369 lines
57 KiB
XML
369 lines
57 KiB
XML
<document id="8ED893325C0FA1C532232EE3C186537B" ENCODING="UTF-8" ID-GBIF-Dataset="caa93092-6ec2-4fc3-a298-722ee61880af" ModsDocID="z01067p001" checkinTime="1247054664313" checkinUser="thomas" docAuthor="Wilson J. E. M. Costa" docDate="2005" docId="F5993F232D2182B14514563125BBA00C" docLanguage="en" docName="2005_Costa2_gg1.xml" docOrigin="Zootaxa 1067" docSource="http://www.zoobank.org/urn:lsid:zoobank.org:pub:3A9C14FE-2B1A-408F-BACA-4F65D98E22FB" docTitle="Pterolebias Garman" docType="treatment" docVersion="13" lastPageNumber="24" masterDocId="2551B8783B24704F60CBEC56C4F62971" masterDocTitle="The Neotropical annual killifish genus Pterolebias Garman (Teleostei: Cyprinodontiformes: Rivulidae): phylogenetic relationships, descriptive morphology, and taxonomic revision." masterLastPageNumber="36" masterPageNumber="1" pageNumber="20" updateTime="1701325845312" updateUser="ExternalLinkService">
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<mods:titleInfo id="C4EF778E56ACBF433719512A4568B5C6">
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<mods:title id="F091A39EEF8BE2D036CB4A15ACCFF12E">The Neotropical annual killifish genus Pterolebias Garman (Teleostei: Cyprinodontiformes: Rivulidae): phylogenetic relationships, descriptive morphology, and taxonomic revision.</mods:title>
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<mods:namePart id="ECC158F0D4CD03993A2E86757D03C9A9">Wilson J. E. M. Costa</mods:namePart>
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<mods:title id="E95E683A27A6CFBEDFAB4B4E1B36DA0E">Zootaxa</mods:title>
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<mods:date id="033CD7A35D633657C7E9049B7CDE0291">2005</mods:date>
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<mods:classification id="0DF4590C8DCC8E5B999507B2AFCB3EFB">journal article</mods:classification>
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<treatment id="F5993F232D2182B14514563125BBA00C" ID-DOI="http://doi.org/10.5281/zenodo.6266694" ID-GBIF-Taxon="100119209" ID-Zenodo-Dep="6266694" LSID="urn:lsid:plazi:treatment:F5993F232D2182B14514563125BBA00C" httpUri="http://treatment.plazi.org/id/F5993F232D2182B14514563125BBA00C" lastPageNumber="24" pageNumber="20">
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<subSubSection id="1938610B96C489B00754CC54DED4867F" type="nomenclature">
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<paragraph id="374CF1EC7D226A500C23E0C8DE28FCB3" pageNumber="20">
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[[ Genus
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<taxonomicName id="83FADC5E9A519A0B3544643F668E16BE" ID-CoL="74RK" ID-ENA="60340" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="genus">Pterolebias Garman</taxonomicName>
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]]
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</paragraph>
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</subSubSection>
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<subSubSection id="052F54B75103706B718539814621A54A" type="discussion">
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<paragraph id="8E8B3BD844D51E39A7972D861AAE7421" pageNumber="20">Discussion</paragraph>
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<paragraph id="C76B12BBFB15FDA04B8D34C0C7067E39" pageNumber="20">
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Taxonomy.
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<taxonomicName id="0C1E37D164DD84638FFEE6BD1A298048" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
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possibly poses the greatest taxonomic problems within the
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<taxonomicName id="3B3067157399610C63CA50FDE96892D9" family="Rivulidae" pageNumber="20" rank="family">Rivulidae</taxonomicName>
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(see History above). This is due to the wide geographic range
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<normalizedToken id="BFECB5F218F38C26AD877A181E54B4E2" originalValue="oîP">of</normalizedToken>
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<taxonomicName id="6AF283E0531E5679758C2E1B79A86FE8" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
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, which is unique for an annual fish, and overlaps the type locality of two similar, controversial species,
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<taxonomicName id="E9F818CB2686BD49EFA4ED3A0C06FE9A" LSID-ZBK="urn:lsid:zoobank.org:act:F53F7AF3-340E-4EFE-93C2-478ACB197C80" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias bokermanni Travassos 1955:35" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="bokermanni">P. bokermanni</taxonomicName>
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and
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<taxonomicName id="EFB0EA41A5E88D98956415B5A59E9EFA" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="luelingi">P. luelingi</taxonomicName>
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.
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</paragraph>
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<paragraph id="5CCDC05E69C81F6EC226A20BBDBCF909" lastPageNumber="21" pageNumber="20">
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Travassos (1955) distinguished
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<taxonomicName id="FB3233801850B22881A614719334C2ED" LSID-ZBK="urn:lsid:zoobank.org:act:F53F7AF3-340E-4EFE-93C2-478ACB197C80" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias bokermanni Travassos 1955:35" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="bokermanni">P. bokermanni</taxonomicName>
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from
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<taxonomicName id="9C413BBBDE90B2C38244688786B80BAA" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
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by the former having pelvic-fin bases medially united, dorsal and anal fins placed more posteriorly, a
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<pageStartToken id="AB4FCB16FC2675325BEB2456F00ED556" pageNumber="21">
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<pageStartToken id="853BAB88C24029B59C9BE191EF27E7CD" pageNumber="21">slender</pageStartToken>
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</pageStartToken>
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body, pelvic fin longer, fewer scales in the transverse series, more scales in the longitudinal series, and greater development of cephalic “pores” (i.e. neuromasts). However, Travassos never examined any specimens of
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<taxonomicName id="262C19BDA71217344B94E818FA2704F8" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
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from the type locality or from any other region, and based comparisons on the limited data provided in Garmans (1895) description. None of these features differs among populations formerly identified as
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<taxonomicName id="FE6FD59A4EA163A2D42FB2C5293FE786" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
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,
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<taxonomicName id="FF7EA188A9FAF359894C77718179B536" LSID-ZBK="urn:lsid:zoobank.org:act:F53F7AF3-340E-4EFE-93C2-478ACB197C80" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias bokermanni Travassos 1955:35" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="bokermanni">P. bokermanni</taxonomicName>
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, and
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<taxonomicName id="322469838DCB288A32710B4BBF62CD9F" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="luelingi">P. luelingi</taxonomicName>
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.
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</paragraph>
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<paragraph id="66CD8AEF615F8BA45388BC6A62991E7F" pageNumber="21">
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The reasons that
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<taxonomicName id="D0F5F628C701A5C3203EF9A6FB42668C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="luelingi">P. luelingi</taxonomicName>
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was first placed in
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<taxonomicName id="9EB922E45EC196284C0BE1B5FA5D0FC9" LSID-ZBK="urn:lsid:zoobank.org:act:368164C2-3E85-4AD8-A9F1-C42A28B13023" family="Rivulidae" genus="Rivulichthys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulichthys Myers 1927:118" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Rivulichthys</taxonomicName>
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by Meinken (1969) and not in
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<taxonomicName id="DD93104D049A2BAA27DE128A86028A5E" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
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are unknown. The few characters presented in the original description of
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<taxonomicName id="3D34C9E52B028B37BE6851FC127D50FA" LSID-ZBK="urn:lsid:zoobank.org:act:A1AFFABB-34D6-41C3-9037-B24480828AC8" family="Rivulidae" genus="Rivulichthys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulichthys luelingi Meinken 1969:423" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="luelingi">R. luelingi</taxonomicName>
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are congruent with those included in the original description of
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<taxonomicName id="81862A2D32DAAC52E00EE8ACC156FFB1" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
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. Thomerson (1984) considered
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<taxonomicName id="B4CCD40EE1C634964F82C833C7332663" LSID-ZBK="urn:lsid:zoobank.org:act:A1AFFABB-34D6-41C3-9037-B24480828AC8" family="Rivulidae" genus="Rivulichthys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulichthys luelingi Meinken 1969:423" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="luelingi">R. luelingi</taxonomicName>
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to be a synonym of
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<taxonomicName id="6AF09C0957DF7B3C7FFAD59359A7BFA2" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
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, although he recorded eight pelvic-fin rays for the types of
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<taxonomicName id="A6EADA2925D8BDAC9611F0E9E5E07528" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
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, in contrast to seven rays recorded by Meinken for
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<taxonomicName id="33515DBA6479CE059C45936A91361ABE" LSID-ZBK="urn:lsid:zoobank.org:act:A1AFFABB-34D6-41C3-9037-B24480828AC8" family="Rivulidae" genus="Rivulichthys" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rivulichthys luelingi Meinken 1969:423" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="luelingi">R. luelingi</taxonomicName>
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. However, Schindler (2004) used this putative difference in number of pelvic-fin rays to distinguish
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<taxonomicName id="7F12FBC3EE89C4B61C211C9B44AB08BA" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="luelingi">P. luelingi</taxonomicName>
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, occurring in the
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<normalizedToken id="4F45F1441E127E168D0627D6C43C3EF5" originalValue="Mamoré">Mamore</normalizedToken>
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River basin, from
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<taxonomicName id="D729923BAE12ECFB4997830893287948" LSID-ZBK="urn:lsid:zoobank.org:act:F53F7AF3-340E-4EFE-93C2-478ACB197C80" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias bokermanni Travassos 1955:35" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="bokermanni">P. bokermanni</taxonomicName>
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, from the
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<normalizedToken id="0747FE68A18C8B5DFCCE9FC2A60C9E8E" originalValue="Guaporé">Guapore</normalizedToken>
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and Paraguay river basins, and from
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<taxonomicName id="24B3D4D850FA71F9B7B0977E23F2332C" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
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. In fact, the type locality of
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<taxonomicName id="F9761F34FF29DFF820A81C36B736B817" LSID-ZBK="urn:lsid:zoobank.org:act:F53F7AF3-340E-4EFE-93C2-478ACB197C80" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias bokermanni Travassos 1955:35" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="bokermanni">P. bokermanni</taxonomicName>
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is
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<normalizedToken id="2F18EDD596733B9B6D84EE322F561F62" originalValue="Guajará-Mirim">Guajara-Mirim</normalizedToken>
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, on floodplains of the
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<normalizedToken id="A4535D986F65D4EB191AB02674A8BF54" originalValue="Mamoré">Mamore</normalizedToken>
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River, although it is also recorded from the
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<normalizedToken id="CEE5007A41BFF36A858A328C0EEF6CF8" originalValue="Guaporé">Guapore</normalizedToken>
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River floodplains (Travassos, 1955). All of the approximately 400 specimens of
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<taxonomicName id="7A88E1F6A0DFF7CFFC2B8F46BFD3A739" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
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examined in the present study have eight pelvic-fin rays, except for a few specimens with nine rays. Schindler (2004) also noted a reduced squamation on the anal-fin base in males, including two or three rows of scales in
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<taxonomicName id="760DA029A48F9ABEBC18AE80F247FB7F" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
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and
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<taxonomicName id="FFA252B49F37918808610DDAFFC80CD3" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="luelingi">P. luelingi</taxonomicName>
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, in contrast to four or more rows in
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<taxonomicName id="012E3870026B21F166E4E2D78C81E681" LSID-ZBK="urn:lsid:zoobank.org:act:F53F7AF3-340E-4EFE-93C2-478ACB197C80" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias bokermanni Travassos 1955:35" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="bokermanni">P. bokermanni</taxonomicName>
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. However, all adult males of
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<taxonomicName id="6DDD64D0D2F06DF480C4F18A7B5D8343" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
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examined during the present study have four to seven rows, except
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<taxonomicName id="51CDE61F4D0E80925589B08B3AF1B1EE" LSID-ZBK="urn:lsid:zoobank.org:act:A68FF2CC-0501-4D84-BF18-691BD5FB08C4" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias phasianus Costa 1988:658" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="phasianus">P. phasianus</taxonomicName>
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, which has a single irregular row. Therefore, diagnostic features recorded by Schindler (2004) are not useful for distinguishing
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<taxonomicName id="64DD1D6E9A56649DB1E093316C50C0A6" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="luelingi">P. luelingi</taxonomicName>
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and
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<taxonomicName id="319F4156B9C9CE931B06931024C49791" LSID-ZBK="urn:lsid:zoobank.org:act:F53F7AF3-340E-4EFE-93C2-478ACB197C80" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias bokermanni Travassos 1955:35" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="bokermanni">P. bokermanni</taxonomicName>
|
|
from
|
|
<taxonomicName id="2175446FFB5B226D9FCA51B44B672C11" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
|
|
.
|
|
</paragraph>
|
|
<paragraph id="FEC89899AFE8DE8140426BACD3A27582" pageNumber="21">
|
|
Characters examined were not useful for recognizing
|
|
<taxonomicName id="BE6B87FFEE7CB45EDD1CD5CFD5924C7B" LSID-ZBK="urn:lsid:zoobank.org:act:F53F7AF3-340E-4EFE-93C2-478ACB197C80" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias bokermanni Travassos 1955:35" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="bokermanni">P. bokermanni</taxonomicName>
|
|
and
|
|
<taxonomicName id="A11C1CE39011D8B0DF905DC8C54DDD95" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="luelingi">P. luelingi</taxonomicName>
|
|
as valid species, nor for distinguishing geographically distant populations previously identified as
|
|
<taxonomicName id="FE31D12AC2A1532C0AFDA6E94A0E98B3" LSID-ZBK="urn:lsid:zoobank.org:act:CDCDF67E-4003-4B96-B765-7E7BA7E46A93" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias longipinnis Garman 1895:142" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="species" species="longipinnis">P. longipinnis</taxonomicName>
|
|
from the
|
|
<normalizedToken id="48607EB3D4A8577329293C3E7645C5D0" originalValue="Paraná">Parana</normalizedToken>
|
|
-Paraguay system. Some distinctive color patterns, such as stripes on male anal fin, were noted in individuals from the
|
|
<normalizedToken id="1DB2496A1A02F009821F90E761882852" originalValue="Mamoré">Mamore</normalizedToken>
|
|
and Paraguay river basins in Brazil (see above description). However, these color features are not constant, are absent from a large percentage of individuals from those regions, and therefore are not useful as diagnostic characters in this particular instance.
|
|
</paragraph>
|
|
<paragraph id="73927C250B31844650B707B1656B59DC" pageNumber="21">
|
|
Phylogenetic relationships. Both monophyly of
|
|
<taxonomicName id="300C913F0005024EA4C25AFC7E3CC5AC" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
and the sister group relationship between
|
|
<taxonomicName id="C124DFFE26AEB60A63D4FDDA90A06D75" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
and
|
|
<taxonomicName id="8357A7B99824162B59B84F7D9518E0C2" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
are strongly supported by bootstrap values (Fig. 1). The clade
|
|
<taxonomicName id="084171C2AB07102E8989B9DEEA680FDD" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
was previously corroborated by morphological data (Costa, 1998a) and molecular studies (Murphy et al., 1999; Hrbek & Larson, 1999). However, the clade
|
|
<taxonomicName id="8E630314E5CD9C5E306514B14FE1E335" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
plus
|
|
<taxonomicName id="3B1056C53168A2E62AD4E59B488FC04F" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
, already reported in morphological analysis (Costa, 1998a), is in conflict with molecular phylogenetic studies, which strongly support the monophyly of a clade including
|
|
<taxonomicName id="E77D5A402D427ADDADC156A6E8516536" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="21" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
and other taxa endemic to northern South America (Murphy et al., 1999; Hrbek & Larson, 1999) (Fig. 11).
|
|
</paragraph>
|
|
<caption id="91F4643AC5A48C5C8427FC004BC8C371">
|
|
<paragraph id="C42E9F796D7FDE924258C1FD3C3AC37B" pageNumber="22">
|
|
<pageStartToken id="7FCFE24B9C989AE21FED9C5BC04E19C3" pageNumber="22">
|
|
<pageStartToken id="8203ABD8731BCE036954CFC59AAC962D" pageNumber="22">FIGURE</pageStartToken>
|
|
</pageStartToken>
|
|
11. Molecular hypotheses of relationships among species of
|
|
<taxonomicName id="EF454024A38CF023BA0AEEABF7899715" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
,
|
|
<taxonomicName id="B8471BE40426FD0D7074E245CA0F88DE" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
and other rivulids: a, modified from Murphy et al. (1999); b, modified from Hrbek & Larson (1999). Numbers left of branches are bootstrap values.
|
|
</paragraph>
|
|
</caption>
|
|
<paragraph id="48FC390B689425570C5C98F49AAE9AF8" pageNumber="22">
|
|
Monophyly of
|
|
<taxonomicName id="52CADD25A512618F691632D1C74D1378" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
is herein supported by four unambiguous synapomorphies: ventral process of angulo-articular narrow and pointed (16.2) (Figs. 3a, c), medial flap of second pharyngobranchial expanded (33.1) (Fig. 3g), small metallic orange spots on humeral region in males (82.1) (Figs. 5, 6, 9), and bars on pectoral fin in males (90.2) (more conspicuous in the example of Fig. 9). The
|
|
<taxonomicName id="C2DDEA6FDBF570710441C5907663EA2C" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
clade is further corroborated by four synapomorphies homoplastically occurring in other rivulids: maxilla greatly twisted (13.2) (Figs. 3a-b), dorsal portion of metapterygoid strongly constricted (19.2) (Fig. 3a), posterior process of quadrate long (20.1) (Fig. 3a), and basihyal long (25.1) (Fig. 3a); and one reversal: pectoral fin rounded (62.0).
|
|
</paragraph>
|
|
<paragraph id="35A2ECB5768F091FC235109DBDF09433" lastPageNumber="23" pageNumber="22">
|
|
Monophyly of the clade comprising
|
|
<taxonomicName id="785D5AF96A824A57E113BA82BF5E291A" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
and
|
|
<taxonomicName id="E9EDC6D3AC6EAFBC0B6FF08EB9434A80" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
is supported by three unambiguous synapomorphies: interarcual cartilage minute (31.1) (Figs. 3d, g), anterior proximal radiais of anal fin curved and posteriorly directed (48.1) (Fig. 4b), and anal-fin base with scales (75.1). This assemblage is also supported by four synapomorphies homoplastically occurring in other rivulid lineages: external median teeth of premaxilla and dentary displaced laterally (17.1) (Figs. 3a-c), ischial process of pelvic girdle vestigial (55.1) (Fig. 4c), eight pelvic-fin rays (56.1), and pelvic fin long in males (64.1) (Figs. 5, 6, 9, 12). However, in a study of mitochondrial DNA by Hrbek & Larson (1999),
|
|
<taxonomicName id="CBB4F2A1F1051610BBB081C45009773C" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="zonatus">Gnatholebias zonatus</taxonomicName>
|
|
was hypothesized to be more closely related to
|
|
<taxonomicName id="D39407741D8797816102CE0B8E77B303" family="Rivulidae" genus="Rachovia" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="stellifer">Rachovia stellifer</taxonomicName>
|
|
than to other rivulids, and
|
|
<taxonomicName id="902863C0089B48F5F6F922A05F24AB99" family="Rivulidae" genus="Terranatos" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="dolichopterus">Terranatos dolichopterus</taxonomicName>
|
|
was hypothesized to be the
|
|
<pageStartToken id="700ABBD66C96215500D15CD17FC5C480" pageNumber="23">
|
|
<pageStartToken id="7D4D851C2C1893E61954EBFA2F67B22F" pageNumber="23">sister</pageStartToken>
|
|
</pageStartToken>
|
|
group of the clade including
|
|
<taxonomicName id="1E11E5104A1FA69D88658BC3188C8DCD" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="zonatus">G. zonatus</taxonomicName>
|
|
plus
|
|
<taxonomicName id="6B439D04366A8D7B8F2E4FA938248A6B" family="Rivulidae" genus="Rachovia" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="stellifer">R. stellifer</taxonomicName>
|
|
(Fig. 11b). A similar hypothesis was obtained from another mitochondrial DNA study (Murphy et al., 1999), in which
|
|
<taxonomicName id="EE7AABA94F5213EF5BF1D8BFC276B4C7" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="zonatus">G. zonatus</taxonomicName>
|
|
was considered to be the sister group of
|
|
<taxonomicName id="D7FC9D58B9FA5D36D67ECEC446E66695" genus="Terratanos" pageNumber="22" rank="species" species="dolichopterus">T. dolichopterus</taxonomicName>
|
|
, but
|
|
<taxonomicName id="FDE8C422F1EAC4C5B64FF56D939C5642" family="Rivulidae" genus="Rachovia" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="stellifer">R. stellifer</taxonomicName>
|
|
was not included in the analysis (Fig. 11a).
|
|
</paragraph>
|
|
<paragraph id="357ED0A1B10DE7B4934676F91A196C47" pageNumber="23">
|
|
I have found no morphological support for an assemblage including
|
|
<taxonomicName id="D79384901110202DE20A3D10D5A4C6FA" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="zonatus">Gnatholebias zonatus</taxonomicName>
|
|
,
|
|
<taxonomicName id="812378240729FBDF0D1570D8F4A695DF" genus="Terratanos" pageNumber="23" rank="species" species="dolichopterus">Terratanos dolichopterus</taxonomicName>
|
|
, and
|
|
<taxonomicName id="61092B2A9B3E767A1B587B5CC5E3B587" family="Rivulidae" genus="Rachovia" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="stellifer">Rachovia stellifer</taxonomicName>
|
|
, since all apomorphic conditions shared by these three genera are present in several other rivulid lineages. Both
|
|
<taxonomicName id="A3DCDAED7B730124958C1F2CAA9934EC" family="Rivulidae" genus="Rachovia" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="stellifer">R. stellifer</taxonomicName>
|
|
and
|
|
<taxonomicName id="CEEED5D9CD1E9FFB9B2184470E842D50" genus="Terratanos" pageNumber="23" rank="species" species="dolichopterus">T. dolichopterus</taxonomicName>
|
|
have the pelvic fin slightly elongated in males, which could be considered homologous to the long pelvic fin of
|
|
<taxonomicName id="1B7245E85C16A69D57D94F49A6C395A6" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="zonatus">G. zonatus</taxonomicName>
|
|
and
|
|
<taxonomicName id="22C2AAD08971DCA84AFE1CD878A4FA78" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="hoignei">G. hoignei</taxonomicName>
|
|
. However, the pelvic fin is much longer in
|
|
<taxonomicName id="6CCB7CF29A9F134B1141FA5E6957435A" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
and
|
|
<taxonomicName id="B19FB21849EC74D5A23DB4FD5779EED6" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
. In addition, in the latter two genera, the pelvic fin elongation is due to a long fourth ray (of eight rays) always occurring in the fin. In
|
|
<taxonomicName id="DF271CE7B9641746329C17CBFF926DE4" genus="Terratanos" pageNumber="23" rank="species" species="dolichopterus">T. dolichopterus</taxonomicName>
|
|
, however, the longer ray is the third (of eight), thus not considered a homologous condition; and in
|
|
<taxonomicName id="33FDE025291D9AE72BE372582C6E87E3" family="Rivulidae" genus="Rachovia" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="stellifer">R. stellifer</taxonomicName>
|
|
the longest ray is the fourth, but since there are only seven pelvic-fin rays in that species homology is doubtful.
|
|
<taxonomicName id="00686EB711E4DDE9EADD006CF45CF018" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="zonatus">Gnatholebias zonatus</taxonomicName>
|
|
and
|
|
<taxonomicName id="DE77D169854D0E0235AE0C16044D75AD" genus="Terratanos" pageNumber="23" rank="species" species="dolichopterus">T. dolichopterus</taxonomicName>
|
|
share a medial fusion on the pelvic fin (65.2), but this is not significant when compared to the many morphological synapomorphies shared by
|
|
<taxonomicName id="724E2B79F429A10E9A8BE2F339E5773A" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
and
|
|
<taxonomicName id="0386B6E3BC5FFDBC2813F7C10D5A82CA" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
(see above).
|
|
<taxonomicName id="80BEB1791F29DD37304DC9338C4D4B6E" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="zonatus">Gnatholebias zonatus</taxonomicName>
|
|
and
|
|
<taxonomicName id="BE49F3B380B5AED30748670BE6785D49" genus="Terratanos" pageNumber="23" rank="species" species="dolichopterus">T. dolichopterus</taxonomicName>
|
|
share reduction or loss of the ischial pelvic process (55.1), but this condition is also present in
|
|
<taxonomicName id="01012A4F122B86C6EE65AE76CD44392B" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
. Like
|
|
<taxonomicName id="2643EBAB471460D99D3E96AA274AC0A5" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
species,
|
|
<taxonomicName id="FA0821EDB727C02B193626CED8AB1BAB" genus="Terratanos" pageNumber="23" rank="species" species="dolichopterus">T. dolichopterus</taxonomicName>
|
|
has filamentous rays on the unpaired fins (67.1, 68.1), but this feature is also present in
|
|
<taxonomicName id="8C2382E34D7EF4E9CCA8EE22A15880B4" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
and in some species of
|
|
<taxonomicName id="95AA0EB3E62691190EC2176952368860" LSID-ZBK="urn:lsid:zoobank.org:act:F12E2023-61ED-419A-BF66-AB71508F9A02" family="Rivulidae" genus="Rachovia" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Rachovia Myers 1927:119" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Rachovia</taxonomicName>
|
|
and
|
|
<taxonomicName id="3F7CCE47BB1CD55A8CFB709BDA2CF59F" LSID-ZBK="urn:lsid:zoobank.org:act:A12D02B2-18D2-450F-B6ED-A28984BAD110" family="Rivulidae" genus="Austrofundulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Austrofundulus Myers 1932:159" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Austrofundulus</taxonomicName>
|
|
(e. g.,
|
|
<taxonomicName id="DAED4A8041A81EB64731FFDDEC0F1B62" family="Rivulidae" genus="Rachovia" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="brevis">R. brevis (Regan)</taxonomicName>
|
|
,
|
|
<taxonomicName id="5D93752E591F9E6F0F6FFEB23E7848CB" LSID-ZBK="urn:lsid:zoobank.org:act:BC5CD6B6-7039-4B80-98D6-8A5CAEA1A54E" family="Rivulidae" genus="Austrofundulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Austrofundulus leohoignei Hrbek, Taphorn & Thomerson 2005:19" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="leohoignei">A. leohoignei Hrbek, Taphorn & Thomerson</taxonomicName>
|
|
,
|
|
<taxonomicName id="B140FCC6EC05E4D0BE4F5481A679E4E5" LSID-ZBK="urn:lsid:zoobank.org:act:1A32491D-43B3-42CF-8DA4-6CDEA02DBC1B" family="Rivulidae" genus="Austrofundulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Austrofundulus leoni Hrbek, Taphorn & Thomerson 2005:25" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="leoni">A. leoni Hrbek, Taphorn & Thomerson</taxonomicName>
|
|
), which would be more closely related to
|
|
<taxonomicName id="291643E8649B08098D05E94C8E3722D7" LSID-ZBK="urn:lsid:zoobank.org:act:79CEE90A-5FE1-406E-BE38-5DF8C249CD32" family="Rivulidae" genus="Terranatos" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Terranatos Taphorn & Thomerson 1978:384" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Terranatos</taxonomicName>
|
|
according to the morphological studies. In addition,
|
|
<taxonomicName id="ACC3759995CB7009461069BB886D77A8" genus="Terratanos" pageNumber="23" rank="species" species="dolichopterus">T. dolichopterus</taxonomicName>
|
|
,
|
|
<taxonomicName id="6AE56F52F974905B3D86556401CD3A80" family="Rivulidae" genus="Rachovia" higherTaxonomySource="CoL" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="maculipinnis">R. maculipinnis</taxonomicName>
|
|
, and
|
|
<taxonomicName id="238779C36526190F421070545145186B" LSID-ZBK="urn:lsid:zoobank.org:act:8DF8C2F2-A5F1-4548-BC19-AD4E00BDC5AB" family="Rivulidae" genus="Austrofundulus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Austrofundulus transilis Myers 1932:160" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="transilis">A. transilis</taxonomicName>
|
|
share four synapomorphies: dorsally directed process on the base of anterior epipleural ribs (40.1), dorsal profile of the anterior portion of body convex (59.1), supraorbital spot adjacent to eye (85.1), and dark pigmentation reduced on flank and fins in females (92.1). The first feature, already described and illustrated by Costa (1998a: fig. 28a), seems to be unique among aplocheiloid fishes.
|
|
</paragraph>
|
|
<paragraph id="5BD209689636679BF782F0E3A82D44A1" pageNumber="23">Although morphological and molecular studies often produce slightly different estimates of phylogenetic relationships among species of the same group, Hillis & Wiens (2000) considered significant incongruence rare, pointing out some common causes, such as undersampling of characters or taxa, differences in phylogenetic methods and differences in rooting techniques. However, none of these causes seems to be applicable, since all studies are based on parsimony analysis, similar outgroup analysis methods, and considerable numbers of characters and representative taxa, with conflicting nodes having high bootstrap values.</paragraph>
|
|
<paragraph id="0BCFFFD6363E40CD4C1802C7A3548E5A" lastPageNumber="24" pageNumber="23">
|
|
It still not possible to satisfactorily understand the incongruence found between morphological and molecular hypotheses of relationships among
|
|
<taxonomicName id="976226D638FC1CB3F95A786E7DF6C519" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
,
|
|
<taxonomicName id="2354DE3DB49A8D98419EC397278F1486" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
and other rivulids, making it difficult to establish which data set is
|
|
<pageStartToken id="9792358A95538AF45B3A8E38BD9940C5" pageNumber="24">
|
|
<pageStartToken id="DF84EA4DCD83A9693A488C9F2A2BEC55" pageNumber="24">misleading</pageStartToken>
|
|
</pageStartToken>
|
|
. It would be desirable to improve estimates of phylogenetic relationships through the addition of characters from unsampled structures in the morphological analysis or introduction of new genes in the molecular analysis, but this is beyond the scope of the present study.
|
|
</paragraph>
|
|
<paragraph id="39940448E31788C6AA2061D8E87A5070" pageNumber="24">
|
|
Classification. Based upon a morphological phylogenetic analysis of the
|
|
<taxonomicName id="A6C8085CFB640108E76AAF61304310F1" family="Rivulidae" pageNumber="24" rank="family">Rivulidae</taxonomicName>
|
|
, Costa (1998a) proposed a new classification, in which
|
|
<taxonomicName id="CB95FB39C65CD7602BDCA915091214C1" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
was divided into four genera (
|
|
<taxonomicName id="D2C3ACCDEA2E0442DC91C66006768BBE" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
,
|
|
<taxonomicName id="2380057FD048D3714C1369BCAE0F8B32" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
,
|
|
<taxonomicName id="FB944E25F4B9671840FB9769C88AE9C8" LSID-ZBK="urn:lsid:zoobank.org:act:F913F3E6-E328-4D4A-BB0B-D7722E5A6786" family="Rivulidae" genus="Aphyolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Aphyolebias Costa 1998:78" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Aphyolebias</taxonomicName>
|
|
, and
|
|
<taxonomicName id="3EBAABC718FC66DBB23F14711E794403" LSID-ZBK="urn:lsid:zoobank.org:act:763FE250-1022-45BD-8F75-E7E164A2172E" family="Rivulidae" genus="Micromoema" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Micromoema Costa 1998:78" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Micromoema</taxonomicName>
|
|
). The groups recognized as genera were corroborated by subsequent molecular phylogenetic studies (Murphy et al, 1999; Hrbek & Larson, 1999), and the general classification has been employed both in scientific and aquaristic literature. However, Schindler (2004) considered
|
|
<taxonomicName id="0C1A481DFB26E3F66A1E86EA6211523B" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
to be a junior synonym of
|
|
<taxonomicName id="EAEBDFDA7273BB5D80B4708146D2F924" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
, claiming that
|
|
<taxonomicName id="9F35AFB31240FEDEDF2C1C440807BDCE" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
plus
|
|
<taxonomicName id="F7BCF92FB4622E4B3C9FD8B4DF388631" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
form a monophyletic group, and that “... it would be better to adopt a more conservative nomenclature than to change the systematic categories after every new analysis ...” (Schindler, 2004: 74). This proposal is not followed here by the following reasons:
|
|
</paragraph>
|
|
<paragraph id="2608983C8BE249A1458F171B2B74CF39" pageNumber="24">
|
|
1)
|
|
<taxonomicName id="71996280A11F053DDF0A49204A89C15C" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
including species placed in
|
|
<taxonomicName id="57394978D853A90CB67095A96514CCA8" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
plus
|
|
<taxonomicName id="65757856E37E63C38EDEBDF9AE83B564" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
sensu Costa (1998a) is not a conservative nomenclature as proposed by Schindler (2004), since this generic composition would be new.
|
|
<taxonomicName id="6737E6A1C0AE1B1CFD39CC12D9F13A28" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
was a more inclusive genus prior to publication of Costas (1998a) work, and included species today placed in
|
|
<taxonomicName id="AD8A18A0A5B5021483E36363C4E8BC84" LSID-ZBK="urn:lsid:zoobank.org:act:F913F3E6-E328-4D4A-BB0B-D7722E5A6786" family="Rivulidae" genus="Aphyolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Aphyolebias Costa 1998:78" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Aphyolebias</taxonomicName>
|
|
,
|
|
<taxonomicName id="897A06807A3E7E303F12CCB1990A9520" LSID-ZBK="urn:lsid:zoobank.org:act:22A92A57-BBC3-4553-B0E3-A2C95E8F6BD4" family="Rivulidae" genus="Moema" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Moema Costa 1989:223" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Moema</taxonomicName>
|
|
, and
|
|
<taxonomicName id="471E6345414124C392C51647E59BFC69" LSID-ZBK="urn:lsid:zoobank.org:act:763FE250-1022-45BD-8F75-E7E164A2172E" family="Rivulidae" genus="Micromoema" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Micromoema Costa 1998:78" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Micromoema</taxonomicName>
|
|
, thus forming a paraphyletic assemblage, as estimated both by morphological and molecular studies (Costa, 1998a; Murphy et al., 1999; Hrbek & Larson, 1999). Therefore,
|
|
<taxonomicName id="3AC3BC7D12E6D93A8B68E16E71F3B0BD" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
including species placed in
|
|
<taxonomicName id="B13F2BB387C2556EE213B622C9CAC31D" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
but excluding species of
|
|
<taxonomicName id="D46192B0B67EA4F4D56FF611C4783366" LSID-ZBK="urn:lsid:zoobank.org:act:F913F3E6-E328-4D4A-BB0B-D7722E5A6786" family="Rivulidae" genus="Aphyolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Aphyolebias Costa 1998:78" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Aphyolebias</taxonomicName>
|
|
,
|
|
<taxonomicName id="D48C9B7224E15C754843C7D988094502" LSID-ZBK="urn:lsid:zoobank.org:act:22A92A57-BBC3-4553-B0E3-A2C95E8F6BD4" family="Rivulidae" genus="Moema" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Moema Costa 1989:223" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Moema</taxonomicName>
|
|
, and
|
|
<taxonomicName id="E872F4F64D3A960DD6A0938FC93FB52F" LSID-ZBK="urn:lsid:zoobank.org:act:763FE250-1022-45BD-8F75-E7E164A2172E" family="Rivulidae" genus="Micromoema" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Micromoema Costa 1998:78" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Micromoema</taxonomicName>
|
|
should not be viewed as conservative, especially considering that Costas classification has been consistently employed during the last seven years.
|
|
</paragraph>
|
|
<paragraph id="F3F485B89932FBA6D2B3E98849773BBB" pageNumber="24">
|
|
2) Although the clade
|
|
<taxonomicName id="E7268D02F305685F5A3EEC61D36AFF4B" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
plus
|
|
<taxonomicName id="2676EBF94E15E9F3F5AD975737E0E624" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
is strongly supported by morphological characters (Costa, 1998a; the present study), it is still controversial, since it is not corroborated in two independent molecular phylogenetic analyses (Murphy et al., 1999; Hrbek & Larson, 1999) (Fig. 11). Thus,
|
|
<taxonomicName id="8CFC0F6EC7EB2B2171D879D40B37EC7B" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
, as proposed by Schindler (2004), would be paraphyletic according to molecular studies.
|
|
</paragraph>
|
|
<paragraph id="D9E6337842979021E565C14D24DEE944" pageNumber="24">
|
|
3) Both
|
|
<taxonomicName id="9A0DF9CC99758E1512CAAA5A21A66A86" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
and
|
|
<taxonomicName id="DEAE8BDDD156067F97FEB21D69FE923A" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
sensu Costa (1998a) are monophyletic assemblages, well corroborated by morphological and molecular studies (Costa, 1998a; the present study). In addition,
|
|
<taxonomicName id="5246C1BCE50DAFBF9CC020DE7D151CA4" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
and
|
|
<taxonomicName id="7C9006C30F61D392F08DC6FC81DCF935" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
are readily distinguished even by external morphological characters, easily accessible both by museum systematists and amateur ichthyologists (e. g., short snout, rounded pectoral fin, and dark bars on the pectoral fin in
|
|
<taxonomicName id="46ACF4FAB362EC94C93605145EEA1EF4" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
|
|
, vs. long snout, pointed pectoral fin, and pectoral fin with black ventral margin in
|
|
<taxonomicName id="1066E04C441EEBCA93197873DA4B8562" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
|
|
) (Fig. 12).
|
|
</paragraph>
|
|
<paragraph id="E7A1CA72CCC62F2D83FDE4CC822E6AED" pageNumber="24">
|
|
4) These two monophyletic groups (i. e.,
|
|
<taxonomicName id="B2E978B2FA277236E19B342F5F7551DF" LSID-ZBK="urn:lsid:zoobank.org:act:EF0F634C-4A82-4B33-86D6-31A498C4DB7C" family="Rivulidae" genus="Pterolebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Pterolebias Garman 1895:141" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Pterolebias</taxonomicName>
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and
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<taxonomicName id="6024A451C69341782CBAA646B3315CD3" LSID-ZBK="urn:lsid:zoobank.org:act:C8DC9E17-42A0-4D0C-A6E4-FBB00CD1ACF1" family="Rivulidae" genus="Gnatholebias" higherTaxonomySource="CoL" kingdom="Animalia" lsidName-ZBK="Gnatholebias Costa 1998:77" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Gnatholebias</taxonomicName>
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) are potential indicators of historic biogeography, since, although not necessarily sister groups, they are closely related taxa with completely disjunct biogeographic distribution patterns.
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</paragraph>
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</subSubSection>
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</treatment>
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</document> |