453 lines
58 KiB
XML
453 lines
58 KiB
XML
<document id="B7492753E69F039AB737472196CB8F85" ID-DOI="10.1093/zoolinnean/zlad034" ID-ISSN="0024-4082" ID-Zenodo-Dep="8326382" IM.bibliography_approvedBy="juliana" IM.illustrations_approvedBy="juliana" IM.materialsCitations_approvedBy="juliana" IM.metadata_approvedBy="felipe" IM.tables_approvedBy="juliana" IM.taxonomicNames_approvedBy="juliana" IM.treatments_approvedBy="juliana" checkinTime="1693804754615" checkinUser="plazi" docAuthor="Jennings, W. Bryan, Souto-Santos, Igor C. A., Buckup, Paulo A. & Zandonà, Eugenia" docDate="2023" docId="03E187BCB06CFFE620FD1155FDBCFA43" docLanguage="en" docName="zlad034.pdf" docOrigin="Zoological Journal of the Linnean Society 199 (1)" docSource="http://dx.doi.org/10.1093/zoolinnean/zlad034" docStyle="DocumentStyle:4F230B9370E98E256D973D6DFB57F36C.1:ZoolJLinnSoc.2023-.journal_article" docStyleId="4F230B9370E98E256D973D6DFB57F36C" docStyleIdOld="36B3BD6A90C22AB4F7F465C853188CC8" docStyleName="ZoolJLinnSoc.2023-.journal_article" docStyleNameOld="ZoolJLinnSoc.2017-.journal_article" docStyleOld="DocumentStyle:36B3BD6A90C22AB4F7F465C853188CC8.7:ZoolJLinnSoc.2017-.journal_article" docStyleVersion="1" docStyleVersionOld="7" docTitle="Phalloceros Eigenmann 1907" docType="treatment" docVersion="1" lastPageNumber="214" masterDocId="FFD8FFC4B06AFFEE2375143CFFC7FFCE" masterDocTitle="Do mismatched genitalia promote mechanical isolation in syntopic species of Phalloceros fishes (Poeciliidae)?" masterLastPageNumber="215" masterPageNumber="206" pageNumber="212" updateTime="1694179649428" updateUser="juliana">
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<mods:title id="CD544CBD9ABF997F793984638C1D2E90">Do mismatched genitalia promote mechanical isolation in syntopic species of Phalloceros fishes (Poeciliidae)?</mods:title>
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<mods:roleTerm id="EF995ACC86EAFB41AF53F58D247CD477">Author</mods:roleTerm>
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<mods:namePart id="B5AA115274DC2D84FB3B7B6EB3752EA8">Jennings, W. Bryan</mods:namePart>
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<mods:affiliation id="2E1ACD9364468012EA88F9089DCAF926">Departamento de Ecologia, Programa de Pós-Graduação em Ecologia e Evolução, Universidade do Estado do Rio de Janeiro, 20550 - 013 Rio de Janeiro, RJ, Brazil & Museu Nacional, Departamento de Vertebrados, Universidade Federal do Rio de Janeiro, ºuinta da Boa Vista, 20490 - 040 Rio de Janeiro, RJ, Brazil & Department of Evolution, Ecology, and Organismic Biology, University of California, Riverside, Riverside, CA 92521, USA</mods:affiliation>
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<mods:nameIdentifier id="E8C542560371461078983FE1B8DB2121" type="email">bryan.jennings@ucr.edu</mods:nameIdentifier>
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<mods:namePart id="63FE1BD4D803E9B2BCE472DE9D96509C">Souto-Santos, Igor C. A.</mods:namePart>
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<mods:affiliation id="17079E2B64D8164D482130365982A533">Museu Nacional, Departamento de Vertebrados, Universidade Federal do Rio de Janeiro, ºuinta da Boa Vista, 20490 - 040 Rio de Janeiro, RJ, Brazil</mods:affiliation>
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<mods:namePart id="E52FFF846B598B92185A1440E5CEBA49">Buckup, Paulo A.</mods:namePart>
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<mods:affiliation id="1A7441E988A10D5887BC50D60071E67F">Museu Nacional, Departamento de Vertebrados, Universidade Federal do Rio de Janeiro, ºuinta da Boa Vista, 20490 - 040 Rio de Janeiro, RJ, Brazil</mods:affiliation>
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<mods:namePart id="E62CA96F1E8651F01CE4A81B29350D74">Zandonà, Eugenia</mods:namePart>
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<mods:affiliation id="F551B4E4B86A30743A1937777F109C10">Departamento de Ecologia, Programa de Pós-Graduação em Ecologia e Evolução, Universidade do Estado do Rio de Janeiro, 20550 - 013 Rio de Janeiro, RJ, Brazil</mods:affiliation>
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<mods:title id="8605FBAF7E0BFAA1B2FD5DFBBC5F8CEE">Zoological Journal of the Linnean Society</mods:title>
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<mods:part id="83EE283DC6AA7136530C04467AF83A2C">
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<mods:date id="2209BBE501AB00FE653CC211ABB1DDFC">2023</mods:date>
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<mods:number id="1E2D05D92547521068322908527015D6">2023-06-29</mods:number>
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<mods:number id="C9DEE266E6C63480DD1EFEEA73C17E3F">199</mods:number>
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<mods:identifier id="C94E55C34F765B3AA21B177F4232E74F" type="DOI">10.1093/zoolinnean/zlad034</mods:identifier>
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<mods:identifier id="79DA39FCD3CCE4521A714B1AF39479E8" type="ISSN">0024-4082</mods:identifier>
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<mods:identifier id="FAC33BFD977BFE7F18497B1EBC11EE6D" type="Zenodo-Dep">8326382</mods:identifier>
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<treatment id="03E187BCB06CFFE620FD1155FDBCFA43" LSID="urn:lsid:plazi:treatment:03E187BCB06CFFE620FD1155FDBCFA43" httpUri="http://treatment.plazi.org/id/03E187BCB06CFFE620FD1155FDBCFA43" lastPageId="8" lastPageNumber="214" pageId="6" pageNumber="212">
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<subSubSection id="C3526521B06CFFE820FD1155FA92FA4D" box="[904,1365,1385,1412]" pageId="6" pageNumber="212" type="nomenclature">
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<paragraph id="8BF736AAB06CFFE820FD1155FA92FA4D" blockId="6.[810,1460,1385,1980]" box="[904,1365,1385,1412]" pageId="6" pageNumber="212">
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<emphasis id="B93CEAB8B06CFFE820FD1155FA92FA4D" bold="true" box="[904,1365,1385,1412]" pageId="6" pageNumber="212">
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Low genetic diversity in
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<taxonomicName id="4C484D29B06CFFE827F21155FB3DFA4A" ID-CoL="6MQG" ID-ENA="55107" authorityName="Eigenmann" authorityYear="1907" box="[1159,1274,1385,1412]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" pageNumber="212" phylum="Chordata" rank="genus">
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<emphasis id="B93CEAB8B06CFFE827F21155FB3DFA4A" bold="true" box="[1159,1274,1385,1412]" italics="true" pageId="6" pageNumber="212">Phalloceros</emphasis>
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species?
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</emphasis>
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</paragraph>
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<subSubSection id="C3526521B06CFFE6205F11ACFDBCFA43" lastPageId="8" lastPageNumber="214" pageId="6" pageNumber="212" type="description">
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<paragraph id="8BF736AAB06CFFE8205F11ACFC21F872" blockId="6.[810,1460,1385,1980]" pageId="6" pageNumber="212">
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Another noteworthy finding from our molecular results concerns the low genetic diversity of the populations we studied. Despite having obtained mtDNA sequences from 31 individuals of
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<taxonomicName id="4C484D29B06CFFE8200011D2FBC2F9C8" box="[885,1029,1518,1542]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" pageNumber="212" phylum="Chordata" rank="species" species="anisophallos">
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<emphasis id="B93CEAB8B06CFFE8200011D2FBC2F9C8" box="[885,1029,1518,1542]" italics="true" pageId="6" pageNumber="212">P. anisophallos</emphasis>
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</taxonomicName>
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and eight of
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<taxonomicName id="4C484D29B06CFFE827FA11D2FACFF9C8" box="[1167,1288,1518,1542]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="6" pageNumber="212" phylum="Chordata" rank="species" species="leptokeras">
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<emphasis id="B93CEAB8B06CFFE827FA11D2FACFF9C8" box="[1167,1288,1518,1542]" italics="true" pageId="6" pageNumber="212">P. leptokeras</emphasis>
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</taxonomicName>
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from the Parati-Mirim River, all sampled individuals within each of these species had the same haplotype sequence (
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<figureCitation id="13732A2FB06CFFE827931210FAE7F98A" box="[1254,1312,1580,1604]" captionStart="Figure 2" captionStartId="4.[114,179,882,906]" captionTargetBox="[149,1426,144,837]" captionTargetId="figure-598@4.[146,1326,170,854]" captionTargetPageId="4" captionText="Figure 2. Bayesian mitochondrial (mtDNA) and nuclear (nDNA) gene trees, showing the inferred relationships among haplotypes of Phalloceros anisophallos and Phalloceros leptokeras from the Parati-Mirim River and other localities.A, mtDNA tree for the COI sequences. B, nDNA tree for the MG1 sequences.Shaded names refer to samples misidentified based on morphological criteria (see Materials and Methods).Numbers in parentheses represent the number of identical haplotypes. Other species of Phalloceros were included as reference samples, and Cnesterodon decemmaculatus and Poecilia vivipara were used to root the trees.Phalloceros anisophallos and Phalloceros leptokeras are sympatric in the Parati-Mirim and Japuíba rivers.São Roque and Paquequer correspond to the type localities of P. anisophallos and P.leptokeras, respectively (Lucinda 2008). Numbers above each branch are the estimated Bayesian posterior probabilities." figureDoi="http://doi.org/10.5281/zenodo.8326386" httpUri="https://zenodo.org/record/8326386/files/figure.png" pageId="6" pageNumber="212">Fig. 2</figureCitation>
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). A perusal of several other drainages that had multiple individuals sampled for the mtDNA gene showed the same pattern (i.e. Itamambuca, Puruba, ºuiririm, Paquequer, Macacu and Perequê Açu Rivers;
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<figureCitation id="13732A2FB06CFFE8205F1296FCBEF90C" box="[810,889,1706,1730]" captionStart="Figure 2" captionStartId="4.[114,179,882,906]" captionTargetBox="[149,1426,144,837]" captionTargetId="figure-598@4.[146,1326,170,854]" captionTargetPageId="4" captionText="Figure 2. Bayesian mitochondrial (mtDNA) and nuclear (nDNA) gene trees, showing the inferred relationships among haplotypes of Phalloceros anisophallos and Phalloceros leptokeras from the Parati-Mirim River and other localities.A, mtDNA tree for the COI sequences. B, nDNA tree for the MG1 sequences.Shaded names refer to samples misidentified based on morphological criteria (see Materials and Methods).Numbers in parentheses represent the number of identical haplotypes. Other species of Phalloceros were included as reference samples, and Cnesterodon decemmaculatus and Poecilia vivipara were used to root the trees.Phalloceros anisophallos and Phalloceros leptokeras are sympatric in the Parati-Mirim and Japuíba rivers.São Roque and Paquequer correspond to the type localities of P. anisophallos and P.leptokeras, respectively (Lucinda 2008). Numbers above each branch are the estimated Bayesian posterior probabilities." figureDoi="http://doi.org/10.5281/zenodo.8326386" httpUri="https://zenodo.org/record/8326386/files/figure.png" pageId="6" pageNumber="212">Fig. 2A</figureCitation>
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). Only the Grande and Indaiá Rivers showed more than one distinctive haplotype sequence (
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<bibRefCitation id="EFD94B5BB06CFFE8279912F5FC99F8CE" author="Souto-Santos ICA & Jennings WB & Buckup PA" pageId="6" pageNumber="212" pagination="514 - 31" refId="ref10329" refString="Souto-Santos ICA, Jennings WB, Buckup PA. Testing palaeodrainage hypotheses in south-eastern Brazil: phylogeography of the sinistral livebearerfishofthegenus Phalloceros (Cyprinodontiformes: Poeciliidae). Zoological Journal of the Linnean Society 2023; 197: 514 - 31." type="journal article" year="2023">
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Souto-Santos
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<emphasis id="B93CEAB8B06CFFE826F412F6FA73F92F" box="[1409,1460,1737,1761]" italics="true" pageId="6" pageNumber="212">et al.</emphasis>
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2023
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</bibRefCitation>
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). These occurrences might have been attributable to gene flow that occurred when the sea level was low enough to join these adjacent rivers into one or they might represent ancestral polymorphisms that originated in the ancestral population for the two rivers. However, the low genetic diversity seen in each of these isolated populations could have arisen via bottlenecks or founder events.
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</paragraph>
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<paragraph id="8BF736AAB06DFFE923BC14ACFD07FF64" blockId="7.[128,778,143,1741]" box="[201,704,143,170]" pageId="7" pageNumber="213">
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<emphasis id="B93CEAB8B06DFFE923BC14ACFD07FF64" bold="true" box="[201,704,143,170]" pageId="7" pageNumber="213">
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Problems with identifying species of
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<taxonomicName id="4C484D29B06DFFE9213814B3FD07FF64" box="[589,704,143,170]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
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<emphasis id="B93CEAB8B06DFFE9213814B3FD07FF64" bold="true" box="[589,704,143,170]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
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</taxonomicName>
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</emphasis>
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</paragraph>
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<paragraph id="8BF736AAB06DFFE923F4148BFEA2FE07" blockId="7.[128,778,143,1741]" pageId="7" pageNumber="213">
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We also found that intraspecific variation in morphology in these species can lead to incorrect species IDs, especially with immature individuals.This latter finding is of importance to researchers who study
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<taxonomicName id="4C484D29B06DFFE923861528FEA6FEE2" box="[243,353,276,300]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
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<emphasis id="B93CEAB8B06DFFE923861528FEA6FEE2" box="[243,353,276,300]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
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</taxonomicName>
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because it shows that morphology alone might be insufficient to identify individuals in systems with multiple co-occurring species of
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<taxonomicName id="4C484D29B06DFFE922C6156FFDE6FEA5" box="[435,545,339,363]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
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<emphasis id="B93CEAB8B06DFFE922C6156FFDE6FEA5" box="[435,545,339,363]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
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</taxonomicName>
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. However, our results show that standard mtDNA barcode sequences and our newly developed nDNA barcode marker provide reliable species IDs for
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<taxonomicName id="4C484D29B06DFFE923D6158DFEA2FE07" box="[163,357,433,457]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="undetermined">
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<emphasis id="B93CEAB8B06DFFE923D6158DFED6FE07" box="[163,273,433,457]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
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species.
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</taxonomicName>
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</paragraph>
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<paragraph id="8BF736AAB06DFFE923E915ECFED3FC13" blockId="7.[128,778,143,1741]" pageId="7" pageNumber="213">
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Of the
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<specimenCount id="9D4EFD23B06DFFE9239215ECFEFAFE26" box="[231,317,464,488]" pageId="7" pageNumber="213" type="adult">30 adult</specimenCount>
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(
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<specimenCount id="9D4EFD23B06DFFE9223915ECFE72FE26" box="[332,437,464,488]" pageId="7" pageNumber="213" type="female">25 female</specimenCount>
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and
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<specimenCount id="9D4EFD23B06DFFE9229F15ECFD8EFE26" box="[490,585,464,488]" pageId="7" pageNumber="213" type="male">five male</specimenCount>
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) specimens in our study, only one (MNTI 14559) was misidentified using genital characters. This female individual was misidentified initially as
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<taxonomicName id="4C484D29B06DFFE923F41613FED5FD88" box="[129,274,558,582]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="anisophallos">
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<emphasis id="B93CEAB8B06DFFE923F41613FED5FD88" box="[129,274,558,582]" italics="true" pageId="7" pageNumber="213">P. anisophallos</emphasis>
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</taxonomicName>
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because it had a right-turned papilla. However, our re-examination of this specimen revealed that the papilla on this individual was aberrant; instead of pointing distally, as is the usual case for
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<taxonomicName id="4C484D29B06DFFE9226616B1FE64FD6A" box="[275,419,652,676]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="anisophallos">
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<emphasis id="B93CEAB8B06DFFE9226616B1FE64FD6A" box="[275,419,652,676]" italics="true" pageId="7" pageNumber="213">P. anisophallos</emphasis>
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</taxonomicName>
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, it was pointed in a latero-anterior direction like the normal condition for mature
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<taxonomicName id="4C484D29B06DFFE921181690FD20FD0D" box="[621,743,683,707]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="leptokeras">
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<emphasis id="B93CEAB8B06DFFE921181690FD20FD0D" box="[621,743,683,707]" italics="true" pageId="7" pageNumber="213">P. leptokeras</emphasis>
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</taxonomicName>
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females. Moreover, the papilla had a fleshy process protruding outwards from its anterior side, which is a unique characteristic of
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<taxonomicName id="4C484D29B06DFFE923BB1736FDA6FCEC" authority="(Souto-Santos et al. 2023)" baseAuthorityName="Souto-Santos" baseAuthorityYear="2023" box="[206,609,777,802]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="leptokeras">
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<emphasis id="B93CEAB8B06DFFE923BB1736FE80FCEF" box="[206,327,777,801]" italics="true" pageId="7" pageNumber="213">P. leptokeras</emphasis>
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(
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<bibRefCitation id="EFD94B5BB06DFFE9222C1736FD92FCEC" author="Souto-Santos ICA & Jennings WB & Buckup PA" box="[345,597,777,802]" pageId="7" pageNumber="213" pagination="514 - 31" refId="ref10329" refString="Souto-Santos ICA, Jennings WB, Buckup PA. Testing palaeodrainage hypotheses in south-eastern Brazil: phylogeography of the sinistral livebearerfishofthegenus Phalloceros (Cyprinodontiformes: Poeciliidae). Zoological Journal of the Linnean Society 2023; 197: 514 - 31." type="journal article" year="2023">
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||
Souto-Santos
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||
<emphasis id="B93CEAB8B06DFFE9229F1736FDDCFCEF" box="[490,539,777,801]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
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2023
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</bibRefCitation>
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)
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</taxonomicName>
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. Although aberrantly shaped papillae have been reported for
|
||
<taxonomicName id="4C484D29B06DFFE9212D1716FD0FFC8F" authorityName="Lucinda" authorityYear="2008" box="[600,712,809,833]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="harpagos">
|
||
<emphasis id="B93CEAB8B06DFFE9212D1716FD0FFC8F" box="[600,712,809,833]" italics="true" pageId="7" pageNumber="213">P. harpagos</emphasis>
|
||
</taxonomicName>
|
||
, there is negligible intraspecific variation in this trait or in the shapes of species-specific gonopodia (
|
||
<bibRefCitation id="EFD94B5BB06DFFE922CE175BFD8EFC4E" author="Lucinda PHF" box="[443,585,871,896]" pageId="7" pageNumber="213" pagination="113 - 58" refId="ref9750" refString="Lucinda PHF. Systematics and biogeography of the genus Phalloceros Eigenmann, 1907 (Cyprinodontiformes: Poeciliidae: Poeciliinae), with the description of twenty-one new species. Neotropical Ichthyology 2008; 6: 113 - 58." type="journal article" year="2008">Lucinda 2008</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFD94B5BB06DFFE921231754FF73FC51" author="Ono RH & Shibaua OA" pageId="7" pageNumber="213" pagination="22 - 5" refId="ref9938" refString="Ono RH, Shibaua OA. Avaliacao de caracteres taxonomicos diagnosticos do aparelho genital de Phalloceros harpagos Lucinda, 2008 (Cyprinodontiformes: Poeciliidae). Boletim Sociedade Brasileira de Ictiologia 2015; 115: 22 - 5." type="journal article" year="2015">Ono and Shibatta 2015</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="EFD94B5BB06DFFE923B517BBFEB9FC51" author="Thomaz AT & Carvalho TP & Malabarba LR" box="[192,382,903,927]" pageId="7" pageNumber="213" pagination="265 - 74" refId="ref10431" refString="Thomaz AT, Carvalho TP, Malabarba LR, et al. Geographic distributions, phenotypes, and phylogenetic relationships of Phalloceros (Cyprinodontiformes: Poeciliidae): insights about diversification among sympatric species pools. Molecular Phylogenetics and Evolution 2019; 132: 265 - 74." type="journal article" year="2019">
|
||
Thomaz
|
||
<emphasis id="B93CEAB8B06DFFE9226D17B4FE81FC51" box="[280,326,903,927]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
|
||
2019
|
||
</bibRefCitation>
|
||
). Thus, our study supports the practice of using genitalia to identify adult individuals of
|
||
<taxonomicName id="4C484D29B06DFFE9211D179BFD19FC70" box="[616,734,934,958]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="leptokeras">
|
||
<emphasis id="B93CEAB8B06DFFE9211D179BFD19FC70" box="[616,734,934,958]" italics="true" pageId="7" pageNumber="213">P. leptokeras</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C484D29B06DFFE923F417FAFEC9FC13" box="[129,270,965,989]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="anisophallos">
|
||
<emphasis id="B93CEAB8B06DFFE923F417FAFEC9FC13" box="[129,270,965,989]" italics="true" pageId="7" pageNumber="213">P. anisophallos</emphasis>
|
||
</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="8BF736AAB06DFFE923E917D9FD62F903" blockId="7.[128,778,143,1741]" pageId="7" pageNumber="213">
|
||
Although our results showed that adult
|
||
<taxonomicName id="4C484D29B06DFFE9213717D9FD77FC33" box="[578,688,997,1021]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE9213717D9FD77FC33" box="[578,688,997,1021]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
individuals from the Parati-Mirim River can be identified reliably as
|
||
<taxonomicName id="4C484D29B06DFFE923F41018FED4FBF5" box="[129,275,1059,1083]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="anisophallos">
|
||
<emphasis id="B93CEAB8B06DFFE923F41018FED4FBF5" box="[129,275,1059,1083]" italics="true" pageId="7" pageNumber="213">P. anisophallos</emphasis>
|
||
</taxonomicName>
|
||
or
|
||
<taxonomicName id="4C484D29B06DFFE9224B1018FE7DFBF5" box="[318,442,1059,1083]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="leptokeras">
|
||
<emphasis id="B93CEAB8B06DFFE9224B1018FE7DFBF5" box="[318,442,1059,1083]" italics="true" pageId="7" pageNumber="213">P. leptokeras</emphasis>
|
||
</taxonomicName>
|
||
based on the genital traits, we could not use these morphological features to identify the immature individuals in our sample because their genitals were either not visible (in juveniles) or they were underdeveloped (in subadults). We therefore identified these individuals provisionally using the presence or absence of dark lateral spots, following
|
||
<bibRefCitation id="EFD94B5BB06DFFE923F410E3FEE1FB39" author="Lucinda PHF" box="[129,294,1247,1271]" pageId="7" pageNumber="213" pagination="113 - 58" refId="ref9750" refString="Lucinda PHF. Systematics and biogeography of the genus Phalloceros Eigenmann, 1907 (Cyprinodontiformes: Poeciliidae: Poeciliinae), with the description of twenty-one new species. Neotropical Ichthyology 2008; 6: 113 - 58." type="journal article" year="2008">Lucinda (2008)</bibRefCitation>
|
||
. However, substantial intraspecific variation in this trait explains the poor correlation between the presence of spots and identities of
|
||
<taxonomicName id="4C484D29B06DFFE922001122FDF2FAF8" box="[373,565,1310,1334]" pageId="7" pageNumber="213">
|
||
<emphasis id="B93CEAB8B06DFFE922001122FE24FAF8" box="[373,483,1310,1334]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
species
|
||
</taxonomicName>
|
||
in the Parati-Mirim River. This practice led us to misidentify four of the
|
||
<specimenCount id="9D4EFD23B06DFFE921E11101FF6DFABA" pageId="7" pageNumber="213" type="immature">nine immature</specimenCount>
|
||
individuals in our sample. Most surprising of all was that 72% of our
|
||
<taxonomicName id="4C484D29B06DFFE923811141FE45FA5A" box="[244,386,1404,1428]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="anisophallos">
|
||
<emphasis id="B93CEAB8B06DFFE923811141FE45FA5A" box="[244,386,1404,1428]" italics="true" pageId="7" pageNumber="213">P. anisophallos</emphasis>
|
||
</taxonomicName>
|
||
specimens lacked spots, which contradicts
|
||
<bibRefCitation id="EFD94B5BB06DFFE923C011A7FEA3FA7D" author="Lucinda PHF" box="[181,356,1435,1459]" pageId="7" pageNumber="213" pagination="113 - 58" refId="ref9750" refString="Lucinda PHF. Systematics and biogeography of the genus Phalloceros Eigenmann, 1907 (Cyprinodontiformes: Poeciliidae: Poeciliinae), with the description of twenty-one new species. Neotropical Ichthyology 2008; 6: 113 - 58." type="journal article" year="2008">Lucinda’s (2008)</bibRefCitation>
|
||
assertion that the presence of lateral spots is a diagnostic character for this species. Although we also discovered that individuals of
|
||
<taxonomicName id="4C484D29B06DFFE922EA11E7FDDEFA3C" box="[415,537,1498,1522]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="leptokeras">
|
||
<emphasis id="B93CEAB8B06DFFE922EA11E7FDDEFA3C" box="[415,537,1498,1522]" italics="true" pageId="7" pageNumber="213">P. leptokeras</emphasis>
|
||
</taxonomicName>
|
||
from the Parati-Mirim River departed from the expected no-spot pattern (
|
||
<bibRefCitation id="EFD94B5BB06DFFE921C011C5FF72F9FF" author="Lucinda PHF" pageId="7" pageNumber="213" pagination="113 - 58" refId="ref9750" refString="Lucinda PHF. Systematics and biogeography of the genus Phalloceros Eigenmann, 1907 (Cyprinodontiformes: Poeciliidae: Poeciliinae), with the description of twenty-one new species. Neotropical Ichthyology 2008; 6: 113 - 58." type="journal article" year="2008">Lucinda 2008</bibRefCitation>
|
||
), our observations that individuals of this species can show variable spotting patterns (see
|
||
<tableCitation id="C6CA0311B06DFFE922C11204FE38F99E" box="[436,511,1592,1616]" captionStart="Table 1" captionStartId="6.[113,168,143,167]" captionTargetPageId="6" captionText="Table 1. Presence of dark lateral spots on individuals of Phalloceros anisophallos and Phalloceros leptokeras from the Parati-Mirim River." pageId="7" pageNumber="213">Table 1</tableCitation>
|
||
) confirms the earlier findings of
|
||
<bibRefCitation id="EFD94B5BB06DFFE923A4126BFE2EF9A1" author="Souto-Santos ICA & Jennings WB & Buckup PA" box="[209,489,1623,1647]" pageId="7" pageNumber="213" pagination="514 - 31" refId="ref10329" refString="Souto-Santos ICA, Jennings WB, Buckup PA. Testing palaeodrainage hypotheses in south-eastern Brazil: phylogeography of the sinistral livebearerfishofthegenus Phalloceros (Cyprinodontiformes: Poeciliidae). Zoological Journal of the Linnean Society 2023; 197: 514 - 31." type="journal article" year="2023">
|
||
Souto-Santos
|
||
<emphasis id="B93CEAB8B06DFFE922111264FE50F9A1" box="[356,407,1623,1647]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
|
||
(2023)
|
||
</bibRefCitation>
|
||
. Thus, an important implication of our study for
|
||
<taxonomicName id="4C484D29B06DFFE92204124AFE18F940" box="[369,479,1654,1678]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE92204124AFE18F940" box="[369,479,1654,1678]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
researchers is that immature individuals of these two species from the Parati-Mirim River can be identified only using molecular barcode sequences.
|
||
</paragraph>
|
||
<paragraph id="8BF736AAB06DFFE923D612C6FD20F8DD" blockId="7.[129,777,1785,1973]" box="[163,743,1785,1812]" pageId="7" pageNumber="213">
|
||
<emphasis id="B93CEAB8B06DFFE923D612C6FD20F8DD" bold="true" box="[163,743,1785,1812]" pageId="7" pageNumber="213">
|
||
How prevalent is syntopy among species of
|
||
<taxonomicName id="4C484D29B06DFFE9211F12C5FD1AF8DA" box="[618,733,1785,1812]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE9211F12C5FD1AF8DA" bold="true" box="[618,733,1785,1812]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
?
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph id="8BF736AAB06DFFE923F4131CFB8FFCB7" blockId="7.[129,777,1785,1973]" lastBlockId="7.[825,1474,144,1452]" pageId="7" pageNumber="213">
|
||
<bibRefCitation id="EFD94B5BB06DFFE923F4131CFED9F8F6" author="Lucinda PHF" box="[129,286,1824,1848]" pageId="7" pageNumber="213" pagination="113 - 58" refId="ref9750" refString="Lucinda PHF. Systematics and biogeography of the genus Phalloceros Eigenmann, 1907 (Cyprinodontiformes: Poeciliidae: Poeciliinae), with the description of twenty-one new species. Neotropical Ichthyology 2008; 6: 113 - 58." type="journal article" year="2008">Lucinda (2008</bibRefCitation>
|
||
: p. 156) noted that in
|
||
<taxonomicName id="4C484D29B06DFFE92168131CFD4CF8F6" box="[541,651,1824,1848]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE92168131CFD4CF8F6" box="[541,651,1824,1848]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
, ‘some species are sympatric or even syntopic’, and then listed the pairs
|
||
<emphasis id="B93CEAB8B06DFFE923F41363FE09F8B9" box="[129,462,1887,1911]" italics="true" pageId="7" pageNumber="213">
|
||
<taxonomicName id="4C484D29B06DFFE923F41363FEF4F8B9" authorityName="Hensel" authorityYear="1868" box="[129,307,1887,1911]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="caudimaculatus">P. caudimaculatus</taxonomicName>
|
||
–
|
||
<taxonomicName id="4C484D29B06DFFE922351363FE09F8B9" authorityName="Lucinda" authorityYear="2008" box="[320,462,1887,1911]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="heptaktinos">P. heptaktinos</taxonomicName>
|
||
</emphasis>
|
||
and
|
||
<emphasis id="B93CEAB8B06DFFE9217F1363FCCEF8B9" box="[522,777,1887,1911]" italics="true" pageId="7" pageNumber="213">
|
||
<taxonomicName id="4C484D29B06DFFE9217F1363FDB8F8B9" authorityName="Lucinda" authorityYear="2008" box="[522,639,1887,1911]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="harpagos">P. harpagos</taxonomicName>
|
||
–
|
||
<taxonomicName id="4C484D29B06DFFE921F91363FCCEF8B9" box="[652,777,1887,1911]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="leptokeras">P. leptokeras</taxonomicName>
|
||
</emphasis>
|
||
as syntopic pairs of
|
||
<taxonomicName id="4C484D29B06DFFE9223A1342FE7AF858" box="[335,445,1918,1942]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE9223A1342FE7AF858" box="[335,445,1918,1942]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
. However,
|
||
<bibRefCitation id="EFD94B5BB06DFFE921471342FCC5F858" author="Thomaz AT & Carvalho TP & Malabarba LR" box="[562,770,1918,1942]" pageId="7" pageNumber="213" pagination="265 - 74" refId="ref10431" refString="Thomaz AT, Carvalho TP, Malabarba LR, et al. Geographic distributions, phenotypes, and phylogenetic relationships of Phalloceros (Cyprinodontiformes: Poeciliidae): insights about diversification among sympatric species pools. Molecular Phylogenetics and Evolution 2019; 132: 265 - 74." type="journal article" year="2019">
|
||
Thomaz
|
||
<emphasis id="B93CEAB8B06DFFE921F91343FD7AF858" box="[652,701,1918,1942]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
|
||
(2019
|
||
</bibRefCitation>
|
||
: p. 270) pointed out that
|
||
<taxonomicName id="4C484D29B06DFFE922E413A2FD83F87B" authorityName="Hensel" authorityYear="1868" box="[401,580,1949,1973]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="caudimaculatus">
|
||
<emphasis id="B93CEAB8B06DFFE922E413A2FD83F87B" box="[401,580,1949,1973]" italics="true" pageId="7" pageNumber="213">P. caudimaculatus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C484D29B06DFFE9210A13A2FCCEF87B" authorityName="Lucinda" authorityYear="2008" box="[639,777,1949,1973]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="heptaktinos">
|
||
<emphasis id="B93CEAB8B06DFFE9210A13A2FCCEF87B" box="[639,777,1949,1973]" italics="true" pageId="7" pageNumber="213">P. heptaktinos</emphasis>
|
||
</taxonomicName>
|
||
occur in different portions of the same river basin and thus might not be syntopic. Indeed, our re-examination of Lucinda’s data suggests that only the
|
||
<emphasis id="B93CEAB8B06DFFE9275314F3FAE7FF29" box="[1062,1312,207,231]" italics="true" pageId="7" pageNumber="213">
|
||
<taxonomicName id="4C484D29B06DFFE9275314F3FB5EFF29" authorityName="Lucinda" authorityYear="2008" box="[1062,1177,207,231]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="harpagos">P. harpagos</taxonomicName>
|
||
–
|
||
<taxonomicName id="4C484D29B06DFFE927D314F3FAE7FF29" box="[1190,1312,207,231]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="leptokeras">P. leptokeras</taxonomicName>
|
||
</emphasis>
|
||
pair represents a valid case of syntopy in his argument.
|
||
<bibRefCitation id="EFD94B5BB06DFFE9279314D2FA7CFEC8" author="Thomaz AT & Carvalho TP & Malabarba LR" box="[1254,1467,238,262]" pageId="7" pageNumber="213" pagination="265 - 74" refId="ref10431" refString="Thomaz AT, Carvalho TP, Malabarba LR, et al. Geographic distributions, phenotypes, and phylogenetic relationships of Phalloceros (Cyprinodontiformes: Poeciliidae): insights about diversification among sympatric species pools. Molecular Phylogenetics and Evolution 2019; 132: 265 - 74." type="journal article" year="2019">
|
||
Thomaz
|
||
<emphasis id="B93CEAB8B06DFFE9263714D3FAB3FEC8" box="[1346,1396,238,262]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
|
||
(2019
|
||
</bibRefCitation>
|
||
: p. 270) performed a genus-wide analysis of species distributions for
|
||
<emphasis id="B93CEAB8B06DFFE9202B1511FC03FE8B" box="[862,964,301,325]" italics="true" pageId="7" pageNumber="213">Phallocero</emphasis>
|
||
s and concluded that ≥ 14 of the 22 then-known species can be found in sympatry with one or more congeners and that most of the sympatric cases represented instances of syntopy. However, our re-analysis of their data shows that
|
||
<bibRefCitation id="EFD94B5BB06DFFE9204C1596FBDBFE0C" author="Thomaz AT & Carvalho TP & Malabarba LR" box="[825,1052,426,450]" pageId="7" pageNumber="213" pagination="265 - 74" refId="ref10431" refString="Thomaz AT, Carvalho TP, Malabarba LR, et al. Geographic distributions, phenotypes, and phylogenetic relationships of Phalloceros (Cyprinodontiformes: Poeciliidae): insights about diversification among sympatric species pools. Molecular Phylogenetics and Evolution 2019; 132: 265 - 74." type="journal article" year="2019">
|
||
Thomaz
|
||
<emphasis id="B93CEAB8B06DFFE920E31597FC0EFE0C" box="[918,969,426,450]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
|
||
(2019)
|
||
</bibRefCitation>
|
||
had uncovered only four valid cases of syntopic species:
|
||
<emphasis id="B93CEAB8B06DFFE9209F15F6FB7BFE2F" box="[1002,1212,457,481]" italics="true" pageId="7" pageNumber="213">
|
||
<taxonomicName id="4C484D29B06DFFE9209F15F6FBA3FE2F" authorityName="Lucinda" authorityYear="2008" box="[1002,1124,457,481]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="alessandre">P. alessandre</taxonomicName>
|
||
–
|
||
<taxonomicName id="4C484D29B06DFFE9270515F5FB7BFE2F" authorityName="Lucinda" authorityYear="2008" box="[1136,1212,457,481]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="pellos">P. pellos</taxonomicName>
|
||
</emphasis>
|
||
,
|
||
<emphasis id="B93CEAB8B06DFFE927B315F6FA7AFE2F" box="[1222,1469,457,481]" italics="true" pageId="7" pageNumber="213">
|
||
<taxonomicName id="4C484D29B06DFFE927B315F6FA8CFE2F" authorityName="Lucinda" authorityYear="2008" box="[1222,1355,457,481]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="malabarbai">P. malabarbai</taxonomicName>
|
||
–
|
||
<taxonomicName id="4C484D29B06DFFE9262D15F5FA7AFE2F" authorityName="Lucinda" authorityYear="2008" box="[1368,1469,457,481]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="spiloura">P. spiloura</taxonomicName>
|
||
</emphasis>
|
||
,
|
||
<emphasis id="B93CEAB8B06DFFE9204C15D5FBDFFDCE" box="[825,1048,488,512]" italics="true" pageId="7" pageNumber="213">
|
||
<taxonomicName id="4C484D29B06DFFE9204C15D5FC7EFDCE" authorityName="Lucinda" authorityYear="2008" box="[825,953,488,512]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="megapolos">P. megapolos</taxonomicName>
|
||
–
|
||
<taxonomicName id="4C484D29B06DFFE920B015D4FBDFFDCE" authorityName="Lucinda" authorityYear="2008" box="[965,1048,488,512]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="pellos">P. pellos</taxonomicName>
|
||
</emphasis>
|
||
and
|
||
<emphasis id="B93CEAB8B06DFFE9272115D5FAFFFDCE" box="[1108,1336,488,512]" italics="true" pageId="7" pageNumber="213">
|
||
<taxonomicName id="4C484D29B06DFFE9272115D5FB14FDCE" authorityName="Lucinda" authorityYear="2008" box="[1108,1235,488,512]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="megapolos">P. megapolos</taxonomicName>
|
||
–
|
||
<taxonomicName id="4C484D29B06DFFE9279515D4FAFFFDCE" authorityName="Lucinda" authorityYear="2008" box="[1248,1336,488,512]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="titthos">P. titthos</taxonomicName>
|
||
</emphasis>
|
||
(Supporting Information, Table S3). Nonetheless, the new data we have presented here (Supporting Information, Table S3) suggest that syntopy is not only widespread in the genus, but that ≥ 14 species of
|
||
<taxonomicName id="4C484D29B06DFFE92008165AFC2CFDB0" box="[893,1003,614,638]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE92008165AFC2CFDB0" box="[893,1003,614,638]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
occur in syntopy with a congener, thereby supporting the conjecture of
|
||
<bibRefCitation id="EFD94B5BB06DFFE9273116B9FAE7FD53" author="Thomaz AT & Carvalho TP & Malabarba LR" box="[1092,1312,645,669]" pageId="7" pageNumber="213" pagination="265 - 74" refId="ref10431" refString="Thomaz AT, Carvalho TP, Malabarba LR, et al. Geographic distributions, phenotypes, and phylogenetic relationships of Phalloceros (Cyprinodontiformes: Poeciliidae): insights about diversification among sympatric species pools. Molecular Phylogenetics and Evolution 2019; 132: 265 - 74." type="journal article" year="2019">
|
||
Thomaz
|
||
<emphasis id="B93CEAB8B06DFFE927EB16BAFB08FD53" box="[1182,1231,645,669]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
|
||
(2019)
|
||
</bibRefCitation>
|
||
. We also found that every case of syntopy involved only two species. Moreover, when we examined these syntopic pairs in light of their likely topological placements in the
|
||
<taxonomicName id="4C484D29B06DFFE9270416DFFAE8FD35" box="[1137,1327,739,763]" pageId="7" pageNumber="213">
|
||
<emphasis id="B93CEAB8B06DFFE9270416DFFB18FD35" box="[1137,1247,739,763]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
species
|
||
</taxonomicName>
|
||
tree (
|
||
<bibRefCitation id="EFD94B5BB06DFFE9261A16DFFC67FCD5" author="Thomaz AT & Carvalho TP & Malabarba LR" pageId="7" pageNumber="213" pagination="265 - 74" refId="ref10431" refString="Thomaz AT, Carvalho TP, Malabarba LR, et al. Geographic distributions, phenotypes, and phylogenetic relationships of Phalloceros (Cyprinodontiformes: Poeciliidae): insights about diversification among sympatric species pools. Molecular Phylogenetics and Evolution 2019; 132: 265 - 74." type="journal article" year="2019">
|
||
Thomaz
|
||
<emphasis id="B93CEAB8B06DFFE9204C173FFCAFFCD4" box="[825,872,770,794]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
|
||
2019
|
||
</bibRefCitation>
|
||
: fig. 3), all these syntopic pairs consisted of non-sister species, similar to the finding reported by
|
||
<bibRefCitation id="EFD94B5BB06DFFE92790171EFA7AFCF4" author="Thomaz AT & Carvalho TP & Malabarba LR" box="[1253,1469,802,826]" pageId="7" pageNumber="213" pagination="265 - 74" refId="ref10431" refString="Thomaz AT, Carvalho TP, Malabarba LR, et al. Geographic distributions, phenotypes, and phylogenetic relationships of Phalloceros (Cyprinodontiformes: Poeciliidae): insights about diversification among sympatric species pools. Molecular Phylogenetics and Evolution 2019; 132: 265 - 74." type="journal article" year="2019">
|
||
Thomaz
|
||
<emphasis id="B93CEAB8B06DFFE9264B171EFAAAFCF4" box="[1342,1389,802,826]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
|
||
(2019)
|
||
</bibRefCitation>
|
||
, although, again, their conclusion was based mainly on records of sympatry and not syntopy.
|
||
</paragraph>
|
||
<paragraph id="8BF736AAB06DFFE9202017BCFAB4FA62" blockId="7.[825,1474,144,1452]" pageId="7" pageNumber="213">
|
||
The independent dataset we obtained for this study also strongly corroborates the conjecture of
|
||
<bibRefCitation id="EFD94B5BB06DFFE927AA17A3FA05FC79" author="Thomaz AT & Carvalho TP & Malabarba LR" box="[1247,1474,927,951]" pageId="7" pageNumber="213" pagination="265 - 74" refId="ref10431" refString="Thomaz AT, Carvalho TP, Malabarba LR, et al. Geographic distributions, phenotypes, and phylogenetic relationships of Phalloceros (Cyprinodontiformes: Poeciliidae): insights about diversification among sympatric species pools. Molecular Phylogenetics and Evolution 2019; 132: 265 - 74." type="journal article" year="2019">
|
||
Thomaz
|
||
<emphasis id="B93CEAB8B06DFFE92649179CFAA8FC79" box="[1340,1391,927,951]" italics="true" pageId="7" pageNumber="213">et al.</emphasis>
|
||
(2019)
|
||
</bibRefCitation>
|
||
concerning the prevalence of mismatched genitalia in syntopic
|
||
<taxonomicName id="4C484D29B06DFFE9204C17E2FC60FC38" box="[825,935,990,1014]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE9204C17E2FC60FC38" box="[825,935,990,1014]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
. Indeed, all new cases of syntopy that we identified involved two coexisting species of
|
||
<taxonomicName id="4C484D29B06DFFE927EF17C1FACFFBDB" box="[1178,1288,1021,1045]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE927EF17C1FACFFBDB" box="[1178,1288,1021,1045]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
had genitalia mismatched to some degree (Supporting Information, Table S3). This study thus confirms that syntopy is widespread in
|
||
<taxonomicName id="4C484D29B06DFFE926211000FA05FB9A" box="[1364,1474,1084,1108]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE926211000FA05FB9A" box="[1364,1474,1084,1108]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
and that these species pairs seem always to consist of species with apparently mismatched genitals. However, if
|
||
<bibRefCitation id="EFD94B5BB06DFFE9267A1046FA05FB5C" author="Lucinda PHF" box="[1295,1474,1146,1170]" pageId="7" pageNumber="213" pagination="113 - 58" refId="ref9750" refString="Lucinda PHF. Systematics and biogeography of the genus Phalloceros Eigenmann, 1907 (Cyprinodontiformes: Poeciliidae: Poeciliinae), with the description of twenty-one new species. Neotropical Ichthyology 2008; 6: 113 - 58." type="journal article" year="2008">Lucinda’s (2008)</bibRefCitation>
|
||
claim that
|
||
<taxonomicName id="4C484D29B06DFFE920DD10A6FB9FFB7C" authorityName="Hensel" authorityYear="1868" box="[936,1112,1178,1202]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="caudimaculatus">
|
||
<emphasis id="B93CEAB8B06DFFE920DD10A6FB9FFB7C" box="[936,1112,1178,1202]" italics="true" pageId="7" pageNumber="213">P. caudimaculatus</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C484D29B06DFFE927FB10A6FAD1FB7C" authorityName="Lucinda" authorityYear="2008" box="[1166,1302,1178,1202]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="heptaktinos">
|
||
<emphasis id="B93CEAB8B06DFFE927FB10A6FAD1FB7C" box="[1166,1302,1178,1202]" italics="true" pageId="7" pageNumber="213">P. heptaktinos</emphasis>
|
||
</taxonomicName>
|
||
can be found in syntopy is eventually confirmed, then this species pair would represent an intriguing exception to the syntopy/mismatched genitalia rule in
|
||
<emphasis id="B93CEAB8B06DFFE920AF10C4FB87FADE" box="[986,1088,1272,1296]" italics="true" pageId="7" pageNumber="213">Phallocero</emphasis>
|
||
s. This is because the mechanical isolation hypothesis implies that co-occurring species of
|
||
<taxonomicName id="4C484D29B06DFFE92621112BFA05FAE1" box="[1364,1474,1303,1327]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE92621112BFA05FAE1" box="[1364,1474,1303,1327]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
having ‘matched’ genital morphologies can interbreed with each other. The existence of such an exceptional case would thus be problematic for this hypothesis unless a future study can show that these two species are interbreeding with each other.
|
||
</paragraph>
|
||
<paragraph id="8BF736AAB06DFFE9203111E1FA70FA39" blockId="7.[825,1475,1501,1970]" box="[836,1463,1501,1527]" pageId="7" pageNumber="213">
|
||
<emphasis id="B93CEAB8B06DFFE9203111E1FA70FA39" bold="true" box="[836,1463,1501,1527]" pageId="7" pageNumber="213">Do all mismatches in genitalia cause reproductive isolation?</emphasis>
|
||
</paragraph>
|
||
<paragraph id="8BF736AAB06DFFE6204C123FFDA3FE8B" blockId="7.[825,1475,1501,1970]" lastBlockId="8.[113,763,144,1421]" lastPageId="8" lastPageNumber="214" pageId="7" pageNumber="213">
|
||
Our results are consistent with the hypothesis that syntopic species of
|
||
<taxonomicName id="4C484D29B06DFFE920D2121FFBD2F9F5" box="[935,1045,1571,1595]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE920D2121FFBD2F9F5" box="[935,1045,1571,1595]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
with mismatched genitalia are unable to hybridize with each other. However, even if this is true for the
|
||
<taxonomicName id="4C484D29B06DFFE9204C125EFC0AF9B7" box="[825,973,1633,1657]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="anisophallos">
|
||
<emphasis id="B93CEAB8B06DFFE9204C125EFC0AF9B7" box="[825,973,1633,1657]" italics="true" pageId="7" pageNumber="213">P. anisophallos</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C484D29B06DFFE9277F125EFB40F9B7" box="[1034,1159,1633,1657]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="species" species="leptokeras">
|
||
<emphasis id="B93CEAB8B06DFFE9277F125EFB40F9B7" box="[1034,1159,1633,1657]" italics="true" pageId="7" pageNumber="213">P. leptokeras</emphasis>
|
||
</taxonomicName>
|
||
populations we studied, this might not necessarily apply to other syntopic pairs of
|
||
<taxonomicName id="4C484D29B06DFFE9263A12BDFA7AF957" box="[1359,1469,1665,1689]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE9263A12BDFA7AF957" box="[1359,1469,1665,1689]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
, because there are at least five different
|
||
<typeStatus id="54F38808B06DFFE92794129CFADFF976" box="[1249,1304,1696,1720]" pageId="7" pageNumber="213">types</typeStatus>
|
||
of ‘mismatched genitalia’ (
|
||
<tableCitation id="C6CA0311B06DFFE920DD1283FC31F919" box="[936,1014,1727,1751]" captionStart="Table 2" captionStartId="6.[113,168,615,639]" captionTargetPageId="6" captionText="Table 2. Relative degree of mismatch in characters of the genitalia between syntopic species of Phalloceros." pageId="7" pageNumber="213">Table 2</tableCitation>
|
||
), which could vary in their efficacies to inhibit cross-species fertilization. Of the three genital characters that show variation at the interspecific level, divergent papillae (with or without accompanying divergences in hook number and appendix shape) characterize 11 of 17 syntopic species pairs (
|
||
<tableCitation id="C6CA0311B06DFFE920311360FC57F8BA" box="[836,912,1884,1908]" captionStart="Table 2" captionStartId="6.[113,168,615,639]" captionTargetPageId="6" captionText="Table 2. Relative degree of mismatch in characters of the genitalia between syntopic species of Phalloceros." pageId="7" pageNumber="213">Table 2</tableCitation>
|
||
), hence orientation of the papillae might be a primary determinant of whether two co-occurring species of
|
||
<taxonomicName id="4C484D29B06DFFE926211347FA05F85D" box="[1364,1474,1915,1939]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="7" pageNumber="213" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B06DFFE926211347FA05F85D" box="[1364,1474,1915,1939]" italics="true" pageId="7" pageNumber="213">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
can interbreed with each other. Interestingly, the remaining six syntopic pairs showed interspecific differences in only hook number (four syntopic pairs) and hook number + shape of appendices (two pairs;
|
||
<tableCitation id="C6CA0311B062FFE6223114F3FE48FF29" box="[324,399,207,231]" captionStart="Table 2" captionStartId="6.[113,168,615,639]" captionTargetPageId="6" captionText="Table 2. Relative degree of mismatch in characters of the genitalia between syntopic species of Phalloceros." pageId="8" pageNumber="214">Table 2</tableCitation>
|
||
). Thus, there might be other factors involved (e.g. species-specific courtship behaviours) that help to maintain the species boundaries in these six exceptional species pairs, or perhaps they interbreed with each other.
|
||
</paragraph>
|
||
<paragraph id="8BF736AAB062FFE623F81570FDBCFA43" blockId="8.[113,763,144,1421]" pageId="8" pageNumber="214">
|
||
The
|
||
<taxonomicName id="4C484D29B062FFE623CB1570FEEBFEAA" box="[190,300,332,356]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" pageNumber="214" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B062FFE623CB1570FEEBFEAA" box="[190,300,332,356]" italics="true" pageId="8" pageNumber="214">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
clade is an exciting study system in which to explore the evolutionary significance of mismatched genitalia to speciation and the subsequent maintenance of species boundaries. Although we presented evidence that is consistent with the mechanical isolation hypothesis for
|
||
<taxonomicName id="4C484D29B062FFE6212C15F5FD00FE2F" box="[601,711,457,481]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" pageNumber="214" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B062FFE6212C15F5FD00FE2F" box="[601,711,457,481]" italics="true" pageId="8" pageNumber="214">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
, our findings suffer from two important limitations. First, we examined hybridization in only a small number of individuals from each population using two molecular markers. Had we sampled a larger number of individuals and/or genomic loci, we might have obtained evidence showing that
|
||
<taxonomicName id="4C484D29B062FFE62148165BFD0AFDB0" box="[573,717,614,638]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" pageNumber="214" phylum="Chordata" rank="species" species="anisophallos">
|
||
<emphasis id="B93CEAB8B062FFE62148165BFD0AFDB0" box="[573,717,614,638]" italics="true" pageId="8" pageNumber="214">P. anisophallos</emphasis>
|
||
</taxonomicName>
|
||
and
|
||
<taxonomicName id="4C484D29B062FFE6230416BAFF2DFD53" box="[113,234,645,669]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" pageNumber="214" phylum="Chordata" rank="species" species="leptokeras">
|
||
<emphasis id="B93CEAB8B062FFE6230416BAFF2DFD53" box="[113,234,645,669]" italics="true" pageId="8" pageNumber="214">P. leptokeras</emphasis>
|
||
</taxonomicName>
|
||
in the Parati-Mirim River have interbred with each other. Second, our findings, and all relevant published works on
|
||
<taxonomicName id="4C484D29B062FFE6230416F8FF18FD12" box="[113,223,708,732]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" pageNumber="214" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B062FFE6230416F8FF18FD12" box="[113,223,708,732]" italics="true" pageId="8" pageNumber="214">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
, have presented only correlational evidence in support of this hypothesis. In the future, researchers should explore the two promising avenues of population genomics and laboratory breeding experiments, which might shed much more light on the functional and evolutionary significance of mismatched genitalia in these fishes. The former approach would allow researchers to determine the levels, if any, of historical gene flow between co-occurring
|
||
<taxonomicName id="4C484D29B062FFE6222217A3FDD4FC79" box="[343,531,927,951]" pageId="8" pageNumber="214">
|
||
<emphasis id="B93CEAB8B062FFE6222217A3FE02FC79" box="[343,453,927,951]" italics="true" pageId="8" pageNumber="214">Phalloceros</emphasis>
|
||
species
|
||
</taxonomicName>
|
||
that exemplify each of the observed mismatch variants listed in
|
||
<tableCitation id="C6CA0311B062FFE621691782FDAEFC19" box="[540,617,958,983]" captionStart="Table 2" captionStartId="6.[113,168,615,639]" captionTargetPageId="6" captionText="Table 2. Relative degree of mismatch in characters of the genitalia between syntopic species of Phalloceros." pageId="8" pageNumber="214">Table 2</tableCitation>
|
||
, and between coexisting species that have matched genitalia. In the latter approach, researchers should conduct laboratory breeding experiments between different species of
|
||
<taxonomicName id="4C484D29B062FFE622831020FDA3FBFA" box="[502,612,1052,1076]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" pageNumber="214" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B062FFE622831020FDA3FBFA" box="[502,612,1052,1076]" italics="true" pageId="8" pageNumber="214">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
to determine whether mismatched genitalia can inhibit cross-species fertilization. If the results confirm that mismatched genitalia do act as mechanical barriers to fertilization, then further experiments can be done to ascertain the relative efficacies of the different mismatch possibilities and to test the hypothesis that co-occurring species having matched genitalia are able to interbreed. Only asser these studies are completed will we be able to understand fully the evolutionary significance of genital morphology variation in
|
||
<taxonomicName id="4C484D29B062FFE623BD110AFEF1FA80" box="[200,310,1334,1358]" family="Poeciliidae" genus="Phalloceros" kingdom="Animalia" order="Cyprinodontiformes" pageId="8" pageNumber="214" phylum="Chordata" rank="genus">
|
||
<emphasis id="B93CEAB8B062FFE623BD110AFEF1FA80" box="[200,310,1334,1358]" italics="true" pageId="8" pageNumber="214">Phalloceros</emphasis>
|
||
</taxonomicName>
|
||
and its possible link to their diversification. This, in turn, might reveal the general importance of mismatched genitalia as a mechanism for reproductive isolation.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |