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<mods:title id="A0C9E917D0F704387FB57444B97EE065">Cutting the ribbon: bathyal Nemertea from seeps along the Costa Rica margin, with descriptions of 2 new genera and 9 new species</mods:title>
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<mods:affiliation id="7BF25C1027EC0CA63DA183C401DE4834">Institute of Evolutionary Biology and Animal Ecology, Rheinische Friedrich-Wilhelms-Universität Bonn, An der Immenburg 1, 53121 Bonn, Germany. &amp; Scripps Institution of Oceanography, 9500 Gilman Drive, La Jolla, California 92093 - 0202, USA.</mods:affiliation>
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<paragraph id="D7198944FFA8FFF5FDC3FC8EFC1F2BED" blockId="10.[568,1020,891,953]" box="[568,1020,891,918]" pageId="10" pageNumber="142">
<heading id="8C513E28FFA8FFF5FDC3FC8EFC1F2BED" box="[568,1020,891,918]" centered="true" fontSize="11" level="2" pageId="10" pageNumber="142" reason="2">
<taxonomicName id="10A6F2C7FFA8FFF5FDC3FC8EFC1F2BED" ID-CoL="59JZ5" authority="Cantell, 2001" authorityName="Cantell" authorityYear="2001" box="[568,1020,891,918]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" isUncertain="true" kingdom="Animalia" order="Tubulaniformes" pageId="10" pageNumber="142" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFA8FFF5FDC3FC8EFD582BED" bold="true" box="[568,699,891,918]" italics="true" pageId="10" pageNumber="142">Tubulanus</emphasis>
cf.
<emphasis id="E5D25556FFA8FFF5FD1CFC8EFCB52BED" bold="true" box="[743,854,891,918]" italics="true" pageId="10" pageNumber="142">lutescens</emphasis>
<bibRefCitation id="B337F4B5FFA8FFF5FCA5FC89FC1F2BED" author="Cantell C. - E." box="[862,1020,892,918]" pageId="10" pageNumber="142" pagination="213 - 221" refId="ref19009" refString="Cantell C. - E. 2001. On the anatomy and taxonomy of Tubulanus lutescens n. sp. (Nemertini) from the West Coast of Sweden. Ophelia 54: 213 - 221. https: // doi. org / 10.1080 / 00785236.2001.10409467" type="journal article" year="2001">Cantell, 2001</bibRefCitation>
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<paragraph id="D7198944FFA8FFF5FD30FC6AFC8A2BC2" blockId="10.[568,1020,891,953]" box="[715,873,927,953]" pageId="10" pageNumber="142">
<figureCitation id="4F9D95C1FFA8FFF5FD30FC6AFCAE2BC2" box="[715,845,927,953]" captionStart="Fig" captionStartId="11.[190,233,1852,1878]" captionTargetBox="[189,1398,1021,1811]" captionTargetId="figure-262@11.[189,1398,1021,1811]" captionTargetPageId="11" captionText="Fig. 3. Live images of palaeonemerteans, heteronemerteans, and polystiliferous hoplonemerteans collected along the Costa Rica margin. A. Tubulanus cf. lutescens (SIO-BIC N107) (Palaeonemertea). B. Dorsal surface of head region of Tubulanus cf. lutescens (SIO-BIC N233). C. Lineidae (SIO-BIC N254) (Heteronemertea). DE. Reptantia (SIO-BIC N263) (Polystilifera) dorsal (D) and ventral view (E)." figureDoi="http://doi.org/10.5281/zenodo.7258860" httpUri="https://zenodo.org/record/7258860/files/figure.png" pageId="10" pageNumber="142">Figs 3AB</figureCitation>
,
<figureCitation id="4F9D95C1FFA8FFF5FCA1FC6AFC8A2BC2" box="[858,873,927,953]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="10" pageNumber="142">4</figureCitation>
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<heading id="8C513E28FFA8FFF5FF46FC15FED52B81" bold="true" box="[189,310,992,1018]" fontSize="11" level="3" pageId="10" pageNumber="142" reason="3">
<emphasis id="E5D25556FFA8FFF5FF46FC15FED52B81" bold="true" box="[189,310,992,1018]" pageId="10" pageNumber="142">Diagnosis</emphasis>
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<taxonomicName id="10A6F2C7FFA8FFF5FF46FBFAFE342C52" authorityName="Cantell" authorityYear="2001" box="[189,471,1039,1065]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" isUncertain="true" kingdom="Animalia" order="Tubulaniformes" pageId="10" pageNumber="142" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFA8FFF5FF46FBFAFED42C52" box="[189,311,1039,1065]" italics="true" pageId="10" pageNumber="142">Tubulanus</emphasis>
cf.
<emphasis id="E5D25556FFA8FFF5FE90FBFAFE342C52" box="[363,471,1039,1065]" italics="true" pageId="10" pageNumber="142">lutescens</emphasis>
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can be clearly identified as a member of the genus
<taxonomicName id="10A6F2C7FFA8FFF5FBA3FBFAFB312C52" authorityName="Renier" authorityYear="1804" box="[1112,1234,1039,1065]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Phyllodocida" pageId="10" pageNumber="142" phylum="Nemertea" rank="genus">
<emphasis id="E5D25556FFA8FFF5FBA3FBFAFB312C52" box="[1112,1234,1039,1065]" italics="true" pageId="10" pageNumber="142">Tubulanus</emphasis>
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based on the amount of body wall, proboscis, and rhynchocoel muscle layers, the position of the nervous system, the composition of the cerebral organs, and several other internal characteristics (
<bibRefCitation id="B337F4B5FFA8FFF5FBB9FBA0FB3E2C0B" author="Gibson R." box="[1090,1245,1109,1136]" pageId="10" pageNumber="142" refId="ref20008" refString="Gibson R. 1982. British Nemerteans. Cambridge University Press, Cambridge." type="book" year="1982">Gibson 1982</bibRefCitation>
;
<bibRefCitation id="B337F4B5FFA8FFF5FB12FBA0FE822CE8" author="Sundberg P. &amp; Hylbom R." pageId="10" pageNumber="142" pagination="347 - 402" refId="ref24023" refString="Sundberg P. &amp; Hylbom R. 1994. Phylogeny of the nemertean Subclass Palaeonemertea (Anopla, Nemertea). Cladistics 10: 347 - 402. https: // doi. org / 10.1006 / clad. 1994.1025" type="journal article" year="1994">Sundberg &amp; Hylbom 1994</bibRefCitation>
;
<bibRefCitation id="B337F4B5FFA8FFF5FE97FB8CFD792CE8" author="Gibson R. &amp; Sundberg P." box="[364,666,1145,1171]" pageId="10" pageNumber="142" pagination="151 - 196" refId="ref20148" refString="Gibson R. &amp; Sundberg P. 1999. Six new species of palaeonemerteans (Nemertea) from Hong Kong. Zoological Journal of the Linnean Society 125: 151 - 196. https: // doi. org / 10.1111 / j. 1096 - 3642.1999. tb 00590. x" type="journal article" year="1999">Gibson &amp; Sundberg 1999</bibRefCitation>
). As in other members of the genus, the body wall is composed of three muscle layers (outer circular, median longitudinal, and inner circular musculature), whereas the proboscis is composed of two layers (outer longitudinal, inner circular) and the rhynchocoel wall of only one circular muscle layer. The nervous system is located between the epidermal basement membrane and the outer circular body wall musculature and the cerebral organs consist of ciliated canals in the epidermis. Moreover, a mid-dorsal blood vessel and eyes are absent. The collected specimens are of bright red coloration and lacks a conspicuous colour pattern. A lower dorsal nerve is absent.
</paragraph>
</subSubSection>
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<emphasis id="E5D25556FFA8FFF5FF46FA78FE492DDC" bold="true" box="[189,426,1421,1447]" pageId="10" pageNumber="142">Material examined</emphasis>
</heading>
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and
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leg.; collected by HOV
<emphasis id="E5D25556FFA8FFF5FD6FF9F7FD332E67" box="[660,720,1538,1564]" italics="true" pageId="10" pageNumber="142">Alvin</emphasis>
, Dive 4907; SIO-BIC N277 (ex SIO-BIC N233) (image series: https://doi.org/10.5281/zenodo.6368041)
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<specimenCount id="C1A042CDFFA8FFF5FCF3F9D3FCBA2E3B" box="[776,857,1574,1600]" pageId="10" pageNumber="142" type="generic">1 spec.</specimenCount>
(anterior fixed in paraformaldehyde, posterior fixed in ethanol); same collection data as for preceding; GenBank:
<accessionNumber id="C8F514A7FFA8FFF5FC30F9BCFBB32E18" box="[971,1104,1609,1635]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON186263" pageId="10" pageNumber="142">ON186263</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FBA7F9BCFB012E18" box="[1116,1250,1609,1635]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON036061" pageId="10" pageNumber="142">ON036061</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FB16F9BCFA912E1F" box="[1261,1394,1609,1636]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON021853" pageId="10" pageNumber="142">ON021853</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FF46F999FEA02EFD" box="[189,323,1644,1670]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON182046" pageId="10" pageNumber="142">ON182046</accessionNumber>
; SIO-BIC N233
</materialsCitation>
<materialsCitation id="67CE8319FFA8FFF5FDEEF999FC312ED2" ID-GBIF-Occurrence="3948783326" accessionNumber="ON186263, ON036061, ON021853, ON182046" collectingDate="2017-05-22" collectionCode="MZUCR" collectorName="Lisa Levin &amp; Charlotte Seid" country="Costa Rica" elevation="995" latitude="8.93" location="Mound" longLatPrecision="77" longitude="-84.313" pageId="10" pageNumber="142" specimenCount="1">
<specimenCount id="C1A042CDFFA8FFF5FDEEF999FD852EFD" box="[533,614,1644,1670]" pageId="10" pageNumber="142" type="generic">1 spec.</specimenCount>
(fixed in paraformaldehyde); same collection data as for preceding;
<collectionCode id="B1B71181FFA8FFF5FF46F97AFEC62ED2" box="[189,293,1679,1705]" pageId="10" pageNumber="142">MZUCR</collectionCode>
Nemertea Collection MZUCR-103-01 (ex SIO-BIC N233)
</materialsCitation>
<materialsCitation id="67CE8319FFA8FFF5FC1CF97AFDE12F68" ID-GBIF-Occurrence="3948783312" accessionNumber="ON186261, ON036062, ON021852, ON182043" collectingDate="2009-02-26" collectorName="Greg Rouse &amp; Mike Skowronski" country="Costa Rica" elevation="1032" latitude="8.92" location="Mound" longLatPrecision="77" longitude="-84.306" pageId="10" pageNumber="142" specimenCount="1">
<specimenCount id="C1A042CDFFA8FFF5FC1CF97AFBD42ED1" box="[999,1079,1679,1706]" pageId="10" pageNumber="142" type="generic">1 spec.</specimenCount>
(piece in ethanol); methane seep
<location id="D279DF9FFFA8FFF5FF06F947FEB12EB7" LSID="urn:lsid:plazi:treatment:5F0F3852FFA8FFF1FDC3FC8EFD2A2912:D279DF9FFFA8FFF5FF06F947FEB12EB7" box="[253,338,1714,1740]" country="Costa Rica" latitude="8.92" longLatPrecision="77" longitude="-84.306" name="Mound" pageId="10" pageNumber="142">Mound</location>
11;
<geoCoordinate id="B292EF83FFA8FFF5FE75F947FDE32EB6" box="[398,512,1714,1741]" degrees="8.920" direction="north" orientation="latitude" pageId="10" pageNumber="142" precision="55" value="8.92">8.920° N</geoCoordinate>
,
<geoCoordinate id="B292EF83FFA8FFF5FDEBF947FD752EB6" box="[528,662,1714,1741]" degrees="84.306" direction="west" orientation="longitude" pageId="10" pageNumber="142" precision="55" value="-84.306">84.306° W</geoCoordinate>
; depth
<quantity id="105E24A1FFA8FFF5FD0FF947FC7D2EB6" box="[756,926,1714,1741]" metricMagnitude="3" metricUnit="m" metricValue="1.032" metricValueMax="1.045" metricValueMin="1.019" pageId="10" pageNumber="142" unit="m" value="1032.0" valueMax="1045.0" valueMin="1019.0">
<elevation id="5C8B6E77FFA8FFF5FD0FF947FC7D2EB6" box="[756,926,1714,1741]" metricMagnitude="3" metricUnit="m" metricValue="1.032" metricValueMax="1.045" metricValueMin="1.019" pageId="10" pageNumber="142" unit="m" value="1032.0" valueMax="1045.0" valueMin="1019.0">10191045 m</elevation>
</quantity>
;
<date id="A318AF84FFA8FFF5FC54F947FBB52EB7" box="[943,1110,1714,1740]" pageId="10" pageNumber="142" value="2009-02-26">
<collectingDate id="B35C566CFFA8FFF5FC54F947FBB52EB7" box="[943,1110,1714,1740]" pageId="10" pageNumber="142" value="2009-02-26">26 Feb. 2009</collectingDate>
</date>
;
<collectorName id="7A53EC92FFA8FFF5FB9DF947FB152EB6" box="[1126,1270,1714,1741]" pageId="10" pageNumber="142">Greg Rouse</collectorName>
and
<collectorName id="7A53EC92FFA8FFF5FAC2F947FEAA2E94" pageId="10" pageNumber="142">Mike Skowronski</collectorName>
leg.; collected by HOV
<emphasis id="E5D25556FFA8FFF5FD94F920FD4F2E94" box="[623,684,1749,1775]" italics="true" pageId="10" pageNumber="142">Alvin</emphasis>
, Dive 4505; GenBank:
<accessionNumber id="C8F514A7FFA8FFF5FC33F920FBAD2E94" box="[968,1102,1749,1775]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON186261" pageId="10" pageNumber="142">ON186261</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FBA1F920FB032E94" box="[1114,1248,1749,1775]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON036062" pageId="10" pageNumber="142">ON036062</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FB16F920FA902E8B" box="[1261,1395,1749,1776]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON021852" pageId="10" pageNumber="142">ON021852</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FF45F90CFEA62F68" box="[190,325,1785,1811]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON182043" pageId="10" pageNumber="142">ON182043</accessionNumber>
; SIO-BIC N107
</materialsCitation>
<materialsCitation id="67CE8319FFA8FFF5FDE1F90CFBCF2F22" ID-GBIF-Occurrence="3948783302" accessionNumber="ON186262, ON036063, ON021854, ON182044" collectingDate="2010-01-07" collectorName="Lisa Levin &amp; Graham Nash" country="Costa Rica" elevation="991" latitude="8.931" location="Mound" longLatPrecision="77" longitude="-84.313" pageId="10" pageNumber="142" specimenCount="1">
<specimenCount id="C1A042CDFFA8FFF5FDE1F90CFD882F68" box="[538,619,1785,1811]" pageId="10" pageNumber="142" type="generic">1 spec.</specimenCount>
(piece in ethanol); methane seep
<location id="D279DF9FFFA8FFF5FC09F90CFBA42F68" LSID="urn:lsid:plazi:treatment:5F0F3852FFA8FFF1FDC3FC8EFD2A2912:D279DF9FFFA8FFF5FC09F90CFBA42F68" box="[1010,1095,1785,1811]" country="Costa Rica" latitude="8.931" longLatPrecision="77" longitude="-84.313" name="Mound" pageId="10" pageNumber="142">Mound</location>
12;
<geoCoordinate id="B292EF83FFA8FFF5FB81F90CFB042F68" box="[1146,1255,1785,1811]" degrees="8.931" direction="north" orientation="latitude" pageId="10" pageNumber="142" precision="55" value="8.931">8.931° N</geoCoordinate>
,
<geoCoordinate id="B292EF83FFA8FFF5FB09F90CFA902F68" box="[1266,1395,1785,1811]" degrees="84.313" direction="west" orientation="longitude" pageId="10" pageNumber="142" precision="55" value="-84.313">84.313° W</geoCoordinate>
; depth
<quantity id="105E24A1FFA8FFF5FEFEF8E9FE792F4D" box="[261,410,1820,1846]" metricMagnitude="2" metricUnit="m" metricValue="9.91" metricValueMax="10.0" metricValueMin="9.82" pageId="10" pageNumber="142" unit="m" value="991.0" valueMax="1000.0" valueMin="982.0">
<elevation id="5C8B6E77FFA8FFF5FEFEF8E9FE792F4D" box="[261,410,1820,1846]" metricMagnitude="2" metricUnit="m" metricValue="9.91" metricValueMax="10.0" metricValueMin="9.82" pageId="10" pageNumber="142" unit="m" value="991.0" valueMax="1000.0" valueMin="982.0">9821000 m</elevation>
</quantity>
;
<date id="A318AF84FFA8FFF5FE5DF8E9FDCD2F4D" box="[422,558,1820,1846]" pageId="10" pageNumber="142" value="2010-01-07">
<collectingDate id="B35C566CFFA8FFF5FE5DF8E9FDCD2F4D" box="[422,558,1820,1846]" pageId="10" pageNumber="142" value="2010-01-07">7 Jan. 2010</collectingDate>
</date>
;
<collectorName id="7A53EC92FFA8FFF5FDC2F8E9FD552F4D" box="[569,694,1820,1846]" pageId="10" pageNumber="142">Lisa Levin</collectorName>
and
<collectorName id="7A53EC92FFA8FFF5FD15F8E9FC6D2F4D" box="[750,910,1820,1846]" pageId="10" pageNumber="142">Graham Nash</collectorName>
leg.; collected by HOV
<emphasis id="E5D25556FFA8FFF5FB53F8E9FB052F4D" box="[1192,1254,1820,1846]" italics="true" pageId="10" pageNumber="142">Alvin</emphasis>
, Dive 4586; GenBank:
<accessionNumber id="C8F514A7FFA8FFF5FEC2F8CAFE5C2F22" box="[313,447,1855,1881]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON186262" pageId="10" pageNumber="142">ON186262</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FE32F8CAFDAC2F22" box="[457,591,1855,1881]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON036063" pageId="10" pageNumber="142">ON036063</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FDA2F8CAFD3D2F21" box="[601,734,1855,1882]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON021854" pageId="10" pageNumber="142">ON021854</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FD11F8CAFC932F22" box="[746,880,1855,1881]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON182044" pageId="10" pageNumber="142">ON182044</accessionNumber>
; SIO-BIC N120
</materialsCitation>
<materialsCitation id="67CE8319FFA8FFF5FBB8F8CAFE972FBF" ID-GBIF-Occurrence="3948783323" accessionNumber="ON186264, ON036060, ON021851, ON182045" collectingDate="2018-10-20" collectorName="Lisa Levin &amp; Kyle Metcalfe" country="Costa Rica" elevation="996" latitude="8.93037" location="Mound" longLatPrecision="1" longitude="-84.31319" pageId="10" pageNumber="142" specimenCount="1">
<specimenCount id="C1A042CDFFA8FFF5FBB8F8CAFB772F21" box="[1091,1172,1855,1882]" pageId="10" pageNumber="142" type="generic">1 spec.</specimenCount>
(ethanol); methane seep
<location id="D279DF9FFFA8FFF5FF02F897FEAD2F07" LSID="urn:lsid:plazi:treatment:5F0F3852FFA8FFF1FDC3FC8EFD2A2912:D279DF9FFFA8FFF5FF02F897FEAD2F07" box="[249,334,1890,1916]" country="Costa Rica" latitude="8.93037" longLatPrecision="1" longitude="-84.31319" name="Mound" pageId="10" pageNumber="142">Mound</location>
12;
<geoCoordinate id="B292EF83FFA8FFF5FE78F897FDED2F06" box="[387,526,1890,1917]" degrees="8.93037" direction="north" orientation="latitude" pageId="10" pageNumber="142" precision="1" value="8.93037">8.93037° N</geoCoordinate>
,
<geoCoordinate id="B292EF83FFA8FFF5FDE0F897FD592F06" box="[539,698,1890,1917]" degrees="84.31319" direction="west" orientation="longitude" pageId="10" pageNumber="142" precision="1" value="-84.31319">84.31319° W</geoCoordinate>
; depth
<quantity id="105E24A1FFA8FFF5FCEBF897FCBD2F06" box="[784,862,1890,1917]" metricMagnitude="2" metricUnit="m" metricValue="9.959999999999999" pageId="10" pageNumber="142" unit="m" value="996.0">
<elevation id="5C8B6E77FFA8FFF5FCEBF897FCBD2F06" box="[784,862,1890,1917]" metricMagnitude="2" metricUnit="m" metricValue="9.959999999999999" pageId="10" pageNumber="142" unit="m" value="996.0">996 m</elevation>
</quantity>
;
<date id="A318AF84FFA8FFF5FC91F897FBE42F07" box="[874,1031,1890,1916]" pageId="10" pageNumber="142" value="2018-10-20">
<collectingDate id="B35C566CFFA8FFF5FC91F897FBE42F07" box="[874,1031,1890,1916]" pageId="10" pageNumber="142" value="2018-10-20">20 Oct. 2018</collectingDate>
</date>
;
<collectorName id="7A53EC92FFA8FFF5FBEFF897FB702F07" box="[1044,1171,1890,1916]" pageId="10" pageNumber="142">Lisa Levin</collectorName>
and
<collectorName id="7A53EC92FFA8FFF5FB36F897FA952F07" box="[1229,1398,1890,1916]" pageId="10" pageNumber="142">Kyle Metcalfe</collectorName>
leg.; collected by HOV
<emphasis id="E5D25556FFA8FFF5FE20F870FDF42FE4" box="[475,535,1925,1951]" italics="true" pageId="10" pageNumber="142">Alvin</emphasis>
, Dive 4974; GenBank:
<accessionNumber id="C8F514A7FFA8FFF5FCCEF870FC592FE4" box="[821,954,1925,1951]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON186264" pageId="10" pageNumber="142">ON186264</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FC3CF870FBAE2FE4" box="[967,1101,1925,1951]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON036060" pageId="10" pageNumber="142">ON036060</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FBA2F870FB3C2FDB" box="[1113,1247,1925,1952]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON021851" pageId="10" pageNumber="142">ON021851</accessionNumber>
,
<accessionNumber id="C8F514A7FFA8FFF5FB17F870FA912FDB" box="[1260,1394,1925,1952]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/ON182045" pageId="10" pageNumber="142">ON182045</accessionNumber>
; SIO-BIC N257
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="9FBCDACFFFA9FFF3FF46FEFDFE6A293D" lastPageId="12" lastPageNumber="144" pageId="11" pageNumber="143" type="description">
<paragraph id="D7198944FFA9FFF4FF46FEFDFEAD2959" blockId="11.[189,334,264,290]" box="[189,334,264,290]" pageId="11" pageNumber="143">
<heading id="8C513E28FFA9FFF4FF46FEFDFEAD2959" bold="true" box="[189,334,264,290]" fontSize="11" level="3" pageId="11" pageNumber="143" reason="3">
<emphasis id="E5D25556FFA9FFF4FF46FEFDFEAD2959" bold="true" box="[189,334,264,290]" pageId="11" pageNumber="143">Description</emphasis>
</heading>
</paragraph>
<paragraph id="D7198944FFA9FFF4FF46FEC2FE8529A5" blockId="11.[189,1399,311,478]" pageId="11" pageNumber="143">
Specimens up to
<quantity id="105E24A1FFA9FFF4FE7CFEC2FE332929" box="[391,464,311,338]" metricMagnitude="-1" metricUnit="m" metricValue="2.0" pageId="11" pageNumber="143" unit="cm" value="20.0">20 cm</quantity>
in length, up to
<quantity id="105E24A1FFA9FFF4FD6AFEC2FD352929" box="[657,726,311,338]" metricMagnitude="-3" metricUnit="m" metricValue="4.0" pageId="11" pageNumber="143" unit="mm" value="4.0">4 mm</quantity>
in diameter. Body more or less cylindrical. Tapered head slightly wider than rest of body (
<figureCitation id="4F9D95C1FFA9FFF4FDB7FEAFFD4E290F" box="[588,685,346,372]" captionStart="Fig" captionStartId="11.[190,233,1852,1878]" captionTargetBox="[189,1398,1021,1811]" captionTargetId="figure-262@11.[189,1398,1021,1811]" captionTargetPageId="11" captionText="Fig. 3. Live images of palaeonemerteans, heteronemerteans, and polystiliferous hoplonemerteans collected along the Costa Rica margin. A. Tubulanus cf. lutescens (SIO-BIC N107) (Palaeonemertea). B. Dorsal surface of head region of Tubulanus cf. lutescens (SIO-BIC N233). C. Lineidae (SIO-BIC N254) (Heteronemertea). DE. Reptantia (SIO-BIC N263) (Polystilifera) dorsal (D) and ventral view (E)." figureDoi="http://doi.org/10.5281/zenodo.7258860" httpUri="https://zenodo.org/record/7258860/files/figure.png" pageId="11" pageNumber="143">Fig. 3A</figureCitation>
). One pair of dorsal, V-shaped cephalic furrows in front of brain lobes (
<figureCitation id="4F9D95C1FFA9FFF4FEB4FE88FE4829E3" box="[335,427,381,408]" captionStart="Fig" captionStartId="11.[190,233,1852,1878]" captionTargetBox="[189,1398,1021,1811]" captionTargetId="figure-262@11.[189,1398,1021,1811]" captionTargetPageId="11" captionText="Fig. 3. Live images of palaeonemerteans, heteronemerteans, and polystiliferous hoplonemerteans collected along the Costa Rica margin. A. Tubulanus cf. lutescens (SIO-BIC N107) (Palaeonemertea). B. Dorsal surface of head region of Tubulanus cf. lutescens (SIO-BIC N233). C. Lineidae (SIO-BIC N254) (Heteronemertea). DE. Reptantia (SIO-BIC N263) (Polystilifera) dorsal (D) and ventral view (E)." figureDoi="http://doi.org/10.5281/zenodo.7258860" httpUri="https://zenodo.org/record/7258860/files/figure.png" pageId="11" pageNumber="143">Fig. 3B</figureCitation>
). Body bright red, trunk slightly translucent. Anterior tip brighter than rest of head (
<figureCitation id="4F9D95C1FFA9FFF4FF3DFE54FEAA29C0" box="[198,329,417,443]" captionStart="Fig" captionStartId="11.[190,233,1852,1878]" captionTargetBox="[189,1398,1021,1811]" captionTargetId="figure-262@11.[189,1398,1021,1811]" captionTargetPageId="11" captionText="Fig. 3. Live images of palaeonemerteans, heteronemerteans, and polystiliferous hoplonemerteans collected along the Costa Rica margin. A. Tubulanus cf. lutescens (SIO-BIC N107) (Palaeonemertea). B. Dorsal surface of head region of Tubulanus cf. lutescens (SIO-BIC N233). C. Lineidae (SIO-BIC N254) (Heteronemertea). DE. Reptantia (SIO-BIC N263) (Polystilifera) dorsal (D) and ventral view (E)." figureDoi="http://doi.org/10.5281/zenodo.7258860" httpUri="https://zenodo.org/record/7258860/files/figure.png" pageId="11" pageNumber="143">Fig. 3AB</figureCitation>
). Epidermal constrictions along whole body length. Gut and gonads visible through body wall (
<figureCitation id="4F9D95C1FFA9FFF4FEFAFE31FEBB29A5" box="[257,344,452,478]" captionStart="Fig" captionStartId="11.[190,233,1852,1878]" captionTargetBox="[189,1398,1021,1811]" captionTargetId="figure-262@11.[189,1398,1021,1811]" captionTargetPageId="11" captionText="Fig. 3. Live images of palaeonemerteans, heteronemerteans, and polystiliferous hoplonemerteans collected along the Costa Rica margin. A. Tubulanus cf. lutescens (SIO-BIC N107) (Palaeonemertea). B. Dorsal surface of head region of Tubulanus cf. lutescens (SIO-BIC N233). C. Lineidae (SIO-BIC N254) (Heteronemertea). DE. Reptantia (SIO-BIC N263) (Polystilifera) dorsal (D) and ventral view (E)." figureDoi="http://doi.org/10.5281/zenodo.7258860" httpUri="https://zenodo.org/record/7258860/files/figure.png" pageId="11" pageNumber="143">Fig 3A</figureCitation>
).
</paragraph>
<paragraph id="D7198944FFA9FFF3FF46FDF0FE6A293D" blockId="11.[187,1400,517,931]" lastBlockId="12.[188,1398,265,326]" lastPageId="12" lastPageNumber="144" pageId="11" pageNumber="143">
Epidermis in foregut region 150200 µm thick. Body wall composed of outer circular, middle longitudinal and inner circular musculature (
<figureCitation id="4F9D95C1FFA9FFF4FD3FFDDDFCFC2A38" box="[708,799,552,579]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4E</figureCitation>
). Rhynchocoel wall only with circular muscle layer (
<figureCitation id="4F9D95C1FFA9FFF4FF3EFDB9FEC32A1D" box="[197,288,588,614]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4E</figureCitation>
). Proboscis with two layers of musculature (outer longitudinal, inner circular), with two nerves, no proboscis armature (
<figureCitation id="4F9D95C1FFA9FFF4FE36FD9AFDCA2AF2" box="[461,553,623,649]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4C</figureCitation>
). Proboscis pore subterminal, ventral (
<figureCitation id="4F9D95C1FFA9FFF4FC1CFD9AFBA62AF2" box="[999,1093,623,649]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4A</figureCitation>
); mouth opening separate, behind cerebral ganglia (
<figureCitation id="4F9D95C1FFA9FFF4FE21FD67FDD62AD7" box="[474,565,658,684]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4B</figureCitation>
). Cerebral ganglia with only outer neurilemma, neurochord cells absent. Dorsal and ventral ganglia fused, located in ventral part of head (
<figureCitation id="4F9D95C1FFA9FFF4FC47FD40FBF82AB4" box="[956,1051,693,719]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4D</figureCitation>
). Lateral nerve cords without accessory nerves, myofibrillae absent (
<figureCitation id="4F9D95C1FFA9FFF4FD77FD2DFD042A88" box="[652,743,728,755]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4F</figureCitation>
). Precerebral nerves numerous, not grouped (
<figureCitation id="4F9D95C1FFA9FFF4FAF3FD2DFA8A2A88" box="[1288,1385,728,755]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4A</figureCitation>
); buccal nerves paired. Upper dorsal nerve present (
<figureCitation id="4F9D95C1FFA9FFF4FD00FD09FCB62B6D" box="[763,853,764,790]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4E</figureCitation>
). Nervous system between epidermal basement membrane and outer circular body wall musculature (
<figureCitation id="4F9D95C1FFA9FFF4FCB2FCEAFC2D2B42" box="[841,974,799,825]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4D, F</figureCitation>
). Eyes lacking. Cerebral organs as simple ciliated canals in epidermis, posterior to ventral ganglia (
<figureCitation id="4F9D95C1FFA9FFF4FC44FCB7FBC32B27" box="[959,1056,834,860]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="11" pageNumber="143">Fig. 4D</figureCitation>
). Apical organ as depression frontal of rhynchopore, cephalic glands in anterior half of head, scattered between muscle fibres of head. Blood vascular system consisting of cephalic loop giving rise to two paired postcerebral, lateral vessels (
<figureCitation id="4F9D95C1FFAEFFF3FEE4FEFCFE432958" box="[287,416,265,291]" captionStart="Fig" captionStartId="12.[189,232,1642,1668]" captionTargetBox="[189,1398,912,1601]" captionTargetId="figure-218@12.[189,1398,912,1602]" captionTargetPageId="12" captionText="Fig. 4. Tubulanus cf. lutescens Cantell, 2001 (SIO-BIC N277), transverse histological sections. A. Anterior region of head with proboscis pore. B. Posterior brain region with mouth opening and buccal nerves (arrowheads). C. Proboscis composition and the two proboscis nerves (arrowheads) in the foregut region. D. Brain region showing the cerebral canal. E. Body wall composition in the foregut region, arrowhead indicates upper dorsal nerve. F. Lateral nerve cord in the foregut region. Abbreviations: br = brain; cc = cerebral canal; cl = cephalic loop; dl = dorsal brain lobe; e = epidermis; es = esophagus; icm = inner circular muscle layer; lb = lateral blood vessel; lm = longitudinal muscle layer; ln = lateral nerve cord; mo = mouth opening; ocm = outer circular muscle layer; p = proboscis; pcm = circular musculature of proboscis; pcn = precerebral nerves; pe = proboscis epithelium; plm = longitudinal musculature of proboscis; pp = proboscis pore; rc = rhynchocoel; rcm = circular musculature of rhynchocoel; vc = ventral commissure; vl = ventral brain lobe. Scale bars: AC, E = 200 µm; D, F = 100 µm." figureDoi="http://doi.org/10.5281/zenodo.7258862" httpUri="https://zenodo.org/record/7258862/files/figure.png" pageId="12" pageNumber="144">Fig. 4AB</figureCitation>
). Mid-dorsal blood vessel absent, smaller vessels splitting from lateral blood vessels in foregut region.
</paragraph>
</subSubSection>
<caption id="83D9D9CCFFA9FFF4FF45F8C9FEC92F98" ID-DOI="http://doi.org/10.5281/zenodo.7258860" ID-Zenodo-Dep="7258860" httpUri="https://zenodo.org/record/7258860/files/figure.png" pageId="11" pageNumber="143" startId="11.[190,233,1852,1878]" targetBox="[189,1398,1021,1811]" targetPageId="11">
<paragraph id="D7198944FFA9FFF4FF45F8C9FEC92F98" blockId="11.[189,1399,1852,2019]" pageId="11" pageNumber="143">
<emphasis id="E5D25556FFA9FFF4FF45F8C9FEF72F2D" bold="true" box="[190,276,1852,1878]" pageId="11" pageNumber="143">Fig. 3.</emphasis>
Live images of palaeonemerteans, heteronemerteans, and polystiliferous hoplonemerteans collected along the Costa Rica margin.
<emphasis id="E5D25556FFA9FFF4FD69F8AAFD452F02" bold="true" box="[658,678,1887,1913]" pageId="11" pageNumber="143">A</emphasis>
.
<taxonomicName id="10A6F2C7FFA9FFF4FD4EF8AAFC282F02" authorityName="Cantell" authorityYear="2001" box="[693,971,1887,1913]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" isUncertain="true" kingdom="Animalia" order="Tubulaniformes" pageId="11" pageNumber="143" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFA9FFF4FD4EF8AAFCCC2F02" box="[693,815,1887,1913]" italics="true" pageId="11" pageNumber="143">Tubulanus</emphasis>
cf.
<emphasis id="E5D25556FFA9FFF4FCA4F8AAFC282F02" box="[863,971,1887,1913]" italics="true" pageId="11" pageNumber="143">lutescens</emphasis>
</taxonomicName>
(SIO-BIC N107) (
<taxonomicName id="10A6F2C7FFA9FFF4FB51F8AAFA882F02" authorityName="Hubrecht" authorityYear="1879" box="[1194,1387,1887,1913]" class="Palaeonemertea" kingdom="Animalia" pageId="11" pageNumber="143" phylum="Nemertea" rank="class">Palaeonemertea</taxonomicName>
).
<emphasis id="E5D25556FFA9FFF4FF45F876FF322FE6" bold="true" box="[190,209,1923,1949]" pageId="11" pageNumber="143">B</emphasis>
. Dorsal surface of head region of
<taxonomicName id="10A6F2C7FFA9FFF4FD8FF876FC6F2FE6" authorityName="Cantell" authorityYear="2001" box="[628,908,1923,1949]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" isUncertain="true" kingdom="Animalia" order="Tubulaniformes" pageId="11" pageNumber="143" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFA9FFF4FD8FF876FD0D2FE6" box="[628,750,1923,1949]" italics="true" pageId="11" pageNumber="143">Tubulanus</emphasis>
cf.
<emphasis id="E5D25556FFA9FFF4FCDBF876FC6F2FE6" box="[800,908,1923,1949]" italics="true" pageId="11" pageNumber="143">lutescens</emphasis>
</taxonomicName>
(SIO-BIC N233).
<emphasis id="E5D25556FFA9FFF4FB94F877FB612FE7" bold="true" box="[1135,1154,1922,1948]" pageId="11" pageNumber="143">C</emphasis>
.
<taxonomicName id="10A6F2C7FFA9FFF4FB69F876FB1A2FE6" box="[1170,1273,1923,1949]" class="Pilidiophora" family="Lineidae" kingdom="Animalia" order="Heteronemertea" pageId="11" pageNumber="143" phylum="Nemertea" rank="family">Lineidae</taxonomicName>
(SIO-BIC N254) (
<taxonomicName id="10A6F2C7FFA9FFF4FED9F853FE062FBB" authorityName="Burger" authorityYear="1892" box="[290,485,1958,1984]" class="Hoplonemertea" kingdom="Animalia" order="Heteronemertea" pageId="11" pageNumber="143" phylum="Nemertea" rank="order">Heteronemertea</taxonomicName>
).
<emphasis id="E5D25556FFA9FFF4FE06F853FDD72FBB" bold="true" box="[509,564,1958,1984]" pageId="11" pageNumber="143">DE</emphasis>
. Reptantia (SIO-BIC N263) (
<taxonomicName id="10A6F2C7FFA9FFF4FC4AF853FBA72FBB" box="[945,1092,1958,1984]" class="Hoplonemertea" kingdom="Animalia" order="Polystilifera" pageId="11" pageNumber="143" phylum="Nemertea" rank="order">Polystilifera</taxonomicName>
) dorsal (D) and ventral view (E).
</paragraph>
</caption>
<subSubSection id="9FBCDACFFFAEFFF1FF46FE9BFD2A2912" lastPageId="14" lastPageNumber="146" pageId="12" pageNumber="144" type="discussion">
<paragraph id="D7198944FFAEFFF3FF46FE9BFECC29F3" blockId="12.[189,303,366,392]" box="[189,303,366,392]" pageId="12" pageNumber="144">
<heading id="8C513E28FFAEFFF3FF46FE9BFECC29F3" bold="true" box="[189,303,366,392]" fontSize="11" level="3" pageId="12" pageNumber="144" reason="3">
<emphasis id="E5D25556FFAEFFF3FF46FE9BFECC29F3" bold="true" box="[189,303,366,392]" pageId="12" pageNumber="144">Remarks</emphasis>
</heading>
</paragraph>
<paragraph id="D7198944FFAEFFF1FF46FE69FD2A2912" blockId="12.[187,1400,412,826]" lastBlockId="14.[189,1398,265,362]" lastPageId="14" lastPageNumber="146" pageId="12" pageNumber="144">
The Costa Rican specimens are attributed to the genus
<taxonomicName id="10A6F2C7FFAEFFF3FCD0FE69FC4629CD" authorityName="Renier" authorityYear="1804" box="[811,933,412,438]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Phyllodocida" pageId="12" pageNumber="144" phylum="Nemertea" rank="genus">
<emphasis id="E5D25556FFAEFFF3FCD0FE69FC4629CD" box="[811,933,412,438]" italics="true" pageId="12" pageNumber="144">Tubulanus</emphasis>
</taxonomicName>
, based on the phylogenetic analyses and the similarity of external and internal characteristics. In the phylogenetic analysis, this species belongs to a highly supported clade containing
<taxonomicName id="10A6F2C7FFAEFFF3FD6DFE17FBA32986" authority="Yamaoka, 1940" authorityName="Yamaoka" authorityYear="1940" box="[662,1088,482,509]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Phyllodocida" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="eozensis">
<emphasis id="E5D25556FFAEFFF3FD6DFE17FC9F2987" box="[662,892,482,508]" italics="true" pageId="12" pageNumber="144">Tubulanus eozensis</emphasis>
Yamaoka, 1940
</taxonomicName>
,
<taxonomicName id="10A6F2C7FFAEFFF3FBAAFE17FA8C2987" authorityName="Renier" authorityYear="1804" box="[1105,1391,482,508]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Phyllodocida" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="polymorphus">
<emphasis id="E5D25556FFAEFFF3FBAAFE17FA8C2987" box="[1105,1391,482,508]" italics="true" pageId="12" pageNumber="144">Tubulanus polymorphus</emphasis>
</taxonomicName>
, and an undescribed tubulanid from the Sea of Okhotsk (
<figureCitation id="4F9D95C1FFAEFFF3FC93FDF3FC512A5B" box="[872,946,518,544]" captionStart="Fig" captionStartId="11.[190,233,1852,1878]" captionTargetBox="[189,1398,1021,1811]" captionTargetId="figure-262@11.[189,1398,1021,1811]" captionTargetPageId="11" captionText="Fig. 3. Live images of palaeonemerteans, heteronemerteans, and polystiliferous hoplonemerteans collected along the Costa Rica margin. A. Tubulanus cf. lutescens (SIO-BIC N107) (Palaeonemertea). B. Dorsal surface of head region of Tubulanus cf. lutescens (SIO-BIC N233). C. Lineidae (SIO-BIC N254) (Heteronemertea). DE. Reptantia (SIO-BIC N263) (Polystilifera) dorsal (D) and ventral view (E)." figureDoi="http://doi.org/10.5281/zenodo.7258860" httpUri="https://zenodo.org/record/7258860/files/figure.png" pageId="12" pageNumber="144">Fig. 3</figureCitation>
). A clade of undescribed abyssal and hadal tubulanids is sister to that clade. The pairwise distances of the COI gene between the four deepsea specimens are below 1%. Interestingly, pairwise distances between
<taxonomicName id="10A6F2C7FFAEFFF3FBFBFDB9FAF02A1D" authorityName="Cantell" authorityYear="2001" box="[1024,1299,588,614]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" isUncertain="true" kingdom="Animalia" order="Tubulaniformes" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFAEFFF3FBFBFDB9FB992A1D" box="[1024,1146,588,614]" italics="true" pageId="12" pageNumber="144">Tubulanus</emphasis>
cf.
<emphasis id="E5D25556FFAEFFF3FB5CFDB9FAF02A1D" box="[1191,1299,588,614]" italics="true" pageId="12" pageNumber="144">lutescens</emphasis>
</taxonomicName>
and two sequences of
<taxonomicName id="10A6F2C7FFAEFFF3FEA5FD9AFDA82AF2" authorityName="Cantell" authorityYear="2001" box="[350,587,623,649]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Tubulaniformes" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFAEFFF3FEA5FD9AFDA82AF2" box="[350,587,623,649]" italics="true" pageId="12" pageNumber="144">Tubulanus lutescens</emphasis>
</taxonomicName>
, a shallow water species collected in
<collectingCountry id="AFB1C9D4FFAEFFF3FBF1FD9AFB892AF2" box="[1034,1130,623,649]" name="Sweden" pageId="12" pageNumber="144">Sweden</collectingCountry>
, are only 11.3%. In a TCS haplotype network analysis of the mitochondrial COI gene
<taxonomicName id="10A6F2C7FFAEFFF3FC41FD67FB4A2AD7" authorityName="Cantell" authorityYear="2001" box="[954,1193,658,684]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Tubulaniformes" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFAEFFF3FC41FD67FB4A2AD7" box="[954,1193,658,684]" italics="true" pageId="12" pageNumber="144">Tubulanus lutescens</emphasis>
</taxonomicName>
is only separated from our specimens by six nucleotide substitutions (
<figureCitation id="4F9D95C1FFAEFFF3FCD4FD43FC9B2AAB" box="[815,888,694,720]" captionStart="Fig" captionStartId="11.[190,233,1852,1878]" captionTargetBox="[189,1398,1021,1811]" captionTargetId="figure-262@11.[189,1398,1021,1811]" captionTargetPageId="11" captionText="Fig. 3. Live images of palaeonemerteans, heteronemerteans, and polystiliferous hoplonemerteans collected along the Costa Rica margin. A. Tubulanus cf. lutescens (SIO-BIC N107) (Palaeonemertea). B. Dorsal surface of head region of Tubulanus cf. lutescens (SIO-BIC N233). C. Lineidae (SIO-BIC N254) (Heteronemertea). DE. Reptantia (SIO-BIC N263) (Polystilifera) dorsal (D) and ventral view (E)." figureDoi="http://doi.org/10.5281/zenodo.7258860" httpUri="https://zenodo.org/record/7258860/files/figure.png" pageId="12" pageNumber="144">Fig. 3</figureCitation>
). However, the two species differ by body coloration as
<taxonomicName id="10A6F2C7FFAEFFF3FE9AFD2CFD962A88" authorityName="Cantell" authorityYear="2001" box="[353,629,729,755]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" isUncertain="true" kingdom="Animalia" order="Tubulaniformes" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFAEFFF3FE9AFD2CFE382A88" box="[353,475,729,755]" italics="true" pageId="12" pageNumber="144">Tubulanus</emphasis>
cf.
<emphasis id="E5D25556FFAEFFF3FDF2FD2CFD962A88" box="[521,629,729,755]" italics="true" pageId="12" pageNumber="144">lutescens</emphasis>
</taxonomicName>
is bright red, whereas
<taxonomicName id="10A6F2C7FFAEFFF3FC70FD2CFB982A88" authorityName="Cantell" authorityYear="2001" box="[907,1147,729,755]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Tubulaniformes" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFAEFFF3FC70FD2CFB982A88" box="[907,1147,729,755]" italics="true" pageId="12" pageNumber="144">Tubulanus lutescens</emphasis>
</taxonomicName>
is yellow. Moreover,
<taxonomicName id="10A6F2C7FFAEFFF3FF44FD09FE2F2B6D" authorityName="Cantell" authorityYear="2001" box="[191,460,764,790]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" isUncertain="true" kingdom="Animalia" order="Tubulaniformes" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFAEFFF3FF44FD09FEDA2B6D" box="[191,313,764,790]" italics="true" pageId="12" pageNumber="144">Tubulanus</emphasis>
cf.
<emphasis id="E5D25556FFAEFFF3FE9BFD09FE2F2B6D" box="[352,460,764,790]" italics="true" pageId="12" pageNumber="144">lutescens</emphasis>
</taxonomicName>
lacks a lower dorsal nerve which is present in
<taxonomicName id="10A6F2C7FFAEFFF3FC20FD09FB242B6D" authorityName="Cantell" authorityYear="2001" box="[987,1223,764,790]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Tubulaniformes" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFAEFFF3FC20FD09FB242B6D" box="[987,1223,764,790]" italics="true" pageId="12" pageNumber="144">Tubulanus lutescens</emphasis>
</taxonomicName>
. The markedly dissimilar habitats of both species (
<taxonomicName id="10A6F2C7FFAEFFF3FDA0FCD5FD002B42" authorityName="Cantell" authorityYear="2001" box="[603,739,799,825]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Tubulaniformes" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFAEFFF3FDA0FCD5FD002B42" box="[603,739,799,825]" italics="true" pageId="12" pageNumber="144">T. lutescens</emphasis>
</taxonomicName>
intertidal from Northern Europe vs
<taxonomicName id="10A6F2C7FFAEFFF3FB72FCD5FAD42B42" authorityName="Cantell" authorityYear="2001" box="[1161,1335,799,825]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" isUncertain="true" kingdom="Animalia" order="Tubulaniformes" pageId="12" pageNumber="144" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFAEFFF3FB72FCD5FB422B42" box="[1161,1185,800,825]" italics="true" pageId="12" pageNumber="144">T.</emphasis>
cf.
<emphasis id="E5D25556FFAEFFF3FB30FCEAFAD42B42" box="[1227,1335,799,825]" italics="true" pageId="12" pageNumber="144">lutescens</emphasis>
</taxonomicName>
from ca
<quantity id="105E24A1FFACFFF1FF25FEFCFED42958" box="[222,311,265,291]" metricMagnitude="3" metricUnit="m" metricValue="1.0" pageId="14" pageNumber="146" unit="m" value="1000.0">1000 m</quantity>
depth in Eastern Pacific) might indicate that they represent different species. Future analyses of nuclear sequences of
<taxonomicName id="10A6F2C7FFACFFF1FE1FFED8FD93293D" authorityName="Cantell" authorityYear="2001" box="[484,624,300,326]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Tubulaniformes" pageId="14" pageNumber="146" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFACFFF1FE1FFED8FD93293D" box="[484,624,300,326]" italics="true" pageId="14" pageNumber="146">T. lutescens</emphasis>
</taxonomicName>
are warranted to determine whether
<taxonomicName id="10A6F2C7FFACFFF1FBD6FED9FAA0293D" authorityName="Cantell" authorityYear="2001" box="[1069,1347,300,326]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" isUncertain="true" kingdom="Animalia" order="Tubulaniformes" pageId="14" pageNumber="146" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFACFFF1FBD6FED9FB44293D" box="[1069,1191,300,326]" italics="true" pageId="14" pageNumber="146">Tubulanus</emphasis>
cf.
<emphasis id="E5D25556FFACFFF1FB2CFED9FAA0293D" box="[1239,1347,300,326]" italics="true" pageId="14" pageNumber="146">lutescens</emphasis>
</taxonomicName>
and
<taxonomicName id="10A6F2C7FFACFFF1FF44FEA5FEAB2912" authorityName="Cantell" authorityYear="2001" box="[191,328,335,361]" class="Palaeonemertea" family="Tubulanidae" genus="Tubulanus" kingdom="Animalia" order="Tubulaniformes" pageId="14" pageNumber="146" phylum="Nemertea" rank="species" species="lutescens">
<emphasis id="E5D25556FFACFFF1FF44FEA5FEAB2912" box="[191,328,335,361]" italics="true" pageId="14" pageNumber="146">T. lutescens</emphasis>
</taxonomicName>
are indeed two different species.
</paragraph>
</subSubSection>
</treatment>
</document>