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<document id="4B651136DBAA0A2F2D0129C68F34058C" ID-DOI="10.5281/zenodo.190845" ID-GBIF-Dataset="4de87e60-ade2-400c-9f1e-9a07122ea13e" ID-ISSN="1175-5326" ID-Zenodo-Dep="190845" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" checkinTime="1461120409804" checkinUser="plazi" docAuthor="Schories, Susanne, Meyer, Manfred K. &amp; Schartl, Manfred" docDate="2009" docId="03EE6834FF968B6EEAF4D3DEFC54404E" docLanguage="en" docName="zt02266p050.pdf" docOrigin="Zootaxa 2266" docStyle="DocumentStyle:890A69B780ED73D6DB8551B71C8AC79E.4:Zootaxa.2009-2012.journal_article" docStyleId="890A69B780ED73D6DB8551B71C8AC79E" docStyleName="Zootaxa.2009-2012.journal_article" docStyleVersion="4" docTitle="Poecilia (Acanthophacelus) obscura Schories, Meyer &amp; Schartl, 2009, new species" docType="treatment" docVersion="11" lastPageNumber="48" masterDocId="FFD7104CFF928B60EA63D535FFC4413D" masterDocTitle="Description of Poecilia (Acanthophacelus) obscura n. sp., (Teleostei: Poeciliidae), a new guppy species from western Trinidad, with remarks on P. w i n g e i and the status of the “ Endlers guppy ”" masterLastPageNumber="50" masterPageNumber="35" pageNumber="39" updateTime="1698595850771" updateUser="plazi">
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<mods:title id="C3E79233FF6C18287BEEE2DF0338BF8E">Description of Poecilia (Acanthophacelus) obscura n. sp., (Teleostei: Poeciliidae), a new guppy species from western Trinidad, with remarks on P. w i n g e i and the status of the “ Endlers guppy ”</mods:title>
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<mods:namePart id="4F7772CC86EE98021C04C427E9BC2679">Schories, Susanne</mods:namePart>
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<mods:namePart id="44AD51DE474232DB4D14D3F24FA07387">Meyer, Manfred K.</mods:namePart>
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<mods:namePart id="2FD21571B9569A3C283B6612C6787472">Schartl, Manfred</mods:namePart>
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<mods:date id="ADAD086DB78806B50ED05F13FEAE2A21">2009</mods:date>
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<treatment id="03EE6834FF968B6EEAF4D3DEFC54404E" ID-DOI="http://doi.org/10.5281/zenodo.5691795" ID-GBIF-Taxon="119625637" ID-Zenodo-Dep="5691795" LSID="urn:lsid:plazi:treatment:03EE6834FF968B6EEAF4D3DEFC54404E" httpUri="http://treatment.plazi.org/id/03EE6834FF968B6EEAF4D3DEFC54404E" lastPageId="14" lastPageNumber="48" pageId="4" pageNumber="39">
<subSubSection id="C35D8AA9FF968B64EAF4D3DEFE324615" pageId="4" pageNumber="39" type="nomenclature">
<paragraph id="8BF8D922FF968B64EAF4D3DEFD2F4638" blockId="4.[151,747,1771,1832]" box="[151,747,1771,1797]" pageId="4" pageNumber="39">
<heading id="D0B06E4EFF968B64EAF4D3DEFD2F4638" bold="true" box="[151,747,1771,1797]" fontSize="11" level="1" pageId="4" pageNumber="39" reason="1">
<emphasis id="B9330530FF968B64EAF4D3DEFD2F4638" bold="true" box="[151,747,1771,1797]" pageId="4" pageNumber="39">
<taxonomicName id="4C47A2A1FF968B64EAF4D3DEFD894638" ID-CoL="77T5M" box="[151,589,1771,1797]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="4" pageNumber="39" phylum="Chordata" rank="species" species="obscura" status="sp. nov." subGenus="Acanthophacelus">
<emphasis id="B9330530FF968B64EAF4D3DEFD894638" bold="true" box="[151,589,1771,1797]" italics="true" pageId="4" pageNumber="39">Poecilia (Acanthophacelus) obscura</emphasis>
</taxonomicName>
,
<taxonomicNameLabel id="A200B84BFF968B64E838D3DEFD2F4638" box="[603,747,1771,1797]" pageId="4" pageNumber="39" rank="species">new species</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph id="8BF8D922FF968B64EAF4D23BFE324615" blockId="4.[151,747,1771,1832]" box="[151,502,1806,1832]" pageId="4" pageNumber="39">
(
<figureCitation id="137CC5A7FF968B64EAFCD23BFF354615" box="[159,241,1806,1832]" captionStart="FIGURE 2. P" captionStartId="5.[151,255,1910,1934]" captionTargetBox="[187,1405,533,1892]" captionTargetId="figure@5.[187,1409,533,1892]" captionTargetPageId="5" captionText="FIGURE 2. P. obscura male (a) and female (b) from locus typicus, Oropuche River (OR). (c) P. obscura, population Rio Seco (RS); (d) P, reticulata, population Caroni Swamp (CS); (e) P. reticulata, population Rio Yaguaracual (RY); (f) P. c f “ wingei ” from El Cordon waterfall (EC); (g) P. w i n g e i from locus typicus, population Campoma (Ca); (h) P. w i n g e i, population Laguna de los Patos (LP)." httpUri="https://zenodo.org/record/190847/files/figure.png" pageId="4" pageNumber="39">Figs. 2</figureCitation>
ac, 3, 4, 68, table 2)
</paragraph>
</subSubSection>
<subSubSection id="C35D8AA9FF968B65EAF4D26EFAC3403F" lastPageId="5" lastPageNumber="40" pageId="4" pageNumber="39" type="materials_examined">
<paragraph id="8BF8D922FF968B64EAF4D26EFB4D46F8" blockId="4.[151,1436,1883,1989]" pageId="4" pageNumber="39">
<typeStatus id="54FC6780FF968B64EAF4D26EFECC4648" box="[151,264,1883,1909]" pageId="4" pageNumber="39" type="holotype">Holotype</typeStatus>
. Male (
<collectionCode id="ED5641E7FF968B64EB3DD26EFE654648" box="[350,417,1883,1909]" httpUri="http://grbio.org/cool/4j3t-w40g" name="Museum of Zoology Senckenberg Dresden" pageId="4" pageNumber="39">MTD</collectionCode>
F31503),
<quantity id="4CBF74C7FF968B64E879D26EFD474648" box="[538,643,1883,1909]" metricMagnitude="-2" metricUnit="m" metricValue="1.51" pageId="4" pageNumber="39" unit="mm" value="15.1">15.1 mm</quantity>
SL, for other measures see table 2; Oropuche River (
<geoCoordinate id="EE73BFE5FF968B64EE90D26EFA5F4648" box="[1267,1435,1883,1909]" direction="north" orientation="latitude" pageId="4" pageNumber="39" precision="1" value="10.717533">N 10° 43.052</geoCoordinate>
<geoCoordinate id="EE73BFE5FF968B64EAF4D2B7FEFD46A1" box="[151,313,1922,1948]" direction="west" orientation="longitude" pageId="4" pageNumber="39" precision="1" value="-61.14785">W 61° 8.871</geoCoordinate>
) below the bridge where the road from Cumaca to Cumaca Caves crosses the river approx
<quantity id="4CBF74C7FF968B64EF14D2B7FF7946F8" metricMagnitude="3" metricUnit="m" metricValue="2.5" pageId="4" pageNumber="39" unit="km" value="2.5">2.5 km</quantity>
past Cumaca,
<collectingCountry id="F35099B2FF968B64EB0AD29EFE1546F8" box="[361,465,1963,1989]" name="Trinidad and Tobago" pageId="4" pageNumber="39">Trinidad</collectingCountry>
;
<date id="FFF9FFE2FF968B64EBBFD29EFD7F46F8" box="[476,699,1961,1989]" pageId="4" pageNumber="39" value="2008-02-13">February 13th 2008</date>
, S. Schories, M. Schartl, P. Fischer leg.
</paragraph>
<paragraph id="8BF8D922FF978B65EAA5D5ADFAC3403F" blockId="5.[151,1436,152,490]" pageId="5" pageNumber="40">
<typeStatus id="54FC6780FF978B65EAA5D5ADFEF8418F" box="[198,316,152,178]" pageId="5" pageNumber="40" type="paratype">Paratypes</typeStatus>
.
<specimenCount id="9D4112ABFF978B65EB2FD5ADFE69418F" box="[332,429,152,178]" pageId="5" pageNumber="40" type="male">3 males</specimenCount>
,
<specimenCount id="9D4112ABFF978B65EBDDD5ADFDF7418F" box="[446,563,152,178]" pageId="5" pageNumber="40" type="female">5 females</specimenCount>
(
<collectionCode id="ED5641E7FF978B65E824D5ADFD4E418F" box="[583,650,152,178]" httpUri="http://grbio.org/cool/4j3t-w40g" name="Museum of Zoology Senckenberg Dresden" pageId="5" pageNumber="40">MTD</collectionCode>
F31504 31511); for measures see table 2; same collection as
<typeStatus id="54FC6780FF978B65EAF4D58BFEC641E5" box="[151,258,190,216]" pageId="5" pageNumber="40" type="holotype">holotype</typeStatus>
.
<specimenCount id="9D4112ABFF978B65EB6ED58BFEAF41E5" box="[269,363,190,216]" pageId="5" pageNumber="40" type="male">4 males</specimenCount>
,
<specimenCount id="9D4112ABFF978B65EB1BD58BFE2E41E5" box="[376,490,190,216]" pageId="5" pageNumber="40" type="female">2 females</specimenCount>
(
<collectionCode id="ED5641E7FF978B65EB99D58BFDF241E5" LSID="urn:lsid:biocol.org:col:34838" box="[506,566,190,216]" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34838" name="Forschungsinstitut und Natur-Museum Senckenberg" pageId="5" pageNumber="40">SMF</collectionCode>
31068); for measures see table 2; Rio Seco, below Río Seco waterfall, at junction with Salybia River,
<collectingCountry id="F35099B2FF978B65EB84D5DDFD8A403F" box="[487,590,232,258]" name="Trinidad and Tobago" pageId="5" pageNumber="40">Trinidad</collectingCountry>
,
<date id="FFF9FFE2FF978B65E839D5DDFCFD403F" box="[602,825,229,258]" pageId="5" pageNumber="40" value="2008-02-17">February 17th 2008</date>
, S. Schories, M. Schartl, P. Fischer leg.
</paragraph>
</subSubSection>
<subSubSection id="C35D8AA9FF978B66EAA5D43BFCFB41C3" lastPageId="6" lastPageNumber="41" pageId="5" pageNumber="40" type="diagnosis">
<paragraph id="8BF8D922FF978B65EAA5D43BFD0C40D7" blockId="5.[151,1436,152,490]" pageId="5" pageNumber="40">
<emphasis id="B9330530FF978B65EAA5D43BFE824015" bold="true" box="[198,326,270,296]" pageId="5" pageNumber="40">Diagnosis.</emphasis>
<taxonomicName id="4C47A2A1FF978B65EB37D43AFDE64015" box="[340,546,271,296]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" pageNumber="40" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF978B65EB37D43AFDE64015" box="[340,546,271,296]" italics="true" pageId="5" pageNumber="40">Poecilia obscura</emphasis>
</taxonomicName>
(
<figureCitation id="137CC5A7FF978B65E85AD43AFD424014" box="[569,646,271,297]" captionStart="FIGURE 2. P" captionStartId="5.[151,255,1910,1934]" captionTargetBox="[187,1405,533,1892]" captionTargetId="figure@5.[187,1409,533,1892]" captionTargetPageId="5" captionText="FIGURE 2. P. obscura male (a) and female (b) from locus typicus, Oropuche River (OR). (c) P. obscura, population Rio Seco (RS); (d) P, reticulata, population Caroni Swamp (CS); (e) P. reticulata, population Rio Yaguaracual (RY); (f) P. c f “ wingei ” from El Cordon waterfall (EC); (g) P. w i n g e i from locus typicus, population Campoma (Ca); (h) P. w i n g e i, population Laguna de los Patos (LP)." httpUri="https://zenodo.org/record/190847/files/figure.png" pageId="5" pageNumber="40">Fig. 2</figureCitation>
ac, 3, table 2) is a small poeciliid species from the subgenus
<emphasis id="B9330530FF978B65EAF4D403FB454072" box="[151,1153,310,335]" italics="true" pageId="5" pageNumber="40">
<taxonomicName id="4C47A2A1FF978B65EAF4D403FEA04072" box="[151,356,310,335]" family="Poeciliidae" genus="Acanthophacelus" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" pageNumber="40" phylum="Chordata" rank="genus">Acanthophacelus</taxonomicName>
(SL in males usually not exceeding
<quantity id="4CBF74C7FF978B65E975D403FCA34072" box="[790,871,310,335]" metricMagnitude="-2" metricUnit="m" metricValue="2.0" pageId="5" pageNumber="40" unit="mm" value="20.0">20 mm</quantity>
and in females
<quantity id="4CBF74C7FF978B65EE45D403FBBC4072" box="[1062,1144,310,335]" metricMagnitude="-2" metricUnit="m" metricValue="3.0" pageId="5" pageNumber="40" unit="mm" value="30.0">30 mm</quantity>
)
</emphasis>
, which is distinguished from all other species of the subgenus by the following characters: comb-like spines of gonopodium ray 3 not numerous (11 to 15), basal massive, vs. numerous (14 to 18), basal slender in
<taxonomicName id="4C47A2A1FF978B65EE49D4B6FB7340A1" box="[1066,1207,387,412]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" pageNumber="40" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF978B65EE49D4B6FB7340A1" box="[1066,1207,387,412]" italics="true" pageId="5" pageNumber="40">P. reticulata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C47A2A1FF978B65EE92D4B6FA9840A1" box="[1265,1372,387,412]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" pageNumber="40" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF978B65EE92D4B6FA9840A1" box="[1265,1372,387,412]" italics="true" pageId="5" pageNumber="40">P. wingei</emphasis>
</taxonomicName>
(Fig. 3). Being genetically divergent from the other species of the subgenus, the definition of the new species is mainly based on evidence from DNA sequence.
</paragraph>
<caption id="DF3889AAFF978B65EAF4D243FDE046CB" httpUri="https://zenodo.org/record/190847/files/figure.png" pageId="5" pageNumber="40" targetBox="[187,1405,533,1892]" targetPageId="5">
<paragraph id="8BF8D922FF978B65EAF4D243FDE046CB" blockId="5.[151,1436,1910,2038]" pageId="5" pageNumber="40">
<emphasis id="B9330530FF978B65EAF4D243FEDC46B3" bold="true" box="[151,280,1910,1934]" pageId="5" pageNumber="40">FIGURE 2.</emphasis>
<taxonomicName id="4C47A2A1FF978B65EB7CD242FE4B46B3" box="[287,399,1911,1934]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" pageNumber="40" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF978B65EB7CD242FE4B46B3" box="[287,399,1911,1934]" italics="true" pageId="5" pageNumber="40">P. obscura</emphasis>
</taxonomicName>
male (a) and female (b) from locus typicus, Oropuche River (OR). (c)
<taxonomicName id="4C47A2A1FF978B65EEE3D242FB3446B3" box="[1152,1264,1911,1934]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" pageNumber="40" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF978B65EEE3D242FB3446B3" box="[1152,1264,1911,1934]" italics="true" pageId="5" pageNumber="40">P. obscura</emphasis>
</taxonomicName>
, population Rio Seco (RS); (d)
<emphasis id="B9330530FF978B65EB5BD2AFFE7C468C" box="[312,440,1946,1969]" italics="true" pageId="5" pageNumber="40">
P,
<taxonomicName id="4C47A2A1FF978B65EB31D2AFFE7C468C" box="[338,440,1946,1969]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" pageNumber="40" phylum="Chordata" rank="species" species="reticulata">reticulata</taxonomicName>
</emphasis>
, population Caroni Swamp (CS); (e)
<taxonomicName id="4C47A2A1FF978B65E929D2AFFC0E468C" box="[842,970,1946,1969]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" pageNumber="40" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF978B65E929D2AFFC0E468C" box="[842,970,1946,1969]" italics="true" pageId="5" pageNumber="40">P. reticulata</emphasis>
</taxonomicName>
, population Rio Yaguaracual (RY); (f)
<emphasis id="B9330530FF978B65EF0DD2AFFF3946EE" italics="true" pageId="5" pageNumber="40">
P. c f “
<taxonomicName id="4C47A2A1FF978B65EAFCD289FF3146EE" box="[159,245,1980,2003]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="5" pageNumber="40" phylum="Chordata" rank="species" species="wingei">wingei</taxonomicName>
</emphasis>
from El Cordon waterfall (EC); (g)
<emphasis id="B9330530FF978B65E8ECD289FD3446EE" box="[655,752,1980,2003]" italics="true" pageId="5" pageNumber="40">P. w i n g e i</emphasis>
from locus typicus, population Campoma (Ca); (h)
<emphasis id="B9330530FF978B65EF52D289FA5546EE" box="[1329,1425,1980,2003]" italics="true" pageId="5" pageNumber="40">P. w i n g e i</emphasis>
, population Laguna de los Patos (LP).
</paragraph>
</caption>
<paragraph id="8BF8D922FF948B66EAA5D5ADFCFB41C3" blockId="6.[151,1437,152,1261]" pageId="6" pageNumber="41">
<taxonomicName id="4C47A2A1FF948B66EAA5D5ADFE81418C" box="[198,325,152,177]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" pageNumber="41" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF948B66EAA5D5ADFE81418C" box="[198,325,152,177]" italics="true" pageId="6" pageNumber="41">P. obscura</emphasis>
</taxonomicName>
is also recognized by morphometrics: caudal peduncle short in females (SL/CPD = 4.8 to 6.15), vs. long (SL/CPD = 6.05 to 7.4) in
<taxonomicName id="4C47A2A1FF948B66E8E1D58AFCD441E5" box="[642,784,191,216]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" pageNumber="41" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF948B66E8E1D58AFCD441E5" box="[642,784,191,216]" italics="true" pageId="6" pageNumber="41">P. reticulata</emphasis>
</taxonomicName>
and
<emphasis id="B9330530FF948B66E92AD58AFC7441E5" box="[841,944,191,216]" italics="true" pageId="6" pageNumber="41">P. w i n g e i</emphasis>
(SL/CPD = 6.5 to 7.3) (
<figureCitation id="137CC5A7FF948B66EEB1D58BFADE41E5" box="[1234,1306,190,216]" captionStart="FIGURE 4" captionStartId="8.[151,255,974,998]" captionTargetBox="[431,1115,204,925]" captionTargetId="figure@8.[415,1171,194,950]" captionTargetPageId="8" captionText="FIGURE 4. CPD / SL / CPD ratios of P. obscura (symbols, population Rio Seco (RS), population Oropuche (OR )) and P. reticulata females (symbols, population Claxton Bay (CB) population Caroni Swamp (CS ))." httpUri="https://zenodo.org/record/190848/files/figure.png" pageId="6" pageNumber="41">Fig. 4</figureCitation>
); usually 6 dorsal fin rays, vs. usually
<quantity id="4CBF74C7FF948B66EBB2D5D0FE3B41C2" box="[465,511,229,255]" metricMagnitude="-1" metricUnit="m" metricValue="1.778" pageId="6" pageNumber="41" unit="in" value="7.0">7 in</quantity>
<taxonomicName id="4C47A2A1FF948B66E865D5D0FD5041C3" box="[518,660,229,254]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" pageNumber="41" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF948B66E865D5D0FD5041C3" box="[518,660,229,254]" italics="true" pageId="6" pageNumber="41">P. reticulata</emphasis>
</taxonomicName>
and
<emphasis id="B9330530FF948B66E8AFD5D0FCFB41C3" box="[716,831,229,254]" italics="true" pageId="6" pageNumber="41">
<taxonomicName id="4C47A2A1FF948B66E8AFD5D0FCFF41C3" box="[716,827,229,254]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" pageNumber="41" phylum="Chordata" rank="species" species="wingei">P. wingei</taxonomicName>
.
</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C35D8AA9FF948B67EAA5D43EFCA1442E" lastPageId="7" lastPageNumber="42" pageId="6" pageNumber="41" type="description">
<paragraph id="8BF8D922FF948B66EAA5D43EFCBB404E" blockId="6.[151,1437,152,1261]" pageId="6" pageNumber="41">
<emphasis id="B9330530FF948B66EAA5D43EFE9B4018" bold="true" box="[198,351,267,293]" pageId="6" pageNumber="41">Description.</emphasis>
Body deep in females, head long and not sharply pointed, 25.526.5% of SL. Longitudinal scale series 24 to 25 (rarely 24); predorsal scale series 12 to 13; scale series around caudal peduncle 14. Number of vertebrae 25 to 27. Gill rakers on first arch 11 to 12.
</paragraph>
<paragraph id="8BF8D922FF948B66EAA6D4B5FE7C43BF" blockId="6.[151,1437,152,1261]" pageId="6" pageNumber="41">Teeth of upper and lower jaws unicuspid and recurved; those of outer row enlarged and spatula-shaped, numerous and widely spaced, the much smaller posterior teeth have a single sharp spine. Upper pharyngeal bones balloon shaped. Teeth of the medial region somewhat enlarged, each side with a series of 8 rows, teeth small and conical. Lower pharyngeal bone (ceratobranchial 5) heart-shaped and with very numerous pointed and slender unicuspid teeth, 17 to 19 on posterior rows, 5 to 6 on middle rows. Teeth of medial region somewhat enlarged. Arms of pharyngeal very short and split at the ends. Ceratobranchial 4 without teeth, hypobranchial 4 present.</paragraph>
<paragraph id="8BF8D922FF948B66EAA6D7BBFA5F43F2" blockId="6.[151,1437,152,1261]" pageId="6" pageNumber="41">Supraorbital canal system well developed, section 1 to 2a and 2b to 4a, usually represented as grooves, 6b to 7 as pits; preopercular canal with 6 pores; preorbital canal with 4 pores (sometimes represented as groove).</paragraph>
<paragraph id="8BF8D922FF948B66EAA6D7E9FA9342AD" blockId="6.[151,1437,152,1261]" pageId="6" pageNumber="41">Gonopodium (Fig. 3) short and compact, 2.9 to 3.1 times in SL; ray 3 broadly expanded, curved distally and with widely spaced, 11 to 13 comb-like spines not terminating in a hook; ray 4a curved distally, with a small serra, well developed broad and long segments; ray 4p distally somewhat curved and with massive developed serrae, proximal serrae with 3 to 5 thorns, rays 5a and 5p straight, distal with a large recurved boomerang-shaped bony hook. Rays 6 and 7 long and somewhat thickened distally and curved dorsally.</paragraph>
<paragraph id="8BF8D922FF948B66EAA6D6A8FDA14516" blockId="6.[151,1437,152,1261]" pageId="6" pageNumber="41">
Gonopodial suspensorium with two well developed gonapophyses, gonapophyses I and II long and slender, each with an uncinus. Ligastyle absent or not well developed. Gonactinost 1 with a small inferior wing-like appendage, gonactinostal complex
<quantity id="4CBF74C7FF948B66E8CED6DFFCCB4539" box="[685,783,1002,1028]" metricMagnitude="-2" metricUnit="m" metricValue="7.62" metricValueMax="10.16" metricValueMin="5.08" pageId="6" pageNumber="41" unit="in" value="3.0" valueMax="4.0" valueMin="2.0">2 to 4 in</quantity>
front with a superior small lateral wing, gonactinosts 5 to 9 without bony plates or outgrowths.
</paragraph>
<paragraph id="8BF8D922FF948B66EAA6D10DFEF745D0" blockId="6.[151,1437,152,1261]" pageId="6" pageNumber="41">Small dorsal fin with 6 to 7 rays (first ray simple, all others branched), origin of dorsal fin posterior to the insertion of anal fin; caudal fin with 22 to 23 rays (1013 rays branched); anal fin 9 rays (6 rays branched); pectoral fin with 14 rays (9 rays branched); ventral fin with 6 rays (first and last ray simple, all others branched), in females not reaching to the anal fin base and in males reaching widely over the base of the gonopodium.</paragraph>
<paragraph id="8BF8D922FF948B67EAF4D2F3FEF04309" blockId="6.[151,573,1990,2014]" lastBlockId="7.[151,1437,152,564]" lastPageId="7" lastPageNumber="42" pageId="6" pageNumber="41">
<emphasis id="B9330530FF948B66EAF4D2F3FEDD46E3" bold="true" box="[151,281,1990,2014]" pageId="6" pageNumber="41">FIGURE 3.</emphasis>
Gonopodium of
<emphasis id="B9330530FF948B66EBB3D2F2FDF946E3" box="[464,573,1991,2014]" italics="true" pageId="6" pageNumber="41">P. o b s c u r a</emphasis>
Males and females with sex specific coloration (
<figureCitation id="137CC5A7FF958B67E899D5ADFC84418F" box="[762,832,152,178]" captionStart="FIGURE 2. P" captionStartId="5.[151,255,1910,1934]" captionTargetBox="[187,1405,533,1892]" captionTargetId="figure@5.[187,1409,533,1892]" captionTargetPageId="5" captionText="FIGURE 2. P. obscura male (a) and female (b) from locus typicus, Oropuche River (OR). (c) P. obscura, population Rio Seco (RS); (d) P, reticulata, population Caroni Swamp (CS); (e) P. reticulata, population Rio Yaguaracual (RY); (f) P. c f “ wingei ” from El Cordon waterfall (EC); (g) P. w i n g e i from locus typicus, population Campoma (Ca); (h) P. w i n g e i, population Laguna de los Patos (LP)." httpUri="https://zenodo.org/record/190847/files/figure.png" pageId="7" pageNumber="42">Fig. 2</figureCitation>
a,c). Body color of adult females grayish brown to yellow, fins hyaline; body sides of adult males with red, blue, orange and yellow bright pigment spots, some reflecting iridescent, usually with 1 to 3 rounded black spots, sometimes with a series of irregularly thin and short or long brown or light black horizontal lines or with very short brown vertical lines sometimes crossing the horizontal bars; the caudal fin base often shows a lower or upper black spot surrounded by small dark and short dashes and yellow pigment, dorsal or ventral caudal fin rays sometimes pigmented and rarely elongated over the caudal margin of the fin, forming a short “sword”; dorsal fin often whitish, dark or polychromatic colored, sometimes flag-like elongated, all other fins hyaline. Male body coloration extremely polymorphic: in natural habitats no two males being alike. Body coloration and caudal appendix phenotype predominantly heritable characters, male offspring of a single male in laboratory crosses being very similar to their father and to each other.
</paragraph>
<caption id="DF3889AAFF958B67EAF4D756FBCE4346" box="[151,1034,611,635]" pageId="7" pageNumber="42">
<paragraph id="8BF8D922FF958B67EAF4D756FBCE4346" blockId="7.[151,1421,611,1243]" box="[151,1034,611,635]" pageId="7" pageNumber="42">
<emphasis id="B9330530FF958B67EAF4D756FECC4346" bold="true" box="[151,264,611,635]" pageId="7" pageNumber="42">TABLE 2.</emphasis>
Measurements of holotype and paratypes of
<taxonomicName id="4C47A2A1FF958B67E889D751FC0C4346" box="[746,968,612,635]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="7" pageNumber="42" phylum="Chordata" rank="species" species="obscura" status="sp. nov." subGenus="Acanthophacelus">
<emphasis id="B9330530FF958B67E889D751FC0C4346" box="[746,968,612,635]" italics="true" pageId="7" pageNumber="42">Poecila (A.) obscura</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A200B84BFF958B67E9ADD756FBCE4346" box="[974,1034,611,635]" pageId="7" pageNumber="42" rank="species">sp. n.</taxonomicNameLabel>
</paragraph>
</caption>
<paragraph id="8BF8D922FF958B67EB65D7A1FB634258" blockId="7.[151,1421,611,1243]" pageId="7" pageNumber="42"># TL SL HL SNL BD IOW GL ED APL CPL CPD PL VL PDL HT(M) MTD F 31503 21.3 15.1 4.2 1.2 4.1 1.9 4.4 1.3 4.9 2.9 3.5 9.9 PT(F) MTD F 31504 33.4 25.9 6.0 1.4 7.3 3.4 2.1 9.6 8.9 4.4 5.6 3.3 16.6 PT(F) MTD F 31505 35.8 28.3 6.2 1.5 7.7 3.6 2.2 9.9 9.3 4.7 5.6 3.4 17.1 PT(F) MTD F 31506 32.6 25.5 5.7 1.4 7.2 3.3 2.1 8.3 8.0 4.3 5.3 3.2 16.3 PT(F) MTD F 31507 28.9 21.7 5.3 1.3 5.8 2.8 1.8 3.8</paragraph>
<paragraph id="8BF8D922FF958B67EAFCD641FAB845E6" blockId="7.[151,1421,611,1243]" pageId="7" pageNumber="42">PT(M) MTD F 31508 24.4 17.8 4.6 1.3 4.8 2.3 4.6 1.5 3.7 6.4 3.3 3.7 10.6 PT(F) MTD F 31509 21.1 16.2 4.3 1.3 4.2 2.1 1.4 3.2 10.2 PT(M) MTD F 31510 21.4 15.3 4.2 1.3 4.2 2.0 4.5 1.4 2.7 4.2 3.6 9.8 PT(M) MTD F 31511 19.2 13.9 4.0 1.1 3.9 1.7 4.2 1.2 4.6 2.7 4.0 3.4 9.6 PT(M) SMF 31068 24.0 17.7 4.5 1.3 4.9 2.3 4.7 1.5 3.8 6.3 3.3 4.9 3.7 10.7 PT(F) SMF 31068 32.7 25.4 6.1 1.4 7.0 3.4 2.1 9.7 9.4 4.2 5.4 3.3 16.4 PT(F) SMF 31068 33.5 26.1 7.2 3.5 2.1 9.5 9.5 4.5 5.5 3.3 16.5 PT(M) SMF 31068 25.6 18.2 4.7 1.3 5.0 2.4 4.9 1.6 3.6 3.9 10.9 PT(M) SMF 31068 24.4 17.9 4.5 1.2 2.3 4.7 1.5 3.8 6.2 3.5 3.8 10.5 PT(M) SMF 31068 20.4 14.7 1.1 4.0 1.8 4.3 1.3 2.8 9.4</paragraph>
<paragraph id="8BF8D922FF958B67EAF4D1C9FCA1442E" blockId="7.[151,869,1276,1299]" box="[151,869,1276,1299]" pageId="7" pageNumber="42">
all values are in mm, HT
<typeStatus id="54FC6780FF958B67EBC4D1C9FDC3442E" box="[423,519,1276,1299]" pageId="7" pageNumber="42" type="holotype">holotype</typeStatus>
, PT
<typeStatus id="54FC6780FF958B67E854D1C9FD52442E" box="[567,662,1276,1299]" pageId="7" pageNumber="42" type="paratype">paratype</typeStatus>
, M male, F female.
</paragraph>
</subSubSection>
<subSubSection id="C35D8AA9FF958B67EAA6D075FC4446D1" pageId="7" pageNumber="42" type="etymology">
<paragraph id="8BF8D922FF958B67EAA6D075FE0E44BF" blockId="7.[151,1437,1344,2028]" pageId="7" pageNumber="42">
<emphasis id="B9330530FF958B67EAA6D075FE964467" bold="true" box="[197,338,1344,1370]" pageId="7" pageNumber="42">Etymology.</emphasis>
The name of the new species is derived from the latin word obscurus (hidden) because of its status as a cryptic species.
</paragraph>
<paragraph id="8BF8D922FF958B67EAA6D0BBFBBD4757" blockId="7.[151,1437,1344,2028]" pageId="7" pageNumber="42">
For the common name we propose to call
<taxonomicName id="4C47A2A1FF958B67E883D0BAFCA34495" box="[736,871,1423,1448]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="7" pageNumber="42" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF958B67E883D0BAFCA34495" box="[736,871,1423,1448]" italics="true" pageId="7" pageNumber="42">P. obscura</emphasis>
</taxonomicName>
the Oropuche guppy due to its main area of distribution. For
<taxonomicName id="4C47A2A1FF958B67EB04D080FD9944F2" authority="Poeser" authorityName="Poeser" box="[359,605,1461,1487]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="42" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF958B67EB04D080FE3844F3" box="[359,508,1461,1486]" italics="true" pageId="7" pageNumber="42">P. reticulata</emphasis>
,
<bibRefCitation id="EFD6A4D3FF958B67E86FD080FD9944F2" box="[524,605,1461,1487]" pageId="7" pageNumber="50" refString="Poeser, F. N., Kempkes, M. &amp; Isbrucker, I. J. H. (2005) Description of Poecilia (Acanthophacelus) wingei n. sp. from the Paria Peninsula, Venezuela, including notes on Acanthophacelus Eigenmann, 1907 and other subgenera of Poecilia Bloch and Schneider, 1801 (Teleostei, Cyprinodontiformes, Poeciliidae). Contributions to Zoology, 74, 97 - 115." type="journal article">Poeser</bibRefCitation>
</taxonomicName>
et al (Poeser
<emphasis id="B9330530FF958B67E964D080FC8244F3" box="[775,838,1461,1486]" italics="true" pageId="7" pageNumber="42">et al.</emphasis>
, 2005) suggested the name Common or Orinoco guppy. For
<emphasis id="B9330530FF958B67EB47D0E9FE5544C8" box="[292,401,1500,1525]" italics="true" pageId="7" pageNumber="42">P. w i n g e i</emphasis>
they proposed the name Campoma guppy; reflecting the narrow range on the Paria peninsula where they found the species. After it is now clear on the basis of our molecular analysis that the species also occurs in the Cumaná region and that it represents Endlers guppy, it makes sense to return to the priority common name of Cumaná or Endlers guppy (
<bibRefCitation id="EFD6A4D3FF958B67E976D365FBA94757" author="Alexander" box="[789,1133,1616,1642]" pageId="7" pageNumber="49" refString="Alexander, H. J. &amp; Breden, F. (2004) Sexual isolation and extreme morphological divergence in the Cumana guppy: a possible case of incipient speciation. Journal of Evolutionary Biology, 17, 1238 - 54." type="journal article" year="2004">Alexander and Breden, 2004</bibRefCitation>
).
</paragraph>
<paragraph id="8BF8D922FF958B67EAA5D343FC4446D1" blockId="7.[151,1437,1344,2028]" pageId="7" pageNumber="42">
<emphasis id="B9330530FF958B67EAA5D343FEBC47AD" bold="true" box="[198,376,1654,1680]" pageId="7" pageNumber="42">Habitat notes.</emphasis>
<taxonomicName id="4C47A2A1FF958B67EBE2D342FE3A47AD" box="[385,510,1655,1680]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="7" pageNumber="42" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF958B67EBE2D342FE3A47AD" box="[385,510,1655,1680]" italics="true" pageId="7" pageNumber="42">P. obscura</emphasis>
</taxonomicName>
was collected from fast flowing small rivers and ditches with generally clear waters. The species also occurs in downstream habitats. Thus it inhabits both so-called low and high predation sites described in the guppy literature (see
<bibRefCitation id="EFD6A4D3FF958B67E8D5D3F1FC4247E3" author="Magurran" box="[694,902,1732,1758]" pageId="7" pageNumber="50" refString="Magurran, A. E. (2005) Evolutionary Ecology - The Trinidadian Guppy. Oxford University Press, New York, 224 pp." type="book" year="2005">Magurran, 2005</bibRefCitation>
). The locus typicus (
<figureCitation id="137CC5A7FF958B67EEEFD3F1FB1C47E3" box="[1164,1240,1732,1758]" captionStart="FIGURE 5" captionStartId="8.[151,255,1875,1899]" captionText="FIGURE 5. Upper Oropuche river, locus typicus for P. obscura." pageId="7" pageNumber="42">Fig. 5</figureCitation>
) is a site in the headwaters of the Oropuche River. The river is here about 5 meters wide and fast flowing with a maximum depth of
<quantity id="4CBF74C7FF958B67EB62D224FE9A4616" box="[257,350,1809,1835]" metricMagnitude="0" metricUnit="m" metricValue="1.0" pageId="7" pageNumber="42" unit="m" value="1.0">1 meter</quantity>
. The riverbed is gravel and sand with some larger rocks in the middle. Submerse plants are absent, but the vegetation on the river banks focally reaches into the water and provides shaded sections throughout most of the day. On the
<date id="FFF9FFE2FF958B67E85AD26BFD6A4645" box="[569,686,1886,1912]" pageId="7" pageNumber="42" value="2008-02-13">13.2.2008</date>
(
<specimenCount id="9D4112ABFF958B67E8A2D26BFD2F4645" box="[705,747,1886,1912]" pageId="7" pageNumber="42" type="generic">1 p.</specimenCount>
m., air temperature 25.5°C) the following parameters were recorded: clear water, water temperature 22.5°C, hardness 510, conductivity 51 mS, pH 7, nitrate 0, nitrite 0. Accompanying fishes were
<taxonomicName id="4C47A2A1FF958B67EBB8D299FDBF46F8" box="[475,635,1964,1989]" class="Actinopterygii" family="Rivulidae" genus="Rivulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="42" phylum="Chordata" rank="species" species="hartii">
<emphasis id="B9330530FF958B67EBB8D299FDBF46F8" box="[475,635,1964,1989]" italics="true" pageId="7" pageNumber="42">Rivulus hartii</emphasis>
</taxonomicName>
,
<emphasis id="B9330530FF958B67E8EAD299FC2A46F8" box="[649,1006,1964,1989]" italics="true" pageId="7" pageNumber="42">
<taxonomicName id="4C47A2A1FF958B67E8EAD299FCF246F8" box="[649,822,1964,1989]" class="Actinopterygii" family="Characidae" genus="Astyanax" kingdom="Animalia" order="Characiformes" pageId="7" pageNumber="42" phylum="Chordata" rank="species" species="spec">Astyanax spec</taxonomicName>
., Ancistrus
<taxonomicName id="4C47A2A1FF958B67E9D9D299FC2A46F8" box="[954,1006,1964,1989]" class="Actinopterygii" family="Characidae" genus="Astyanax" kingdom="Animalia" order="Characiformes" pageId="7" pageNumber="42" phylum="Chordata" rank="species" species="spec">spec</taxonomicName>
</emphasis>
. and small cichlids. The inclusion of cichlids and characins defines the habitat as high predation site.
</paragraph>
</subSubSection>
<caption id="DF3889AAFF9A8B68EAF4D6FBFBAB4536" httpUri="https://zenodo.org/record/190848/files/figure.png" pageId="8" pageNumber="43" targetBox="[431,1115,204,925]" targetPageId="8">
<paragraph id="8BF8D922FF9A8B68EAF4D6FBFBAB4536" blockId="8.[151,1434,974,1035]" pageId="8" pageNumber="43">
<emphasis id="B9330530FF9A8B68EAF4D6FBFEDD42DB" bold="true" box="[151,281,974,998]" pageId="8" pageNumber="43">FIGURE 4.</emphasis>
CPD/SL/CPD ratios of
<taxonomicName id="4C47A2A1FF9A8B68E87DD6FBFD4B42D8" box="[542,655,974,997]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="8" pageNumber="43" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9A8B68E87DD6FBFD4B42D8" box="[542,655,974,997]" italics="true" pageId="8" pageNumber="43">P. obscura</emphasis>
</taxonomicName>
(symbols, population Rio Seco (RS), population Oropuche (OR)) and
<taxonomicName id="4C47A2A1FF9A8B68EFEBD6FBFF394536" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" pageNumber="43" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9A8B68EFEBD6FBFF394536" italics="true" pageId="8" pageNumber="43">P. reticulata</emphasis>
</taxonomicName>
females (symbols, population Claxton Bay (CB) population Caroni Swamp (CS)).
</paragraph>
</caption>
<caption id="DF3889AAFF9A8B68EAF4D266FC834657" box="[151,839,1875,1899]" pageId="8" pageNumber="43">
<paragraph id="8BF8D922FF9A8B68EAF4D266FC834657" blockId="8.[151,839,1875,1899]" box="[151,839,1875,1899]" pageId="8" pageNumber="43">
<emphasis id="B9330530FF9A8B68EAF4D266FEDD4656" bold="true" box="[151,281,1875,1899]" pageId="8" pageNumber="43">FIGURE 5.</emphasis>
Upper Oropuche river, locus typicus for
<taxonomicName id="4C47A2A1FF9A8B68E8B3D266FC844657" box="[720,832,1875,1898]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="8" pageNumber="43" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9A8B68E8B3D266FC844657" box="[720,832,1875,1898]" italics="true" pageId="8" pageNumber="43">P. obscura</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<subSubSection id="C35D8AA9FF9A8B6DEAA5D2AEFA4844BA" lastPageId="13" lastPageNumber="48" pageId="8" pageNumber="43" type="description">
<paragraph id="8BF8D922FF9A8B69EAA5D2AEFB234071" blockId="8.[151,1437,1947,2012]" lastBlockId="9.[151,1436,152,332]" lastPageId="9" lastPageNumber="44" pageId="8" pageNumber="43">
<emphasis id="B9330530FF9A8B68EAA5D2AEFCC54688" bold="true" box="[198,769,1947,1973]" pageId="8" pageNumber="43">Comparison, relationships and discussion.</emphasis>
On the basis of synapomorphies,
<taxonomicName id="4C47A2A1FF9A8B68EE8BD2A9FAB64688" box="[1256,1394,1948,1973]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="8" pageNumber="43" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9A8B68EE8BD2A9FAB64688" box="[1256,1394,1948,1973]" italics="true" pageId="8" pageNumber="43">P. obscura</emphasis>
</taxonomicName>
is unequivocally attached to the subgenus
<emphasis id="B9330530FF9A8B68E8EAD2F7FCA146E6" box="[649,869,1986,2011]" italics="true" pageId="8" pageNumber="43">
<taxonomicName id="4C47A2A1FF9A8B68E8EAD2F7FCA546E6" box="[649,865,1986,2011]" family="Poeciliidae" genus="Acanthophacelus" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" pageNumber="43" phylum="Chordata" rank="genus">Acanthophacelus</taxonomicName>
.
</emphasis>
By morphological criteria
<taxonomicName id="4C47A2A1FF9A8B68EEDBD2F7FAFE46E6" box="[1208,1338,1986,2011]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="8" pageNumber="43" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9A8B68EEDBD2F7FAFE46E6" box="[1208,1338,1986,2011]" italics="true" pageId="8" pageNumber="43">P. obscura</emphasis>
</taxonomicName>
is most closely related to
<taxonomicName id="4C47A2A1FF9B8B69EB0FD5ADFE39418C" box="[364,509,152,177]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9B8B69EB0FD5ADFE39418C" box="[364,509,152,177]" italics="true" pageId="9" pageNumber="44">P. reticulata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C47A2A1FF9B8B69E854D5ADFD60418C" box="[567,676,152,177]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9B8B69E854D5ADFD60418C" box="[567,676,152,177]" italics="true" pageId="9" pageNumber="44">P. wingei</emphasis>
</taxonomicName>
. There are several synapomorphies that unite the three species, namely: large recurved hook present on tip of gonopodium ray 5 and serrae of ray 4p and 4a well developed, long and pointed comb-like spines of gonopodium ray 3 starting very close to the tip. Male pigmentation colorful and extremely polymorphic, more brilliant, metallic and less variable in
<emphasis id="B9330530FF9B8B69EEE3D439FB294018" box="[1152,1261,268,293]" italics="true" pageId="9" pageNumber="44">P. w i n g e i</emphasis>
. We noted no difference in the morphology of the genital pore of the females from the three species (
<figureCitation id="137CC5A7FF9B8B69EEF3D407FB1C4071" box="[1168,1240,306,332]" captionStart="FIGURE 6" captionStartId="9.[151,255,784,808]" captionTargetBox="[151,1435,390,758]" captionTargetId="figure@9.[151,1436,390,759]" captionTargetPageId="9" captionText="FIGURE 6. Scanning electron microscopy images of female genital pores of (a) P. w i n g e i (Laguna de los Patos), (b) P. reticulata (Caroni Swamp), (c) P. obscura, (Upper Oropuche). Bar represents 200 mm." httpUri="https://zenodo.org/record/190849/files/figure.png" pageId="9" pageNumber="44">Fig. 6</figureCitation>
).
</paragraph>
<caption id="DF3889AAFF9B8B69EAF4D625FBF14277" httpUri="https://zenodo.org/record/190849/files/figure.png" pageId="9" pageNumber="44" targetBox="[151,1435,390,758]" targetPageId="9">
<paragraph id="8BF8D922FF9B8B69EAF4D625FBF14277" blockId="9.[151,1434,784,842]" pageId="9" pageNumber="44">
<emphasis id="B9330530FF9B8B69EAF4D625FEDE4215" bold="true" box="[151,282,784,808]" pageId="9" pageNumber="44">FIGURE 6.</emphasis>
Scanning electron microscopy images of female genital pores of (a)
<emphasis id="B9330530FF9B8B69EE62D624FB9B4215" box="[1025,1119,785,808]" italics="true" pageId="9" pageNumber="44">P. w i n g e i</emphasis>
(Laguna de los Patos), (b)
<taxonomicName id="4C47A2A1FF9B8B69EFEBD624FE7C4277" authority="Caroni Swamp" authorityName="Caroni Swamp" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9B8B69EFEBD624FF394277" italics="true" pageId="9" pageNumber="44">P. reticulata</emphasis>
(Caroni Swamp)
</taxonomicName>
, (c)
<taxonomicName id="4C47A2A1FF9B8B69EB84D606FD934277" box="[487,599,819,842]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9B8B69EB84D606FD934277" box="[487,599,819,842]" italics="true" pageId="9" pageNumber="44">P. obscura</emphasis>
</taxonomicName>
, (Upper Oropuche). Bar represents 200 mm.
</paragraph>
</caption>
<paragraph id="8BF8D922FF9B8B69EAA6D64DFACA4547" blockId="9.[151,1436,888,1146]" pageId="9" pageNumber="44">
However,
<taxonomicName id="4C47A2A1FF9B8B69EB5ED64CFE7342AF" box="[317,439,889,914]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9B8B69EB5ED64CFE7342AF" box="[317,439,889,914]" italics="true" pageId="9" pageNumber="44">P. obscura</emphasis>
</taxonomicName>
is recognized as a separate species, because it does not share the following characters with
<taxonomicName id="4C47A2A1FF9B8B69EAB7D6AAFEA14285" box="[212,357,927,952]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9B8B69EAB7D6AAFEA14285" box="[212,357,927,952]" italics="true" pageId="9" pageNumber="44">P. reticulata</emphasis>
</taxonomicName>
and
<emphasis id="B9330530FF9B8B69EBC3D6AAFDCE4285" box="[416,522,927,952]" italics="true" pageId="9" pageNumber="44">P. w i n g e i</emphasis>
: fewer subdistal comb-like spines of gonopodium ray 3 (1115, vs. 1418), higher and shorter caudal peduncle in females, apparent as lower SL/CPD values (
<figureCitation id="137CC5A7FF9B8B69EE0FD6F3FB7142DD" box="[1132,1205,966,992]" captionStart="FIGURE 4" captionStartId="8.[151,255,974,998]" captionTargetBox="[431,1115,204,925]" captionTargetId="figure@8.[415,1171,194,950]" captionTargetPageId="8" captionText="FIGURE 4. CPD / SL / CPD ratios of P. obscura (symbols, population Rio Seco (RS), population Oropuche (OR )) and P. reticulata females (symbols, population Claxton Bay (CB) population Caroni Swamp (CS ))." httpUri="https://zenodo.org/record/190848/files/figure.png" pageId="9" pageNumber="44">Fig. 4</figureCitation>
). In
<taxonomicName id="4C47A2A1FF9B8B69EE8FD6F3FAAD42E2" box="[1260,1385,966,991]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9B8B69EE8FD6F3FAAD42E2" box="[1260,1385,966,991]" italics="true" pageId="9" pageNumber="44">P. obscura</emphasis>
</taxonomicName>
SL/ CPD values of males are in the same range as females while there is a sexual dimorphism for this value in
<taxonomicName id="4C47A2A1FF9B8B69EFE5D6D8FECF4511" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9B8B69EFE5D6D8FECF4511" italics="true" pageId="9" pageNumber="44">P. reticulata</emphasis>
</taxonomicName>
. In this species the SL/CPD ratios of males are lower than for females and in the same range as males of
<taxonomicName id="4C47A2A1FF9B8B69EB6AD10FFE4F456E" box="[265,395,1082,1107]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9B8B69EB6AD10FFE4F456E" box="[265,395,1082,1107]" italics="true" pageId="9" pageNumber="44">P. obscura</emphasis>
</taxonomicName>
. Whether also
<emphasis id="B9330530FF9B8B69E824D10FFD71456E" box="[583,693,1082,1107]" italics="true" pageId="9" pageNumber="44">P. w i n g e i</emphasis>
is polymorphic for this character could not be conclusively determined from the available specimen and requires inspection of the
<typeStatus id="54FC6780FF9B8B69E9B5D155FBCD4547" box="[982,1033,1120,1146]" pageId="9" pageNumber="44">type</typeStatus>
material in the future.
</paragraph>
<caption id="DF3889AAFF9B8B69EAA6D3E7FA92466F" httpUri="https://zenodo.org/record/190850/files/figure.png" pageId="9" pageNumber="44" targetBox="[375,1211,1206,1717]" targetPageId="9">
<paragraph id="8BF8D922FF9B8B69EAA6D3E7FA92466F" blockId="9.[151,1437,1746,1874]" pageId="9" pageNumber="44">
<emphasis id="B9330530FF9B8B69EAA6D3E7FE8D47D7" bold="true" box="[197,329,1746,1770]" pageId="9" pageNumber="44">FIGURE 7.</emphasis>
Phylogramm of
<emphasis id="B9330530FF9B8B69E860D3E6FDB847D7" box="[515,636,1747,1770]" italics="true" pageId="9" pageNumber="44">
<taxonomicName id="4C47A2A1FF9B8B69E860D3E6FDBC47D7" box="[515,632,1747,1770]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="obscura">P. obscura</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C47A2A1FF9B8B69E8D1D3E6FCF347D7" box="[690,823,1747,1770]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9B8B69E8D1D3E6FCF347D7" box="[690,823,1747,1770]" italics="true" pageId="9" pageNumber="44">P. reticulata</emphasis>
</taxonomicName>
from different localities and
<emphasis id="B9330530FF9B8B69EE14D3E6FB1347D7" box="[1143,1239,1747,1770]" italics="true" pageId="9" pageNumber="44">P. w i n g e i</emphasis>
based on approx. 2200bp of mitochondrial DNA sequences. 50% majority rule consensus tree rooted on
<taxonomicName id="4C47A2A1FF9B8B69EE32D3C0FB684631" box="[1105,1196,1781,1804]" class="Actinopterygii" family="Poeciliidae" genus="Micropoecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="picta">
<emphasis id="B9330530FF9B8B69EE32D3C0FB684631" box="[1105,1196,1781,1804]" italics="true" pageId="9" pageNumber="44">M. picta</emphasis>
</taxonomicName>
as outgroup. Average bootstrap values obtained using different types of analysis (see Materials and Methods) are indicated above the branches. For abbreviations of origins see table 1. Numbers in brackets indicate different individuals from the same locality.
</paragraph>
</caption>
<paragraph id="8BF8D922FF9B8B6AEAA6D2B1FE4C4071" blockId="9.[151,1437,1924,2027]" lastBlockId="10.[151,1437,152,2014]" lastPageId="10" lastPageNumber="45" pageId="9" pageNumber="44">
The female of the
<typeStatus id="54FC6780FF9B8B69EBC5D2B1FDDE46A3" box="[422,538,1924,1950]" pageId="9" pageNumber="44" type="paratype">paratypes</typeStatus>
of “
<taxonomicName id="4C47A2A1FF9B8B69E831D2B1FCE946A0" box="[594,813,1924,1949]" class="Actinopterygii" family="Poeciliidae" genus="Girardinus" kingdom="Animalia" order="Cyprinodontiformes" pageId="9" pageNumber="44" phylum="Chordata" rank="species" species="guppii">
<emphasis id="B9330530FF9B8B69E831D2B1FCE946A0" box="[594,813,1924,1949]" italics="true" pageId="9" pageNumber="44">Girardinus guppii</emphasis>
</taxonomicName>
” (ZMB 6081) that were sent by Dr. Günther to his colleague at the Museum in Berlin was also measured. These fish are designated as” origin
<collectingCountry id="F35099B2FF9B8B69EF7FD29FFA4946F9" box="[1308,1421,1962,1988]" name="Trinidad and Tobago" pageId="9" pageNumber="44">Trinidad</collectingCountry>
”, however, no further information is available on the exact collection place or even river system. The SL/CPD value of this fish is close to the average for
<taxonomicName id="4C47A2A1FF988B6AE8F5D5ADFCE0418C" box="[662,804,152,177]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF988B6AE8F5D5ADFCE0418C" box="[662,804,152,177]" italics="true" pageId="10" pageNumber="45">P. reticulata</emphasis>
</taxonomicName>
. It has to be considered, however, that with respect to morphometrics substantial shape differences have been noted between different populations (
<bibRefCitation id="EFD6A4D3FF988B6AEEB4D58BFF1241C2" author="Alexander" pageId="10" pageNumber="49" refString="Alexander, H. J., Taylor, J. S., Wu, S. S. &amp; Breden, F. (2006) Parallel evolution and vicariance in the guppy (Poecilia reticulata) over multiple spatial and temporal scales. Evolution, 60, 2352 - 69." type="journal article" year="2006">
Alexander
<emphasis id="B9330530FF988B6AEF39D58AFA5041E5" box="[1370,1428,191,216]" italics="true" pageId="10" pageNumber="45">et al.</emphasis>
, 2006
</bibRefCitation>
;
<bibRefCitation id="EFD6A4D3FF988B6AEA80D5D0FE1541C2" author="Hendry" box="[227,465,229,255]" pageId="10" pageNumber="49" refString="Hendry, A. P., Kelly, M. L., Kinnison, M. T. &amp; Reznick, D. N. (2006) Parallel evolution of the sexes? Effects of predation and habitat features on the size and shape of wild guppies. Journal of Evolutionary Biology, 19, 741 - 754." type="journal article" year="2006">
Hendry
<emphasis id="B9330530FF988B6AEB27D5D0FE4541C3" box="[324,385,229,254]" italics="true" pageId="10" pageNumber="45">et al.</emphasis>
, 2006
</bibRefCitation>
). Only when more populations of
<emphasis id="B9330530FF988B6AE90FD5D0FC2141C3" box="[876,997,229,254]" italics="true" pageId="10" pageNumber="45">P. o b s c u r a</emphasis>
can be analyzed in the future it will become clear how robust these characters are and how suitable they will be for determining the taxonomic status of the species.
</paragraph>
<paragraph id="8BF8D922FF988B6AEAA6D46CFEC240DA" blockId="10.[151,1437,152,2014]" pageId="10" pageNumber="45">
A molecular phylogenetic analysis was performed using mitochondrial D-loop and cytochrome
<emphasis id="B9330530FF988B6AEFEED46CFA58404F" box="[1421,1436,345,370]" italics="true" pageId="10" pageNumber="45">b</emphasis>
sequences. The molecular data were analyzed with maximum likelihood, parsimony and neighbor-joining methods which yielded almost identical phylogenetic results. The topology of the resulting trees was always the same.
</paragraph>
<paragraph id="8BF8D922FF988B6AEAA6D4C1FB6743F2" blockId="10.[151,1437,152,2014]" pageId="10" pageNumber="45">
The phylogram rooted on
<taxonomicName id="4C47A2A1FF988B6AE863D4C1FD284330" box="[512,748,500,525]" class="Actinopterygii" family="Poeciliidae" genus="Micropoecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="picta">
<emphasis id="B9330530FF988B6AE863D4C1FD284330" box="[512,748,500,525]" italics="true" pageId="10" pageNumber="45">Micropoecilia picta</emphasis>
</taxonomicName>
has three well-separated branches (
<figureCitation id="137CC5A7FF988B6AEEF9D4C1FB344333" box="[1178,1264,500,526]" captionStart="FIGURE 7" captionStartId="9.[197,302,1746,1770]" captionTargetBox="[375,1211,1206,1717]" captionTargetId="figure@9.[371,1217,1201,1722]" captionTargetPageId="9" captionText="FIGURE 7. Phylogramm of P. obscura, and P. reticulata from different localities and P. w i n g e i based on approx. 2200 bp of mitochondrial DNA sequences. 50 % majority rule consensus tree rooted on M. picta as outgroup. Average bootstrap values obtained using different types of analysis (see Materials and Methods) are indicated above the branches. For abbreviations of origins see table 1. Numbers in brackets indicate different individuals from the same locality." httpUri="https://zenodo.org/record/190850/files/figure.png" pageId="10" pageNumber="45">Figs. 7</figureCitation>
,
<figureCitation id="137CC5A7FF988B6AEE9DD4C1FACA4333" box="[1278,1294,500,526]" captionStart="FIGURE 8" captionStartId="11.[151,255,1910,1934]" captionTargetBox="[412,1180,964,1898]" captionTargetId="figure@11.[402,1201,946,1905]" captionTargetPageId="11" captionText="FIGURE 8. Phylogramm of P. o b s c u r a and P. reticulata and P. w i n g e i from different localities based on approximately 800 bp of cytochrome b DNA sequences. 50 % majority rule consensus tree rooted on M. picta as outgroup. Average bootstrap values obtained using different types of analysis (see Materials and Methods) are indicated above the branches. For abbreviations of origins see table 1. Numbers in brackets indicate different individuals from the same locality." httpUri="https://zenodo.org/record/190851/files/figure.png" pageId="10" pageNumber="45">8</figureCitation>
), which are supported by high bootstrap values. One branch is built by the sequences of
<taxonomicName id="4C47A2A1FF988B6AEE43D72FFB4F430E" box="[1056,1163,538,563]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF988B6AEE43D72FFB4F430E" box="[1056,1163,538,563]" italics="true" pageId="10" pageNumber="45">P. wingei</emphasis>
</taxonomicName>
from the locus typicus and several collections from the city of Cumana. A second branch consists of the
<taxonomicName id="4C47A2A1FF988B6AEE00D774FB1B4367" box="[1123,1247,577,602]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF988B6AEE00D774FB1B4367" box="[1123,1247,577,602]" italics="true" pageId="10" pageNumber="45">P. obscura</emphasis>
</taxonomicName>
sequences. The third branch represents
<taxonomicName id="4C47A2A1FF988B6AEBD5D75DFD8E43BC" box="[438,586,616,641]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF988B6AEBD5D75DFD8E43BC" box="[438,586,616,641]" italics="true" pageId="10" pageNumber="45">P. reticulata</emphasis>
</taxonomicName>
. Here all other guppy sequences cluster that span the entire mainland range of the species from
<collectingCountry id="F35099B2FF988B6AEBACD7BBFD804395" box="[463,580,654,680]" name="United States of America" pageId="10" pageNumber="45">Columbia</collectingCountry>
to
<collectingCountry id="F35099B2FF988B6AE808D7BBFD704395" box="[619,692,654,680]" name="Brazil" pageId="10" pageNumber="45">Brazil</collectingCountry>
and the islands of
<collectingCountry id="F35099B2FF988B6AE9FBD7BBFC354395" box="[920,1009,654,680]" name="Trinidad and Tobago" pageId="10" pageNumber="45">Tobago</collectingCountry>
and
<collectingCountry id="F35099B2FF988B6AEE4FD7BBFB934395" box="[1068,1111,654,680]" name="Iceland" pageId="10" pageNumber="45">Isla</collectingCountry>
Margarita. Importantly the West Trinidadian guppies from the Caroni drainage are also firmly nested in this branch.
</paragraph>
<paragraph id="8BF8D922FF988B6AEAA6D7E9FE9742AD" blockId="10.[151,1437,152,2014]" pageId="10" pageNumber="45">
We also obtained sequences from a specimen of
<taxonomicName id="4C47A2A1FF988B6AE97DD7E9FC7743C8" box="[798,947,732,757]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF988B6AE97DD7E9FC7743C8" box="[798,947,732,757]" italics="true" pageId="10" pageNumber="45">P. reticulata</emphasis>
</taxonomicName>
(ZMB 6299) of the collection of Julius Gollmer from which the
<typeStatus id="54FC6780FF988B6AEBB0D637FDCD4221" box="[467,521,770,796]" pageId="10" pageNumber="45">type</typeStatus>
material for the first description was taken. This fish was collected and preserved on the
<date id="FFF9FFE2FF988B6AEB04D61CFE1B427E" box="[359,479,809,835]" pageId="10" pageNumber="45" value="1856-05-14">
14.5.
<quantity id="4CBF74C7FF988B6AEBC3D61CFE1B427E" box="[416,479,809,835]" metricMagnitude="1" metricUnit="m" metricValue="4.714239999999999" pageId="10" pageNumber="45" unit="in" value="1856.0">1856</quantity>
</date>
in the Rio Guayre near the city of Caracas (
<collectingCountry id="F35099B2FF988B6AE995D61CFBBD427E" box="[1014,1145,809,835]" name="Venezuela" pageId="10" pageNumber="45">Venezuela</collectingCountry>
). This sequence clusters with all present-day
<taxonomicName id="4C47A2A1FF988B6AEBF0D665FDE34254" box="[403,551,848,873]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF988B6AEBF0D665FDE34254" box="[403,551,848,873]" italics="true" pageId="10" pageNumber="45">P. reticulata</emphasis>
</taxonomicName>
including a fish taken in 2008 from a tributary to the Guayre (RG) near Caracas (
<figureCitation id="137CC5A7FF988B6AEB60D643FE8742AD" box="[259,323,886,912]" captionStart="FIGURE 8" captionStartId="11.[151,255,1910,1934]" captionTargetBox="[412,1180,964,1898]" captionTargetId="figure@11.[402,1201,946,1905]" captionTargetPageId="11" captionText="FIGURE 8. Phylogramm of P. o b s c u r a and P. reticulata and P. w i n g e i from different localities based on approximately 800 bp of cytochrome b DNA sequences. 50 % majority rule consensus tree rooted on M. picta as outgroup. Average bootstrap values obtained using different types of analysis (see Materials and Methods) are indicated above the branches. For abbreviations of origins see table 1. Numbers in brackets indicate different individuals from the same locality." httpUri="https://zenodo.org/record/190851/files/figure.png" pageId="10" pageNumber="45">Fig 8</figureCitation>
).
</paragraph>
<paragraph id="8BF8D922FF988B6AEAA6D6A8FE6A45A2" blockId="10.[151,1437,152,2014]" pageId="10" pageNumber="45">
It is a well-known fact that guppies evolve extremely rapidly in response to natural selection (
<bibRefCitation id="EFD6A4D3FF988B6AEF7CD6A8FF1242E3" author="Magurran" pageId="10" pageNumber="50" refString="Magurran, A. E. (1998) Population differentiation without speciation. Philosophical Transactions of The Royal Society London B Biological Sciences, 353, 275 - 286." type="journal article" year="1998">Magurran, 1998</bibRefCitation>
;
<bibRefCitation id="EFD6A4D3FF988B6AEA80D6F1FE1C42E3" author="Reznick" box="[227,472,964,990]" pageId="10" pageNumber="50" refString="Reznick, D. N., Shaw, F. H., Rodd, F. H. &amp; Shaw, R. G. (1997) Evaluation of the Rate of Evolution in Natural Populations of Guppies (Poecilia reticulata). Science, 275, 1934 - 1937." type="journal article" year="1997">
Reznick
<emphasis id="B9330530FF988B6AEB2FD6F1FE4D42E0" box="[332,393,964,989]" italics="true" pageId="10" pageNumber="45">et al.</emphasis>
, 1997
</bibRefCitation>
). This phenomenon could be partially explained by a very high natural mutation rate, which in turn could compromise the significance of the molecular analysis. We therefore included in our dataset five different ornamental guppy strains. These fish independently have been under the most extreme selection for several polygenic traits under enforced short generation times by guppy breeders in
<collectingCountry id="F35099B2FF988B6AEF20D10DFA53456F" box="[1347,1431,1080,1106]" name="Russia" pageId="10" pageNumber="45">Russia</collectingCountry>
,
<collectingCountry id="F35099B2FF988B6AEAF4D16BFF364545" box="[151,242,1118,1144]" name="Taiwan" pageId="10" pageNumber="45">Taiwan</collectingCountry>
,
<collectingCountry id="F35099B2FF988B6AEA9DD16BFEAD4545" box="[254,361,1118,1144]" name="Thailand" pageId="10" pageNumber="45">Thailand</collectingCountry>
and
<collectingCountry id="F35099B2FF988B6AEBC6D16BFDE64545" box="[421,546,1118,1144]" name="Singapore" pageId="10" pageNumber="45">Singapore</collectingCountry>
. On the molecular level they were, however, indistinguishable from wildcaught guppies (
<figureCitation id="137CC5A7FF988B6AEB34D1B0FE5B45A2" box="[343,415,1157,1183]" captionStart="FIGURE 8" captionStartId="11.[151,255,1910,1934]" captionTargetBox="[412,1180,964,1898]" captionTargetId="figure@11.[402,1201,946,1905]" captionTargetPageId="11" captionText="FIGURE 8. Phylogramm of P. o b s c u r a and P. reticulata and P. w i n g e i from different localities based on approximately 800 bp of cytochrome b DNA sequences. 50 % majority rule consensus tree rooted on M. picta as outgroup. Average bootstrap values obtained using different types of analysis (see Materials and Methods) are indicated above the branches. For abbreviations of origins see table 1. Numbers in brackets indicate different individuals from the same locality." httpUri="https://zenodo.org/record/190851/files/figure.png" pageId="10" pageNumber="45">Fig. 8</figureCitation>
).
</paragraph>
<paragraph id="8BF8D922FF988B6AEAA6D199FDD54407" blockId="10.[151,1437,152,2014]" pageId="10" pageNumber="45">
We included also a laboratory strain of
<emphasis id="B9330530FF988B6AE8F6D199FC9D45F8" box="[661,857,1196,1221]" italics="true" pageId="10" pageNumber="45">
<taxonomicName id="4C47A2A1FF988B6AE8F6D199FCE045F8" box="[661,804,1196,1221]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="reticulata">P. reticulata</taxonomicName>
that
</emphasis>
is kept as closely inbred line and displays a very homogeneous male pigmentation phenotype. This strain, designated maculatus-zebrinus (ma-ze), was established by Ojvind Winge in the twenties of the last century. Its mitochondrial DNA sequence is nested within all the other
<taxonomicName id="4C47A2A1FF988B6AEB1FD015FDCE4404" box="[380,522,1312,1337]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF988B6AEB1FD015FDCE4404" box="[380,522,1312,1337]" italics="true" pageId="10" pageNumber="45">P. reticulata</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BF8D922FF988B6AEAA6D073FCBA47C6" blockId="10.[151,1437,152,2014]" pageId="10" pageNumber="45">
The new species is described mainly on the basis of molecular data. Although the differentiation of the three branches of guppies is unequivocal and indicates separated (mitochondrial) gene pools it may be questioned whether the genetic distance is in the range that usually is found between species that are more classically defined by morphological criteria. To evaluate this, the genetic distances and branch lengths between the three guppy species were compared to the values obtained in studies of other poeciliid fish groups where also mitochondrial control region and the cytochrome b gene were used for phylogeny construction. The genetic distance separating the three guppy taxa is 0,0 29. This is well in the range of values obtained for morphologically very different species in the genus
<taxonomicName id="4C47A2A1FF988B6AE963D379FC5C4758" box="[768,920,1612,1637]" class="Actinopterygii" family="Poeciliidae" genus="Xiphophorus" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="genus">
<emphasis id="B9330530FF988B6AE963D379FC5C4758" box="[768,920,1612,1637]" italics="true" pageId="10" pageNumber="45">Xiphophorus</emphasis>
</taxonomicName>
(0.029 between Southern platyfish species,
<taxonomicName id="4C47A2A1FF988B6AEAF4D344FEF447B7" box="[151,304,1649,1674]" class="Actinopterygii" family="Poeciliidae" genus="Xiphophorus" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="maculatus">
<emphasis id="B9330530FF988B6AEAF4D344FEF447B7" box="[151,304,1649,1674]" italics="true" pageId="10" pageNumber="45">X. maculatus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C47A2A1FF988B6AEB5CD344FE6A47B7" box="[319,430,1649,1674]" class="Actinopterygii" family="Poeciliidae" genus="Xiphophorus" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="milleri">
<emphasis id="B9330530FF988B6AEB5CD344FE6A47B7" box="[319,430,1649,1674]" italics="true" pageId="10" pageNumber="45">X. milleri</emphasis>
</taxonomicName>
, and Southern swordtails, e.g
<taxonomicName id="4C47A2A1FF988B6AE971D344FC4347B7" box="[786,903,1649,1674]" class="Actinopterygii" family="Poeciliidae" genus="Xiphophorus" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="hellerii">
<emphasis id="B9330530FF988B6AE971D344FC4347B7" box="[786,903,1649,1674]" italics="true" pageId="10" pageNumber="45">X. hellerii</emphasis>
</taxonomicName>
,
<taxonomicName id="4C47A2A1FF988B6AE9F4D344FA8947B6" authority="Meyer and Schartl, 2002" authorityName="Meyer and Schartl" authorityYear="2002" box="[919,1357,1649,1675]" class="Actinopterygii" family="Poeciliidae" genus="Xiphophorus" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="signum">
<emphasis id="B9330530FF988B6AE9F4D344FBC847B7" box="[919,1036,1649,1674]" italics="true" pageId="10" pageNumber="45">X. signum</emphasis>
(
<bibRefCitation id="EFD6A4D3FF988B6AEE7FD344FA8247B6" author="Meyer" box="[1052,1350,1649,1675]" pageId="10" pageNumber="50" refString="Meyer, M. &amp; Schartl, M. (2002) Xiphophorus mayae, a new swordtail from Guatemala (Teleostei: Poeciliidae). Ichthyological Exploration of Freshwaters, 13, 59 - 66." type="journal article" year="2002">Meyer and Schartl, 2002</bibRefCitation>
)
</taxonomicName>
; 0,0 18 between the Southern swordtail species, e.g.
<taxonomicName id="4C47A2A1FF988B6AE8DBD3A2FCF5478D" box="[696,817,1687,1712]" class="Actinopterygii" family="Poeciliidae" genus="Xiphophorus" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="hellerii">
<emphasis id="B9330530FF988B6AE8DBD3A2FCF5478D" box="[696,817,1687,1712]" italics="true" pageId="10" pageNumber="45">X. hellerii</emphasis>
</taxonomicName>
,
<taxonomicName id="4C47A2A1FF988B6AE922D3A2FC7E478D" box="[833,954,1687,1712]" class="Actinopterygii" family="Poeciliidae" genus="Xiphophorus" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="signum">
<emphasis id="B9330530FF988B6AE922D3A2FC7E478D" box="[833,954,1687,1712]" italics="true" pageId="10" pageNumber="45">X. signum</emphasis>
</taxonomicName>
,
<taxonomicName id="4C47A2A1FF988B6AE9A9D3A2FA5C478D" authority="Meyer and Schartl, 2003" authorityName="Meyer and Schartl" authorityYear="2003" box="[970,1432,1686,1712]" class="Actinopterygii" family="Poeciliidae" genus="Xiphophorus" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="alvarezi">
<emphasis id="B9330530FF988B6AE9A9D3A2FB88478D" box="[970,1100,1687,1712]" italics="true" pageId="10" pageNumber="45">X. alvarezi</emphasis>
(
<bibRefCitation id="EFD6A4D3FF988B6AEE3FD3A3FA4B478D" author="Meyer" box="[1116,1423,1686,1712]" pageId="10" pageNumber="50" refString="Meyer, M. &amp; Schartl, M. (2003) Xiphophorus kallmani sp. n. - a new species of swordtail from Mexico (Teleostei, Cyprinodontiformes, Poeciliidae). Zoologische Abhandlungen Museum fur Tierkunde Dresden, 53, 57 - 64." type="journal article" year="2003">Meyer and Schartl, 2003</bibRefCitation>
)
</taxonomicName>
; 0,0 25 between
<taxonomicName id="4C47A2A1FF988B6AEB29D389FDE647E8" box="[330,546,1724,1749]" class="Actinopterygii" family="Poeciliidae" genus="Priapella" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="olmecae">
<emphasis id="B9330530FF988B6AEB29D389FDE647E8" box="[330,546,1724,1749]" italics="true" pageId="10" pageNumber="45">Priapella olmecae</emphasis>
</taxonomicName>
,
<taxonomicName id="4C47A2A1FF988B6AE852D389FD6747E8" box="[561,675,1724,1749]" class="Actinopterygii" family="Poeciliidae" genus="Priapella" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="species" species="chamulae">
<emphasis id="B9330530FF988B6AE852D389FD6747E8" box="[561,675,1724,1749]" italics="true" pageId="10" pageNumber="45">chamulae</emphasis>
</taxonomicName>
,
<emphasis id="B9330530FF988B6AE8D1D389FCEA47E8" box="[690,814,1724,1749]" italics="true" pageId="10" pageNumber="45">compressa</emphasis>
and
<emphasis id="B9330530FF988B6AE904D389FC2247E8" box="[871,998,1724,1749]" italics="true" pageId="10" pageNumber="45">intermedia</emphasis>
(
<bibRefCitation id="EFD6A4D3FF988B6AE995D389FB1847EB" author="Schartl" box="[1014,1244,1724,1750]" pageId="10" pageNumber="50" refString="Schartl, M., Meyer, M. K. &amp; Wilde, B. (2006) Description of Priapella chamulae sp. n. - a new poeciliid fish from the upper rio Grijalva system, Tabasco, Mexico (Teleostei: Cyprinodontiformes: Poeciliidae). Zoologische Abhandlungen Museum fur Tierkunde Dresden, 55, 59 - 67." type="journal article" year="2006">
Schartl
<emphasis id="B9330530FF988B6AEE30D389FB4A47E8" box="[1107,1166,1724,1749]" italics="true" pageId="10" pageNumber="45">et al.</emphasis>
, 2006
</bibRefCitation>
); 0,0 38 between the shortfin mollies of the genus
<taxonomicName id="4C47A2A1FF988B6AE87BD3D4FCBE47C6" authority="Meyer et al., 2004" authorityName="Meyer et al." authorityYear="2004" box="[536,890,1761,1787]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="genus">
<emphasis id="B9330530FF988B6AE87BD3D4FDBD47C7" box="[536,633,1761,1786]" italics="true" pageId="10" pageNumber="45">Poecilia</emphasis>
(
<bibRefCitation id="EFD6A4D3FF988B6AE8EAD3D4FCA347C6" author="Meyer" box="[649,871,1761,1787]" pageId="10" pageNumber="50" refString="Meyer, M. K., Schneider, K., Radda, A. C., Wilde, B. &amp; Schartl, M. (2004) A new species of Poecilia, subgenus Mollienesia, from upper Rio Cahabon system, Guatemala, with remarks on the Nomenclature of Mollienesia petenensis Gunther, 1866 (Teleostei: Cyprinodontiformes: Poeciliidae). Zoologische Abhandlungen Museum fur Tierkunde Dresden, 54, 145 - 154." type="journal article" year="2004">
Meyer
<emphasis id="B9330530FF988B6AE8BCD3D4FCDE47C7" box="[735,794,1761,1786]" italics="true" pageId="10" pageNumber="45">et al.</emphasis>
, 2004
</bibRefCitation>
))
</taxonomicName>
.
</paragraph>
<paragraph id="8BF8D922FF988B6AEAA6D233FBF94656" blockId="10.[151,1437,152,2014]" pageId="10" pageNumber="45">
Because the three species of the subgenus
<taxonomicName id="4C47A2A1FF988B6AE8D0D233FCB9461D" box="[691,893,1798,1824]" family="Poeciliidae" genus="Acanthophacelus" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageId="10" pageNumber="45" phylum="Chordata" rank="genus">Acanthophacelus</taxonomicName>
are not readily or reliably distinguishable on a morphological basis but can be separated using non-morphological data, such as from DNA sequence analysis, they can be regarded as another example of a cryptic species complex.
</paragraph>
<paragraph id="8BF8D922FF988B6BEAA6D243FE974071" blockId="10.[151,1437,152,2014]" lastBlockId="11.[151,1437,152,912]" lastPageId="11" lastPageNumber="46" pageId="10" pageNumber="45">
Although the validity of estimating divergence times from DNA sequence variations has been debated (e.g.
<bibRefCitation id="EFD6A4D3FF988B6AEABBD2A8FE09468A" author="Howell" box="[216,461,1949,1975]" pageId="10" pageNumber="49" refString="Howell, N., Howell, C. &amp; Elson, J. L. (2008) Molecular clock debate: Time dependency of molecular rate estimates for mtDNA: this is not the time for wishful thinking. Heredity, 101, 107 - 108." type="journal article" year="2008">
Howell
<emphasis id="B9330530FF988B6AEB5FD2A8FEB8468B" box="[316,380,1949,1974]" italics="true" pageId="10" pageNumber="45">et al.</emphasis>
, 2008
</bibRefCitation>
) it can give at least a reasonable estimate, in particular when calibrations are available from related groups of organisms as is the case for the mitochondrial control region and cytochrome b genes of actinopterygian fishes (for a recent compilation see
<bibRefCitation id="EFD6A4D3FF998B6BE9EFD5ADFB48418F" author="Burridge" box="[908,1164,152,178]" pageId="11" pageNumber="49" refString="Burridge, C. P., Craw, D., Jack, D. C., King, T. M. &amp; Waters, J. M. (2008) Does fish ecology predict dispersal across a river drainage divide? Evolution, 62, 1484 - 99." type="journal article" year="2008">
Burridge
<emphasis id="B9330530FF998B6BE99DD5ADFBFF418C" box="[1022,1083,152,177]" italics="true" pageId="11" pageNumber="46">et al.</emphasis>
, 2008
</bibRefCitation>
). Using the fastest and slowest rates of all known calibrated molecular clocks in teleosts the separation of the three guppy species should have occurred between 2.5 and 5 million years ago based on the cytochrome b sequences (0.0076 0.0036 changes/site/Myr) and between 0.4 and 4.2 million years based on the sequences of the control region (0.044 0.004).
</paragraph>
<paragraph id="8BF8D922FF998B6BEAA6D46CFD1242AD" blockId="11.[151,1437,152,912]" pageId="11" pageNumber="46">
<emphasis id="B9330530FF998B6BEAA6D46CFEFA404F" box="[197,318,345,370]" italics="true" pageId="11" pageNumber="46">P. o b s c u r a</emphasis>
is endemic to the island of
<collectingCountry id="F35099B2FF998B6BE8EDD46CFD33404E" box="[654,759,345,371]" name="Trinidad and Tobago" pageId="11" pageNumber="46">Trinidad</collectingCountry>
. A genetic divergence of the guppies from the Oropuche drainage has been noted earlier using mitochondrial DNA sequences or allozyme data, (
<bibRefCitation id="EFD6A4D3FF998B6BEE87D4B5FEFC40FD" author="Alexander" pageId="11" pageNumber="49" refString="Alexander, H. J. &amp; Breden, F. (2004) Sexual isolation and extreme morphological divergence in the Cumana guppy: a possible case of incipient speciation. Journal of Evolutionary Biology, 17, 1238 - 54." type="journal article" year="2004">Alexander and Breden, 2004</bibRefCitation>
; Carvalho
<emphasis id="B9330530FF998B6BEBD6D492FE2B40FD" box="[437,495,423,448]" italics="true" pageId="11" pageNumber="46">et al.</emphasis>
, 1991;
<bibRefCitation id="EFD6A4D3FF998B6BE824D493FC5140FD" author="Russell" box="[583,917,422,448]" pageId="11" pageNumber="50" refString="Russell, S. T. &amp; Magurran, A. E. (2006) Intrinsic reproductive isolation between Trinidadian populations of the guppy, Poecilia reticulata. Journal of Evolutionary Biology, 19, 1294 - 1303." type="journal article" year="2006">Russell and Magurran, 2006</bibRefCitation>
); (
<bibRefCitation id="EFD6A4D3FF998B6BE9D2D493FB7840FD" author="Alexander" box="[945,1212,422,448]" pageId="11" pageNumber="49" refString="Alexander, H. J., Taylor, J. S., Wu, S. S. &amp; Breden, F. (2006) Parallel evolution and vicariance in the guppy (Poecilia reticulata) over multiple spatial and temporal scales. Evolution, 60, 2352 - 69." type="journal article" year="2006">
Alexander
<emphasis id="B9330530FF998B6BEE57D492FBAA40FD" box="[1076,1134,423,448]" italics="true" pageId="11" pageNumber="46">et al.</emphasis>
, 2006
</bibRefCitation>
). In all phylogentic trees published so far by these authors the so-called “Oropuche guppies” formed a separate branch. But this was never discussed on basis of recognizing these fish as a separate species. Moreover the differentiation between the fish from the Caroni and the Oropuche drainages was interpreted as the two subgroups being on their way to establish different species, but not having completed the whole process. Our analyses presents a different view, namely of two species that have been separated since a long time. The Caroni drainage, which drains west into the gulf of Paria is separated by a watershed from the Oropuche drainage although the minimum distance between the two river systems during the rainy season can be as close as
<quantity id="4CBF74C7FF998B6BEEB8D780FACB43F2" box="[1243,1295,693,719]" metricMagnitude="1" metricUnit="m" metricValue="7.0" pageId="11" pageNumber="46" unit="m" value="70.0">70m</quantity>
(
<bibRefCitation id="EFD6A4D3FF998B6BEF7CD780FF1C43CB" author="Magurran" pageId="11" pageNumber="50" refString="Magurran, A. E. (2005) Evolutionary Ecology - The Trinidadian Guppy. Oxford University Press, New York, 224 pp." type="book" year="2005">Magurran, 2005</bibRefCitation>
). The Oropuche and some other rivers with
<taxonomicName id="4C47A2A1FF998B6BE88FD7E9FCAE43C8" box="[748,874,732,757]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="11" pageNumber="46" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF998B6BE88FD7E9FCAE43C8" box="[748,874,732,757]" italics="true" pageId="11" pageNumber="46">P. obscura</emphasis>
</taxonomicName>
flow east and drain into the Atlantic. Notably, there is a third drainage, namely those rivers that drain north into the Caribbean Sea. Interestingly these fish, although clearly representing the species
<taxonomicName id="4C47A2A1FF998B6BE8F4D61CFCE9427F" box="[663,813,809,834]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" pageNumber="46" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF998B6BE8F4D61CFCE9427F" box="[663,813,809,834]" italics="true" pageId="11" pageNumber="46">P. reticulata</emphasis>
</taxonomicName>
somehow cluster in the phylogenetic trees with
<taxonomicName id="4C47A2A1FF998B6BEFE5D61CFECA4254" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" pageNumber="46" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF998B6BEFE5D61CFECA4254" italics="true" pageId="11" pageNumber="46">P. reticulata</emphasis>
</taxonomicName>
from the northward-located island of
<collectingCountry id="F35099B2FF998B6BE884D665FC864257" box="[743,834,848,874]" name="Trinidad and Tobago" pageId="11" pageNumber="46">Tobago</collectingCountry>
(
<figureCitation id="137CC5A7FF998B6BE937D665FC5B4257" box="[852,927,848,874]" captionStart="FIGURE 8" captionStartId="11.[151,255,1910,1934]" captionTargetBox="[412,1180,964,1898]" captionTargetId="figure@11.[402,1201,946,1905]" captionTargetPageId="11" captionText="FIGURE 8. Phylogramm of P. o b s c u r a and P. reticulata and P. w i n g e i from different localities based on approximately 800 bp of cytochrome b DNA sequences. 50 % majority rule consensus tree rooted on M. picta as outgroup. Average bootstrap values obtained using different types of analysis (see Materials and Methods) are indicated above the branches. For abbreviations of origins see table 1. Numbers in brackets indicate different individuals from the same locality." httpUri="https://zenodo.org/record/190851/files/figure.png" pageId="11" pageNumber="46">Fig. 8</figureCitation>
), indicating a possible colonization from
<collectingCountry id="F35099B2FF998B6BEAF4D643FF2B42AD" box="[151,239,886,912]" name="Trinidad and Tobago" pageId="11" pageNumber="46">Tobago</collectingCountry>
towards Northern
<collectingCountry id="F35099B2FF998B6BEBA8D643FDEB42AD" box="[459,559,886,912]" name="Trinidad and Tobago" pageId="11" pageNumber="46">Trinidad</collectingCountry>
or vice-versa.
</paragraph>
<caption id="DF3889AAFF998B6BEAF4D243FA9346C8" httpUri="https://zenodo.org/record/190851/files/figure.png" pageId="11" pageNumber="46" targetBox="[412,1180,964,1898]" targetPageId="11">
<paragraph id="8BF8D922FF998B6BEAF4D243FA9346C8" blockId="11.[151,1437,1910,2037]" pageId="11" pageNumber="46">
<emphasis id="B9330530FF998B6BEAF4D243FEDE46B3" bold="true" box="[151,282,1910,1934]" pageId="11" pageNumber="46">FIGURE 8.</emphasis>
Phylogramm of
<emphasis id="B9330530FF998B6BEBB3D243FDFA46B0" box="[464,574,1910,1933]" italics="true" pageId="11" pageNumber="46">P. o b s c u r a</emphasis>
and
<taxonomicName id="4C47A2A1FF998B6BE816D243FD3346B0" box="[629,759,1910,1933]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" pageNumber="46" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF998B6BE816D243FD3346B0" box="[629,759,1910,1933]" italics="true" pageId="11" pageNumber="46">P. reticulata</emphasis>
</taxonomicName>
and
<emphasis id="B9330530FF998B6BE94ED243FC4F46B0" box="[813,907,1910,1933]" italics="true" pageId="11" pageNumber="46">P. w i n g e i</emphasis>
from different localities based on approximately 800 bp of cytochrome b DNA sequences. 50% majority rule consensus tree rooted on
<taxonomicName id="4C47A2A1FF998B6BEE2DD2ACFB6E468D" box="[1102,1194,1945,1968]" class="Actinopterygii" family="Poeciliidae" genus="Micropoecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="11" pageNumber="46" phylum="Chordata" rank="species" species="picta">
<emphasis id="B9330530FF998B6BEE2DD2ACFB6E468D" box="[1102,1194,1945,1968]" italics="true" pageId="11" pageNumber="46">M. picta</emphasis>
</taxonomicName>
as outgroup. Average bootstrap values obtained using different types of analysis (see Materials and Methods) are indicated above the branches. For abbreviations of origins see table 1. Numbers in brackets indicate different individuals from the same locality.
</paragraph>
</caption>
<paragraph id="8BF8D922FF9E8B6CEAA6D5ADFBBC4333" blockId="12.[151,1437,152,2034]" pageId="12" pageNumber="47">
The hypothesis that the Caroni (including the Northern rivers) and Oropuche drainages have an independent freshwater fauna from different ancestral lineages is supported by studies on reptiles (
<bibRefCitation id="EFD6A4D3FF9E8B6CEF36D58BFF1341C2" author="Boos" pageId="12" pageNumber="49" refString="Boos, H. E. A. (1984) A consideration of the terrestrial reptile fauna on some offshore islands north west of Trinidad. Living World Journal Trinidad and Tobago Field Naturalists' Club, 19 - 26." type="book chapter" year="1984">Boos, 1984</bibRefCitation>
), the cyprinodont fish
<taxonomicName id="4C47A2A1FF9E8B6CEB89D5D0FC5741C2" authority="Jowers et al., 2008" authorityName="Jowers et al." authorityYear="2008" box="[490,915,229,255]" class="Actinopterygii" family="Rivulidae" genus="Rivulus" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="hartii">
<emphasis id="B9330530FF9E8B6CEB89D5D0FD5541C3" box="[490,657,229,254]" italics="true" pageId="12" pageNumber="47">Rivulus hartii</emphasis>
(
<bibRefCitation id="EFD6A4D3FF9E8B6CE8C2D5D0FC4F41C2" author="Jowers" box="[673,907,229,255]" pageId="12" pageNumber="49" refString="Jowers, M. J., Cohen, B. L. &amp; Downie, J. R. (2008) The cyprinodont fish Rivulus (Aplocheiloidei: Rivulidae) in Trinidad and Tobago: molecular evidence for marine dispersal, genetic isolation and local differentiation. Journal of Zoological Systematics and Evolutionary Research, 46, 48 - 55." type="journal article" year="2008">
Jowers
<emphasis id="B9330530FF9E8B6CE89CD5D0FCF841C3" box="[767,828,229,254]" italics="true" pageId="12" pageNumber="47">et al.</emphasis>
, 2008
</bibRefCitation>
)
</taxonomicName>
and paleogeographic evidence. During the glaciation periods of the Pleistocene sea levels were up to
<quantity id="4CBF74C7FF9E8B6CE938D439FC6C401B" box="[859,936,268,294]" metricMagnitude="2" metricUnit="m" metricValue="1.3" pageId="12" pageNumber="47" unit="m" value="130.0">130 m</quantity>
lower and
<collectingCountry id="F35099B2FF9E8B6CEE53D439FB52401B" box="[1072,1174,268,294]" name="Trinidad and Tobago" pageId="12" pageNumber="47">Trinidad</collectingCountry>
was connected to the mainland (
<bibRefCitation id="EFD6A4D3FF9E8B6CEB77D407FE744071" author="Kenny" box="[276,432,306,332]" pageId="12" pageNumber="49" refString="Kenny, J. S. (1988) Hermatypic scleractinian corals of Trinidad. Studies of the fauna of Curacao and other Caribbean islands, 123, 83 - 100." type="journal article" year="1988">Kenny, 1988</bibRefCitation>
). The Orinoco, the main distribution area of
<taxonomicName id="4C47A2A1FF9E8B6CE9DCD407FB894076" box="[959,1101,306,331]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9E8B6CE9DCD407FB894076" box="[959,1101,306,331]" italics="true" pageId="12" pageNumber="47">P. reticulata</emphasis>
</taxonomicName>
, at that time reached further northward and was probably connected to the Caroni drainage, discharging to the Gulf of Paria. The Oropuche drainage had most likely confluence with a separate more easterly river system and was colonized from there. Thus
<emphasis id="B9330530FF9E8B6CEAB9D492FE9040FD" box="[218,340,423,448]" italics="true" pageId="12" pageNumber="47">P. o b s c u r a</emphasis>
and
<taxonomicName id="4C47A2A1FF9E8B6CEBF0D492FDE640FD" box="[403,546,423,448]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9E8B6CEBF0D492FDE640FD" box="[403,546,423,448]" italics="true" pageId="12" pageNumber="47">P. reticulata</emphasis>
</taxonomicName>
arrived from different regions to
<collectingCountry id="F35099B2FF9E8B6CE9D5D493FBDF40FD" box="[950,1051,422,448]" name="Trinidad and Tobago" pageId="12" pageNumber="47">Trinidad</collectingCountry>
and did not cross the watershed that still separates them in the Northern mountain range. We hypothesize that
<taxonomicName id="4C47A2A1FF9E8B6CEE4FD4F8FB5340DB" box="[1068,1175,461,486]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9E8B6CEE4FD4F8FB5340DB" box="[1068,1175,461,486]" italics="true" pageId="12" pageNumber="47">P. wingei</emphasis>
</taxonomicName>
was the most western species but recently was circled by northwest colonizing populations of
<taxonomicName id="4C47A2A1FF9E8B6CE987D4C1FBB44330" box="[996,1136,500,525]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9E8B6CE987D4C1FBB44330" box="[996,1136,500,525]" italics="true" pageId="12" pageNumber="47">P. reticulata</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BF8D922FF9E8B6CEAA6D72FFBB34395" blockId="12.[151,1437,152,2034]" pageId="12" pageNumber="47">
<bibRefCitation id="EFD6A4D3FF9E8B6CEAA6D72FFE6B4309" box="[197,431,538,564]" pageId="12" pageNumber="50" refString="Poeser, F. N., Kempkes, M. &amp; Isbrucker, I. J. H. (2005) Description of Poecilia (Acanthophacelus) wingei n. sp. from the Paria Peninsula, Venezuela, including notes on Acanthophacelus Eigenmann, 1907 and other subgenera of Poecilia Bloch and Schneider, 1801 (Teleostei, Cyprinodontiformes, Poeciliidae). Contributions to Zoology, 74, 97 - 115." type="journal article">
Poeser
<emphasis id="B9330530FF9E8B6CEB7FD72FFE9D430E" box="[284,345,538,563]" italics="true" pageId="12" pageNumber="47">et al.</emphasis>
(2005)
</bibRefCitation>
argued that the Eastern
<collectingCountry id="F35099B2FF9E8B6CE8B0D72FFCFC4309" box="[723,824,538,564]" name="Trinidad and Tobago" pageId="12" pageNumber="47">Trinidad</collectingCountry>
guppies, now
<taxonomicName id="4C47A2A1FF9E8B6CE98BD72FFBA1430E" box="[1000,1125,538,563]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9E8B6CE98BD72FFBA1430E" box="[1000,1125,538,563]" italics="true" pageId="12" pageNumber="47">P. obscura</emphasis>
</taxonomicName>
, originally were the same species as
<emphasis id="B9330530FF9E8B6CEB79D774FE434367" box="[282,391,577,602]" italics="true" pageId="12" pageNumber="47">P. w i n g e i</emphasis>
and have been “despeciated” through hybridization and genetic introgression by the Common (or Orinoco) guppy,
<taxonomicName id="4C47A2A1FF9E8B6CE86BD75DFD5E43BC" box="[520,666,616,641]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9E8B6CE86BD75DFD5E43BC" box="[520,666,616,641]" italics="true" pageId="12" pageNumber="47">P. reticulata</emphasis>
</taxonomicName>
. Our molecular analysis does not support this interpretation, as there is absolutely no indication of a closer relationship of
<taxonomicName id="4C47A2A1FF9E8B6CE924D7BAFC064395" box="[839,962,655,680]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9E8B6CE924D7BAFC064395" box="[839,962,655,680]" italics="true" pageId="12" pageNumber="47">P. obscura</emphasis>
</taxonomicName>
with
<taxonomicName id="4C47A2A1FF9E8B6CEE66D7BAFBB44395" box="[1029,1136,655,680]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9E8B6CEE66D7BAFBB44395" box="[1029,1136,655,680]" italics="true" pageId="12" pageNumber="47">P. wingei</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BF8D922FF9E8B6CEAA6D780FD33427E" blockId="12.[151,1437,152,2034]" pageId="12" pageNumber="47">
Given the strong variation in male pigmentation the description of
<taxonomicName id="4C47A2A1FF9E8B6CE9ACD780FBFE43F3" box="[975,1082,693,718]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9E8B6CE9ACD780FBFE43F3" box="[975,1082,693,718]" italics="true" pageId="12" pageNumber="47">P. wingei</emphasis>
</taxonomicName>
on the basis of coloration was not conclusive. But our molecular data now define the species unequivocally and show that it is a separated taxon from
<taxonomicName id="4C47A2A1FF9E8B6CEB42D637FE6B4226" box="[289,431,770,795]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9E8B6CEB42D637FE6B4226" box="[289,431,770,795]" italics="true" pageId="12" pageNumber="47">P. reticulata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C47A2A1FF9E8B6CEB8AD637FDA14226" box="[489,613,770,795]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9E8B6CEB8AD637FDA14226" box="[489,613,770,795]" italics="true" pageId="12" pageNumber="47">P. obscura</emphasis>
</taxonomicName>
. Clearly, all fish earlier classified and recognized as Endlers guppies from localities in and around Cumana are
<taxonomicName id="4C47A2A1FF9E8B6CE8E6D61CFD34427F" box="[645,752,809,834]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9E8B6CE8E6D61CFD34427F" box="[645,752,809,834]" italics="true" pageId="12" pageNumber="47">P. wingei</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph id="8BF8D922FF9E8B6CEAA6D665FD96442E" blockId="12.[151,1437,152,2034]" pageId="12" pageNumber="47">
The molecular barcode from mitochondrial sequences will help to define the actual species ranges and hybrid zones. Evidently fish from collecting station 6 of
<bibRefCitation id="EFD6A4D3FF9E8B6CE94ED643FBD642AD" box="[813,1042,886,912]" pageId="12" pageNumber="50" refString="Poeser, F. N., Kempkes, M. &amp; Isbrucker, I. J. H. (2005) Description of Poecilia (Acanthophacelus) wingei n. sp. from the Paria Peninsula, Venezuela, including notes on Acanthophacelus Eigenmann, 1907 and other subgenera of Poecilia Bloch and Schneider, 1801 (Teleostei, Cyprinodontiformes, Poeciliidae). Contributions to Zoology, 74, 97 - 115." type="journal article">
Poeser
<emphasis id="B9330530FF9E8B6CE9E0D642FC7942AD" box="[899,957,887,912]" italics="true" pageId="12" pageNumber="47">et al.</emphasis>
(2005)
</bibRefCitation>
were designated by these authors as
<taxonomicName id="4C47A2A1FF9E8B6CEAD4D6A8FE81428B" box="[183,325,925,950]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9E8B6CEAD4D6A8FE81428B" box="[183,325,925,950]" italics="true" pageId="12" pageNumber="47">P. reticulata</emphasis>
</taxonomicName>
on the basis of their male coloration phenotype. We revisited this place (El Cordon, EC). The molecular analysis revealed clearly that these fish have the
<taxonomicName id="4C47A2A1FF9E8B6CE90FD6F1FC1842E0" box="[876,988,964,989]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9E8B6CE90FD6F1FC1842E0" box="[876,988,964,989]" italics="true" pageId="12" pageNumber="47">P. wingei</emphasis>
</taxonomicName>
mitochondrial sequence (
<figureCitation id="137CC5A7FF9E8B6CEF76D6F1FA9B42E3" box="[1301,1375,964,990]" captionStart="FIGURE 8" captionStartId="11.[151,255,1910,1934]" captionTargetBox="[412,1180,964,1898]" captionTargetId="figure@11.[402,1201,946,1905]" captionTargetPageId="11" captionText="FIGURE 8. Phylogramm of P. o b s c u r a and P. reticulata and P. w i n g e i from different localities based on approximately 800 bp of cytochrome b DNA sequences. 50 % majority rule consensus tree rooted on M. picta as outgroup. Average bootstrap values obtained using different types of analysis (see Materials and Methods) are indicated above the branches. For abbreviations of origins see table 1. Numbers in brackets indicate different individuals from the same locality." httpUri="https://zenodo.org/record/190851/files/figure.png" pageId="12" pageNumber="47">Fig. 8</figureCitation>
). No further data exist to clarify whether this discrepancy can be explained by interspecific hybridization. We also noted as reported by
<bibRefCitation id="EFD6A4D3FF9E8B6CEBEDD124FDB14516" box="[398,629,1041,1067]" pageId="12" pageNumber="50" refString="Poeser, F. N., Kempkes, M. &amp; Isbrucker, I. J. H. (2005) Description of Poecilia (Acanthophacelus) wingei n. sp. from the Paria Peninsula, Venezuela, including notes on Acanthophacelus Eigenmann, 1907 and other subgenera of Poecilia Bloch and Schneider, 1801 (Teleostei, Cyprinodontiformes, Poeciliidae). Contributions to Zoology, 74, 97 - 115." type="journal article">
Poeser
<emphasis id="B9330530FF9E8B6CEB87D124FDE44517" box="[484,544,1041,1066]" italics="true" pageId="12" pageNumber="47">et al.</emphasis>
(2005)
</bibRefCitation>
that in the river below the El Cordon waterfall several males had an unusual long gonopodium. However, closer inspection revealed that all of these were immature males and that the gonopodium, once fully differentiated is not longer than of any other guppy. Only in the transition period of the anal fin to the intromittent male genitalia a prolongation of the fin happens. This can be seen - although in a less extreme degree - in any other guppy strain. It is unclear whether this is an environmental effect, because such an extreme lengthening of the transforming anal fin was not observed anymore in the subsequent captivity reared offspring.
</paragraph>
<paragraph id="8BF8D922FF9E8B6CEAA6D015FC4C44C6" blockId="12.[151,1437,152,2034]" pageId="12" pageNumber="47">
Male coloration is much more uniform in
<taxonomicName id="4C47A2A1FF9E8B6CE8D9D015FCE24404" box="[698,806,1312,1337]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9E8B6CE8D9D015FCE24404" box="[698,806,1312,1337]" italics="true" pageId="12" pageNumber="47">P. wingei</emphasis>
</taxonomicName>
from Cumana and is much alike the pigmentation of the fish from Campoma and all earlier known Endlers guppies (
<figureCitation id="137CC5A7FF9E8B6CE9F0D073FC1E445D" box="[915,986,1350,1376]" captionStart="FIGURE 2. P" captionStartId="5.[151,255,1910,1934]" captionTargetBox="[187,1405,533,1892]" captionTargetId="figure@5.[187,1409,533,1892]" captionTargetPageId="5" captionText="FIGURE 2. P. obscura male (a) and female (b) from locus typicus, Oropuche River (OR). (c) P. obscura, population Rio Seco (RS); (d) P, reticulata, population Caroni Swamp (CS); (e) P. reticulata, population Rio Yaguaracual (RY); (f) P. c f “ wingei ” from El Cordon waterfall (EC); (g) P. w i n g e i from locus typicus, population Campoma (Ca); (h) P. w i n g e i, population Laguna de los Patos (LP)." httpUri="https://zenodo.org/record/190847/files/figure.png" pageId="12" pageNumber="47">
Fig.
<quantity id="4CBF74C7FF9E8B6CE9A9D073FC1E445D" box="[970,986,1350,1376]" metricMagnitude="-3" metricUnit="kg" metricValue="2.0" pageId="12" pageNumber="47" unit="g" value="2.0">2</quantity>
</figureCitation>
g,h). We suspect that fish (see
<figureCitation id="137CC5A7FF9E8B6CEF20D073FA4C445D" box="[1347,1416,1350,1376]" captionStart="FIGURE 2. P" captionStartId="5.[151,255,1910,1934]" captionTargetBox="[187,1405,533,1892]" captionTargetId="figure@5.[187,1409,533,1892]" captionTargetPageId="5" captionText="FIGURE 2. P. obscura male (a) and female (b) from locus typicus, Oropuche River (OR). (c) P. obscura, population Rio Seco (RS); (d) P, reticulata, population Caroni Swamp (CS); (e) P. reticulata, population Rio Yaguaracual (RY); (f) P. c f “ wingei ” from El Cordon waterfall (EC); (g) P. w i n g e i from locus typicus, population Campoma (Ca); (h) P. w i n g e i, population Laguna de los Patos (LP)." httpUri="https://zenodo.org/record/190847/files/figure.png" pageId="12" pageNumber="47">Fig. 2</figureCitation>
f) collected by
<bibRefCitation id="EFD6A4D3FF9E8B6CEB53D058FDD844BA" box="[304,540,1389,1415]" pageId="12" pageNumber="50" refString="Poeser, F. N., Kempkes, M. &amp; Isbrucker, I. J. H. (2005) Description of Poecilia (Acanthophacelus) wingei n. sp. from the Paria Peninsula, Venezuela, including notes on Acanthophacelus Eigenmann, 1907 and other subgenera of Poecilia Bloch and Schneider, 1801 (Teleostei, Cyprinodontiformes, Poeciliidae). Contributions to Zoology, 74, 97 - 115." type="journal article">
Poeser
<emphasis id="B9330530FF9E8B6CEBEBD058FE0044BB" box="[392,452,1389,1414]" italics="true" pageId="12" pageNumber="47">et al.</emphasis>
(2005)
</bibRefCitation>
including those from the El Cordon waterfall, which have a “similar color pattern as the Orinoco variety” (=
<taxonomicName id="4C47A2A1FF9E8B6CE850D0A1FD034490" box="[563,711,1428,1453]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9E8B6CE850D0A1FD034490" box="[563,711,1428,1453]" italics="true" pageId="12" pageNumber="47">P. reticulata</emphasis>
</taxonomicName>
), but have additionally some metallic pigment spots like
<taxonomicName id="4C47A2A1FF9E8B6CEFE5D0A1FF2C44E9" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9E8B6CEFE5D0A1FF2C44E9" italics="true" pageId="12" pageNumber="47">P. wingei</emphasis>
</taxonomicName>
from the Locus typicus and the Cumana populations (CC, LP) may be hybrids. Future studies that include nuclear genetic markers will be necessary to clarify this.
</paragraph>
<paragraph id="8BF8D922FF9E8B6DEAA6D33DFD884071" blockId="12.[151,1437,152,2034]" lastBlockId="13.[151,1437,152,2034]" lastPageId="13" lastPageNumber="48" pageId="12" pageNumber="47">
All other fish that based on molecular data could be unequivocally assigned to the
<taxonomicName id="4C47A2A1FF9E8B6CEEEFD33DFB33471C" box="[1164,1271,1544,1569]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9E8B6CEEEFD33DFB33471C" box="[1164,1271,1544,1569]" italics="true" pageId="12" pageNumber="47">P. wingei</emphasis>
</taxonomicName>
clade were of a typical coloration distinct (see also
<figureCitation id="137CC5A7FF9E8B6CE82DD31BFD504775" box="[590,660,1582,1608]" captionStart="FIGURE 2. P" captionStartId="5.[151,255,1910,1934]" captionTargetBox="[187,1405,533,1892]" captionTargetId="figure@5.[187,1409,533,1892]" captionTargetPageId="5" captionText="FIGURE 2. P. obscura male (a) and female (b) from locus typicus, Oropuche River (OR). (c) P. obscura, population Rio Seco (RS); (d) P, reticulata, population Caroni Swamp (CS); (e) P. reticulata, population Rio Yaguaracual (RY); (f) P. c f “ wingei ” from El Cordon waterfall (EC); (g) P. w i n g e i from locus typicus, population Campoma (Ca); (h) P. w i n g e i, population Laguna de los Patos (LP)." httpUri="https://zenodo.org/record/190847/files/figure.png" pageId="12" pageNumber="47">
Fig.
<quantity id="4CBF74C7FF9E8B6CE8E6D31BFD504775" box="[645,660,1582,1608]" metricMagnitude="-3" metricUnit="kg" metricValue="2.0" pageId="12" pageNumber="47" unit="g" value="2.0">2</quantity>
</figureCitation>
g,h) from
<emphasis id="B9330530FF9E8B6CE968D31AFC474775" box="[779,899,1583,1608]" italics="true" pageId="12" pageNumber="47">P. o b s c u r a</emphasis>
and
<taxonomicName id="4C47A2A1FF9E8B6CE9A3D31AFB8B4775" box="[960,1103,1583,1608]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9E8B6CE9A3D31AFB8B4775" box="[960,1103,1583,1608]" italics="true" pageId="12" pageNumber="47">P. reticulata</emphasis>
</taxonomicName>
, which we describe here as: longitudinal broad red band extending maximally from the operculum to the caudal fin base, along or above the lateral midline in the trunk and along or below the lateral midline in the peduncle, often interrupted by a vertical black bar originating around the anal fin base and extending to the dorsal fin base or anterior thereof. The red band often dissolved into oval or rarely round red spots, especially in the peduncle. A black stripe above the red band extending maximally to the eyes and the caudal fin base, sometimes missing in the trunk area. Additional dark black coloration of the ventral margin of the peduncle in many males. Caudal fin often with ventral and dorsal sword-like coloration, red, yellow or white, often with a black margin, frequently upper and/or lower rays of caudal fin prolonged to form colored swords or double-swords. Some males with large black shoulder spot with fuzzy margin, similar to the vertical bar. Perfectly round dark black body spots with sharp borders, which are typical for
<taxonomicName id="4C47A2A1FF9E8B6CE818D284FCCC46F7" box="[635,776,1969,1994]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9E8B6CE818D284FCCC46F7" box="[635,776,1969,1994]" italics="true" pageId="12" pageNumber="47">P. reticulata</emphasis>
</taxonomicName>
and
<taxonomicName id="4C47A2A1FF9E8B6CE921D284FC7946F7" box="[834,957,1969,1994]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="12" pageNumber="47" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9E8B6CE921D284FC7946F7" box="[834,957,1969,1994]" italics="true" pageId="12" pageNumber="47">P. obscura</emphasis>
</taxonomicName>
males, very rare. Large roundish or oval bright metallic blotches of, green, or more rarely yellow or light blue iridescent color following the basic longitudinal pattern interrupting the red band in the peduncle. White markings rare on the body sides. Rarely small black irregular spots on the belly. Dorsal fin hyaline, yellow and black coloration, sometimes anterior and dorsal margin black. This typical coloration has been documented extensively by
<bibRefCitation id="EFD6A4D3FF9F8B6DEEC1D5D0FF22401B" author="Alexander" pageId="13" pageNumber="49" refString="Alexander, H. J. &amp; Breden, F. (2004) Sexual isolation and extreme morphological divergence in the Cumana guppy: a possible case of incipient speciation. Journal of Evolutionary Biology, 17, 1238 - 54." type="journal article" year="2004">Alexander &amp; Breden (2004)</bibRefCitation>
for the populations from Central Cumana and we found this also for the Laguna de los Patos (West Cumana) and Campoma populations.
</paragraph>
<paragraph id="8BF8D922FF9F8B6DEAA6D46CFD784395" blockId="13.[151,1437,152,2034]" pageId="13" pageNumber="48">
<emphasis id="B9330530FF9F8B6DEAA6D46CFE86404F" box="[197,322,345,370]" italics="true" pageId="13" pageNumber="48">P. o b s c u r a</emphasis>
was most probably not recognized so far as a separate species, because these fish are not different from
<taxonomicName id="4C47A2A1FF9F8B6DEB39D4B5FE3F40A4" box="[346,507,384,409]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9F8B6DEB39D4B5FE3F40A4" box="[346,507,384,409]" italics="true" pageId="13" pageNumber="48">P. reticulata</emphasis>
</taxonomicName>
in the most obvious character of guppies, namely male coloration and pigmentation pattern (
<figureCitation id="137CC5A7FF9F8B6DEBFFD493FE2040FD" box="[412,484,422,448]" captionStart="FIGURE 2. P" captionStartId="5.[151,255,1910,1934]" captionTargetBox="[187,1405,533,1892]" captionTargetId="figure@5.[187,1409,533,1892]" captionTargetPageId="5" captionText="FIGURE 2. P. obscura male (a) and female (b) from locus typicus, Oropuche River (OR). (c) P. obscura, population Rio Seco (RS); (d) P, reticulata, population Caroni Swamp (CS); (e) P. reticulata, population Rio Yaguaracual (RY); (f) P. c f “ wingei ” from El Cordon waterfall (EC); (g) P. w i n g e i from locus typicus, population Campoma (Ca); (h) P. w i n g e i, population Laguna de los Patos (LP)." httpUri="https://zenodo.org/record/190847/files/figure.png" pageId="13" pageNumber="48">Fig. 2</figureCitation>
a, ce), most of them being linked to sex chromosomes. This is different from
<taxonomicName id="4C47A2A1FF9F8B6DEFE5D492FF2140DB" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="wingei">
<emphasis id="B9330530FF9F8B6DEFE5D492FF2140DB" italics="true" pageId="13" pageNumber="48">P. wingei</emphasis>
</taxonomicName>
, which because of its more bright pigmentation became suspicious to John Endler and was an issue of debate since then. Nevertheless
<emphasis id="B9330530FF9F8B6DE875D4C1FD444330" box="[534,640,500,525]" italics="true" pageId="13" pageNumber="48">P. w i n g e i</emphasis>
males also share many components of their pigmentation patterns with males from the two other species, suggesting that this is an ancestral trait. This is comparable to the situation in the genus
<taxonomicName id="4C47A2A1FF9F8B6DEBFBD774FDEB4367" box="[408,559,577,602]" class="Actinopterygii" family="Poeciliidae" genus="Xiphophorus" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="genus">
<emphasis id="B9330530FF9F8B6DEBFBD774FDEB4367" box="[408,559,577,602]" italics="true" pageId="13" pageNumber="48">Xiphophorus</emphasis>
</taxonomicName>
, were the (mostly sex chromosome linked) macromelanophore pattern can be found in similar expression phenotype in different species; or in the genus
<taxonomicName id="4C47A2A1FF9F8B6DEE0FD75DFB1043BC" box="[1132,1236,616,641]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="genus">
<emphasis id="B9330530FF9F8B6DEE0FD75DFB1043BC" box="[1132,1236,616,641]" italics="true" pageId="13" pageNumber="48">Poecilia</emphasis>
</taxonomicName>
where the male coloration is almost alike in the sailfin mollies.
</paragraph>
<paragraph id="8BF8D922FF9F8B6DEAA6D780FA4844BA" blockId="13.[151,1437,152,2034]" pageId="13" pageNumber="48">
A mentioned above the genetic difference between
<taxonomicName id="4C47A2A1FF9F8B6DE94AD780FC6243F3" box="[809,934,693,718]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9F8B6DE94AD780FC6243F3" box="[809,934,693,718]" italics="true" pageId="13" pageNumber="48">P. obscura</emphasis>
</taxonomicName>
from East
<collectingCountry id="F35099B2FF9F8B6DEE48D780FB4B43F2" box="[1067,1167,693,719]" name="Trinidad and Tobago" pageId="13" pageNumber="48">Trinidad</collectingCountry>
and
<taxonomicName id="4C47A2A1FF9F8B6DEEA8D780FA9E43F3" box="[1227,1370,693,718]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9F8B6DEEA8D780FA9E43F3" box="[1227,1370,693,718]" italics="true" pageId="13" pageNumber="48">P. reticulata</emphasis>
</taxonomicName>
from West
<collectingCountry id="F35099B2FF9F8B6DEABBD7E9FEF843CB" box="[216,316,732,758]" name="Trinidad and Tobago" pageId="13" pageNumber="48">Trinidad</collectingCountry>
has been realized in numerous studies using allozyme, microsatellites and mitochondrial DNA sequences (Carvalho
<emphasis id="B9330530FF9F8B6DEBF0D637FE0A4226" box="[403,462,770,795]" italics="true" pageId="13" pageNumber="48">et al.</emphasis>
, 1991;
<bibRefCitation id="EFD6A4D3FF9F8B6DE84BD637FCB24221" author="Russell" box="[552,886,770,796]" pageId="13" pageNumber="50" refString="Russell, S. T. &amp; Magurran, A. E. (2006) Intrinsic reproductive isolation between Trinidadian populations of the guppy, Poecilia reticulata. Journal of Evolutionary Biology, 19, 1294 - 1303." type="journal article" year="2006">Russell and Magurran, 2006</bibRefCitation>
;
<bibRefCitation id="EFD6A4D3FF9F8B6DE919D637FB124221" author="Alexander" box="[890,1238,770,796]" pageId="13" pageNumber="49" refString="Alexander, H. J. &amp; Breden, F. (2004) Sexual isolation and extreme morphological divergence in the Cumana guppy: a possible case of incipient speciation. Journal of Evolutionary Biology, 17, 1238 - 54." type="journal article" year="2004">Alexander and Breden, 2004</bibRefCitation>
). The absence of obvious morphological criteria and the fact that hybridization between both forms occurs readily in the Turure River as a consequence of the Haskins transplant has led to the conclusion that both forms are rather in an early stage of speciation.
<bibRefCitation id="EFD6A4D3FF9F8B6DEBAED643FCF542AD" author="Russell" box="[461,817,886,912]" pageId="13" pageNumber="50" refString="Russell, S. T. &amp; Magurran, A. E. (2006) Intrinsic reproductive isolation between Trinidadian populations of the guppy, Poecilia reticulata. Journal of Evolutionary Biology, 19, 1294 - 1303." type="journal article" year="2006">Russell and Magurran (2006)</bibRefCitation>
noted that laboratory produced hybrids between
<taxonomicName id="4C47A2A1FF9F8B6DEFE5D642FEC9428B" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9F8B6DEFE5D642FEC9428B" italics="true" pageId="13" pageNumber="48">P. reticulata</emphasis>
</taxonomicName>
from West
<collectingCountry id="F35099B2FF9F8B6DEBFED6A8FDC1428A" box="[413,517,925,951]" name="Trinidad and Tobago" pageId="13" pageNumber="48">Trinidad</collectingCountry>
and
<emphasis id="B9330530FF9F8B6DE822D6A8FD79428B" box="[577,701,925,950]" italics="true" pageId="13" pageNumber="48">P. o b s c u r a</emphasis>
display male behavioral sterility and hybrid breakdown for embryo viability, brood size and sperm counts. This indicates some genetic separation which was interpreted as an initial stage rather than what can be expected from long separated species. The extent to which interspecific hybrids show features of hybrid breakdown can vary, however, and is not especially prominent in other poeciliid species (
<bibRefCitation id="EFD6A4D3FF9F8B6DEBD2D10DFD93456F" author="Schartl" box="[433,599,1080,1106]" pageId="13" pageNumber="50" refString="Schartl, M. (2008) Evolution of Xmrk: an oncogene, but also a speciation gene? Bioessays, 30, 822 - 32." type="journal article" year="2008">Schartl, 2008</bibRefCitation>
). Moreover, interspecific hybrids can be produced in the laboratory of most poeciliids tested so far. Prezygotic isolation mechanisms are especially efficient to prevent interspecific hybridization of naturally co-occurring species of the same genus, but may not exist in allopatric species. Thus the Turure explant might have created an artificial situation of “sympatry” like enforced breeding of allopatric species in the laboratory. A similar situation exists for
<emphasis id="B9330530FF9F8B6DE9C8D1E6FBD245D1" box="[939,1046,1235,1260]" italics="true" pageId="13" pageNumber="48">P. w i n g e i</emphasis>
, which was not recognized as a species for a long time although strong genetic differentiation and differences in male display traits, body shape and coloration were clearly documented (
<bibRefCitation id="EFD6A4D3FF9F8B6DE8AAD015FBE74407" author="Alexander" box="[713,1059,1312,1338]" pageId="13" pageNumber="49" refString="Alexander, H. J. &amp; Breden, F. (2004) Sexual isolation and extreme morphological divergence in the Cumana guppy: a possible case of incipient speciation. Journal of Evolutionary Biology, 17, 1238 - 54." type="journal article" year="2004">Alexander and Breden, 2004</bibRefCitation>
). The presence of hybrids in the Cumana area as a result of human activities showing the Y-chromosome dominated phenotype of
<emphasis id="B9330530FF9F8B6DEF4DD072FA5D445D" box="[1326,1433,1351,1376]" italics="true" pageId="13" pageNumber="48">P. w i n g e i</emphasis>
but having the mitochondrial genome of
<taxonomicName id="4C47A2A1FF9F8B6DE816D058FCC744BB" box="[629,771,1389,1414]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9F8B6DE816D058FCC744BB" box="[629,771,1389,1414]" italics="true" pageId="13" pageNumber="48">P. reticulata</emphasis>
</taxonomicName>
might have confused the interpretation of the analyses.
</paragraph>
</subSubSection>
<subSubSection id="C35D8AA9FF9F8B6EEAA6D0A6FC54404E" lastPageId="14" lastPageNumber="49" pageId="13" pageNumber="48" type="distribution">
<paragraph id="8BF8D922FF9F8B6DEAA6D0A6FEE7466A" blockId="13.[151,1437,152,2034]" pageId="13" pageNumber="48">
<emphasis id="B9330530FF9F8B6DEAA6D0A6FEAE4490" bold="true" box="[197,362,1427,1453]" pageId="13" pageNumber="48">Distribution.</emphasis>
<taxonomicName id="4C47A2A1FF9F8B6DEB11D0A1FE364490" box="[370,498,1428,1453]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9F8B6DEB11D0A1FE364490" box="[370,498,1428,1453]" italics="true" pageId="13" pageNumber="48">P. obscura</emphasis>
</taxonomicName>
occurs in freshwaters of the Oropuche River system, North-eastern
<collectingCountry id="F35099B2FF9F8B6DEF4DD0A1FA5D4493" box="[1326,1433,1428,1454]" name="Trinidad and Tobago" pageId="13" pageNumber="48">Trinidad</collectingCountry>
. The species was found (confirmed by DNA sequencing) in the headwaters and upper regions of the northern tributaries to the Oropuche River including the Quare River and La Seiva River. It was also found in the Matura River, the Salybia /Rio Seco Rivers and the LEbranche River, which are not tributaries to the Oropuche River but flow independently next north and south into the Atlantic Ocean at the West Coast of
<collectingCountry id="F35099B2FF9F8B6DEAF4D360FEC14752" box="[151,261,1621,1647]" name="Trinidad and Tobago" pageId="13" pageNumber="48">Trinidad</collectingCountry>
. The southern and northern boarder of the range of the new species have not been determined exactly, but there are guppies in the Tompire River that flows north of the Matura River and in the Mission River, which flows into the Caribbean Sea (Reznick, pers. comm.). Further west on the northern coastline all the rivers that flow into the Caribbean Sea have no guppies until the Madamas River is reached (Reznick, pers. comm.). This river belongs to the northwestern province, the guppies of which - according to the molecular phylogeny - clearly are
<taxonomicName id="4C47A2A1FF9F8B6DE85CD222FD10460D" box="[575,724,1815,1840]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9F8B6DE85CD222FD10460D" box="[575,724,1815,1840]" italics="true" pageId="13" pageNumber="48">P. reticulata</emphasis>
</taxonomicName>
. No information is available for the situation south of the LEbranche.
</paragraph>
<paragraph id="8BF8D922FF9F8B6EEAA6D251FC54404E" blockId="13.[151,1437,152,2034]" lastBlockId="14.[151,1436,152,371]" lastPageId="14" lastPageNumber="49" pageId="13" pageNumber="48">
In 1957 C.P. Haskins released
<specimenCount id="9D4112ABFF9F8B6DE859D251FD424640" box="[570,646,1892,1918]" pageId="13" pageNumber="48" type="generic">
200
<taxonomicName id="4C47A2A1FF9F8B6DE813D251FD424640" box="[624,646,1892,1917]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9F8B6DE813D251FD424640" box="[624,646,1892,1917]" italics="true" pageId="13" pageNumber="48">P.</emphasis>
</taxonomicName>
</specimenCount>
reticulata from the lower Guanapo River of the Caroni drainage (West
<collectingCountry id="F35099B2FF9F8B6DEA82D2BEFE834698" box="[225,327,1931,1957]" name="Trinidad and Tobago" pageId="13" pageNumber="48">Trinidad</collectingCountry>
) into a site of the upper Turure (
<bibRefCitation id="EFD6A4D3FF9F8B6DE8DED2BEFCBB4698" author="Magurran" box="[701,895,1931,1957]" pageId="13" pageNumber="50" refString="Magurran, A. E. (2005) Evolutionary Ecology - The Trinidadian Guppy. Oxford University Press, New York, 224 pp." type="book" year="2005">Magurran, 2005</bibRefCitation>
), a northern tributary to the lower Quare River that flows into the lower Oropuche. These fish spread out from there downstream in the Turure (
<bibRefCitation id="EFD6A4D3FF9F8B6DEF6DD284FF1746CF" author="Shaw" pageId="13" pageNumber="50" refString="Shaw, P. W., Carvalho, G. R., Seghers, B. H. &amp; Magurran, A. E. (1992) Genetic consequences of an artificial introduction of guppies (Poecilia reticulata) in N. Trinidad. Philosophical Transactions of The Royal Society London B Biological Sciences, 248, 111 - 116." type="journal article" year="1992">
Shaw
<emphasis id="B9330530FF9F8B6DEF3AD284FA5146F7" box="[1369,1429,1969,1994]" italics="true" pageId="13" pageNumber="48">et al.</emphasis>
, 1992
</bibRefCitation>
) (
<bibRefCitation id="EFD6A4D3FF9F8B6DEA88D2EDFE6A46CF" author="Magurran" box="[235,430,2008,2034]" pageId="13" pageNumber="50" refString="Magurran, A. E. (2005) Evolutionary Ecology - The Trinidadian Guppy. Oxford University Press, New York, 224 pp." type="book" year="2005">Magurran, 2005</bibRefCitation>
). We noted by presence of the
<taxonomicName id="4C47A2A1FF9F8B6DE97AD2EDFC6346CC" box="[793,935,2008,2033]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="13" pageNumber="48" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9F8B6DE97AD2EDFC6346CC" box="[793,935,2008,2033]" italics="true" pageId="13" pageNumber="48">P. reticulata</emphasis>
</taxonomicName>
mitochondrial haplotype (data not shown) that the invaders have already reached the main river and could threat the newly described species either by displacement or hybridization. Hybrids between
<taxonomicName id="4C47A2A1FF9C8B6EE8BAD58AFCAD41E5" box="[729,873,191,216]" class="Actinopterygii" family="Poeciliidae" genus="Poecilia" kingdom="Animalia" order="Cyprinodontiformes" pageId="14" pageNumber="49" phylum="Chordata" rank="species" species="reticulata">
<emphasis id="B9330530FF9C8B6EE8BAD58AFCAD41E5" box="[729,873,191,216]" italics="true" pageId="14" pageNumber="49">P. reticulata</emphasis>
</taxonomicName>
from the Caroni system (West
<collectingCountry id="F35099B2FF9C8B6EEEBED58BFA8041E5" box="[1245,1348,190,216]" name="Trinidad and Tobago" pageId="14" pageNumber="49">Trinidad</collectingCountry>
) and
<taxonomicName id="4C47A2A1FF9C8B6EEFE5D58AFF3F41C3" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="14" pageNumber="49" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9C8B6EEFE5D58AFF3F41C3" italics="true" pageId="14" pageNumber="49">P. obscura</emphasis>
</taxonomicName>
are easily produced in the laboratory (
<bibRefCitation id="EFD6A4D3FF9C8B6EE886D5D0FB8941C2" author="Russell" box="[741,1101,229,255]" pageId="14" pageNumber="50" refString="Russell, S. T. &amp; Magurran, A. E. (2006) Intrinsic reproductive isolation between Trinidadian populations of the guppy, Poecilia reticulata. Journal of Evolutionary Biology, 19, 1294 - 1303." type="journal article" year="2006">Russell and Magurran, 2006</bibRefCitation>
) and a gene flow from the transplantation site into the native population of the Turure was already noted in a microsatellite study (after Russel 2004, cited in
<bibRefCitation id="EFD6A4D3FF9C8B6EEBFAD407FD984071" author="Magurran" box="[409,604,306,332]" pageId="14" pageNumber="50" refString="Magurran, A. E. (2005) Evolutionary Ecology - The Trinidadian Guppy. Oxford University Press, New York, 224 pp." type="book" year="2005">Magurran, 2005</bibRefCitation>
)). Due to this problem the dispersal of
<taxonomicName id="4C47A2A1FF9C8B6EEE53D407FB684076" box="[1072,1196,306,331]" class="Aves" family="Paridae" genus="Poecila" higherTaxonomySource="GBIF" kingdom="Animalia" order="Passeriformes" pageId="14" pageNumber="49" phylum="Chordata" rank="species" species="obscura">
<emphasis id="B9330530FF9C8B6EEE53D407FB684076" box="[1072,1196,306,331]" italics="true" pageId="14" pageNumber="49">P. obscura</emphasis>
</taxonomicName>
in the main river of the Oropuche system including its southern tributaries is unclear.
</paragraph>
</subSubSection>
</treatment>
</document>