554 lines
72 KiB
XML
554 lines
72 KiB
XML
<document id="A0EB12A269F2BD7947C1404F7F695121" ID-DOI="10.11646/zootaxa.1780.1.2" ID-GBIF-Dataset="bd903eb2-5814-48fa-8d32-73c0e469415f" ID-ISSN="1175-5326" ID-Zenodo-Dep="182322" IM.bibliography_approvedBy="diego" IM.illustrations_approvedBy="diego" IM.materialsCitations_approvedBy="diego" IM.metadata_approvedBy="diego" IM.taxonomicNames_approvedBy="diego" IM.treatments_approvedBy="diego" checkinTime="1459859264701" checkinUser="plazi" docAuthor="Evans, Ben J., Carter, Timothy F., Tobias, Martha L. & Kelley, Darcy B." docDate="2008" docId="03E78791DE67790EFF58253988CDFAB3" docLanguage="en" docName="zt01780p068.pdf" docOrigin="Zootaxa 1780" docStyle="DocumentStyle:6581673A57F01A3145754A1E615EDFF0.4:Zootaxa.2007-2008.journal_article" docStyleId="6581673A57F01A3145754A1E615EDFF0" docStyleName="Zootaxa.2007-2008.journal_article" docStyleVersion="4" docTitle="Xenopus itombwensis Evans, Carter, Tobias & Kelley, 2008, new species" docType="treatment" docVersion="13" lastPageNumber="64" masterDocId="FFDEFFE9DE637907FFCF26448A3CFFF2" masterDocTitle="A new species of clawed frog (genus Xenopus) from the Itombwe Massif, Democratic Republic of the Congo: implications for DNA barcodes and biodiversity conservation." masterLastPageNumber="68" masterPageNumber="55" pageNumber="59" updateTime="1738663638323" updateUser="ExternalLinkService">
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<mods:mods id="72055CB10FFE46E45973365EA2CEA15C" xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo id="9F9CF540829B897EE4380F06C415610C">
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<mods:title id="D0878D0362F5F344E9FF0086CF7AB240">A new species of clawed frog (genus Xenopus) from the Itombwe Massif, Democratic Republic of the Congo: implications for DNA barcodes and biodiversity conservation.</mods:title>
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<mods:name id="56E014301832F1105AE021DD59377933" type="personal">
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<mods:roleTerm id="255BA883601C0CF33D62B1E32E934903">Author</mods:roleTerm>
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<mods:namePart id="3F14C9DBF0A86CC5BF1CD5DCC71EE1A0">Evans, Ben J.</mods:namePart>
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<mods:roleTerm id="65CA8508570F8D23D53DD7228FCC8D9E">Author</mods:roleTerm>
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<mods:namePart id="D35989C5643D90C3751F98CB18C9CE0B">Carter, Timothy F.</mods:namePart>
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<mods:roleTerm id="96DC9B3296CC7E0E2814207B5584B9EA">Author</mods:roleTerm>
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</mods:role>
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<mods:namePart id="530717EA236F078DB1C2FDD26F1AE98D">Tobias, Martha L.</mods:namePart>
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</mods:name>
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<mods:name id="E538DE83A81FA90F301B9CBF0271D110" type="personal">
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<mods:roleTerm id="9159733FDC86912F020FD715BBD5F991">Author</mods:roleTerm>
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</mods:role>
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<mods:namePart id="01E4519EA920A6F9A788FFBC6839E29D">Kelley, Darcy B.</mods:namePart>
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<mods:typeOfResource id="BAD5EB46F5CF5A8A78DD7BC081BA2445">text</mods:typeOfResource>
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<mods:titleInfo id="68C4F7EAE64A45F46BB89E4E35C9FE6E">
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<mods:title id="8CF783449D8DDF10497F689DB7AAE9EE">Zootaxa</mods:title>
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</mods:titleInfo>
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<mods:part id="43D28ADEF8A9B344B231DCB351341D93">
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<mods:date id="33591B3D07A4D6627FE0EC487E3BB1E9">2008</mods:date>
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<mods:detail id="404A5A0FEAF29B2DA38AAF782AF5171E" type="pubDate">
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<mods:number id="D2B15E2D938B67C12C1303DB3D284945">2008-05-28</mods:number>
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<mods:detail id="1D164761E9816998462F781F8B62FCBF" type="volume">
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<mods:number id="4AC013152C83D031B63F71859364A6CC">1780</mods:number>
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<mods:classification id="D55749545F8D93BE66598872588C0F16">journal article</mods:classification>
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<mods:identifier id="731BFD9F0F50AED0B979A843B095ACA8" type="DOI">10.11646/zootaxa.1780.1.2</mods:identifier>
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<mods:identifier id="307E01EBB3B1B98F7F9C6D1BF5966952" type="ISSN">1175-5326</mods:identifier>
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<mods:identifier id="7222C6729BBECA352DBA02908EC79596" type="Zenodo-Dep">182322</mods:identifier>
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<treatment id="03E78791DE67790EFF58253988CDFAB3" ID-DOI="http://doi.org/10.5281/zenodo.6228252" ID-GBIF-Taxon="119367954" ID-Zenodo-Dep="6228252" LSID="urn:lsid:zoobank.org:act:F0AEF218-76FC-4AB7-8FB9-02D8115B10D1" httpUri="http://treatment.plazi.org/id/03E78791DE67790EFF58253988CDFAB3" lastPageId="9" lastPageNumber="64" pageId="4" pageNumber="59">
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<subSubSection id="C354650CDE677903FF5825398807FC65" box="[151,571,893,919]" pageId="4" pageNumber="59" type="nomenclature">
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<paragraph id="8BF13687DE677903FF5825398807FC65" blockId="4.[151,571,893,994]" box="[151,571,893,919]" pageId="4" pageNumber="59">
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<heading id="D0B981EBDE677903FF5825398807FC65" bold="true" box="[151,571,893,919]" fontSize="11" level="1" pageId="4" pageNumber="59" reason="1">
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<emphasis id="B93AEA95DE677903FF5825398807FC65" bold="true" box="[151,571,893,919]" pageId="4" pageNumber="59">
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<taxonomicName id="4C4E4D04DE677903FF5825398BA0FC65" ID-CoL="5CBC6" authorityName="Evans, Carter, Tobias, Kelley, Hanner & Tinsley" authorityYear="2008" box="[151,412,893,919]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="itombwensis" status="sp. nov.">
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<emphasis id="B93AEA95DE677903FF5825398BA0FC65" bold="true" box="[151,412,893,919]" italics="true" pageId="4" pageNumber="59">Xenopus itombwensis</emphasis>
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</taxonomicName>
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,
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<taxonomicNameLabel id="A20957EEDE677903FE6425398807FC65" box="[427,571,893,919]" pageId="4" pageNumber="59" rank="species">new species</taxonomicNameLabel>
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</emphasis>
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||
</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="C354650CDE677903FF5825E48BD5FC48" box="[151,489,928,954]" pageId="4" pageNumber="59" type="vernacular_names">
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<paragraph id="8BF13687DE677903FF5825E48BD5FC48" blockId="4.[151,571,893,994]" box="[151,489,928,954]" pageId="4" pageNumber="59">Itombwe Massif clawed frog</paragraph>
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</subSubSection>
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<subSubSection id="C354650CDE677903FF58258C8AD3FC10" box="[151,239,968,994]" pageId="4" pageNumber="59" type="description">
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<paragraph id="8BF13687DE677903FF58258C8AD3FC10" blockId="4.[151,571,893,994]" box="[151,239,968,994]" pageId="4" pageNumber="59">
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(
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<figureCitation id="13752A02DE677903FF50258C8AD9FC10" box="[159,229,968,994]" captionStart="FIGURE 2" captionStartId="5.[151,255,612,636]" captionTargetBox="[186,1402,197,611]" captionTargetId="figure@5.[151,1436,162,618]" captionTargetPageId="5" captionText="FIGURE 2. Type specimen and variation of X. itombwensis. (A) Holotype specimen MCZ A- 138192 (field number BJE 0275), (B) Paratype MCZ A- 138193 (field no. BJE 0276), and (C) a live unvouchered male individual. Arrows indicate dorsal spots that are not found in X. wittei. Scale bars are 5 mm. Photo credit for B: Jon Woodward." figureDoi="http://doi.org/10.5281/zenodo.182324" httpUri="https://zenodo.org/record/182324/files/figure.png" pageId="4" pageNumber="59">Fig. 2</figureCitation>
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)
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</paragraph>
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</subSubSection>
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<subSubSection id="C354650CDE677903FF58225C8B9CFB08" pageId="4" pageNumber="59" type="materials_examined">
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<paragraph id="8BF13687DE677903FF58225C8B0BFB70" blockId="4.[151,1437,1048,2034]" pageId="4" pageNumber="59">
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<materialsCitation id="3B263CDADE677903FF58225C8B0BFB70" collectingDate="2006-04-21" collectionCode="MCZ" collectorName="Ben J. Evans" country="Democratic Republic of the Congo" elevation="2200" latitude="-3.35679" location="Miki Town" longLatPrecision="1" longitude="23.69011" municipality="Miki Town" pageId="4" pageNumber="59" specimenCode="MCZ A-138192" specimenCount="2" specimenCount-adult="1" specimenCount-male="1" stateProvince="South Kivu Province" typeStatus="holotype">
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<emphasis id="B93AEA95DE677903FF58225C8B31FBC0" bold="true" box="[151,269,1048,1074]" italics="true" pageId="4" pageNumber="59">
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<typeStatus id="54F58825DE677903FF58225C8B3BFBC0" box="[151,263,1048,1074]" pageId="4" pageNumber="59" type="holotype">Holotype</typeStatus>
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:
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</emphasis>
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<specimenCode id="DBE89EFCDE677903FED8225C8BEAFBC0" box="[279,470,1048,1074]" pageId="4" pageNumber="59">MCZ A-138192</specimenCode>
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(field number BJE 0275),
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||
<specimenCount id="9D48FD0EDE677903FCD6225C896FFBC0" box="[793,851,1048,1074]" pageId="4" pageNumber="59" type="adult">adult</specimenCount>
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<specimenCount id="9D48FD0EDE677903FC93225C89A4FBC0" box="[860,920,1048,1074]" pageId="4" pageNumber="59" type="male">male</specimenCount>
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,
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<collectingCountry id="F3597617DE677903FC69225C8F76FBC0" box="[934,1354,1048,1074]" name="Democratic Republic of the Congo" pageId="4" pageNumber="59">Democratic Republic of the Congo</collectingCountry>
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,
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<collectingRegion id="498AF865DE677903FA98225C8B78FBA8" pageId="4" pageNumber="59">South Kivu Province</collectingRegion>
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,
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<collectingMunicipality id="6B95ACFDDE677903FE9F22048BE9FBA8" box="[336,469,1088,1114]" pageId="4" pageNumber="59">Miki Town</collectingMunicipality>
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||
,
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||
<geoCoordinate id="EE7A5040DE677903FE2E2204885FFBA8" box="[481,611,1088,1114]" direction="south" orientation="latitude" pageId="4" pageNumber="59" precision="1" value="-3.35679">3.35679º S</geoCoordinate>
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||
,
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||
<geoCoordinate id="EE7A5040DE677903FDBE22048936FBA8" box="[625,778,1088,1114]" direction="east" orientation="longitude" pageId="4" pageNumber="59" precision="1" value="23.69011">023.69011ºE</geoCoordinate>
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, approximately
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||
<quantity id="4CB69B62DE677903FC0922048E23FBA8" box="[966,1055,1088,1114]" metricMagnitude="3" metricUnit="m" metricValue="2.2" pageId="4" pageNumber="59" unit="m" value="2200.0">
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<elevation id="0063D1B4DE677903FC0922048E23FBA8" box="[966,1055,1088,1114]" metricMagnitude="3" metricUnit="m" metricValue="2.2" pageId="4" pageNumber="59" unit="m" value="2200.0">2200 m</elevation>
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</quantity>
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||
above sea level.
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||
<collectingDate id="EFB4E9AFDE677903FB2622048FA4FBA8" box="[1257,1432,1088,1114]" pageId="4" pageNumber="59">
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<date id="FFF01047DE677903FB2622048F72FBA8" box="[1257,1358,1088,1114]" pageId="4" pageNumber="59" value="1921-04">21 April</date>
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, 2006
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</collectingDate>
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||
,
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<collectorName id="26BB5351DE677903FF58222C8B0EFB70" box="[151,306,1128,1154]" pageId="4" pageNumber="59">Ben J. Evans</collectorName>
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||
.
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||
</materialsCitation>
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||
</paragraph>
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||
<paragraph id="8BF13687DE677903FF0922D48B9CFB08" blockId="4.[151,1437,1048,2034]" pageId="4" pageNumber="59">
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||
<materialsCitation id="3B263CDADE677903FF0922D48B14FB20" collectingDate="2006-04-21" collectionCode="MCZ" collectorName="Ben J. Evans" country="Democratic Republic of the Congo" elevation="2200" latitude="-3.35679" location="Miki Town" longLatPrecision="1" longitude="23.69011" municipality="Miki Town" pageId="4" pageNumber="59" specimenCode="MCZ A-138195, MCZ A-138196, MCZ A-138197" specimenCount="3" specimenCount-adult="3" specimenCount-male="3" stateProvince="South Kivu Province" typeStatus="paratype">
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||
<emphasis id="B93AEA95DE677903FF0922D48B7BFB58" bold="true" box="[198,327,1168,1194]" italics="true" pageId="4" pageNumber="59">
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||
<typeStatus id="54F58825DE677903FF0922D48B7CFB58" box="[198,320,1168,1194]" pageId="4" pageNumber="59" type="paratype">Paratypes</typeStatus>
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:
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</emphasis>
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<specimenCount id="9D48FD0EDE677903FE8122D4881FFB58" box="[334,547,1168,1194]" pageId="4" pageNumber="59">Three adult males</specimenCount>
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:
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<specimenCode id="DBE89EFCDE677903FDE122D488D7FB58" box="[558,747,1168,1194]" pageId="4" pageNumber="59">MCZ A-138195</specimenCode>
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(BJE 0278),
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<specimenCode id="DBE89EFCDE677903FC4922D48E7FFB58" box="[902,1091,1168,1194]" pageId="4" pageNumber="59">MCZ A-138196</specimenCode>
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(BJE 0283),
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<specimenCode id="DBE89EFCDE677903FB1022D48FA0FB58" box="[1247,1436,1168,1194]" pageId="4" pageNumber="59">MCZ A-138197</specimenCode>
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(BJE 0284),
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</materialsCitation>
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<materialsCitation id="3B263CDADE677903FEFB22FC8B9CFB08" collectingDate="2006-04-21" collectionCode="MCZ" collectorName="Ben J. Evans" country="Democratic Republic of the Congo" elevation="2200" latitude="-3.35679" location="Miki Town" longLatPrecision="1" longitude="23.69011" municipality="Miki Town" pageId="4" pageNumber="59" specimenCode="MCZ A-138193, MCZ A-138194" specimenCount="2" stateProvince="South Kivu Province" typeStatus="paratype">
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<specimenCount id="9D48FD0EDE677903FEFB22FC8BE5FB20" box="[308,473,1208,1234]" pageId="4" pageNumber="59">two juveniles</specimenCount>
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, sex undetermined:
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<specimenCode id="DBE89EFCDE677903FD1F22FC89AFFB20" box="[720,915,1208,1234]" pageId="4" pageNumber="59">MCZ A-138193</specimenCode>
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(BJE 0276),
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<specimenCode id="DBE89EFCDE677903FBF322FC8EC2FB20" box="[1084,1278,1208,1234]" pageId="4" pageNumber="59">MCZ A-138194</specimenCode>
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(BJE 0277), same data as holotype.
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</materialsCitation>
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</paragraph>
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||
</subSubSection>
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||
<subSubSection id="C354650CDE677902FF09234C891CFCC4" lastPageId="5" lastPageNumber="60" pageId="4" pageNumber="59" type="diagnosis">
|
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<paragraph id="8BF13687DE677903FF09234C8975F940" blockId="4.[151,1437,1048,2034]" pageId="4" pageNumber="59">
|
||
<emphasis id="B93AEA95DE677903FF09234C8870FAD3" box="[198,588,1288,1314]" italics="true" pageId="4" pageNumber="59">
|
||
<emphasis id="B93AEA95DE677903FF09234C8B74FAD0" bold="true" box="[198,328,1288,1314]" italics="true" pageId="4" pageNumber="59">Diagnosis:</emphasis>
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<taxonomicName id="4C4E4D04DE677903FE80234C8870FAD3" box="[335,588,1288,1313]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="itombwensis">Xenopus itombwensis</taxonomicName>
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</emphasis>
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is a member of the
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<emphasis id="B93AEA95DE677903FCFD234C89E9FAD3" box="[818,981,1288,1313]" italics="true" pageId="4" pageNumber="59">vestitus-wittei</emphasis>
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subgroup of clawed frogs (
|
||
<bibRefCitation id="EFDF4B76DE677903FADC234C8AEFFAB8" author="Kobel, H. R. & Loumont, C. & Tinsley, R. C." editor="Tinsley, R. C. & Kobel, H. R." etAl="et al." firstAuthor="Kobel" journalOrPublisher="Clarendon Press, Oxford" pageId="4" pageNumber="59" pagination="9 - 33" refId="ref6605" refString="Kobel, H. R., Loumont, C. & Tinsley, R. C. (1996) The extant species. In: Tinsley, R. C. and Kobel, H. R., Eds). The Biology of Xenopus. Clarendon Press, Oxford. pp. 9 - 33" title="The extant species" type="book chapter" volumeTitle="The Biology of Xenopus" year="1996">Kobel et al. 1996</bibRefCitation>
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) and can be distinguished from other members of this group by (1) unique but variable morphological coloration and smaller size (Table 1,
|
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<figureCitation id="13752A02DE677903FD8A231C88B7FA80" box="[581,651,1368,1394]" captionStart="FIGURE 2" captionStartId="5.[151,255,612,636]" captionTargetBox="[186,1402,197,611]" captionTargetId="figure@5.[151,1436,162,618]" captionTargetPageId="5" captionText="FIGURE 2. Type specimen and variation of X. itombwensis. (A) Holotype specimen MCZ A- 138192 (field number BJE 0275), (B) Paratype MCZ A- 138193 (field no. BJE 0276), and (C) a live unvouchered male individual. Arrows indicate dorsal spots that are not found in X. wittei. Scale bars are 5 mm. Photo credit for B: Jon Woodward." figureDoi="http://doi.org/10.5281/zenodo.182324" httpUri="https://zenodo.org/record/182324/files/figure.png" pageId="4" pageNumber="59">Fig. 2</figureCitation>
|
||
), by (2) numerous temporal and spectral characteristics of the male advertisement call (Table 2,
|
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<figureCitation id="13752A02DE677903FE2723C48813FA68" box="[488,559,1408,1434]" captionStart="FIGURE 3" captionStartId="7.[151,255,1939,1963]" captionTargetBox="[420,1151,214,1938]" captionTargetId="figure@7.[420,1165,194,1955]" captionTargetPageId="7" captionText="FIGURE 3. Male advertisement vocalization of (A) X. wittei, (B) X. vestitus, and (C) X. itombwensis. The slow trill portion of the X. itombwensis call (beginning at about 400 milliseconds) is a unique feature within the “ vestitus-wittei ” group." figureDoi="http://doi.org/10.5281/zenodo.182325" httpUri="https://zenodo.org/record/182325/files/figure.png" pageId="4" pageNumber="59">Fig. 3</figureCitation>
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), and by (3) divergent mitochondrial and autosomal genes (
|
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<figureCitation id="13752A02DE677903FB3923C48F7CFA68" box="[1270,1344,1408,1434]" captionStart="FIGURE 4" captionStartId="8.[151,255,1722,1746]" captionTargetBox="[200,1372,186,1705]" captionTargetId="figure@8.[200,1372,186,1707]" captionTargetPageId="8" captionText="FIGURE 4. Evolutionary relationships of (A) combined data from two tightly linked nuclear loci (RAG 1 and RAG 2) and (B) mitochondrial DNA illustrate a divergent but sister relationship of X. itombwensis and X. wittei. Nuclear loci but not mitochondrial loci illustrate a close relationship between (X. itombwensis + X. wittei) to half of the allopolyploid genome of X. vestitus. For clarity most posterior probabilities are omitted because they are similar or identical to those found elsewhere (Evans 2007; Evans et al. 2004). However, with reference to X. itombwensis, in (A and B) the red clades have over 95 % posterior probability and the blue clade has over 80 % posterior probability. (C) A species phylogeny illustrating bifurcating and reticulating evolutionary relationships in clawed frogs. The most recent common ancestor of X. wittei and X. itombwensis evolved through allopolyploidization of two tetraploid species. The number of chromosomes in each species is indicated in parentheses after each species name. (A) and (C) are modified from (Evans 2007)." figureDoi="http://doi.org/10.5281/zenodo.182326" httpUri="https://zenodo.org/record/182326/files/figure.png" pageId="4" pageNumber="59">Fig. 4</figureCitation>
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).
|
||
<taxonomicName id="4C4E4D04DE677903FA9923C48B34FA33" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE677903FA9923C48B34FA33" italics="true" pageId="4" pageNumber="59">Xenopus wittei</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE677903FE8D23EC8B87FA33" box="[322,443,1448,1473]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE677903FE8D23EC8B87FA33" box="[322,443,1448,1473]" italics="true" pageId="4" pageNumber="59">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
are both medium sized clawed frogs, with female snout-vent length (SVL) typically 46 and
|
||
<quantity id="4CB69B62DE677903FF2123948B78FA18" box="[238,324,1488,1514]" metricMagnitude="-2" metricUnit="m" metricValue="4.7" pageId="4" pageNumber="59" unit="mm" value="47.0">47 mm</quantity>
|
||
, respectively, and a maximum SVL of 61 and
|
||
<quantity id="4CB69B62DE677903FCA723948983FA18" box="[872,959,1488,1514]" metricMagnitude="-2" metricUnit="m" metricValue="5.5" pageId="4" pageNumber="59" unit="mm" value="55.0">55 mm</quantity>
|
||
, respectively (
|
||
<bibRefCitation id="EFDF4B76DE677903FBA723948F05FA18" author="Kobel, H. R. & Loumont, C. & Tinsley, R. C." box="[1128,1337,1488,1514]" editor="Tinsley, R. C. & Kobel, H. R." etAl="et al." firstAuthor="Kobel" journalOrPublisher="Clarendon Press, Oxford" pageId="4" pageNumber="59" pagination="9 - 33" refId="ref6605" refString="Kobel, H. R., Loumont, C. & Tinsley, R. C. (1996) The extant species. In: Tinsley, R. C. and Kobel, H. R., Eds). The Biology of Xenopus. Clarendon Press, Oxford. pp. 9 - 33" title="The extant species" type="book chapter" volumeTitle="The Biology of Xenopus" year="1996">Kobel et al. 1996</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFDF4B76DE677903FA8A23948B2BF9E0" author="Tinsley, R. C. & Kobel, H. - R. & Fischberg, M." etAl="et al." firstAuthor="Tinsley" journalOrPublisher="Journal Zoology, London" pageId="4" pageNumber="68" pagination="69" part="188" refId="ref7310" refString="Tinsley, R. C., Kobel, H. - R. & Fischberg, M. (1979) The biology and systematics of a new species of Xenopus (Anura: Pipidae) from the highlands of Central Africa. Journal Zoology, London, 188, 69 - 102." title="The biology and systematics of a new species of Xenopus (Anura: Pipidae) from the highlands of Central Africa" type="journal article" year="1979">Tinsley et al. 1979</bibRefCitation>
|
||
). Based on a small sample size of two females,
|
||
<taxonomicName id="4C4E4D04DE677903FC8823BC89C5F9E3" box="[839,1017,1528,1553]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE677903FC8823BC89C5F9E3" box="[839,1017,1528,1553]" italics="true" pageId="4" pageNumber="59">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
appears smaller, averaging approximately
|
||
<quantity id="4CB69B62DE677903FF3720648B70F9C8" box="[248,332,1568,1594]" metricMagnitude="-2" metricUnit="m" metricValue="3.5" pageId="4" pageNumber="59" unit="mm" value="35.0">35 mm</quantity>
|
||
(Table 1). Dorsal coloration of
|
||
<taxonomicName id="4C4E4D04DE677903FD0520648912F9CB" box="[714,814,1568,1593]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE677903FD0520648912F9CB" box="[714,814,1568,1593]" italics="true" pageId="4" pageNumber="59">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
is a uniform dark brown to chocolate with no spots and of
|
||
<taxonomicName id="4C4E4D04DE677903FF23200C8B5BF993" box="[236,359,1608,1633]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE677903FF23200C8B5BF993" box="[236,359,1608,1633]" italics="true" pageId="4" pageNumber="59">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
is a marbeling of light silver-golden to bronze chromatophores over a brown background (
|
||
<bibRefCitation id="EFDF4B76DE677903FF5020348B4FF978" author="Kobel, H. R. & Loumont, C. & Tinsley, R. C." box="[159,371,1648,1674]" editor="Tinsley, R. C. & Kobel, H. R." etAl="et al." firstAuthor="Kobel" journalOrPublisher="Clarendon Press, Oxford" pageId="4" pageNumber="59" pagination="9 - 33" refId="ref6605" refString="Kobel, H. R., Loumont, C. & Tinsley, R. C. (1996) The extant species. In: Tinsley, R. C. and Kobel, H. R., Eds). The Biology of Xenopus. Clarendon Press, Oxford. pp. 9 - 33" title="The extant species" type="book chapter" volumeTitle="The Biology of Xenopus" year="1996">Kobel et al. 1996</bibRefCitation>
|
||
). In contrast, some
|
||
<taxonomicName id="4C4E4D04DE677903FD922034892CF97B" box="[605,784,1648,1673]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE677903FD922034892CF97B" box="[605,784,1648,1673]" italics="true" pageId="4" pageNumber="59">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
individuals have a mottled pattern of brown spots that are slightly darker than the brown background (
|
||
<figureCitation id="13752A02DE677903FD0B20DC8937F940" box="[708,779,1688,1714]" captionStart="FIGURE 2" captionStartId="5.[151,255,612,636]" captionTargetBox="[186,1402,197,611]" captionTargetId="figure@5.[151,1436,162,618]" captionTargetPageId="5" captionText="FIGURE 2. Type specimen and variation of X. itombwensis. (A) Holotype specimen MCZ A- 138192 (field number BJE 0275), (B) Paratype MCZ A- 138193 (field no. BJE 0276), and (C) a live unvouchered male individual. Arrows indicate dorsal spots that are not found in X. wittei. Scale bars are 5 mm. Photo credit for B: Jon Woodward." figureDoi="http://doi.org/10.5281/zenodo.182324" httpUri="https://zenodo.org/record/182324/files/figure.png" pageId="4" pageNumber="59">Fig. 2</figureCitation>
|
||
B,C).
|
||
</paragraph>
|
||
<paragraph id="8BF13687DE677902FF092084891CFCC4" blockId="4.[151,1437,1048,2034]" lastBlockId="5.[151,1437,756,2022]" lastPageId="5" lastPageNumber="60" pageId="4" pageNumber="59">
|
||
The male advertisement call of
|
||
<taxonomicName id="4C4E4D04DE677903FDF2208488CCF92B" box="[573,752,1728,1753]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE677903FDF2208488CCF92B" box="[573,752,1728,1753]" italics="true" pageId="4" pageNumber="59">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
differs from the male advertisement calls of
|
||
<taxonomicName id="4C4E4D04DE677903FAC920848F55F92B" box="[1286,1385,1728,1753]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE677903FAC920848F55F92B" box="[1286,1385,1728,1753]" italics="true" pageId="4" pageNumber="59">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE677903FF5820AC8B2CF8F3" box="[151,272,1768,1793]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE677903FF5820AC8B2CF8F3" box="[151,272,1768,1793]" italics="true" pageId="4" pageNumber="59">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
in that the call of the new species is much shorter (~600 milliseconds), and consists of two distinct components including a “fast trill” and a “slow trill” (Table 2,
|
||
<figureCitation id="13752A02DE677903FC43215489EAF8D8" box="[908,982,1808,1834]" captionStart="FIGURE 3" captionStartId="7.[151,255,1939,1963]" captionTargetBox="[420,1151,214,1938]" captionTargetId="figure@7.[420,1165,194,1955]" captionTargetPageId="7" captionText="FIGURE 3. Male advertisement vocalization of (A) X. wittei, (B) X. vestitus, and (C) X. itombwensis. The slow trill portion of the X. itombwensis call (beginning at about 400 milliseconds) is a unique feature within the “ vestitus-wittei ” group." figureDoi="http://doi.org/10.5281/zenodo.182325" httpUri="https://zenodo.org/record/182325/files/figure.png" pageId="4" pageNumber="59">Fig. 3</figureCitation>
|
||
;
|
||
<bibRefCitation id="EFDF4B76DE677903FC2A21548E85F8D8" author="Kobel, H. R. & Loumont, C. & Tinsley, R. C." box="[997,1209,1808,1834]" editor="Tinsley, R. C. & Kobel, H. R." etAl="et al." firstAuthor="Kobel" journalOrPublisher="Clarendon Press, Oxford" pageId="4" pageNumber="59" pagination="9 - 33" refId="ref6605" refString="Kobel, H. R., Loumont, C. & Tinsley, R. C. (1996) The extant species. In: Tinsley, R. C. and Kobel, H. R., Eds). The Biology of Xenopus. Clarendon Press, Oxford. pp. 9 - 33" title="The extant species" type="book chapter" volumeTitle="The Biology of Xenopus" year="1996">Kobel et al. 1996</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFDF4B76DE677903FB0821548F64F8D8" author="Vigny, C." box="[1223,1368,1808,1834]" firstAuthor="Vigny" journalOrPublisher="Journal Zoology, London" pageId="4" pageNumber="68" pagination="103" part="188" refId="ref7475" refString="Vigny, C. (1979) The mating calls of 12 species and sub-species of the genus Xenopus (Amphibia: Anura). Journal Zoology, London, 188, 103 - 122." title="The mating calls of 12 species and sub-species of the genus Xenopus (Amphibia: Anura)" type="journal article" year="1979">Vigny 1979</bibRefCitation>
|
||
). The dominant frequency of the fast trill component is similar to that of
|
||
<taxonomicName id="4C4E4D04DE677903FC60217C8E2EF8A3" box="[943,1042,1848,1873]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE677903FC60217C8E2EF8A3" box="[943,1042,1848,1873]" italics="true" pageId="4" pageNumber="59">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
but other acoustic characteristics of this part of the vocalization are different, and all parameters we measured from the slow trill component are distinct (Table 2). The most obvious difference between these vocalizations is the slow trill that follows vocalizations of the new species but not
|
||
<taxonomicName id="4C4E4D04DE677903FDFE21F488AEF83B" box="[561,658,1968,1993]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE677903FDFE21F488AEF83B" box="[561,658,1968,1993]" italics="true" pageId="4" pageNumber="59">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
. Other aspects of the slow trill are distinct from call parameters of
|
||
<taxonomicName id="4C4E4D04DE677903FF58219C8AC4F803" box="[151,248,2008,2033]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="4" pageNumber="59" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE677903FF58219C8AC4F803" box="[151,248,2008,2033]" italics="true" pageId="4" pageNumber="59">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
including the lower dominant frequencies, longer interclick interval, and lower number of clicks (Table 2). The fast portion of the call of the new species is also unique in that the interclick interval is shorter than that of
|
||
<taxonomicName id="4C4E4D04DE667902FEEB25598BB9FCC4" box="[292,389,797,822]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FEEB25598BB9FCC4" box="[292,389,797,822]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
and there are fewer clicks per call.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<caption id="DF31660FDE667902FF5824208E87FD33" ID-DOI="http://doi.org/10.5281/zenodo.182324" httpUri="https://zenodo.org/record/182324/files/figure.png" pageId="5" pageNumber="60" targetBox="[186,1402,197,611]" targetPageId="5">
|
||
<paragraph id="8BF13687DE667902FF5824208E87FD33" blockId="5.[151,1436,612,705]" pageId="5" pageNumber="60">
|
||
<emphasis id="B93AEA95DE667902FF5824208B25FD8E" bold="true" box="[151,281,612,636]" pageId="5" pageNumber="60">FIGURE 2.</emphasis>
|
||
Type specimen and variation of
|
||
<emphasis id="B93AEA95DE667902FDBA24218922FD8E" box="[629,798,613,636]" italics="true" pageId="5" pageNumber="60">
|
||
<taxonomicName id="4C4E4D04DE667902FDBA24218926FD8E" box="[629,794,613,636]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">X. itombwensis</taxonomicName>
|
||
.
|
||
</emphasis>
|
||
(A) Holotype specimen MCZ A-138192 (field number BJE 0275), (B) Paratype MCZ A-138193 (field no. BJE 0276), and (C) a live unvouchered male individual. Arrows indicate dorsal spots that are not found in
|
||
<taxonomicName id="4C4E4D04DE667902FE3424EE8868FD33" box="[507,596,682,705]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FE3424EE8868FD33" box="[507,596,682,705]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
. Scale bars are 5 mm. Photo credit for B: Jon Woodward.
|
||
</paragraph>
|
||
</caption>
|
||
<subSubSection id="C354650CDE667902FF0925008985FA44" pageId="5" pageNumber="60" type="discussion">
|
||
<paragraph id="8BF13687DE667902FF0925008985FA44" blockId="5.[151,1437,756,2022]" pageId="5" pageNumber="60">
|
||
<emphasis id="B93AEA95DE667902FF0925008EB3FCAC" box="[198,1167,836,862]" italics="true" pageId="5" pageNumber="60">
|
||
<emphasis id="B93AEA95DE667902FF09250089EAFCAC" bold="true" box="[198,982,836,862]" italics="true" pageId="5" pageNumber="60">Evolutionary relationships to other clawed frogs; DNA barcodes.</emphasis>
|
||
<taxonomicName id="4C4E4D04DE667902FC1225018EB3FCAC" box="[989,1167,837,862]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">X. itombwensis</taxonomicName>
|
||
</emphasis>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FB0725018F16FCAC" box="[1224,1322,837,862]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FB0725018F16FCAC" box="[1224,1322,837,862]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
are probably sister species derived from the same allo-octoploid ancestor. The paternal ancestor of
|
||
<taxonomicName id="4C4E4D04DE667902FB0D25298F07FC74" box="[1218,1339,877,902]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE667902FB0D25298F07FC74" box="[1218,1339,877,902]" italics="true" pageId="5" pageNumber="60">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
, in contrast, shares recent common ancestry with the maternal ancestor of (
|
||
<emphasis id="B93AEA95DE667902FC2D25D18F16FC5C" box="[994,1322,917,942]" italics="true" pageId="5" pageNumber="60">
|
||
<taxonomicName id="4C4E4D04DE667902FC2D25D18EA4FC5C" box="[994,1176,917,942]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">X. itombwensis</taxonomicName>
|
||
+
|
||
<taxonomicName id="4C4E4D04DE667902FB0B25D18F16FC5C" box="[1220,1322,917,942]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">X. wittei</taxonomicName>
|
||
</emphasis>
|
||
), but the maternal ancestor of
|
||
<taxonomicName id="4C4E4D04DE667902FE5F25F98836FC24" box="[400,522,957,982]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE667902FE5F25F98836FC24" box="[400,522,957,982]" italics="true" pageId="5" pageNumber="60">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
is not as closely related to the paternal ancestor of (
|
||
<emphasis id="B93AEA95DE667902FBB925F98A88FC0C" italics="true" pageId="5" pageNumber="60">
|
||
<taxonomicName id="4C4E4D04DE667902FBB925F98F14FC24" box="[1142,1320,957,982]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">X. itombwensis</taxonomicName>
|
||
+
|
||
<taxonomicName id="4C4E4D04DE667902FA8325F98A88FC0C" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">X. wittei</taxonomicName>
|
||
</emphasis>
|
||
).
|
||
<collectingCountry id="F3597617DE667902FF0225A08AD2FC0C" box="[205,238,996,1022]" name="American Samoa" pageId="5" pageNumber="60">As</collectingCountry>
|
||
a result, mitochondrial DNA of the
|
||
<emphasis id="B93AEA95DE667902FD5425A18902FC0C" box="[667,830,997,1022]" italics="true" pageId="5" pageNumber="60">vestitus-wittei</emphasis>
|
||
group is not a clade (
|
||
<figureCitation id="13752A02DE667902FBF125A08EB4FC0C" box="[1086,1160,996,1022]" captionStart="FIGURE 4" captionStartId="8.[151,255,1722,1746]" captionTargetBox="[200,1372,186,1705]" captionTargetId="figure@8.[200,1372,186,1707]" captionTargetPageId="8" captionText="FIGURE 4. Evolutionary relationships of (A) combined data from two tightly linked nuclear loci (RAG 1 and RAG 2) and (B) mitochondrial DNA illustrate a divergent but sister relationship of X. itombwensis and X. wittei. Nuclear loci but not mitochondrial loci illustrate a close relationship between (X. itombwensis + X. wittei) to half of the allopolyploid genome of X. vestitus. For clarity most posterior probabilities are omitted because they are similar or identical to those found elsewhere (Evans 2007; Evans et al. 2004). However, with reference to X. itombwensis, in (A and B) the red clades have over 95 % posterior probability and the blue clade has over 80 % posterior probability. (C) A species phylogeny illustrating bifurcating and reticulating evolutionary relationships in clawed frogs. The most recent common ancestor of X. wittei and X. itombwensis evolved through allopolyploidization of two tetraploid species. The number of chromosomes in each species is indicated in parentheses after each species name. (A) and (C) are modified from (Evans 2007)." figureDoi="http://doi.org/10.5281/zenodo.182326" httpUri="https://zenodo.org/record/182326/files/figure.png" pageId="5" pageNumber="60">Fig. 4</figureCitation>
|
||
). On a molecular level,
|
||
<taxonomicName id="4C4E4D04DE667902FF5822498B70FBD4" box="[151,332,1037,1062]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE667902FF5822498B70FBD4" box="[151,332,1037,1062]" italics="true" pageId="5" pageNumber="60">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
is diverged from
|
||
<taxonomicName id="4C4E4D04DE667902FDE5224988B3FBD4" box="[554,655,1037,1062]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FDE5224988B3FBD4" box="[554,655,1037,1062]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
and from
|
||
<taxonomicName id="4C4E4D04DE667902FCDC224989B3FBD4" box="[787,911,1037,1062]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE667902FCDC224989B3FBD4" box="[787,911,1037,1062]" italics="true" pageId="5" pageNumber="60">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
. Because of the polyphyletic origin of this group, molecular divergence of mitochondrial DNA data (~3000bp) between
|
||
<taxonomicName id="4C4E4D04DE667902FBEF22718EEDFBBC" box="[1056,1233,1077,1102]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE667902FBEF22718EEDFBBC" box="[1056,1233,1077,1102]" italics="true" pageId="5" pageNumber="60">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FAC622718FBEFBBC" box="[1289,1410,1077,1102]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE667902FAC622718FBEFBBC" box="[1289,1410,1077,1102]" italics="true" pageId="5" pageNumber="60">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
is large – on the order of about 9% (uncorrected pairwise distance; hereafter
|
||
<emphasis id="B93AEA95DE667902FBDC22198E1EFB84" box="[1043,1058,1117,1142]" italics="true" pageId="5" pageNumber="60">p</emphasis>
|
||
-distance). Mitochondrial divergence between
|
||
<taxonomicName id="4C4E4D04DE667902FE8022C1883EFB6C" box="[335,514,1157,1182]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE667902FE8022C1883EFB6C" box="[335,514,1157,1182]" italics="true" pageId="5" pageNumber="60">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FDF122C1889DFB6C" box="[574,673,1157,1182]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FDF122C1889DFB6C" box="[574,673,1157,1182]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
is about 4%
|
||
<emphasis id="B93AEA95DE667902FC8E22C1896CFB6C" box="[833,848,1157,1182]" italics="true" pageId="5" pageNumber="60">p</emphasis>
|
||
-distance. When only the “barcode” region of the mitochondria (
|
||
<emphasis id="B93AEA95DE667902FE8B22E98B46FB34" box="[324,378,1197,1222]" italics="true" pageId="5" pageNumber="60">CO1</emphasis>
|
||
) is considered, interspecific divergence between
|
||
<taxonomicName id="4C4E4D04DE667902FC7522E98E57FB34" box="[954,1131,1197,1222]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE667902FC7522E98E57FB34" box="[954,1131,1197,1222]" italics="true" pageId="5" pageNumber="60">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FB6C22E98F27FB34" box="[1187,1307,1197,1222]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE667902FB6C22E98F27FB34" box="[1187,1307,1197,1222]" italics="true" pageId="5" pageNumber="60">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FA9C22E98B35FB1C" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE667902FA9C22E98B35FB1C" italics="true" pageId="5" pageNumber="60">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FE8A22918B95FB1C" box="[325,425,1237,1262]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FE8A22918B95FB1C" box="[325,425,1237,1262]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
is even higher – 17.2% and 8.3%
|
||
<emphasis id="B93AEA95DE667902FC8522918964FB1C" box="[842,856,1237,1262]" italics="true" pageId="5" pageNumber="60">p</emphasis>
|
||
-distance, respectively. Paralogs of
|
||
<emphasis id="B93AEA95DE667902FB3022918F7BFB1C" box="[1279,1351,1237,1262]" italics="true" pageId="5" pageNumber="60">RAG1</emphasis>
|
||
and of
|
||
<emphasis id="B93AEA95DE667902FF5822B98AE2FAE4" box="[151,222,1277,1302]" italics="true" pageId="5" pageNumber="60">RAG2</emphasis>
|
||
are also diverged among these species. Over both genes, the average divergence between the most closely related orthologous paralogs of
|
||
<taxonomicName id="4C4E4D04DE667902FDA8236188E3FACC" box="[615,735,1317,1342]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE667902FDA8236188E3FACC" box="[615,735,1317,1342]" italics="true" pageId="5" pageNumber="60">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FCD6236189F6FACC" box="[793,970,1317,1342]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE667902FCD6236189F6FACC" box="[793,970,1317,1342]" italics="true" pageId="5" pageNumber="60">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
is 1.6%
|
||
<emphasis id="B93AEA95DE667902FBFE23618E7CFACC" box="[1073,1088,1317,1342]" italics="true" pageId="5" pageNumber="60">p</emphasis>
|
||
-distance and between orthologous paralogs of
|
||
<taxonomicName id="4C4E4D04DE667902FEA023098BEDFA94" box="[367,465,1357,1382]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FEA023098BEDFA94" box="[367,465,1357,1382]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FDC523098880FA94" box="[522,700,1357,1382]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE667902FDC523098880FA94" box="[522,700,1357,1382]" italics="true" pageId="5" pageNumber="60">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
is also 1.6%
|
||
<emphasis id="B93AEA95DE667902FC9623098954FA94" box="[857,872,1357,1382]" italics="true" pageId="5" pageNumber="60">p</emphasis>
|
||
-distance. These levels of divergence are typical for closely related species of clawed frog (
|
||
<figureCitation id="13752A02DE667902FD49233088F2FA7C" box="[646,718,1396,1422]" captionStart="FIGURE 4" captionStartId="8.[151,255,1722,1746]" captionTargetBox="[200,1372,186,1705]" captionTargetId="figure@8.[200,1372,186,1707]" captionTargetPageId="8" captionText="FIGURE 4. Evolutionary relationships of (A) combined data from two tightly linked nuclear loci (RAG 1 and RAG 2) and (B) mitochondrial DNA illustrate a divergent but sister relationship of X. itombwensis and X. wittei. Nuclear loci but not mitochondrial loci illustrate a close relationship between (X. itombwensis + X. wittei) to half of the allopolyploid genome of X. vestitus. For clarity most posterior probabilities are omitted because they are similar or identical to those found elsewhere (Evans 2007; Evans et al. 2004). However, with reference to X. itombwensis, in (A and B) the red clades have over 95 % posterior probability and the blue clade has over 80 % posterior probability. (C) A species phylogeny illustrating bifurcating and reticulating evolutionary relationships in clawed frogs. The most recent common ancestor of X. wittei and X. itombwensis evolved through allopolyploidization of two tetraploid species. The number of chromosomes in each species is indicated in parentheses after each species name. (A) and (C) are modified from (Evans 2007)." figureDoi="http://doi.org/10.5281/zenodo.182326" httpUri="https://zenodo.org/record/182326/files/figure.png" pageId="5" pageNumber="60">Fig. 4</figureCitation>
|
||
;
|
||
<bibRefCitation id="EFDF4B76DE667902FD142330895BFA7C" author="Evans, B. J." box="[731,871,1396,1422]" firstAuthor="Evans" journalOrPublisher="Genetics" pageId="5" pageNumber="67" pagination="1119 - 1130" part="176" refId="ref5963" refString="Evans, B. J. (2007) Ancestry influences the fate of duplicated genes millions of years after duplication in allopolyploid clawed frogs (Xenopus). Genetics, 176, 1119 - 1130." title="Ancestry influences the fate of duplicated genes millions of years after duplication in allopolyploid clawed frogs (Xenopus)" type="journal article" year="2007">Evans 2007</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFDF4B76DE667902FCBC23308E7CFA7C" author="Evans, B. J. & Kelley, D. B. & Melnick, D. J. & Cannatella, D. C." box="[883,1088,1396,1422]" etAl="et al." firstAuthor="Evans" journalOrPublisher="Molecular Biology and Evolution" pageId="5" pageNumber="67" pagination="1193 - 1207" part="22" refId="ref6030" refString="Evans, B. J., Kelley, D. B., Melnick, D. J. & Cannatella, D. C. (2005) Evolution of RAG- 1 in polyploid clawed frogs. Molecular Biology and Evolution, 22, 1193 - 1207." title="Evolution of RAG- 1 in polyploid clawed frogs" type="journal article" year="2005">Evans et al. 2005</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFDF4B76DE667902FB8423308F26FA7C" author="Evans, B. J. & Kelley, D. B. & Tinsley, R. C. & Melnick, D. J. & Cannatella, D. C." box="[1099,1306,1396,1422]" etAl="et al." firstAuthor="Evans" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="5" pageNumber="67" pagination="197 - 213" part="33" refId="ref6080" refString="Evans, B. J., Kelley, D. B., Tinsley, R. C., Melnick, D. J. & Cannatella, D. C. (2004) A mitochondrial DNA phylogeny of clawed frogs: phylogeography on sub-Saharan Africa and implications for polyploid evolution. Molecular Phylogenetics and Evolution, 33, 197 - 213." title="A mitochondrial DNA phylogeny of clawed frogs: phylogeography on sub-Saharan Africa and implications for polyploid evolution" type="journal article" year="2004">Evans et al. 2004</bibRefCitation>
|
||
). The Genbank accession number of the barcode of the
|
||
<typeStatus id="54F58825DE667902FD6623D88933FA44" box="[681,783,1436,1462]" pageId="5" pageNumber="60" type="holotype">holotype</typeStatus>
|
||
is
|
||
<accessionNumber id="941DAB64DE667902FCFD23D88989FA44" box="[818,949,1436,1462]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/EU594660" pageId="5" pageNumber="60">EU594660</accessionNumber>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C354650CDE66790EFF0923808EFFFBA3" lastPageId="9" lastPageNumber="64" pageId="5" pageNumber="60" type="description">
|
||
<paragraph id="8BF13687DE667902FF09238088BEF98C" blockId="5.[151,1437,756,2022]" pageId="5" pageNumber="60">
|
||
<emphasis id="B93AEA95DE667902FF0923808821FA2C" bold="true" box="[198,541,1476,1502]" italics="true" pageId="5" pageNumber="60">
|
||
Description of the
|
||
<typeStatus id="54F58825DE667902FE622380882AFA2C" box="[429,534,1476,1502]" pageId="5" pageNumber="60" type="holotype">holotype</typeStatus>
|
||
:
|
||
</emphasis>
|
||
<typeStatus id="54F58825DE667902FDE7238088A9FA2C" box="[552,661,1476,1502]" pageId="5" pageNumber="60" type="holotype">Holotype</typeStatus>
|
||
an adult male, small subocular tentacle present, comprising less than one third of the length of the eye. Claws present on toes I-III, prehallux prominent but without a claw. Size of new species slightly smaller than
|
||
<taxonomicName id="4C4E4D04DE667902FD4C205188D9F9DC" box="[643,741,1557,1582]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FD4C205188D9F9DC" box="[643,741,1557,1582]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FCEF205189A6F9DC" box="[800,922,1557,1582]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE667902FCEF205189A6F9DC" box="[800,922,1557,1582]" italics="true" pageId="5" pageNumber="60">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
(Table 1;
|
||
<bibRefCitation id="EFDF4B76DE667902FBDB20508EDAF9DC" author="Kobel, H. R. & Loumont, C. & Tinsley, R. C." box="[1044,1254,1556,1582]" editor="Tinsley, R. C. & Kobel, H. R." etAl="et al." firstAuthor="Kobel" journalOrPublisher="Clarendon Press, Oxford" pageId="5" pageNumber="60" pagination="9 - 33" refId="ref6605" refString="Kobel, H. R., Loumont, C. & Tinsley, R. C. (1996) The extant species. In: Tinsley, R. C. and Kobel, H. R., Eds). The Biology of Xenopus. Clarendon Press, Oxford. pp. 9 - 33" title="The extant species" type="book chapter" volumeTitle="The Biology of Xenopus" year="1996">Kobel et al. 1996</bibRefCitation>
|
||
). Like
|
||
<taxonomicName id="4C4E4D04DE667902FAF620518FA0F9DC" box="[1337,1436,1557,1582]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FAF620518FA0F9DC" box="[1337,1436,1557,1582]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE667902FF0620798B7FF9A4" box="[201,323,1597,1622]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="vestitus" status="sp. nov.">
|
||
<emphasis id="B93AEA95DE667902FF0620798B7FF9A4" box="[201,323,1597,1622]" italics="true" pageId="5" pageNumber="60">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
,
|
||
<taxonomicNameLabel id="A20957EEDE667902FE9C20788BDDF9A4" box="[339,481,1596,1622]" pageId="5" pageNumber="60" rank="species">new species</taxonomicNameLabel>
|
||
is octoploid with
|
||
<date id="FFF01047DE667902FD77207888EFF9A4" box="[696,723,1596,1622]" pageId="5" pageNumber="60">8x</date>
|
||
=72 chromosomes (
|
||
<figureCitation id="13752A02DE667902FC7020788E34F9A4" box="[959,1032,1596,1622]" captionStart="FIGURE 5" captionStartId="9.[151,255,604,628]" captionTargetId="figure@9.[499,1097,200,581]" captionTargetPageId="9" captionText="FIGURE 5. Karyotype of X. itombwensis illustrating octoploidy, 8 x = 72." pageId="5" pageNumber="60">Fig. 5</figureCitation>
|
||
). Ventral surface of forelimbs and forearm with scattered black nuptial pads.
|
||
</paragraph>
|
||
<paragraph id="8BF13687DE667902FF0920C88E51F8EC" blockId="5.[151,1437,756,2022]" pageId="5" pageNumber="60">
|
||
<emphasis id="B93AEA95DE667902FF0920C888B8F954" bold="true" box="[198,644,1676,1702]" italics="true" pageId="5" pageNumber="60">
|
||
Color of the
|
||
<typeStatus id="54F58825DE667902FE9220C88BFFF954" box="[349,451,1676,1702]" pageId="5" pageNumber="60" type="holotype">holotype</typeStatus>
|
||
in preservative:
|
||
</emphasis>
|
||
Dorsum homogeneous dark brown, transitioning laterally on flanks to a cream-colored venter; dorsal surface of head dark brown; dorsal surface of limbs dark brown; underside of head speckled with gray; venter cream colored with sparse small brown spots on ventral surface of hind limbs; ventral surface of hind feet cream-colored;
|
||
<typeStatus id="54F58825DE667902FD2C21408976F8EC" box="[739,842,1796,1822]" pageId="5" pageNumber="60" type="holotype">holotype</typeStatus>
|
||
color in life unrecorded.
|
||
</paragraph>
|
||
<paragraph id="8BF13687DE66790EFF0921688BE5FBF3" blockId="5.[151,1437,756,2022]" lastBlockId="9.[151,1437,719,1345]" lastPageId="9" lastPageNumber="64" pageId="5" pageNumber="60">
|
||
<emphasis id="B93AEA95DE667902FF0921688832F8B4" bold="true" box="[198,526,1836,1862]" italics="true" pageId="5" pageNumber="60">Variation and color in life:</emphasis>
|
||
There are two dorsal color patterns evident in our sample of
|
||
<taxonomicName id="4C4E4D04DE667902FB2621698FA0F8B4" box="[1257,1436,1837,1862]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE667902FB2621698FA0F8B4" box="[1257,1436,1837,1862]" italics="true" pageId="5" pageNumber="60">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
from the
|
||
<typeStatus id="54F58825DE667902FEC821108B05F89C" box="[263,313,1876,1902]" pageId="5" pageNumber="60">type</typeStatus>
|
||
locality; the difference between these patterns is more subtle in preserved than in live specimens. The first pattern, which is present in the
|
||
<typeStatus id="54F58825DE667902FD0121388904F864" box="[718,824,1916,1942]" pageId="5" pageNumber="60" type="holotype">holotype</typeStatus>
|
||
, is a uniform brown to dark brown coloration (
|
||
<figureCitation id="13752A02DE667902FAA221388A9AF84C" captionStart="FIGURE 2" captionStartId="5.[151,255,612,636]" captionTargetBox="[186,1402,197,611]" captionTargetId="figure@5.[151,1436,162,618]" captionTargetPageId="5" captionText="FIGURE 2. Type specimen and variation of X. itombwensis. (A) Holotype specimen MCZ A- 138192 (field number BJE 0275), (B) Paratype MCZ A- 138193 (field no. BJE 0276), and (C) a live unvouchered male individual. Arrows indicate dorsal spots that are not found in X. wittei. Scale bars are 5 mm. Photo credit for B: Jon Woodward." figureDoi="http://doi.org/10.5281/zenodo.182324" httpUri="https://zenodo.org/record/182324/files/figure.png" pageId="5" pageNumber="60">Fig. 2</figureCitation>
|
||
A) that is similar to
|
||
<taxonomicName id="4C4E4D04DE667902FE5821E18BC6F84C" box="[407,506,1957,1982]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="5" pageNumber="60" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE667902FE5821E18BC6F84C" box="[407,506,1957,1982]" italics="true" pageId="5" pageNumber="60">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
. The second pattern is a brown dorsal pattern with darker brown spots, sometimes with a dark dorsal band that is perpendicular to the body axis and situated caudal to the eyes but rostral with respect to the forearms (
|
||
<figureCitation id="13752A02DE6A790EFE21248B8809FD1B" box="[494,565,719,745]" captionStart="FIGURE 2" captionStartId="5.[151,255,612,636]" captionTargetBox="[186,1402,197,611]" captionTargetId="figure@5.[151,1436,162,618]" captionTargetPageId="5" captionText="FIGURE 2. Type specimen and variation of X. itombwensis. (A) Holotype specimen MCZ A- 138192 (field number BJE 0275), (B) Paratype MCZ A- 138193 (field no. BJE 0276), and (C) a live unvouchered male individual. Arrows indicate dorsal spots that are not found in X. wittei. Scale bars are 5 mm. Photo credit for B: Jon Woodward." figureDoi="http://doi.org/10.5281/zenodo.182324" httpUri="https://zenodo.org/record/182324/files/figure.png" pageId="9" pageNumber="64">Fig. 2</figureCitation>
|
||
B, C). The second color pattern is evident in three of the
|
||
<typeStatus id="54F58825DE6A790EFB1E248B8F7DFD1B" box="[1233,1345,719,745]" pageId="9" pageNumber="64" type="paratype">paratypes</typeStatus>
|
||
including one adult male: MCZ A-138194 (BJE 0277), and both of the juveniles: MCZ A-138193 (BJE 0276) and MCZ A-138195 (BJE 0278). In life, both the gular region and belly are cream colored and the leg and inguinal region are yellowish; sometimes there are small brown spots on the leg and inguinal region (
|
||
<figureCitation id="13752A02DE6A790EFB1E25038F24FC93" box="[1233,1304,839,865]" captionStart="FIGURE 2" captionStartId="5.[151,255,612,636]" captionTargetBox="[186,1402,197,611]" captionTargetId="figure@5.[151,1436,162,618]" captionTargetPageId="5" captionText="FIGURE 2. Type specimen and variation of X. itombwensis. (A) Holotype specimen MCZ A- 138192 (field number BJE 0275), (B) Paratype MCZ A- 138193 (field no. BJE 0276), and (C) a live unvouchered male individual. Arrows indicate dorsal spots that are not found in X. wittei. Scale bars are 5 mm. Photo credit for B: Jon Woodward." figureDoi="http://doi.org/10.5281/zenodo.182324" httpUri="https://zenodo.org/record/182324/files/figure.png" pageId="9" pageNumber="64">Fig. 2</figureCitation>
|
||
C). Coloration at the margins of the lower jaw tends to be slightly darker than the venter, and varies from a dull gray under the head to a thin gray/brown line near the mouth. The barcode sequence of one of the
|
||
<typeStatus id="54F58825DE6A790EFAE325D38FA1FC43" box="[1324,1437,919,945]" pageId="9" pageNumber="64" type="paratype">paratypes</typeStatus>
|
||
(museum accession number MCZ A-138193, field number BJE 0 276, barcode accession number
|
||
<accessionNumber id="941DAB64DE6A790EFADC25FB8FA9FC2B" box="[1299,1429,959,985]" httpUri="https://www.ebi.ac.uk/ena/browser/api/embl/EU566832" pageId="9" pageNumber="64">EU566832</accessionNumber>
|
||
) is the same as the
|
||
<typeStatus id="54F58825DE6A790EFEA425A38BE9FBF3" box="[363,469,999,1025]" pageId="9" pageNumber="64" type="holotype">holotype</typeStatus>
|
||
.
|
||
</paragraph>
|
||
<caption id="DF31660FDE647900FF5821D78AE1F802" ID-DOI="http://doi.org/10.5281/zenodo.182325" httpUri="https://zenodo.org/record/182325/files/figure.png" pageId="7" pageNumber="62" targetBox="[420,1151,214,1938]" targetPageId="7">
|
||
<paragraph id="8BF13687DE647900FF5821D78AE1F802" blockId="7.[151,1436,1939,2032]" pageId="7" pageNumber="62">
|
||
<emphasis id="B93AEA95DE647900FF5821D78B24F859" bold="true" box="[151,280,1939,1963]" pageId="7" pageNumber="62">FIGURE 3.</emphasis>
|
||
Male advertisement vocalization of (A)
|
||
<taxonomicName id="4C4E4D04DE647900FD0521D7891EF858" box="[714,802,1939,1962]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="7" pageNumber="62" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE647900FD0521D7891EF858" box="[714,802,1939,1962]" italics="true" pageId="7" pageNumber="62">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
, (B)
|
||
<taxonomicName id="4C4E4D04DE647900FC9521D789FBF858" box="[858,967,1939,1962]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="7" pageNumber="62" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE647900FC9521D789FBF858" box="[858,967,1939,1962]" italics="true" pageId="7" pageNumber="62">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
, and (C)
|
||
<taxonomicName id="4C4E4D04DE647900FBE421D78EF0F858" box="[1067,1228,1939,1962]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="7" pageNumber="62" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE647900FBE421D78EF0F858" box="[1067,1228,1939,1962]" italics="true" pageId="7" pageNumber="62">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
. The slow trill portion of the
|
||
<taxonomicName id="4C4E4D04DE647900FEDD21F28B8BF83F" box="[274,439,1974,1997]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="7" pageNumber="62" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE647900FEDD21F28B8BF83F" box="[274,439,1974,1997]" italics="true" pageId="7" pageNumber="62">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
call (beginning at about 400 milliseconds) is a unique feature within the “vestitus-wittei” group.
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF31660FDE6B790FFF5820FE8EEEF814" ID-DOI="http://doi.org/10.5281/zenodo.182326" httpUri="https://zenodo.org/record/182326/files/figure.png" pageId="8" pageNumber="63" targetBox="[200,1372,186,1705]" targetPageId="8">
|
||
<paragraph id="8BF13687DE6B790FFF5820FE8EEEF814" blockId="8.[151,1436,1722,2022]" pageId="8" pageNumber="63">
|
||
<emphasis id="B93AEA95DE6B790FFF5820FE8B24F920" bold="true" box="[151,280,1722,1746]" pageId="8" pageNumber="63">FIGURE 4.</emphasis>
|
||
Evolutionary relationships of (A) combined data from two tightly linked nuclear loci (
|
||
<emphasis id="B93AEA95DE6B790FFB7E20FE8ECEF923" box="[1201,1266,1722,1745]" italics="true" pageId="8" pageNumber="63">RAG1</emphasis>
|
||
and
|
||
<emphasis id="B93AEA95DE6B790FFAEA20FE8F5AF923" box="[1317,1382,1722,1745]" italics="true" pageId="8" pageNumber="63">RAG2</emphasis>
|
||
) and (B) mitochondrial DNA illustrate a divergent but sister relationship of
|
||
<taxonomicName id="4C4E4D04DE6B790FFC4520998E10F906" box="[906,1068,1757,1780]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="8" pageNumber="63" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE6B790FFC4520998E10F906" box="[906,1068,1757,1780]" italics="true" pageId="8" pageNumber="63">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE6B790FFBAF20998E85F906" box="[1120,1209,1757,1780]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="8" pageNumber="63" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE6B790FFBAF20998E85F906" box="[1120,1209,1757,1780]" italics="true" pageId="8" pageNumber="63">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
. Nuclear loci but not mitochondrial loci illustrate a close relationship between (
|
||
<taxonomicName id="4C4E4D04DE6B790FFCCF2144899DF8E5" box="[768,929,1792,1815]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="8" pageNumber="63" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE6B790FFCCF2144899DF8E5" box="[768,929,1792,1815]" italics="true" pageId="8" pageNumber="63">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
+
|
||
<taxonomicName id="4C4E4D04DE6B790FFC7221448E29F8E5" box="[957,1045,1792,1815]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="8" pageNumber="63" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE6B790FFC7221448E29F8E5" box="[957,1045,1792,1815]" italics="true" pageId="8" pageNumber="63">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
) to half of the allopolyploid genome of
|
||
<taxonomicName id="4C4E4D04DE6B790FFF7C21668B1CF8CB" box="[179,288,1826,1849]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="8" pageNumber="63" phylum="Chordata" rank="species" species="vestitus">
|
||
<emphasis id="B93AEA95DE6B790FFF7C21668B1CF8CB" box="[179,288,1826,1849]" italics="true" pageId="8" pageNumber="63">X. vestitus</emphasis>
|
||
</taxonomicName>
|
||
. For clarity most posterior probabilities are omitted because they are similar or identical to those found elsewhere (
|
||
<bibRefCitation id="EFDF4B76DE6B790FFF2A21018B5BF8AE" author="Evans, B. J." box="[229,359,1861,1884]" firstAuthor="Evans" journalOrPublisher="Genetics" pageId="8" pageNumber="63" pagination="1119 - 1130" part="176" refId="ref5963" refString="Evans, B. J. (2007) Ancestry influences the fate of duplicated genes millions of years after duplication in allopolyploid clawed frogs (Xenopus). Genetics, 176, 1119 - 1130." title="Ancestry influences the fate of duplicated genes millions of years after duplication in allopolyploid clawed frogs (Xenopus)" type="journal article" year="2007">Evans 2007</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFDF4B76DE6B790FFEBE21018810F8AE" author="Evans, B. J. & Kelley, D. B. & Tinsley, R. C. & Melnick, D. J. & Cannatella, D. C." box="[369,556,1861,1884]" etAl="et al." firstAuthor="Evans" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="8" pageNumber="63" pagination="197 - 213" part="33" refId="ref6080" refString="Evans, B. J., Kelley, D. B., Tinsley, R. C., Melnick, D. J. & Cannatella, D. C. (2004) A mitochondrial DNA phylogeny of clawed frogs: phylogeography on sub-Saharan Africa and implications for polyploid evolution. Molecular Phylogenetics and Evolution, 33, 197 - 213." title="A mitochondrial DNA phylogeny of clawed frogs: phylogeography on sub-Saharan Africa and implications for polyploid evolution" type="journal article" year="2004">Evans et al. 2004</bibRefCitation>
|
||
). However, with reference to
|
||
<taxonomicName id="4C4E4D04DE6B790FFCAB21018E39F8AE" box="[868,1029,1861,1884]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="8" pageNumber="63" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE6B790FFCAB21018E39F8AE" box="[868,1029,1861,1884]" italics="true" pageId="8" pageNumber="63">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
, in (A and B) the red clades have over 95% posterior probability and the blue clade has over 80% posterior probability. (C) A species phylogeny illustrating bifurcating and reticulating evolutionary relationships in clawed frogs. The most recent common ancestor of
|
||
<taxonomicName id="4C4E4D04DE6B790FFAD821CE8F53F853" box="[1303,1391,1930,1953]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="8" pageNumber="63" phylum="Chordata" rank="species" species="wittei">
|
||
<emphasis id="B93AEA95DE6B790FFAD821CE8F53F853" box="[1303,1391,1930,1953]" italics="true" pageId="8" pageNumber="63">X. wittei</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C4E4D04DE6B790FFF5821E98B04F836" box="[151,312,1965,1988]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="8" pageNumber="63" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE6B790FFF5821E98B04F836" box="[151,312,1965,1988]" italics="true" pageId="8" pageNumber="63">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
evolved through allopolyploidization of two tetraploid species. The number of chromosomes in each species is indicated in parentheses after each species name. (A) and (C) are modified from (
|
||
<bibRefCitation id="EFDF4B76DE6B790FFB8C218B8EFAF814" author="Evans, B. J." box="[1091,1222,1999,2022]" firstAuthor="Evans" journalOrPublisher="Genetics" pageId="8" pageNumber="63" pagination="1119 - 1130" part="176" refId="ref5963" refString="Evans, B. J. (2007) Ancestry influences the fate of duplicated genes millions of years after duplication in allopolyploid clawed frogs (Xenopus). Genetics, 176, 1119 - 1130." title="Ancestry influences the fate of duplicated genes millions of years after duplication in allopolyploid clawed frogs (Xenopus)" type="journal article" year="2007">Evans 2007</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</caption>
|
||
<caption id="DF31660FDE6A790EFF58241889A3FD81" box="[151,927,604,628]" pageId="9" pageNumber="64" startId="9.[151,255,604,628]" targetBox="[521,1065,243,559]" targetPageId="9">
|
||
<paragraph id="8BF13687DE6A790EFF58241889A3FD81" blockId="9.[151,927,604,628]" box="[151,927,604,628]" pageId="9" pageNumber="64">
|
||
<emphasis id="B93AEA95DE6A790EFF5824188B26FD86" bold="true" box="[151,282,604,628]" pageId="9" pageNumber="64">FIGURE 5.</emphasis>
|
||
Karyotype of
|
||
<taxonomicName id="4C4E4D04DE6A790EFE7B24188869FD81" box="[436,597,604,627]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="9" pageNumber="64" phylum="Chordata" rank="species" species="itombwensis">
|
||
<emphasis id="B93AEA95DE6A790EFE7B24188869FD81" box="[436,597,604,627]" italics="true" pageId="9" pageNumber="64">X. itombwensis</emphasis>
|
||
</taxonomicName>
|
||
illustrating octoploidy, 8x=72.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8BF13687DE6A790EFF09224B8EFFFBA3" blockId="9.[151,1437,719,1345]" pageId="9" pageNumber="64">
|
||
<emphasis id="B93AEA95DE6A790EFF09224B8BA6FBDB" bold="true" box="[198,410,1039,1065]" italics="true" pageId="9" pageNumber="64">Size dimorphism:</emphasis>
|
||
Females are larger than males; we suspect that the females we measured are not fully grown and that size dimorphism is greater than our measurements would suggest (Table 1).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C354650CDE6A790EFF09221B8B24FB03" pageId="9" pageNumber="64" type="biology_ecology">
|
||
<paragraph id="8BF13687DE6A790EFF09221B8B24FB03" blockId="9.[151,1437,719,1345]" pageId="9" pageNumber="64">
|
||
<emphasis id="B93AEA95DE6A790EFF09221B893FFB8A" box="[198,771,1119,1145]" italics="true" pageId="9" pageNumber="64">
|
||
<emphasis id="B93AEA95DE6A790EFF09221B8BC0FB8B" bold="true" box="[198,508,1119,1145]" italics="true" pageId="9" pageNumber="64">Ecology and distribution:</emphasis>
|
||
<taxonomicName id="4C4E4D04DE6A790EFDCB221B893FFB8A" box="[516,771,1119,1144]" class="Amphibia" family="Pipidae" genus="Xenopus" higherTaxonomySource="CoL" kingdom="Animalia" order="Anura" pageId="9" pageNumber="64" phylum="Chordata" rank="species" species="itombwensis">Xenopus itombwensis</taxonomicName>
|
||
</emphasis>
|
||
was collected only at the
|
||
<typeStatus id="54F58825DE6A790EFBF6221B8E50FB8B" box="[1081,1132,1119,1145]" pageId="9" pageNumber="64">type</typeStatus>
|
||
locality and the extent of its distribution is unknown beyond this locality. These animals were locally abundant in standing water associated with mineral extraction in a region that was surrounded by mature forest and also mixed use agricultural areas.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection id="C354650CDE6A790EFF0922BB88CDFAB3" pageId="9" pageNumber="64" type="etymology">
|
||
<paragraph id="8BF13687DE6A790EFF0922BB88CDFAB3" blockId="9.[151,1437,719,1345]" pageId="9" pageNumber="64">
|
||
<emphasis id="B93AEA95DE6A790EFF0922BB8B6CFAEB" bold="true" box="[198,336,1279,1305]" italics="true" pageId="9" pageNumber="64">Etymology:</emphasis>
|
||
The new species is named after the plateau where it occurs – the Itombwe Massif of South Kivu Province,
|
||
<collectingCountry id="F3597617DE6A790EFE80236388D1FAB3" box="[335,749,1319,1345]" name="Democratic Republic of the Congo" pageId="9" pageNumber="64">Democratic Republic of the Congo</collectingCountry>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |