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<mods:title id="944C8A5573DD3C5BC94B10EBE7AB52C2">Toward a resolution of the cosmopolitan Botryllus schlosseri species complex (Ascidiacea, Styelidae): mitogenomics and morphology of clade E (Botryllus gaiae)</mods:title>
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<mods:namePart id="6BEE001E1E9C90EC58D624CFB3C39984">Brunetti, Riccardo</mods:namePart>
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<mods:namePart id="BEC446942BB4720F9D483DD58A84B525">Griggio, Francesca</mods:namePart>
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<mods:namePart id="354035D7ED6D1A0582EDE9000B491206">Mastrototaro, Francesco</mods:namePart>
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<mods:namePart id="52292C25F53665D4CF1083D8394E6B5B">Gasparini, Fabio</mods:namePart>
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<mods:namePart id="E69E72190FBA88528AA362D0DF6E5D64">Gissi, Carmela</mods:namePart>
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<mods:title id="5326E7F15600AE3E09E9D0A658D6F896">Zoological Journal of the Linnean Society</mods:title>
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<mods:date id="7CA13B86B9C01D5CD3BA13F4867B590F">2020</mods:date>
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<mods:detail id="D8B21BC3AA214E456D0E104BA9394A04" type="volume">
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<mods:number id="15B77BCCD381134D112066C33EC8D5D5">190</mods:number>
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<mods:start id="57D14DB262651F7E0008B445AACA6B6F">1175</mods:start>
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<mods:classification id="24C3E8EE4142083C962A3EA27B4C2BA0">journal article</mods:classification>
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<mods:identifier id="5CB0F16CE56DE528F73351AD2C1BD174" type="ISSN">0024-4082</mods:identifier>
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<mods:identifier id="A7E246F10499BF1042AD8B5FB5F54F1E" type="ZooBank">B18D5A7-0E96-458E-B3C0-5E93327359AC</mods:identifier>
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<treatment id="5947A847E948FFE83CF3C7F6F902FF73" LSID="urn:lsid:plazi:treatment:5947A847E948FFE83CF3C7F6F902FF73" httpUri="http://treatment.plazi.org/id/5947A847E948FFE83CF3C7F6F902FF73" lastPageId="11" lastPageNumber="1184" pageId="7" pageNumber="1182">
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<paragraph id="D1511951E948FFE43CF3C7F6FD69FB8C" blockId="7.[907,1362,1006,1030]" box="[907,1362,1006,1030]" pageId="7" pageNumber="1182">
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<heading id="8A19AE3DE948FFE43CF3C7F6FD69FB8C" box="[907,1362,1006,1030]" centered="true" fontSize="9" level="2" pageId="7" pageNumber="1182" reason="2">
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<taxonomicName id="16EE62D2E948FFE43CF3C7F6FCD3FB8F" ID-CoL="8473B" authority="BRUNETTI" authorityName="Brunetti & Griggio & Mastrototaro & Gasparini & Gissi" authorityYear="2020" box="[907,1256,1006,1030]" class="Ascidiacea" family="Styelidae" genus="Botryllus" kingdom="Animalia" order="Stolidobranchia" pageId="7" pageNumber="1182" phylum="Chordata" rank="species" species="gaiae" status="sp. nov.">
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<emphasis id="E39AC543E948FFE43CF3C7F6FC5DFB8F" bold="true" box="[907,1126,1006,1029]" italics="true" pageId="7" pageNumber="1182">
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<smallCapsWord id="D7B78F8DE948FFE43B67C7EAFC5DFB8F" baselines="1024" box="[1055,1126,1010,1029]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="gaiae" pageId="7" pageNumber="1182">GAIAE</smallCapsWord>
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</emphasis>
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</taxonomicName>
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<taxonomicNameLabel id="F8A97838E948FFE43B97C7E9FD69FB8C" box="[1263,1362,1006,1030]" pageId="7" pageNumber="1182" rank="species">
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.
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<smallCapsWord id="D7B78F8DE948FFE43A62C7E9FD70FB8E" baselines="1023" box="[1306,1355,1009,1028]" lowerCaseFontSize="8" mainFontSize="10" normCase="lower" normString="nov" pageId="7" pageNumber="1182">NOV</smallCapsWord>
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.
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||||
</taxonomicNameLabel>
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</emphasis>
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</heading>
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||||
</paragraph>
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<paragraph id="D1511951E948FFE43C43C00EFC4DFBC1" blockId="7.[827,1444,1046,1160]" pageId="7" pageNumber="1182">
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LSID:
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<uri id="A57F1553E948FFE43CFEC00EFC4DFBC1" pageId="7" pageNumber="1182">
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urn:lsid:zoobank.org:pub:
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<uuid id="A5482384E948FFE43BC6C00EFC4DFBC1" pageId="7" pageNumber="1182">DB18D5A7-0E96- 458E-B3C0-5E93327359AC</uuid>
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<subSubSection id="99F44ADAE948FFE83C43C04DF902FF73" lastPageId="11" lastPageNumber="1186" pageId="7" pageNumber="1182" type="description">
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<paragraph id="D1511951E948FFE43C43C04DFD9FFB02" blockId="7.[827,1444,1046,1160]" pageId="7" pageNumber="1182">
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<emphasis id="E39AC543E948FFE43C43C04DFBF7FBE0" box="[827,972,1108,1130]" italics="true" pageId="7" pageNumber="1182">E t y m o l o g y:</emphasis>
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||||
Fr o m G a i a, t h e n a m e o f t h e f i r s t granddaughter of the author of this species, R. Brunetti.
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</paragraph>
|
||||
<paragraph id="D1511951E948FFE43C43C0A9FD9FFAF5" blockId="7.[827,1444,1201,1407]" pageId="7" pageNumber="1182">
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<emphasis id="E39AC543E948FFE43C43C0A9FC19FB4C" box="[827,1058,1201,1222]" italics="true" pageId="7" pageNumber="1182">Material examined:</emphasis>
|
||||
<materialsCitation id="6186130CE948FFE43B57C0A9FC8EFB6C" collectionCode="MSNVE" pageId="7" pageNumber="1182" specimenCode="MSNVE 25086" specimenCount="1" typeStatus="syntype">
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<typeStatus id="0E55A7F3E948FFE43B57C0A9FCA5FB4C" box="[1071,1182,1201,1222]" pageId="7" pageNumber="1182" type="syntype">Syntypes</typeStatus>
|
||||
:
|
||||
<specimenCode id="8148B12AE948FFE43BD3C0A9FD65FB4D" box="[1195,1374,1201,1223]" collectionCode="MSNVE" pageId="7" pageNumber="1182">MSNVE 25086</specimenCode>
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, BT2, colony fragment in 10% formalin
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</materialsCitation>
|
||||
;
|
||||
<materialsCitation id="6186130CE948FFE43BB9C0C8FCECFA8E" collectionCode="MSNVE" pageId="7" pageNumber="1182" specimenCode="MSNVE 2508" specimenCount="1" typeStatus="syntype">
|
||||
<specimenCode id="8148B12AE948FFE43BB9C0C8FD5AFB6C" box="[1217,1377,1232,1254]" collectionCode="MSNVE" pageId="7" pageNumber="1182">MSNVE 2508</specimenCode>
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, BT4, colony fragment in 10% formalin
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</materialsCitation>
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||||
;
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||||
<materialsCitation id="6186130CE948FFE43B9EC0F7FC39FAC8" collectionCode="MSNVE" pageId="7" pageNumber="1182" specimenCode="MSNVE 25088" specimenCount="1" typeStatus="syntype">
|
||||
<specimenCode id="8148B12AE948FFE43B9EC0F7FDA6FA8E" box="[1254,1437,1262,1284]" collectionCode="MSNVE" pageId="7" pageNumber="1182">MSNVE 25088</specimenCode>
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, BT2A, another fragment of the BT2 colony preserved in 99% ethanol
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</materialsCitation>
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;
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<materialsCitation id="6186130CE948FFE43B6CC134FD9BFAF5" collectionCode="MSNVE" collectorName="All & F. Mastrotorato" pageId="7" pageNumber="1182" specimenCode="MSNVE 25089" specimenCount="1" typeStatus="syntype">
|
||||
<specimenCode id="8148B12AE948FFE43B6CC134FCEFFAC8" box="[1044,1236,1324,1346]" collectionCode="MSNVE" pageId="7" pageNumber="1182">MSNVE 25089</specimenCode>
|
||||
, BT4A, another fragment of the BT4 colony preserved in 99% ethanol.
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||||
<collectorName id="7C1B7C87E948FFE43C43C171FB67FAF5" box="[827,860,1385,1407]" pageId="7" pageNumber="1182">All</collectorName>
|
||||
collected by
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||||
<collectorName id="7C1B7C87E948FFE43C96C172FCA5FAF5" box="[1006,1182,1386,1407]" pageId="7" pageNumber="1182">F. Mastrotorato</collectorName>
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||||
during clam sampling
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||||
</materialsCitation>
|
||||
.
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||||
</paragraph>
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||||
<paragraph id="D1511951E948FFE43C43C1B0FC9FFA71" blockId="7.[827,1443,1448,1531]" pageId="7" pageNumber="1182">
|
||||
<materialsCitation id="6186130CE948FFE43C43C1B0FC9BFA71" country="Italy" latitude="41.382122" location="Barletta" longLatPrecision="1" longitude="16.168495" pageId="7" pageNumber="1182" specimenCount="1" typeStatus="holotype">
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||||
<emphasis id="E39AC543E948FFE43C43C1B0FBDDFA37" box="[827,998,1448,1469]" italics="true" pageId="7" pageNumber="1182">
|
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<typeStatus id="0E55A7F3E948FFE43C43C1B0FB4EFA37" box="[827,885,1448,1469]" pageId="7" pageNumber="1182">Type</typeStatus>
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locality:
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||||
</emphasis>
|
||||
<location id="D4314F8AE948FFE43B79C1B0FC50FA34" LSID="urn:lsid:plazi:treatment:5947A847E948FFE83CF3C7F6F902FF73:D4314F8AE948FFE43B79C1B0FC50FA34" box="[1025,1131,1448,1470]" country="Italy" latitude="41.382122" longLatPrecision="1" longitude="16.168495" name="Barletta" pageId="7" pageNumber="1182">Barletta</location>
|
||||
(
|
||||
<collectingCountry id="A9F959C1E948FFE43BF8C1B0FCFBFA34" box="[1152,1216,1448,1470]" name="Italy" pageId="7" pageNumber="1182">Italy</collectingCountry>
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),
|
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<geoCoordinate id="B4DA7F96E948FFE43BA2C1B0FDA6FA34" box="[1242,1437,1448,1470]" degrees="41" direction="north" minutes="22" orientation="latitude" pageId="7" pageNumber="1182" precision="1" seconds="55.646" value="41.382122">41°22’55.646”N</geoCoordinate>
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||||
,
|
||||
<geoCoordinate id="B4DA7F96E948FFE43C43C1DEFBD2FA56" box="[827,1001,1478,1500]" degrees="16" direction="east" minutes="10" orientation="longitude" pageId="7" pageNumber="1182" precision="1" seconds="06.586" value="16.168495">16°10’06.586”E</geoCoordinate>
|
||||
, 2.5–4.0 m depth
|
||||
<collectedFrom id="62F25AACE948FFE43BD5C1DFFC01FA70" pageId="7" pageNumber="1182">
|
||||
on
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||||
<taxonomicName id="16EE62D2E948FFE43BB5C1DEFC01FA70" authority="(Ucria) Asch. Seagrass" authorityName="Asch. Seagrass" baseAuthorityName="Ucria" class="Malacostraca" family="Sphaeromatidae" genus="Cymodocea" kingdom="Animalia" order="Isopoda" pageId="7" pageNumber="1182" phylum="Arthropoda" rank="species" species="nodosa">
|
||||
<emphasis id="E39AC543E948FFE43BB5C1DEFD98FA51" box="[1229,1443,1478,1499]" italics="true" pageId="7" pageNumber="1182">Cymodocea nodosa</emphasis>
|
||||
(Ucria) Asch. Seagrass
|
||||
</taxonomicName>
|
||||
</collectedFrom>
|
||||
meadow
|
||||
</materialsCitation>
|
||||
.
|
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</paragraph>
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<paragraph id="D1511951E948FFEB3C43C23CFA4DF9B7" blockId="7.[827,1444,1572,1900]" lastBlockId="8.[145,761,1452,1903]" lastPageId="8" lastPageNumber="1183" pageId="7" pageNumber="1182">
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||||
<emphasis id="E39AC543E948FFE43C43C23CFBFCF9B3" box="[827,967,1572,1593]" italics="true" pageId="7" pageNumber="1182">Description:</emphasis>
|
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In both colonies from Barletta, labelled BT2 and BT4, the test surface is hard but, once it is torn, zooids are easily pulled out. Sample BT4 consists of two pieces of colony, each of them with 2–3 systems at the beginning of the generation change (developmental stage 9–11/8–9/6–7/1) (
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<bibRefCitation id="B57F64A0E948FFE43A79C2A5FDAFF958" author="Berrill NJ" box="[1281,1428,1725,1747]" pageId="7" pageNumber="1182" pagination="169 - 184" refId="ref11450" refString="Berrill NJ. 1941. The development of the bud in Botryllus. Biological Bulletin 80: 169-184." type="journal article" year="1941">Berrill, 1941</bibRefCitation>
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). Therefore, the older generation is at the beginning of regression (stage 9–11) and the new generation presents syphons still closed and all internal organs completely developed (stage 8–9). The regressing zooids are clearly arranged in
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<taxonomicName id="16EE62D2E948FFE43BF6C34EFD52F8E1" box="[1166,1385,1878,1899]" class="Ascidiacea" family="Styelidae" genus="Botryllus" kingdom="Animalia" order="Stolidobranchia" pageId="7" pageNumber="1182" phylum="Chordata" rank="species" species="schlosseri">
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<emphasis id="E39AC543E948FFE43BF6C34EFD52F8E1" box="[1166,1385,1878,1899]" italics="true" pageId="7" pageNumber="1182">Botryllus schlosseri</emphasis>
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</taxonomicName>
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-
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<typeStatus id="0E55A7F3E948FFE43A09C34FFD98F8E6" box="[1393,1443,1879,1900]" pageId="7" pageNumber="1182">type</typeStatus>
|
||||
systems. Sample BT2 contains a piece of colony of about
|
||||
<quantity id="1616B4B4E947FFEB3FA5C1D3F92DFA6A" box="[221,278,1483,1504]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="1183" unit="cm" value="1.0">1 cm</quantity>
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||||
<superScript id="269BB419E947FFEB3E6EC1D2F925FA5C" attach="left" box="[278,286,1482,1494]" fontSize="5" pageId="8" pageNumber="1183">2</superScript>
|
||||
of surface with two systems consisting of 15 zooids. As shown in
|
||||
<figureCitation id="49D505D4E947FFEB3EB0C1F1FA16FA75" box="[456,557,1513,1535]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="8" pageNumber="1183">Figure 5</figureCitation>
|
||||
A-B, in life these two colonies are green (BT4) and green-yellow (BT2), whereas in preservative there are whitish.
|
||||
</paragraph>
|
||||
<caption id="859149D9E947FFEB3FE9C10AFA6FFA0A" pageId="8" pageNumber="1183" startId="8.[145,223,1298,1320]" targetBox="[318,1251,195,1258]" targetPageId="8" targetType="figure">
|
||||
<paragraph id="D1511951E947FFEB3FE9C10AFA6FFA0A" blockId="8.[145,1425,1298,1408]" pageId="8" pageNumber="1183">
|
||||
<emphasis id="E39AC543E947FFEB3FE9C10AF8C3FAAD" bold="true" box="[145,248,1298,1320]" pageId="8" pageNumber="1183">Figure 3.</emphasis>
|
||||
Non-synonymous substitution rates (dN) of intraspecies and congeneric ascidian pairs. Red: intra and interclade comparisons between specimens of the
|
||||
<taxonomicName id="16EE62D2E947FFEB3D39C137FB2DFACF" box="[577,790,1327,1349]" class="Ascidiacea" family="Styelidae" genus="Botryllus" kingdom="Animalia" order="Stolidobranchia" pageId="8" pageNumber="1183" phylum="Chordata" rank="species" species="schlosseri">
|
||||
<emphasis id="E39AC543E947FFEB3D39C137FB2DFACF" box="[577,790,1327,1349]" italics="true" pageId="8" pageNumber="1183">Botryllus schlosseri</emphasis>
|
||||
</taxonomicName>
|
||||
species complex; blue: comparisons between ex-cryptic or morphologically hard-to-discriminate species; black: comparisons between all other analysed ascidians. The AC numbers of the analysed mtDNAs are listed in Table 3.
|
||||
</paragraph>
|
||||
</caption>
|
||||
<paragraph id="D1511951E947FFEA3FD1C25DFDA3FAEC" blockId="8.[145,761,1452,1903]" lastBlockId="9.[827,1444,1145,1382]" lastPageId="9" lastPageNumber="1184" pageId="8" pageNumber="1183">
|
||||
The following description is chiefly based on the observation of the first order buds of sample BT2, which have virtually finalized their development (only the syphons have to be completed) (
|
||||
<figureCitation id="49D505D4E947FFEB3D79C2B9FA48F93D" box="[513,627,1697,1719]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="8" pageNumber="1183">Fig. 5C–E</figureCitation>
|
||||
). The adult regressing zooids are about
|
||||
<quantity id="1616B4B4E947FFEB3EB3C2D8FA1BF95C" box="[459,544,1728,1750]" metricMagnitude="-3" metricUnit="m" metricValue="1.5" pageId="8" pageNumber="1183" unit="mm" value="1.5">1.5 mm</quantity>
|
||||
high and the same height is recorded in first order buds. Therefore, it is justifiable to deduce a height up to
|
||||
<quantity id="1616B4B4E947FFEB3D76C2E6FA76F899" box="[526,589,1790,1811]" metricMagnitude="-3" metricUnit="m" metricValue="2.0" pageId="8" pageNumber="1183" unit="mm" value="2.0">2 mm</quantity>
|
||||
for the filtering zooid, because an increase in size is observed when the bud opens the syphons. The body wall presents fine muscle fibres running from the anterior-dorsal side to the ventral one. In the buds there are eight tentacles: four large and four intercalated short. The atrial syphons, still visible in some regressing zooids, is conic with a dorsal tongue. The branchial sac has nine rows of stigmata on the right and eight on the left side; the second row is dorsally incomplete (
|
||||
<figureCitation id="49D505D4E947FFEB3BDEC25DFD2FF9D1" box="[1190,1300,1605,1627]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="8" pageNumber="1183">Fig. 5D–E</figureCitation>
|
||||
). The zooid has a cylindrical shape having about 20 stigmata in the first half row and about
|
||||
<quantity id="1616B4B4E947FFEB3B10C29BFC9AF913" box="[1128,1185,1667,1689]" metricMagnitude="-1" metricUnit="m" metricValue="3.81" pageId="8" pageNumber="1183" unit="in" value="15.0">15 in</quantity>
|
||||
the seventh one. The branchial formula at the level of fourth row of stigmata is usually E 5.5.5.5 DL. Along the transversal branchial vessels, no muscle fibres are detected. A simple edged dorsal lamina rises from the dorsal vessel at level of fourth to fifth row of stigmata and reaches a height of three times the diameter of the dorsal vessel (
|
||||
<figureCitation id="49D505D4E947FFEB3A63C323FDB3F8DB" box="[1307,1416,1851,1873]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="8" pageNumber="1183">Fig. 5D–E</figureCitation>
|
||||
) at the oesophageal opening which is at level of the last row of stigmata. The intestinal loop is horizontally arranged (
|
||||
<figureCitation id="49D505D4E946FFEA3E5DC080F944FB24" box="[293,383,1176,1198]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="9" pageNumber="1184">Fig. 5D</figureCitation>
|
||||
). The stomach is almost totally below the branchial sac. It is slightly cylindrical, with nine folds slightly spirally arranged and all posteriorly closed. These folds have a smooth surface (
|
||||
<figureCitation id="49D505D4E946FFEA3D06C0ECFAC6FA80" box="[638,765,1268,1290]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="9" pageNumber="1184">Fig. 5F, I–L</figureCitation>
|
||||
), without the longitudinal groove observed in
|
||||
<taxonomicName id="16EE62D2E946FFEA3DD9C10BF934FACD" class="Ascidiacea" family="Styelidae" genus="Botryllus" kingdom="Animalia" order="Stolidobranchia" pageId="9" pageNumber="1184" phylum="Chordata" rank="species" species="schlosseri">
|
||||
<emphasis id="E39AC543E946FFEA3DD9C10BF934FACD" italics="true" pageId="9" pageNumber="1184">Botryllus schlosseri</emphasis>
|
||||
</taxonomicName>
|
||||
(Brunetti
|
||||
<emphasis id="E39AC543E946FFEA3EFEC12AF985FACD" box="[390,446,1330,1351]" italics="true" pageId="9" pageNumber="1184">et al.</emphasis>
|
||||
, 2017: fig. 1). They are longer on the mesial side (folds
|
||||
<quantity id="1616B4B4E946FFEA3EB0C148FA2AFAEC" box="[456,529,1360,1382]" metricMagnitude="-1" metricUnit="m" metricValue="1.651" metricValueMax="2.032" metricValueMin="1.27" pageId="9" pageNumber="1184" unit="in" value="6.5" valueMax="8.0" valueMin="5.0">5–8 in</quantity>
|
||||
<figureCitation id="49D505D4E946FFEA3D62C148FA57FAEC" box="[538,620,1360,1382]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="9" pageNumber="1184">Fig. 5L</figureCitation>
|
||||
) and without swellings at their cardiac end. The pyloric caecum is long, about half of the stomach, and has a rounded dilated tip; it rises from about the posterior third part of the typhlosole, the anterior part of which is much larger than the posterior one (
|
||||
<figureCitation id="49D505D4E946FFEA3ECAC1F2FA2CFA75" box="[434,535,1513,1536]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="9" pageNumber="1184">Fig. 5F, I</figureCitation>
|
||||
). The intestine has a peculiar shape, with a hairpin bend (indicated with an arrow head in
|
||||
<figureCitation id="49D505D4E946FFEA3E3FC23FF982F9B7" box="[327,441,1575,1597]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="9" pageNumber="1184">Fig. 5C–D</figureCitation>
|
||||
) at level of the first intestinal loop. This aspect is not due to pressure of the gonads, as it is present also in zooids without developed gonads. The rectum is strongly stretched by a connection with the branchial sac through a trabecula at the level of the transversal vessel between the seventh and eighth rows of stigmata (
|
||||
<figureCitation id="49D505D4E946FFEA3E4CC2C7F9F1F97E" box="[308,458,1759,1781]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="9" pageNumber="1184">Fig. 5D–E, H</figureCitation>
|
||||
). The anus opens at level of the seventh row of stigmata, that is one or two rows above the oesophageal opening. The anal opening has four lobes, each of them splits into three minor ones (
|
||||
<figureCitation id="49D505D4E946FFEA3FD3C341F8C0F8E5" box="[171,251,1881,1903]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="9" pageNumber="1184">Fig. 5G</figureCitation>
|
||||
). On the rectum surface, there are four grooves going from the anus to the pyloric gland. In sample BT4, well-developed gonads are present only in buds of the first order, which are ready to substitute the parents. There are up to
|
||||
<specimenCount id="C7E8D2D8E946FFEA3C8EC0CDFC5BFB61" box="[1014,1120,1237,1259]" count="4" pageId="9" pageNumber="1184" type="egg">four eggs</specimenCount>
|
||||
(diameter of about 140 μm) on each side, anterior and dorsal to the testis follicles, which partially embrace them as a cup (
|
||||
<figureCitation id="49D505D4E946FFEA3A41C10BFDA9FAA3" box="[1337,1426,1299,1321]" captionStart="Figure 5" captionStartId="10.[145,226,1678,1700]" captionTargetBox="[302,1267,198,1637]" captionTargetId="figure-25@10.[300,1269,195,1638]" captionTargetPageId="10" captionText="Figure 5. Botryllus gaiae. A, BT2 living colony, syntype; B, BT4 living colony, syntype; C–E, first order bud ready to substitute the regressing parental zooid (stage 8→9); C–D, left side; E, right side; F, intestinal loop; G–H, rectum and anus; I, parietal side of the stomach; L, mesial side of the stomach. Symbols: arrow head, hairpin bend of the first intestinal loop; an, anus; dl, dorsal lamina; dv, dorsal vessel; end, endostyle; nc, neural complex (dorsal tubercle and neural gland); oes, oesophagus; oo, oocyte; pas, primordia of the atrial syphon; pc, pyloric caecum; pg, pyloric gland; pos, primordia of the oral syphon; r, rectum; st, stomach; t, testis; tr, trabecula; ty, typhlosole." pageId="9" pageNumber="1184">Fig. 5E</figureCitation>
|
||||
). Gonadal primordia are present also in buds of the second order. No larvae and embryos have been found.
|
||||
</paragraph>
|
||||
<caption id="859149D9E946FFEA3FDBC002FDA0FBC7" pageId="9" pageNumber="1184" startId="9.[163,243,1050,1072]" targetBox="[163,1444,195,1010]" targetPageId="9" targetType="figure">
|
||||
<paragraph id="D1511951E946FFEA3FDBC002FDA0FBC7" blockId="9.[163,1443,1050,1101]" pageId="9" pageNumber="1184">
|
||||
<emphasis id="E39AC543E946FFEA3FDBC002F936FBA5" bold="true" box="[163,269,1050,1072]" pageId="9" pageNumber="1184">Figure 4.</emphasis>
|
||||
Phylogeny of the
|
||||
<taxonomicName id="16EE62D2E946FFEA3EAAC002FA71FBBA" box="[466,586,1050,1072]" class="Ascidiacea" family="Botryllinae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Stolidobranchia" pageId="9" pageNumber="1184" phylum="Chordata" rank="family">Botryllinae</taxonomicName>
|
||||
inferred from the concatenation of the 13 mitochondrial-encoded proteins. Node values correspond to ML bootstrap and BPP. Nodes having both ML bootstrap ≤ 55% and BPP ≤ 0.50 have been collapsed.
|
||||
</paragraph>
|
||||
</caption>
|
||||
<paragraph id="D1511951E946FFE83C43C197F902FF73" blockId="9.[827,1443,1423,1905]" lastBlockId="11.[163,778,197,250]" lastPageId="11" lastPageNumber="1186" pageId="9" pageNumber="1184">
|
||||
<emphasis id="E39AC543E946FFEA3C43C197FB92FA2E" box="[827,937,1423,1444]" italics="true" pageId="9" pageNumber="1184">Remarks:</emphasis>
|
||||
Based on the structure of the cloaca, this species clearly belongs to the genus
|
||||
<emphasis id="E39AC543E946FFEA3A7EC1B5FD44FA48" box="[1286,1407,1453,1474]" italics="true" pageId="9" pageNumber="1184">
|
||||
<taxonomicName id="16EE62D2E946FFEA3A7EC1B5FD42FA48" authorityName="Gaertner" authorityYear="1774" box="[1286,1401,1453,1474]" class="Ascidiacea" family="Styelidae" genus="Botryllus" kingdom="Animalia" order="Stolidobranchia" pageId="9" pageNumber="1184" phylum="Chordata" rank="genus">Botryllus</taxonomicName>
|
||||
,
|
||||
</emphasis>
|
||||
as defined by
|
||||
<bibRefCitation id="B57F64A0E946FFEA3CC6C1D4FC4EFA6B" author="Brunetti R" box="[958,1141,1484,1506]" pageId="9" pageNumber="1184" pagination="18 - 32" refId="ref11534" refString="Brunetti R. 2009. Botryllid species (Tunicata, Ascidiacea) from the Mediterranean coast of Israel, with some considerations on the systematics of Botryllinae. Zootaxa 2289: 18-32." type="journal article" year="2009">Brunetti (2009)</bibRefCitation>
|
||||
. The main morphological features distinguishing
|
||||
<taxonomicName id="16EE62D2E946FFEA3B33C1F3FCCCF98A" authorityName="Brunetti & Griggio & Mastrototaro & Gasparini & Gissi" authorityYear="2020" box="[1099,1271,1515,1536]" class="Ascidiacea" family="Styelidae" genus="Botryllus" kingdom="Animalia" order="Stolidobranchia" pageId="9" pageNumber="1184" phylum="Chordata" rank="species" species="gaiae">
|
||||
<emphasis id="E39AC543E946FFEA3B33C1F3FCCCF98A" box="[1099,1271,1515,1536]" italics="true" pageId="9" pageNumber="1184">Botryllus gaiae</emphasis>
|
||||
</taxonomicName>
|
||||
from
|
||||
<taxonomicName id="16EE62D2E946FFEA3A42C1F3FB91F994" class="Ascidiacea" family="Styelidae" genus="Botryllus" kingdom="Animalia" order="Stolidobranchia" pageId="9" pageNumber="1184" phylum="Chordata" rank="species" species="schlosseri">
|
||||
<emphasis id="E39AC543E946FFEA3A42C1F3FB91F994" italics="true" pageId="9" pageNumber="1184">Botryllus schlosseri</emphasis>
|
||||
</taxonomicName>
|
||||
are reported in
|
||||
<tableCitation id="9C6C2CEAE946FFEA3B1DC211FC81F995" box="[1125,1210,1545,1567]" captionStart="Table 4" captionStartId="11.[163,227,835,856]" captionText="Table 4. Comparison between the new species Botryllus gaiae and the neotype of Botryllus schlosseri (Brunetti et al., 2017). The discriminant characters are reported in bold" pageId="9" pageNumber="1184">Table 4</tableCitation>
|
||||
and include several traits currently considered valid diagnostic features to discriminate species in the genus
|
||||
<taxonomicName id="16EE62D2E946FFEA3BB9C25FFD11F9D6" authorityName="Gaertner" authorityYear="1774" box="[1217,1322,1607,1628]" class="Ascidiacea" family="Styelidae" genus="Botryllus" kingdom="Animalia" order="Stolidobranchia" pageId="9" pageNumber="1184" phylum="Chordata" rank="genus">
|
||||
<emphasis id="E39AC543E946FFEA3BB9C25FFD11F9D6" box="[1217,1322,1607,1628]" italics="true" pageId="9" pageNumber="1184">Botryllus</emphasis>
|
||||
</taxonomicName>
|
||||
(
|
||||
<bibRefCitation id="B57F64A0E946FFEA3A42C25FFB4FF9F1" author="Brunetti R" pageId="9" pageNumber="1184" pagination="18 - 32" refId="ref11534" refString="Brunetti R. 2009. Botryllid species (Tunicata, Ascidiacea) from the Mediterranean coast of Israel, with some considerations on the systematics of Botryllinae. Zootaxa 2289: 18-32." type="journal article" year="2009">Brunetti, 2009</bibRefCitation>
|
||||
;
|
||||
<bibRefCitation id="B57F64A0E946FFEA3CFBC27DFCCFF9F1" author="Brunetti R & Mastrototaro F" box="[899,1268,1637,1659]" pageId="9" pageNumber="1184" refId="ref11672" refString="Brunetti R, Mastrototaro F. 2017. Ascidiacea of the European waters. Milan: Edagricole - Edizioni Agricole di New Business Media Srl." type="book" year="2017">Brunetti & Mastrototaro, 2017</bibRefCitation>
|
||||
). These traits, highlighted in bold in
|
||||
<tableCitation id="9C6C2CEAE946FFEA3B40C29CFCB7F913" box="[1080,1164,1668,1690]" captionStart="Table 4" captionStartId="11.[163,227,835,856]" captionText="Table 4. Comparison between the new species Botryllus gaiae and the neotype of Botryllus schlosseri (Brunetti et al., 2017). The discriminant characters are reported in bold" pageId="9" pageNumber="1184">Table 4</tableCitation>
|
||||
, concern peculiarities of the stomach and the intestine, such as the shape of the first and second intestinal curve, the shape and position of the anus opening, the number and surface appearance of the stomach folds and the shape of the typhlosole. All these morphological differences are highly significant and support the hypothesis that clade E is a new species, distinct from
|
||||
<taxonomicName id="16EE62D2E946FFE83A40C343FA59FF50" authority="sensu Brunetti et al. (2017)" authorityName="Brunetti" authorityYear="2017" class="Ascidiacea" family="Styelidae" genus="Botryllus" kingdom="Animalia" lastPageId="11" lastPageNumber="1186" order="Stolidobranchia" pageId="9" pageNumber="1184" phylum="Chordata" rank="species" species="schlosseri">
|
||||
<emphasis id="E39AC543E946FFE83A40C343F965FF51" italics="true" lastPageId="11" lastPageNumber="1186" pageId="9" pageNumber="1184">Botryllus schlosseri sensu</emphasis>
|
||||
Brunetti
|
||||
<emphasis id="E39AC543E944FFE83EACC4DEFA34FF50" box="[468,527,197,219]" italics="true" pageId="11" pageNumber="1186">et al.</emphasis>
|
||||
(2017)
|
||||
</taxonomicName>
|
||||
and therefore from clade A.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
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