D</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B46FF91AAA25D00FB10F835" box="[1179,1215,1899,1925]" captionStart="FIGURE 8" captionStartId="12.[152,255,936,960]" captionTargetBox="[482,1106,192,868]" captionTargetId="figure-35@12.[472,1114,181,877]" captionTargetPageId="12" captionText="FIGURE 8. Circumoral spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329755" httpUri="https://zenodo.org/record/7329755/files/figure.png" pageId="10" pageNumber="389">8A</figureCitation>
|
||
) may have a wider tip or the tip is tapering and more pointed with some granules present on the shaft of the spine (
|
||
<figureCitation id="91E92A041B46FF91AACD5DE4FAE0F819" box="[1268,1359,1935,1961]" captionStart="FIGURE 8" captionStartId="12.[152,255,936,960]" captionTargetBox="[482,1106,192,868]" captionTargetId="figure-35@12.[472,1114,181,877]" captionTargetPageId="12" captionText="FIGURE 8. Circumoral spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329755" httpUri="https://zenodo.org/record/7329755/files/figure.png" pageId="10" pageNumber="389">Fig. 8B</figureCitation>
|
||
). Both shapes are only found in this spine
|
||
<typeStatus id="D66988231B46FF91AC105DD8FDF5F87D" box="[553,602,1971,1997]" pageId="10" pageNumber="389">type</typeStatus>
|
||
of
|
||
<taxonomicName id="CED24D021B46FF91ACB85DD8FD5EF87D" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[641,753,1971,1997]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="10" pageNumber="389" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B46FF91ACB85DD8FD5EF87D" box="[641,753,1971,1997]" italics="true" pageId="10" pageNumber="389">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
. The other oral spines are very similar to the corresponding spines of specimens examined from the other three species (Supp.
|
||
<figureCitation id="91E92A041B46FF91ADAA5DBCFBBDF841" box="[915,1042,2007,2033]" captionStart-0="FIGURE 2" captionStart-1="FIGURE 3" captionStart-2="FIGURE 4" captionStartId-0="5.[152,255,1323,1347]" captionStartId-1="6.[152,255,1199,1223]" captionStartId-2="8.[152,255,1803,1827]" captionTargetBox-0="[151,1436,181,1300]" captionTargetBox-1="[395,1195,188,1170]" captionTargetBox-2="[151,1436,1065,1779]" captionTargetId-0="figure-19@5.[151,1436,181,1300]" captionTargetId-1="figure-35@6.[374,1212,181,1175]" captionTargetId-2="figure-448@8.[151,1436,1065,1779]" captionTargetPageId-0="5" captionTargetPageId-1="6" captionTargetPageId-2="8" captionText-0="FIGURE 2. Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm." captionText-1="FIGURE 3. A: Indo-Pacific ‘Acanthaster planci’ species complex, COI ML tree based on the haplotype alignment of Vogler et al. (2008), supplemented with five samples from Israel (see text for details) extracted from full mitochondrial genome sequences from Yuasa et al. (2021) and four sequences of the type series of Acanthaster benziei sp. nov. (GW4xxx, highlighted in bold). The tree was rooted with Acanthaster brevispinus (accession number AB231476), showing the deep divergence among, and little diversity within, species/geographic clades. ML bootstrap values are above branches, bootstrap values of the NJ clustering of haplotypes below branches. B: Geographic distribution of COI-barcoded clades and of type localities of names (Figure 1 from Haszprunar et al. 2017): red—Red Sea (RS) species; blue—Southern Indian Ocean (SIO) species (A. mauritiensis); yellow—Northern Indian Ocean (NIO) species (A. planci); green—Pacific Ocean (PO) species (A. cf. solaris). Location of type localities of nominal Acanthaster species: asterisk—A. planci; cross—A. echinites; triangle—A. solaris, square—A. mauritiensis; circle—A. ellisii pseudoplanci; “?” - the type locality of A. ellisii was not specified: in South American waters of the East Pacific." captionText-2="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi-0="http://doi.org/10.5281/zenodo.7329743" figureDoi-1="http://doi.org/10.5281/zenodo.7329745" figureDoi-2="http://doi.org/10.5281/zenodo.7329747" httpUri-0="https://zenodo.org/record/7329743/files/figure.png" httpUri-1="https://zenodo.org/record/7329745/files/figure.png" httpUri-2="https://zenodo.org/record/7329747/files/figure.png" pageId="10" pageNumber="389">Figs. S2–4</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="5DAD66091B47FF90AEA15FE1FD22FA75" ID-DOI="http://doi.org/10.5281/zenodo.7329753" ID-Zenodo-Dep="7329753" httpUri="https://zenodo.org/record/7329753/files/figure.png" pageId="11" pageNumber="390" startId="11.[152,255,1418,1442]" targetBox="[479,1108,204,1389]" targetPageId="11">
|
||
<paragraph id="096D36811B47FF90AEA15FE1FD22FA75" blockId="11.[151,1437,1418,1478]" pageId="11" pageNumber="390">
|
||
<emphasis id="3BA6EA931B47FF90AEA15FE1FEB9FA12" bold="true" box="[152,278,1418,1442]" pageId="11" pageNumber="390">FIGURE 7</emphasis>
|
||
. Typical latero-oral spines, showing species-specific variation among (A)
|
||
<taxonomicName id="CED24D021B47FF90AA1C5FE1FB5EFA12" authority="Wörheide & Kaltenbacher & Cowan & Haszprunar, 2022" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[1061,1265,1418,1442]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="benziei" status="sp. nov.">
|
||
<emphasis id="3BA6EA931B47FF90AA1C5FE1FB5EFA12" box="[1061,1265,1418,1442]" italics="true" pageId="11" pageNumber="390">Acanthaster benziei</emphasis>
|
||
</taxonomicName>
|
||
<taxonomicNameLabel id="209557E81B47FF90AAC35FE1FAE1FA11" box="[1274,1358,1418,1441]" pageId="11" pageNumber="390" rank="species">sp. nov.</taxonomicNameLabel>
|
||
, (B)
|
||
<taxonomicName id="CED24D021B47FF90ABBF5FE1FF79FA76" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B47FF90ABBF5FE1FF79FA76" italics="true" pageId="11" pageNumber="390">A. planci</emphasis>
|
||
</taxonomicName>
|
||
, (C)
|
||
<taxonomicName id="CED24D021B47FF90AF325FC5FE0BFA76" box="[267,420,1454,1478]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B47FF90AF325FC5FE0BFA76" box="[267,420,1454,1478]" italics="true" pageId="11" pageNumber="390">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
, and (D)
|
||
<taxonomicName id="CED24D021B47FF90AC3F5FC5FD28FA76" box="[518,647,1454,1478]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" isUncertain="true" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="solaris">
|
||
<emphasis id="3BA6EA931B47FF90AC3F5FC5FDBAFA75" box="[518,533,1454,1477]" italics="true" pageId="11" pageNumber="390">A</emphasis>
|
||
. cf.
|
||
<emphasis id="3BA6EA931B47FF90AC7B5FC5FD28FA76" box="[578,647,1454,1478]" italics="true" pageId="11" pageNumber="390">solaris</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="096D36811B47FF90AEFE5C73FC8AF956" blockId="11.[151,1437,1560,1910]" pageId="11" pageNumber="390">
|
||
Aboral pedicellariae are more numerous in
|
||
<taxonomicName id="CED24D021B47FF90AC805C72FC86F982" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[697,809,1560,1587]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B47FF90AC805C72FC86F982" box="[697,809,1560,1587]" italics="true" pageId="11" pageNumber="390">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
than in the two Indian Ocean species
|
||
<taxonomicName id="CED24D021B47FF90AADA5C72FAE5F982" box="[1251,1354,1560,1587]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B47FF90AADA5C72FAE5F982" box="[1251,1354,1560,1587]" italics="true" pageId="11" pageNumber="390">A. planci</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="CED24D021B47FF90ABBD5C72FE8EF9E6" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B47FF90ABBD5C72FE8EF9E6" italics="true" pageId="11" pageNumber="390">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
. The oral pedicellariae of
|
||
<taxonomicName id="CED24D021B47FF90AC7B5C56FD09F9E6" box="[578,678,1596,1623]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="planci">
|
||
<emphasis id="3BA6EA931B47FF90AC7B5C56FD09F9E6" box="[578,678,1596,1623]" italics="true" pageId="11" pageNumber="390">A. planci</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="91E92A041B47FF90AC8D5C57FCA5F9E6" box="[692,778,1596,1622]" captionStart="FIGURE 9" captionStartId="12.[152,255,1709,1733]" captionTargetBox="[477,1110,1096,1681]" captionTargetId="figure-72@12.[472,1114,1093,1686]" captionTargetPageId="12" captionText="FIGURE 9. Oral pedicellariae, showing species-specific variation among (A) Acanthaster benziei sp. nov., (B) A. planci, (C) A. mauritiensis, and (D) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329757" httpUri="https://zenodo.org/record/7329757/files/figure.png" pageId="11" pageNumber="390">Fig. 9B</figureCitation>
|
||
; Supp.
|
||
<figureCitation id="91E92A041B47FF90AD625C57FC6FF9E6" box="[859,960,1596,1622]" captionStart="FIGURE 2" captionStartId="5.[152,255,1323,1347]" captionTargetBox="[151,1436,181,1300]" captionTargetId="figure-19@5.[151,1436,181,1300]" captionTargetPageId="5" captionText="FIGURE 2. Ethanol-preserved specimens of the type series. GW4202 (A) is the holotype, all the others (B–D) are paratypes. Note that individual GW4266 is a juvenile specimen. Size of labels 6 × 2 cm." figureDoi="http://doi.org/10.5281/zenodo.7329743" httpUri="https://zenodo.org/record/7329743/files/figure.png" pageId="11" pageNumber="390">Fig. S2H</figureCitation>
|
||
) are mostly wider and straighter than those of
|
||
<taxonomicName id="CED24D021B47FF90AE8E5C0AFE88F9CA" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[183,295,1632,1659]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B47FF90AE8E5C0AFE88F9CA" box="[183,295,1632,1659]" italics="true" pageId="11" pageNumber="390">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="91E92A041B47FF90AF005C0BFE30F9CA" box="[313,415,1632,1658]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="11" pageNumber="390">Figs. 4H</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B47FF90AF955C0BFE7EF9CA" box="[428,465,1632,1658]" captionStart="FIGURE 9" captionStartId="12.[152,255,1709,1733]" captionTargetBox="[477,1110,1096,1681]" captionTargetId="figure-72@12.[472,1114,1093,1686]" captionTargetPageId="12" captionText="FIGURE 9. Oral pedicellariae, showing species-specific variation among (A) Acanthaster benziei sp. nov., (B) A. planci, (C) A. mauritiensis, and (D) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329757" httpUri="https://zenodo.org/record/7329757/files/figure.png" pageId="11" pageNumber="390">9A</figureCitation>
|
||
), which are the smallest among the four species, and are not as deeply curved as the pedicellariae of
|
||
<taxonomicName id="CED24D021B47FF90AF725CEEFE5BF92E" box="[331,500,1669,1695]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="mauritiensis">
|
||
<emphasis id="3BA6EA931B47FF90AF725CEEFE5BF92E" box="[331,500,1669,1695]" italics="true" pageId="11" pageNumber="390">A. mauritiensis</emphasis>
|
||
</taxonomicName>
|
||
(
|
||
<figureCitation id="91E92A041B47FF90AC3A5CEFFDF4F92E" box="[515,603,1668,1694]" captionStart="FIGURE 9" captionStartId="12.[152,255,1709,1733]" captionTargetBox="[477,1110,1096,1681]" captionTargetId="figure-72@12.[472,1114,1093,1686]" captionTargetPageId="12" captionText="FIGURE 9. Oral pedicellariae, showing species-specific variation among (A) Acanthaster benziei sp. nov., (B) A. planci, (C) A. mauritiensis, and (D) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329757" httpUri="https://zenodo.org/record/7329757/files/figure.png" pageId="11" pageNumber="390">Fig. 9C</figureCitation>
|
||
; Supp.
|
||
<figureCitation id="91E92A041B47FF90AC975CEFFCB9F92E" box="[686,790,1668,1694]" captionStart="FIGURE 3" captionStartId="6.[152,255,1199,1223]" captionTargetBox="[395,1195,188,1170]" captionTargetId="figure-35@6.[374,1212,181,1175]" captionTargetPageId="6" captionText="FIGURE 3. A: Indo-Pacific ‘Acanthaster planci’ species complex, COI ML tree based on the haplotype alignment of Vogler et al. (2008), supplemented with five samples from Israel (see text for details) extracted from full mitochondrial genome sequences from Yuasa et al. (2021) and four sequences of the type series of Acanthaster benziei sp. nov. (GW4xxx, highlighted in bold). The tree was rooted with Acanthaster brevispinus (accession number AB231476), showing the deep divergence among, and little diversity within, species/geographic clades. ML bootstrap values are above branches, bootstrap values of the NJ clustering of haplotypes below branches. B: Geographic distribution of COI-barcoded clades and of type localities of names (Figure 1 from Haszprunar et al. 2017): red—Red Sea (RS) species; blue—Southern Indian Ocean (SIO) species (A. mauritiensis); yellow—Northern Indian Ocean (NIO) species (A. planci); green—Pacific Ocean (PO) species (A. cf. solaris). Location of type localities of nominal Acanthaster species: asterisk—A. planci; cross—A. echinites; triangle—A. solaris, square—A. mauritiensis; circle—A. ellisii pseudoplanci; “?” - the type locality of A. ellisii was not specified: in South American waters of the East Pacific." figureDoi="http://doi.org/10.5281/zenodo.7329745" httpUri="https://zenodo.org/record/7329745/files/figure.png" pageId="11" pageNumber="390">Fig. S3H</figureCitation>
|
||
). The valves of the flat oral pedicellariae, which resemble the shape of a rhombus, with a widening at the middle, a pointed tip and teeth on the inner surface, were additionally only found in specimens from the Red Sea (
|
||
<figureCitation id="91E92A041B47FF90ACBA5CA7FD48F956" box="[643,743,1740,1766]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="11" pageNumber="390">Figs. 4H</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B47FF90ACCA5CA7FCB7F956" box="[755,792,1740,1766]" captionStart="FIGURE 9" captionStartId="12.[152,255,1709,1733]" captionTargetBox="[477,1110,1096,1681]" captionTargetId="figure-72@12.[472,1114,1093,1686]" captionTargetPageId="12" captionText="FIGURE 9. Oral pedicellariae, showing species-specific variation among (A) Acanthaster benziei sp. nov., (B) A. planci, (C) A. mauritiensis, and (D) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329757" httpUri="https://zenodo.org/record/7329757/files/figure.png" pageId="11" pageNumber="390">9A</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="096D36811B47FF90AEFE5C9AFE68F8C6" blockId="11.[151,1437,1560,1910]" pageId="11" pageNumber="390">
|
||
In summary, the main distinguishing morphological characters of
|
||
<taxonomicName id="CED24D021B47FF90AD8B5C9AFB8EF8BA" authorityName="Wörheide & Kaltenbacher & Cowan & Haszprunar" authorityYear="2022" box="[946,1057,1776,1803]" class="Asteroidea" family="Acanthasteridae" genus="Acanthaster" kingdom="Animalia" order="Valvatida" pageId="11" pageNumber="390" phylum="Echinodermata" rank="species" species="benziei">
|
||
<emphasis id="3BA6EA931B47FF90AD8B5C9AFB8EF8BA" box="[946,1057,1776,1803]" italics="true" pageId="11" pageNumber="390">A. benziei</emphasis>
|
||
</taxonomicName>
|
||
considered to be species-specific are the fanned spine tips in primary (
|
||
<figureCitation id="91E92A041B47FF90AC0C5D7FFD35F89E" box="[565,666,1812,1838]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="11" pageNumber="390">Figs. 4A</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B47FF90AC9C5D7FFD1BF89E" box="[677,692,1812,1838]" captionStart="FIGURE 6" captionStartId="10.[200,303,1266,1290]" captionTargetBox="[479,1103,191,1235]" captionTargetId="figure-35@10.[472,1115,181,1242]" captionTargetPageId="10" captionText="FIGURE 6. Typical primary spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329751" httpUri="https://zenodo.org/record/7329751/files/figure.png" pageId="11" pageNumber="390">6</figureCitation>
|
||
) and latero-oral spines (
|
||
<figureCitation id="91E92A041B47FF90ADFC5D7FFB87F89E" box="[965,1064,1812,1838]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="11" pageNumber="390">Figs. 4C</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B47FF90AA0D5D7FFBEBF89E" box="[1076,1092,1812,1838]" captionStart="FIGURE 7" captionStartId="11.[152,255,1418,1442]" captionTargetBox="[479,1108,204,1389]" captionTargetId="figure-19@11.[472,1115,199,1394]" captionTargetPageId="11" captionText="FIGURE 7. Typical latero-oral spines, showing species-specific variation among (A) Acanthaster benziei sp. nov., (B) A. planci, (C) A. mauritiensis, and (D) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329753" httpUri="https://zenodo.org/record/7329753/files/figure.png" pageId="11" pageNumber="390">7</figureCitation>
|
||
), the wider tip or the taperingpointed shape in circumoral spines (
|
||
<figureCitation id="91E92A041B47FF90AC1E5D53FD25F8E2" box="[551,650,1848,1874]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="11" pageNumber="390">Figs. 4D</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B47FF90ACAC5D53FD0BF8E2" box="[661,676,1848,1874]" captionStart="FIGURE 8" captionStartId="12.[152,255,936,960]" captionTargetBox="[482,1106,192,868]" captionTargetId="figure-35@12.[472,1114,181,877]" captionTargetPageId="12" captionText="FIGURE 8. Circumoral spines, showing species-specific variation among (A, B) Acanthaster benziei sp. nov., (C) A. planci, (D) A. mauritiensis, and (E) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329755" httpUri="https://zenodo.org/record/7329755/files/figure.png" pageId="11" pageNumber="390">8</figureCitation>
|
||
), as well as the rhombus-shaped oral pedicellariae with occasionally internal teeth (
|
||
<figureCitation id="91E92A041B47FF90AF005D37FE31F8C6" box="[313,414,1884,1910]" captionStart="FIGURE 4" captionStartId="8.[152,255,1803,1827]" captionTargetBox="[151,1436,1065,1779]" captionTargetId="figure-448@8.[151,1436,1065,1779]" captionTargetPageId="8" captionText="FIGURE 4. Acanthaster benziei sp. nov. (holotype GW4202) (A-B) Aboral spines, (C-F) oral spines and (G-H) pedicellariae of four adult specimens of Acanthaster benziei sp. nov.: (A) Primary spines, (B) Secondary spines, (C) Latero-oral spines, (D) Circumoral spines, (E) Oral spines, (F) Subambulacral spines, (G) Aboral pedicellariae, (H) Oral pedicellariae." figureDoi="http://doi.org/10.5281/zenodo.7329747" httpUri="https://zenodo.org/record/7329747/files/figure.png" pageId="11" pageNumber="390">Figs. 4H</figureCitation>
|
||
,
|
||
<figureCitation id="91E92A041B47FF90AF905D37FE16F8C6" box="[425,441,1884,1910]" captionStart="FIGURE 9" captionStartId="12.[152,255,1709,1733]" captionTargetBox="[477,1110,1096,1681]" captionTargetId="figure-72@12.[472,1114,1093,1686]" captionTargetPageId="12" captionText="FIGURE 9. Oral pedicellariae, showing species-specific variation among (A) Acanthaster benziei sp. nov., (B) A. planci, (C) A. mauritiensis, and (D) A. cf. solaris." figureDoi="http://doi.org/10.5281/zenodo.7329757" httpUri="https://zenodo.org/record/7329757/files/figure.png" pageId="11" pageNumber="390">9</figureCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |