571 lines
92 KiB
XML
571 lines
92 KiB
XML
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<mods:title id="FCB6CE1B57F0057F969859282B2EC684">Description of three new species of the genus Chromaphyosemion Radda, 1971 (Cyprinodontiformes: Nothobranchiidae) from the coastal plains of Cameroon with a preliminary review of the Chromaphyosemion splendopleure complex.</mods:title>
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<mods:namePart id="221AD82F30FEFD67A22E98FAA5572880">Rainer Sonnenberg</mods:namePart>
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<mods:date id="5E75E4313E23EAB9A216215EFDC1D699">2007</mods:date>
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<treatment id="D7557997832E8C722633D886A898C941" ID-DOI="http://doi.org/10.5281/zenodo.6248389" ID-GBIF-Taxon="125136716" ID-Zenodo-Dep="6248389" LSID="urn:lsid:plazi:treatment:D7557997832E8C722633D886A898C941" httpUri="http://treatment.plazi.org/id/D7557997832E8C722633D886A898C941" lastPageNumber="28" pageNumber="20">
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<subSubSection id="F69C7B5D932E502A60DA94114ACB1C8B" type="nomenclature">
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<paragraph id="34EBBE42CFC580B2874C50CF452A2AF6" pageNumber="20">
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[[ Genus
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<taxonomicName id="7C56BD4755D3097D9B20AFB872FE1DEE" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="genus">Chromaphyosemion Radda</taxonomicName>
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]]
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</paragraph>
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</subSubSection>
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<subSubSection id="5617A8B4042E8D6944D37A8C01819C74" type="discussion">
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<paragraph id="BFE46A0BBACAD1CC2C80B245BF0D2E3D" pageNumber="20">DNA analysis</paragraph>
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<paragraph id="4B63D234F13742F14B425A3CD826F2D4" pageNumber="20">
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The alignment contains 795 bp for a set of 19 specimens and 713 bp for
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<taxonomicName id="34A8E6D9216BD70D821895001F4B919F" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="alpha">C. alpha</taxonomicName>
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. There were 238 variable positions of which 155 are parsimoniously informative. The average amount of uncorrected pair-wise sequence divergence was 8.08%, the maximum observed value 18.82% between
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<taxonomicName id="5CCC83FDC3EAD4A7036BE7E23CAC87E6" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="alpha">C. alpha</taxonomicName>
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and
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<taxonomicName id="5596C6BEDA3741715C1DBA3962A139C5" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="riggenbachi">C. riggenbachi</taxonomicName>
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. The sequences were translated into 264 amino acid positions of which 21 were variable and eight were parsimoniously informative. Average nucelotide frequencies were A = 24.9 %, C = 29.2 %, G = 15.7 %, and T(U) = 30.3 %, showing the characteristic anti-G bias of the mitochondrial genome base composition (
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<bibRefCitation id="A505E60BD6ECF81DE39422E6BE7D18D9" author="Zhang, D. - X. & Hewitt, G. M." pagination="247 - 251" refId="ref19801" refString="Zhang, D. - X. & Hewitt, G. M. (1996) Nuclear integrations: challenges for mitochondrial DNA markers. Trends in Ecology and Evolution, 11, 247 - 251." type="journal article" year="1996">Zhang & Hewitt 1996</bibRefCitation>
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).
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</paragraph>
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<paragraph id="EB3AF65144A6FE972932654F64086B18" lastPageNumber="21" pageNumber="20">
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The MP (Fig. 15) and Bayesian analyses (Fig. 16) yielded nearly identical topologies. Numbers on branches represent bootstrap values (MP topology) or posterior probabilities (Bayesian topology). The Bayesian tree shows more polytomies but does not contradict the MP tree.
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<taxonomicName id="D929E02E6072D787C7106A1049690555" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="alpha">Chromaphyosemion alpha</taxonomicName>
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is the phylogenetically basal species within
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<taxonomicName id="96FBDC78AB3908238748A166C1F04F62" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
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according to a larger dataset with different outgroup species (unpubl. data and
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<bibRefCitation id="12AE4F58D9868776B67539B2203D0889" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
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<normalizedToken id="6A61EF8E68BBFF6F39FF7C440D46FC55" originalValue="Agnèse">Agnese</normalizedToken>
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||
et al. [2006]
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||
</bibRefCitation>
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). The newly described
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<taxonomicName id="2B3CBDD687FB63AE17E4A86F3B2FB631" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="melinoeides">C. melinoeides</taxonomicName>
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is the sister species to
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<taxonomicName id="66B946BD44A89CFA59BAAEF5DE030993" LSID-ZBK="urn:lsid:zoobank.org:act:F722B043-3B19-4507-B897-8F3C3146B9B0" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion lugens Amiet 1991:85" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="lugens">C. lugens</taxonomicName>
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. One well supported clade in both trees contains, besides
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<taxonomicName id="CE879BEA41E56B460A7C61598D01D74F" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="loennbergii">C. loennbergii</taxonomicName>
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and
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<taxonomicName id="212309A11B5D4CDDE10D2A75D367B10E" LSID-ZBK="urn:lsid:zoobank.org:act:58293C95-476D-42FC-B52D-62A8938702E3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion punctulatum Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="punctulatum">C. punctulatum</taxonomicName>
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, two equally well supported groups of closely related species, marked as NC and SC. NC is the northern clade, a group which has its distribution approximately north of the Wouri. SC is the southern clade which is found south of the Wouri (Map 1). Within the northern clade,
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<taxonomicName id="F836655FA9AFBA4F2E82980B696FC5CD" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
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,
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<taxonomicName id="2A0377AC772B7CD62DFDB20FD4124363" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
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and
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<taxonomicName id="AAF2B61521E8A42C431DF4A6C8A851EE" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="volcanum">C. volcanum</taxonomicName>
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have been described by earlier authors. The southern clade contains, besides the undescribed populations from Bioko and Bimbia, the newly described species
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<taxonomicName id="1C26C26CA826E0D7F40001D2F1570E37" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="koungueense">C. koungueense</taxonomicName>
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and
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<taxonomicName id="742C6429DC0E47E9ABDF5FD9EF592811" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="omega">C. omega</taxonomicName>
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. Both had been formerly included in
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<taxonomicName id="6EBFF1107BBFC5319337D95AB815099F" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="20" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
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(
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<bibRefCitation id="2C06FD994F460AA5799A19EF43998985" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
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<normalizedToken id="B09959FEAAADC0E50A87541FAA0AAC9B" originalValue="Agnèse">Agnese</normalizedToken>
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et al. 2006
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</bibRefCitation>
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;
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<bibRefCitation id="C9E3A0840B4001FBAAAD74A5F2431A02" author="Amiet, J. L." refId="ref16267" refString="Amiet, J. L. (1987) Le Genre Aphyosemion Myers (Pisces, Teleostei, Cyprinodontiformes). Faune du Cameroun, Volume 2. Sciences Nat, Compiegne, France, 262 pp." type="book" year="1987">Amiet 1987</bibRefCitation>
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;
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<bibRefCitation id="3DC05F3F59601E0EE7ED2AA43D2CEC4C" author="Huber, J. H." pagination="15 - 23" refId="ref17026" refString="Huber, J. H. (1998 a) Aphyosemion alpha n. sp. - Eine neue Art der Untergattung Chromaphyosemion mit einem ausgepraegtem Faerbungsmuster und einer besonders suedlichen Verbreitung. Das Aquarium, 350, 15 - 23." type="journal article" year="1998">Huber 1998a</bibRefCitation>
|
||
;
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<bibRefCitation id="A7340B0D23F547F87463F43ED7637CC4" author="Legros, O. & Zentz, F." pagination="1 - 28" refId="ref17585" refString="Legros, O. & Zentz, F. (2007) Aphyosemion malumbresi n. sp., un nouveau Chromaphyosemion (Teleostei: Aplocheilidae) originaire de Guinee Equatoriale continentale. AKFB Killi-Contact, no. 6, decembre 2006, 1 - 28." type="journal article" year="2007">Legros & Zentz 2007</bibRefCitation>
|
||
;
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<bibRefCitation id="F09E960E48E44C30024219EB35601FE5" author="Wildekamp, R. H." refId="ref19762" refString="Wildekamp, R. H. (1993) A World of Killies. Atlas of the Oviparous Cyprinodontiform Fishes of the World. Volume 1. The American Killifish Association, Mishawaka, Indiana, 311 pp." type="book" year="1993">Wildekamp1993</bibRefCitation>
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). Within both groups genetic differentiation is low.
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</paragraph>
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<paragraph id="CC6050F4C95FBDBB94A83399F6918289" pageNumber="22">
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The mitochondrial cytochrome b sequences and the corresponding diagnostic region of a nuclear LSU fragment were compared for a subset of specimens, including two additional samples from populations with relevance for conflicting taxonomic hypotheses. This comparison shows incongruence in the grouping of
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<taxonomicName id="95A63D4DE874F47C3DB29C163D7514E0" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
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collected near Tiko (the presumed type locality) and the population of Mboro (CMM 18, as
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<taxonomicName id="132FB37569D2A191922451F06F5A630D" authority="in Agnese et al." authorityName="in Agnese" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" phylum="Chordata" rank="species" species="splendopleure">
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C. splendopleure in
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<bibRefCitation id="F894400E4204AB9CC148957019357A6B" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
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<normalizedToken id="7FF86327F4527CB3449DFDBD8BE33D0E" originalValue="Agnèse">Agnese</normalizedToken>
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et al. [2006]
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</bibRefCitation>
|
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</taxonomicName>
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) (Figs. 17 and 18). The nuclear marker does not support the inclusion of these populations in the southern clade as indicated by the mitochondrial fragment. Instead, the
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<taxonomicName id="00B7B70E80D4466C9B2A0E7F30FDB262" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
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from the area of Tiko is grouped into the northern clade (
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<taxonomicName id="06E39105C28E2FDE87D752C6EF85BB17" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
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and
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<taxonomicName id="9D96BD65A4AD4FEEEE19E8923AEA3964" authority="Agnese et al." authorityName="Agnese" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" phylum="Chordata" rank="species" species="volcanum">
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C. volcanum sensu
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<bibRefCitation id="C0047094803F3D8CE5038C5DF47E310D" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="54E60AD002E3C54E07048DDE49CCAE9B" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. [2006]
|
||
</bibRefCitation>
|
||
</taxonomicName>
|
||
), and the population from Mboro into
|
||
<taxonomicName id="B53D429D9B11938D0D4067EE5829350F" LSID-ZBK="urn:lsid:zoobank.org:act:58293C95-476D-42FC-B52D-62A8938702E3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion punctulatum Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="22" phylum="Chordata" rank="species" species="punctulatum">C. punctulatum</taxonomicName>
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||
, which in both cases is congruent with diagnostic color pattern characters.
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</paragraph>
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<paragraph id="45246DEB5B9A16F7EB8F2CF5C7C5CDD0" pageNumber="23">Discussion</paragraph>
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<paragraph id="8CE2E5138F130DDAC66B8838A41C5C6C" pageNumber="23">
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Research on the genus
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<taxonomicName id="C5DBD93C39A1801E0069988BDB742F6F" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
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was hampered until recently by a missing phylogenetic hypothesis and some confusion caused by insufficiently diagnosed and delimited taxa. Earlier attempts based on morphology, crossing experiments, karyotypes and color patterns could not resolve the relationships within this genus (
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<bibRefCitation id="CF2622BD5D761CC6956396021130A081" author="Huber, J. H." pagination="15 - 23" refId="ref17026" refString="Huber, J. H. (1998 a) Aphyosemion alpha n. sp. - Eine neue Art der Untergattung Chromaphyosemion mit einem ausgepraegtem Faerbungsmuster und einer besonders suedlichen Verbreitung. Das Aquarium, 350, 15 - 23." type="journal article" year="1998">Huber 1998a</bibRefCitation>
|
||
;
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||
<bibRefCitation id="65991C805565D0CB81F7A835C27C1995" author="Scheel, J. J." pagination="261 - 278" refId="ref18640" refString="Scheel, J. J. (1966) Notes on phenotypy, distribution and systematics of Aphyosemion bivittatum (Loennberg), with remarks on the chromosome number in the Rivulinae. Ichthyologica 37, 261 - 278." type="journal article" year="1966">Scheel 1966</bibRefCitation>
|
||
,
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||
<bibRefCitation id="745AE87849DDC4021EABD77FF779F0AC" author="Scheel, J. J." refId="ref18680" refString="Scheel, J. J. (1968) Rivulins of the Old World. TFH Publications, New Jersey, 480 pp." type="book" year="1968">1968</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D270AE53F320DE71E70D5090694EAAEB" author="Scheel, J. J." refId="ref18704" refString="Scheel, J. J. (1974) Rivuline Studies. Taxonomic Studies of Rivuline Cyprinodonts from Tropical Atlantic Africa (Rivulinae, Cyprinodontidae, Atheriniformes, Pisces). Annales du Musee royal de l’Afrique centrale, no. 211, Tervuren, Belgique, 150 pp." type="book" year="1974">1974</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="9768936741458C32E343120946F8B6CF" author="Scheel, J. J." refId="ref18754" refString="Scheel, J. J. (1990) Atlas of the Killifishes of the Old World. TFH Publication, Neptune City, New Jersey, 448 pp." type="book" year="1990">1990</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="091842315FD78DC41051F707B1006916" author="Sonnenberg, R." pagination="79 - 94" refId="ref19032" refString="Sonnenberg, R. (2000) The Distribution of Chromaphyosemion Radda, 1971 (Teleostei: Cyprinodontiformes) on the Coastal Plains of West and Central Africa. In Rheinwald, G. (Ed.) Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium, Bonn. Bonner zoologische Monographien 46, pp. 79 - 94." type="journal article" year="2000">Sonnenberg 2000</bibRefCitation>
|
||
). In his last publication,
|
||
<bibRefCitation id="142F94AA4123AFF03B5C93F78FAF6A1D" author="Scheel, J. J." refId="ref18754" refString="Scheel, J. J. (1990) Atlas of the Killifishes of the Old World. TFH Publication, Neptune City, New Jersey, 448 pp." type="book" year="1990">Scheel (1990)</bibRefCitation>
|
||
underestimated the diversity of
|
||
<taxonomicName id="22704ED472A308C8D578454A788D9207" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
|
||
as now presented by the molecular phylogenies (Figs. 15- 16,
|
||
<bibRefCitation id="D99022B712E5EFE355F940984E8F5B25" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="FFE5E238B5A9D9008EC0EFF7EA2A519D" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. 2006
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="747EA326C2489FA347BD6B48D0573E26" author="Legros, O. & Zentz, F." pagination="1 - 28" refId="ref17585" refString="Legros, O. & Zentz, F. (2007) Aphyosemion malumbresi n. sp., un nouveau Chromaphyosemion (Teleostei: Aplocheilidae) originaire de Guinee Equatoriale continentale. AKFB Killi-Contact, no. 6, decembre 2006, 1 - 28." type="journal article" year="2007">Legros & Zentz 2007</bibRefCitation>
|
||
). Especially
|
||
<taxonomicName id="AF694B91A7C9B9819C09179575661C63" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="bitaeniatum">C. bitaeniatum</taxonomicName>
|
||
in his sense is clearly a mixture of several different lineages with respect to the cladograms in Figures 15-16. It includes
|
||
<taxonomicName id="C329D6EA0E3C8F801FBDC2B32FE3DDD4" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="alpha">C. alpha</taxonomicName>
|
||
,
|
||
<taxonomicName id="0C7D4575F3592A08BA78C52E1ECA7E2C" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="bitaeniatum">C. bitaeniatum</taxonomicName>
|
||
,
|
||
<taxonomicName id="B81FCD0236DC5CCAA33736A557A0A9CF" LSID-ZBK="urn:lsid:zoobank.org:act:6DD323F2-FE35-4ED3-B8CD-FB129111161F" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion kouamense Legros 1999:33" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="kouamense">C. kouamense</taxonomicName>
|
||
,
|
||
<taxonomicName id="35852AB30687AD60590DAE9F3FED85A5" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="koungueense">C. koungueense</taxonomicName>
|
||
,
|
||
<taxonomicName id="D641FDDAFC1618047B761B6A48BD7570" LSID-ZBK="urn:lsid:zoobank.org:act:494F6568-0395-45FD-B6C4-99E11F027244" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion melanogaster Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="melanogaster">C. melanogaster</taxonomicName>
|
||
,
|
||
<taxonomicName id="527DE010DD83B0BB963A48ABC2EC590D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="omega">C. omega</taxonomicName>
|
||
,
|
||
<taxonomicName id="9F89EF823CA425F4EDE56AA0A94FE5AF" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
,
|
||
<taxonomicName id="C57EC06B7315699F99D3431B7DA9000D" LSID-ZBK="urn:lsid:zoobank.org:act:58293C95-476D-42FC-B52D-62A8938702E3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion punctulatum Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="punctulatum">C. punctulatum</taxonomicName>
|
||
, and
|
||
<taxonomicName id="A0EFEACAB78105AD78D0B52E957C3C82" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="BDB7A9474934E113CD4BDC8973A6EA21" pageNumber="23">
|
||
The value of color patterns as phylogenetic characters besides their doubtless taxonomic utility is very limited, and the grouping of
|
||
<taxonomicName id="B7D93E4DEF067A358E1FFD910BD74DC4" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
|
||
species given by
|
||
<bibRefCitation id="C01BA2F4C16BD619C7A1FAE67E851614" author="Huber, J. H." pagination="15 - 23" refId="ref17026" refString="Huber, J. H. (1998 a) Aphyosemion alpha n. sp. - Eine neue Art der Untergattung Chromaphyosemion mit einem ausgepraegtem Faerbungsmuster und einer besonders suedlichen Verbreitung. Das Aquarium, 350, 15 - 23." type="journal article" year="1998">Huber (1998a)</bibRefCitation>
|
||
is only partially compatible with the currently published molecular phylogenies. His groups turn out as para- or polyphyletic in the molecular phylogenies (Figs. 15-16,
|
||
<bibRefCitation id="8CBA13DF94D9D545D5EB15F3B04248D1" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="CF6A1458469FDBCAB8EDE3270CDBF202" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. 2006
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="6801986A4BAC185B05A91EC9C45F871F" author="Legros, O. & Zentz, F." pagination="1 - 28" refId="ref17585" refString="Legros, O. & Zentz, F. (2007) Aphyosemion malumbresi n. sp., un nouveau Chromaphyosemion (Teleostei: Aplocheilidae) originaire de Guinee Equatoriale continentale. AKFB Killi-Contact, no. 6, decembre 2006, 1 - 28." type="journal article" year="2007">Legros & Zentz 2007</bibRefCitation>
|
||
). Several characters used in his assignments intergrade: e.g., the absence of red or dark markings on anal fin in the
|
||
<taxonomicName id="9C9ECE78A102D132330D7F238EB63125" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="bitaeniatum">C. bitaeniatum</taxonomicName>
|
||
subgroup and the more or less streaked or spotted anal fin in the
|
||
<taxonomicName id="10E0BE09697C1F470612E289C1955E0E" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="loennbergii">C. loennbergii</taxonomicName>
|
||
subgroup. Streaks or dots in caudal and dorsal fins have no or only limited phylogenetic value because they occur in all groups and their presence can sometimes even vary within one species (e.g.,
|
||
<taxonomicName id="31FE729853D7C69404E2E2687A92310E" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="loennbergii">C. loennbergii</taxonomicName>
|
||
, Figs. 19-20). In addition red dots, red reticulation, or red blotches on sides are basically the same character and differ only quantitatively.
|
||
<bibRefCitation id="D4ED40E7E9486AE0E2FA15F9327E6250" author="Huber, J. H." pagination="15 - 23" refId="ref17026" refString="Huber, J. H. (1998 a) Aphyosemion alpha n. sp. - Eine neue Art der Untergattung Chromaphyosemion mit einem ausgepraegtem Faerbungsmuster und einer besonders suedlichen Verbreitung. Das Aquarium, 350, 15 - 23." type="journal article" year="1998">Huber's (1998a)</bibRefCitation>
|
||
<taxonomicName id="3C6B5909CEEE940F2661FE75BB4D5DB5" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="bitaeniatum">C. bitaeniatum</taxonomicName>
|
||
group is similar in its composition with Scheel's species
|
||
<taxonomicName id="19989EE1788F853D22B15E55915A7690" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="bitaeniatum">C. bitaeniatum</taxonomicName>
|
||
, and thus also not monophyletic (Figs. 15-16). It contains, besides the basal
|
||
<taxonomicName id="5759CED48FB0C4CE2006FAA3EA8183E2" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="alpha">C. alpha</taxonomicName>
|
||
, all species of the northern and southern clades and
|
||
<taxonomicName id="0D68DB79A1FC44AC7C0E469D3AD2F24B" LSID-ZBK="urn:lsid:zoobank.org:act:58293C95-476D-42FC-B52D-62A8938702E3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion punctulatum Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="punctulatum">C. punctulatum</taxonomicName>
|
||
, but not
|
||
<taxonomicName id="5DF28D3DF1AF3A81FEECEF911191BE64" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="loennbergii">C. loennbergii</taxonomicName>
|
||
and
|
||
<taxonomicName id="7E40888250CE03AA04C986F7F9ED0256" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
. The inclusion of
|
||
<taxonomicName id="747060F86EA535A46AF72C4544984A59" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
into his third group with
|
||
<taxonomicName id="F7436F24D4ADDFED4DA98B6D97245530" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="bivittatum">C. bivittatum</taxonomicName>
|
||
and
|
||
<taxonomicName id="1CFC5D220FB14456E33FC51C5E442597" LSID-ZBK="urn:lsid:zoobank.org:act:F722B043-3B19-4507-B897-8F3C3146B9B0" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion lugens Amiet 1991:85" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="species" species="lugens">C. lugens</taxonomicName>
|
||
is not supported by the molecular phylogeny. In general, this subdivision of
|
||
<taxonomicName id="C65A9F96C2568AC9022615A27FD289A3" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="23" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
|
||
conflicts in several points with the published molecular phylogenies (
|
||
<bibRefCitation id="9326A229F3E7C10169AFDD3887B44291" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="F2A24FD19DF235881E494F94E9B37D17" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. 2006
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="5F36CBE5CF5988C0ADFB0662BB402402" author="Legros, O. & Zentz, F." pagination="1 - 28" refId="ref17585" refString="Legros, O. & Zentz, F. (2007) Aphyosemion malumbresi n. sp., un nouveau Chromaphyosemion (Teleostei: Aplocheilidae) originaire de Guinee Equatoriale continentale. AKFB Killi-Contact, no. 6, decembre 2006, 1 - 28." type="journal article" year="2007">Legros & Zentz 2007</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="69B71C2B7EB35AD7CB1F7F1EDA8D0BB1" pageNumber="24">
|
||
The cladograms presented here are compatible with recently published results (
|
||
<bibRefCitation id="6794A9FD47BED2B08F3BF384881D5C02" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="695243952821E9250F414B11014C9860" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. 2006
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="8C9DC927B41E00568B160DF7808CAB52" author="Legros, O. & Zentz, F. & Agnese, J. F." pagination="3 - 30" refId="ref17632" refString="Legros, O., Zentz, F. & Agnese, J. F. (2005) Description de deux nouveaux Chromaphyosemion (Teleostei: Aplocheilidae) du sud de la Plaine littorale camerounaise. AKFB Killi-Contact, 33, 3 - 30." type="journal article" year="2005">Legros et al. 2005</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="63DE4CB165169D768D7DC3E560E20D0A" author="Legros, O. & Zentz, F." pagination="1 - 28" refId="ref17585" refString="Legros, O. & Zentz, F. (2007) Aphyosemion malumbresi n. sp., un nouveau Chromaphyosemion (Teleostei: Aplocheilidae) originaire de Guinee Equatoriale continentale. AKFB Killi-Contact, no. 6, decembre 2006, 1 - 28." type="journal article" year="2007">Legros & Zentz 2007</bibRefCitation>
|
||
). In cases of incongruences, these involve nodes with no significant bootstrap support or posterior probability and might be caused by the different mutation rates of the considered mitochondrial genes or unsufficient information content of the fragments.
|
||
</paragraph>
|
||
<paragraph id="4246D1585EF37A9E169DE898D082AE7F" pageNumber="24">
|
||
The analysis of the mitochondrial DNA data including all described species currently accepted as valid and most undescribed forms gives an overview of the relationships of this diverse genus. It allows the comparison of color patterns between the most closely related species and the evaluation of the distribution of certain patterns within a phylogenetic framework. Whereas earlier authors were overwhelmed by the diversity of male coloration and had mostly only sparse distribution data, it is now possible to recognize patterns in the distribution of the different color forms. Most of these patterns appear to be geographically localized groups of populations that correspond to already-named species. They are distinct from their surrounding congeners and represent well-separated units and are, with regard to the molecular data (
|
||
<bibRefCitation id="E1502055A807A8D0E841CCE1BF29057C" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="12D78E0C5D2D7D3C1035B6AEA296C1EF" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. 2006
|
||
</bibRefCitation>
|
||
; data presented here), well differentiated, supporting the value of color pattern characters in
|
||
<taxonomicName id="0B2C5AC1DDA7B4404481D63E8BCB833C" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
|
||
taxonomy.
|
||
</paragraph>
|
||
<paragraph id="E8CEEB633BD1582BB14E5400F717ACA0" pageNumber="24">
|
||
<taxonomicName id="9640154264DD5A20B849B9016ACF6D0E" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="species" species="melinoeides">Chromaphyosemion melinoeides</taxonomicName>
|
||
has a more basal position in the phylogenies (Figs. 15-16) and is distinguished from its closest relative,
|
||
<taxonomicName id="1BA526E597318FC6AD0A16E7122BD61D" LSID-ZBK="urn:lsid:zoobank.org:act:F722B043-3B19-4507-B897-8F3C3146B9B0" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion lugens Amiet 1991:85" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="species" species="lugens">C. lugens</taxonomicName>
|
||
, by consistent male and female diagnostic color patterns, which are stable over the known distribution area. It is assumed that the differences are not within the normal variation of these species and they are regarded as distinct evolutionary units, separated by stable, diagnostic characters which most probably belong to a complex of characters employed in mate-choice and/or species recognition. In addition, the currently known karyotypes of the two species differ considerably, with diploid chromosome numbers of 2n=36 in
|
||
<taxonomicName id="4D84FDF92D35CC914FC3E4E0E897F1FA" LSID-ZBK="urn:lsid:zoobank.org:act:F722B043-3B19-4507-B897-8F3C3146B9B0" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion lugens Amiet 1991:85" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="species" species="lugens">C. lugens</taxonomicName>
|
||
and 2n=28 in
|
||
<taxonomicName id="E693D086F0712D8681774251A3DE6640" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="species" species="melinoeides">C. melinoeides</taxonomicName>
|
||
(
|
||
<normalizedToken id="F9A2DE4EB2001C79BECB742EBA28C8CF" originalValue="Vôlker">Volker</normalizedToken>
|
||
et al. 2005; M.
|
||
<normalizedToken id="E8917F672555CFAF2E6BB20A20C70B88" originalValue="Vôlker">Volker</normalizedToken>
|
||
pers. comm.). The different karyotypes might have an influence on reproductive isolation between the two species. However, as mentioned before, karyotype variation is common in
|
||
<taxonomicName id="4D6145023A594B04500297DFA0E0A40F" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="24" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
|
||
(
|
||
<normalizedToken id="8255216973D5277A98C2D183DEE25476" originalValue="Vôlker">Volker</normalizedToken>
|
||
et al. 2005, 2006, 2007a,b, in press;
|
||
<bibRefCitation id="7B6E16638385D8217345EF5B9572AF6B" author="Scheel, J. J." refId="ref18704" refString="Scheel, J. J. (1974) Rivuline Studies. Taxonomic Studies of Rivuline Cyprinodonts from Tropical Atlantic Africa (Rivulinae, Cyprinodontidae, Atheriniformes, Pisces). Annales du Musee royal de l’Afrique centrale, no. 211, Tervuren, Belgique, 150 pp." type="book" year="1974">Scheel 1974</bibRefCitation>
|
||
), but the impact of karyotypic differences on reproductive incompatibilities is not known in detail and difficult to prove (
|
||
<bibRefCitation id="993CAFAB75E75267349F22144CD86956" author="King, M." refId="ref17188" refString="King, M. (1993) Species Evolution. The Role of Chromosome Change. Cambridge University Press, Cambridge, 336 pp." type="book" year="1993">King 1993</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="1BB0BFEFEB3CF2BFD6096DE9DF97CFA8" author="Rieseberg, L. H." pagination="351 - 358" refId="ref18510" refString="Rieseberg, L. H. (2001) Chromosomal rearrangements and speciation. Trends in Ecology and Evolution, 16, 351 - 358." type="journal article" year="2001">Rieseberg 2001</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="B625CC34F73BA6468A89D4D471D0B9DB" pageNumber="25">
|
||
However, within the most recent radiations (NC and SC, Figs. 15-16) molecular and karyotypic differentiation is weak despite pronounced differences of male color pattern between the included species (see e.g.,
|
||
<taxonomicName id="B94BD7AE52B7D4431292BF9BA7680603" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="25" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
, Fig. 21 and
|
||
<taxonomicName id="F1622C6E36A0C2E93CBDAE1FD21288DC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="25" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
, Figs. 22-23) (
|
||
<bibRefCitation id="DE0EFECFD406655B9B84659160C4A403" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="1F92E0279F2F1BAC886B1F6D3DADEF01" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. 2006
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="1B29DA6CC27B575F851D621FB86E5D16" author="Voelker, M. & Rab, P. & Kullmann, H." pagination="1 - 8" refId="ref19665" refString="Voelker, M., Rab, P. & Kullmann, H. (2007 b) Banded karyotypes of Chromaphyosemion poliaki and C. volcanum (Cyprinodontiformes, Nothobranchiidae), with a discussion of the validity of C. poliaki. Ichthyological Exploration of Freshwaters, 18, 1 - 8." type="journal article" year="2007">
|
||
<normalizedToken id="E71C9CAC2B9A92D66BEE3C5328275A2A" originalValue="Völker">Voelker</normalizedToken>
|
||
et al. 2007b
|
||
</bibRefCitation>
|
||
, in press;
|
||
<normalizedToken id="40F10C219F73FED0D38C16145AF5B88C" originalValue="Völker">Voelker</normalizedToken>
|
||
pers. comm.). This is in correspondence with the observation that characters under sexual selection can evolve at higher speed than others (e.g.,
|
||
<bibRefCitation id="BF9D8E3A383581AEC419F478AB495D98" author="Pomiankowski, A. & Iwasa, Y." pagination="5106 - 5111" refId="ref18212" refString="Pomiankowski, A. & Iwasa, Y. (1998) Runaway ornament diversity caused by Fisherian sexual selection. Proceedings of the National Academy of Science, USA, 95, 5106 - 5111." type="journal article" year="1998">Pomiankowski & Iwasa 1998</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D36E95BE5A868DF9B20D24F6552A3C61" author="Uy, J. A. C. & Borgia, G." pagination="273 - 278" refId="ref19393" refString="Uy, J. A. C. & Borgia, G. (2000) Sexual selection drives rapid divergence in bowerbird display traits. Evolution, 54, 273 - 278." type="journal article" year="2000">Uy & Borgia 2000</bibRefCitation>
|
||
), which is a possible explanation for the rapid speciation of cichlids in the great African lakes (
|
||
<bibRefCitation id="4FAC103017E5B48706A52BF515EC1AC1" author="Maan, M. E. & Seehausen, O. & Soederberg, L. & Johnson, L. & Ripmeester, E. A. & Mrosso, H. D. J. & Taylor, M. I. & van Dooren, T. J. M. & van Alphen, J. J. M." pagination="2445 - 2452" refId="ref17711" refString="Maan, M. E., Seehausen, O., Soederberg, L., Johnson, L., Ripmeester, E. A. P, Mrosso, H. D. J., Taylor, M. I., van Dooren, T. J. M. & van Alphen, J. J. M. (2004) Intraspecific sexual selection on a speciation trait, male coloration, in the Lake Victoria cichlid Pundamilia nyererei. Proceedings of the Royal Society, London, B, 271, 2445 - 2452." type="journal article" year="2004">Maan et al. 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="E6FB8F3CE51F78971F1547059091C184" author="Salzburger, W. & Niederstaetter, H. & Brandstaetter, A. & Berger, B. & Parson, W. & Snoeks, J. & Sturmbauer, C." pagination="257 - 266" refId="ref18569" refString="Salzburger, W., Niederstaetter, H., Brandstaetter, A., Berger, B., Parson, W., Snoeks, J. & Sturmbauer, C. (2006) Colourassortative mating among populations of Tropheus moorii, a cichlid fish from Lake Tanganyika, East Africa. Proceedings of the Royal Society, London, B, 273, 257 - 266." type="journal article" year="2006">Salzburger et al. 2006</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="821E18579D441850C87F2FF8C4A6204C" author="Seehausen, O." refId="ref19000" refString="Seehausen, O. (1999) Speciation and Species Richness in African Cichlids. Effects of Sexual Selection by Mate Choice. Proefschrift Universiteit Leiden, Amsterdam, 284 pp." type="book" year="1999">Seehausen 1999</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="4960FF6BCB6A18E1A91A31C94250003B" pageNumber="26">
|
||
Available names for population groups within the northern and southern clades were
|
||
<taxonomicName id="14D743C95A562C65B122F57C9F53336F" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
,
|
||
<taxonomicName id="78BB5CAD3F9D396F4AA9CBF031E652CA" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="volcanum">C. volcanum</taxonomicName>
|
||
and
|
||
<taxonomicName id="F54F96BBC3E64FEADE7E1426F77FA28E" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
. Whereas
|
||
<taxonomicName id="2DB629086D2628230B24318FDBB7E7C2" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
, which is endemic to the southern and southeastern flanks of Mount Cameroon, is easily distinguished from all surrounding populations by male color pattern (Fig. 21), it is difficult to find diagnostic and constant differences between the
|
||
<taxonomicName id="9AFF59883F003DD0DA4F51F5F7B28254" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
|
||
populations from Tiko and Kumba, the areas around the type localities of
|
||
<taxonomicName id="6750D9AD1076F2868CF0BE17BD5D8928" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
(Fig. 22) and
|
||
<taxonomicName id="9E1DD043F3F07CA5CADFC9F9AB7F4D1C" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="volcanum">C. volcanum</taxonomicName>
|
||
(see
|
||
<bibRefCitation id="73F6D56BBB811CA324C584087A4BD62D" author="Eberl, W." refId="ref16675" refString="Eberl, W. (1996) Die Untergattung Chromaphyosemion. Deutsche Killifisch Gemeinschaft Journal, Supplementheft 4, 88 pp." type="book" year="1996">Eberl 1996</bibRefCitation>
|
||
, p.52 for a picture of a specimen from the type locality), respectively.
|
||
<taxonomicName id="1E784574EE97A7B6430BB9E8679CE804" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="volcanum">Chromaphyosemion volcanum</taxonomicName>
|
||
was therefore until recently regarded as a synonym of
|
||
<taxonomicName id="FC70E60B7C05AB3D094B31355AAFBC95" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
(
|
||
<bibRefCitation id="B2810ED01A57F487EF9B1ACA31689DB0" author="Legros, O." pagination="56 - 61" refId="ref17515" refString="Legros, O. (1991) Diskussion ueber die Gueltigkeit der Art Aphyosemion volcanum. Deutsche Killifisch Gemeinschaft Journal, 23, 56 - 61." type="journal article" year="1991">Legros 1991</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="B4D18DC99FE4AE7B9FC2E5B984E3D1A8" author="Sonnenberg, R." pagination="79 - 94" refId="ref19032" refString="Sonnenberg, R. (2000) The Distribution of Chromaphyosemion Radda, 1971 (Teleostei: Cyprinodontiformes) on the Coastal Plains of West and Central Africa. In Rheinwald, G. (Ed.) Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium, Bonn. Bonner zoologische Monographien 46, pp. 79 - 94." type="journal article" year="2000">Sonnenberg 2000</bibRefCitation>
|
||
). The mitochondrial DNA analyses groups populations which are determined as
|
||
<taxonomicName id="3E8852E39B469D073EBFFF9D2DB80C4B" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
and
|
||
<taxonomicName id="24427191623043DCD1BE626DC3190836" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
based on coloration characters, together with some unnamed forms, into the northern clade. Additionally the distinct status of some of the populations of the southern clade was often recognized by earlier authors (
|
||
<bibRefCitation id="F00873FBC327B2419EC572707AE9C389" author="Eberl, W." refId="ref16675" refString="Eberl, W. (1996) Die Untergattung Chromaphyosemion. Deutsche Killifisch Gemeinschaft Journal, Supplementheft 4, 88 pp." type="book" year="1996">Eberl 1996</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="4FD0E9B0A283D42717AF93008B9C4727" author="Eigelshofen, W." pagination="8 - 13" refId="ref16697" refString="Eigelshofen, W. (1998) Chromaphyosemion. Beeindruckend in Form und Farbe. Aquaristik Aktuell, 9 - 10, 8 - 13." type="journal article" year="1998">Eigelshofen 1998</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="70E0C1BA77DF3A64AB7900E33FB7AB01" author="Legros, O. & Zentz, F. & Agnese, J. F." pagination="3 - 30" refId="ref17632" refString="Legros, O., Zentz, F. & Agnese, J. F. (2005) Description de deux nouveaux Chromaphyosemion (Teleostei: Aplocheilidae) du sud de la Plaine littorale camerounaise. AKFB Killi-Contact, 33, 3 - 30." type="journal article" year="2005">Legros et al. 2005</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="D677575093245450B03A087829004F91" author="Sonnenberg, R." pagination="79 - 94" refId="ref19032" refString="Sonnenberg, R. (2000) The Distribution of Chromaphyosemion Radda, 1971 (Teleostei: Cyprinodontiformes) on the Coastal Plains of West and Central Africa. In Rheinwald, G. (Ed.) Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium, Bonn. Bonner zoologische Monographien 46, pp. 79 - 94." type="journal article" year="2000">Sonnenberg 2000</bibRefCitation>
|
||
) but until now without taxonomic consequences.
|
||
</paragraph>
|
||
<paragraph id="62BB733B540E4FB662640E5589D001CC" pageNumber="26">
|
||
After submission of this manuscript,
|
||
<bibRefCitation id="CDF07752A7FF928F7E5D6C1E9E8D3336" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="045F86A9F89C3C6098A996EF477BA09F" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. (2006)
|
||
</bibRefCitation>
|
||
published a molecular study of
|
||
<taxonomicName id="D8AD6A837060638E791408FDB6394215" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
|
||
including a population from the vicinity of the proposed type locality of
|
||
<taxonomicName id="BCA192741C43B118A3D03EAC59C276CB" authority="Tiko" authorityName="Tiko" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure, Tiko</taxonomicName>
|
||
in Cameroon. Whereas the resulting phylogeny is largely congruent with that presented here, some of his taxonomic conclusions are different: the mitochondrial sequences of the Tiko population (CBL 01/25, entrance of Tiko) cluster within what is here named southern clade (Figs. 15-16). It includes the newly described species
|
||
<taxonomicName id="67CEE48BE6AF3E93B36A229CFBDA5B62" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="koungueense">C. koungueense</taxonomicName>
|
||
(Figs. 3-5),
|
||
<taxonomicName id="AE0E12A8B7A25A8D8F6DD76D33CE67AA" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="omega">C. omega</taxonomicName>
|
||
(Figs. 9-10) and the northernmost populations of
|
||
<taxonomicName id="5DE557006689C1AC0936F799357F5484" LSID-ZBK="urn:lsid:zoobank.org:act:58293C95-476D-42FC-B52D-62A8938702E3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion punctulatum Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="punctulatum">C. punctulatum</taxonomicName>
|
||
(Fig. 24), which are all clearly distinguished from
|
||
<taxonomicName id="AD8676A9911C8D70C699A3CDA5720228" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
(Figs. 22-23) by color pattern.
|
||
<bibRefCitation id="35D31BB02826A664B54CB742173C13B9" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="6080B06F87B09E4F003D540B083C4DCB" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. (2006)
|
||
</bibRefCitation>
|
||
assigned the name
|
||
<taxonomicName id="140C3C7F7AD085FE385420D2572AA2FA" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
to the Tiko population and the species mentioned in the previous sentence, based on mitochondrial DNA cluster, and state that this taxon 'exhibits the highest degree of phenotypic variability' and transferred the populations of the northern clade to
|
||
<taxonomicName id="8375544F8E36C92657EEE6FC50C0DEF1" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
and
|
||
<taxonomicName id="F19437AECE103D8EFFDB5B4D2206EAD8" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="volcanum">C. volcanum</taxonomicName>
|
||
. This taxonomic decision based on mitochondrial DNA did not take into account the diagnostic coloration characters and the incongruence was regarded as high phenotypic variability without further discussion and was followed in a later paper by
|
||
<bibRefCitation id="E7FA1CE49786CDC34F6C2B0034E83989" author="Legros, O. & Zentz, F." pagination="1 - 28" refId="ref17585" refString="Legros, O. & Zentz, F. (2007) Aphyosemion malumbresi n. sp., un nouveau Chromaphyosemion (Teleostei: Aplocheilidae) originaire de Guinee Equatoriale continentale. AKFB Killi-Contact, no. 6, decembre 2006, 1 - 28." type="journal article" year="2007">Legros and Zentz (2007)</bibRefCitation>
|
||
.
|
||
</paragraph>
|
||
<paragraph id="245BE411439789BF1A2162B845B2B204" pageNumber="26">
|
||
In order to clarify the incongruences, I sequenced a variable part of the nuclear large ribosomal subunit (LSU) to study the discrepancies with an additional independent marker. The comparison of the mitochondrial clusters and an excerption of the alignment of the nuclear gene fragment showing the most informative region (Figs. 17-18) indicates incongruences in two cases that are important for taxonomic conclusions: a) the clustering of the
|
||
<taxonomicName id="9F747312A6040B0FF90D4457B61B591F" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
collected near Tiko within the southern clade (SC) by the mitochondrial data is not congruent with the presented diagnostic LSU sequence fragment and b) a similar incongruence of the Mboro population. In both cases the grouping by diagnostic coloration characters is compatible with the LSU fragment but not with the clustering of the mitochondrial DNA. The inclusion of
|
||
<taxonomicName id="0AFF552264CFA73290CC8A16CF0BDB4F" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="koungueense">C. koungueense</taxonomicName>
|
||
,
|
||
<taxonomicName id="F355DD6F08474E2D98A7A5167C5C7244" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="omega">C. omega</taxonomicName>
|
||
and the northern populations of
|
||
<taxonomicName id="4EE6D3FC257B31CCA0C2AAC0926F6E46" LSID-ZBK="urn:lsid:zoobank.org:act:58293C95-476D-42FC-B52D-62A8938702E3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion punctulatum Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="punctulatum">C. punctulatum</taxonomicName>
|
||
in a 'polymorphic' taxon
|
||
<taxonomicName id="773102F5627CF780697DBF8C6D756334" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
based on mitochondrial DNA cluster is not supported by the nuclear and coloration pattern data; therefore, the taxonomic conclusions of
|
||
<bibRefCitation id="18B6C2D4441318F1D35A473193329C12" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="81C1F42B47EC69BA096604803489CF6F" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. (2006)
|
||
</bibRefCitation>
|
||
and
|
||
<bibRefCitation id="971C4B708F29A5ADEEB3B3FC34E6D1EF" author="Legros, O. & Zentz, F." pagination="1 - 28" refId="ref17585" refString="Legros, O. & Zentz, F. (2007) Aphyosemion malumbresi n. sp., un nouveau Chromaphyosemion (Teleostei: Aplocheilidae) originaire de Guinee Equatoriale continentale. AKFB Killi-Contact, no. 6, decembre 2006, 1 - 28." type="journal article" year="2007">Legros and Zentz (2007)</bibRefCitation>
|
||
are not followed here. The populations around Tiko belong, according to nuclear DNA and coloration characters, to the northern clade and
|
||
<taxonomicName id="264F53CFDCEA6DB4AEB61C5C5DA5A1C5" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="volcanum">C. volcanum</taxonomicName>
|
||
is again regarded as a synonym of
|
||
<taxonomicName id="9EE9AD5A4A4642B2096E9D864F624348" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
. The southern clade includes
|
||
<taxonomicName id="2F3977AB02174FB4AB0105993D960A1D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="koungueense">C. koungueense</taxonomicName>
|
||
and
|
||
<taxonomicName id="3FD8A55B2897049DE89503B89DBD3A3A" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="omega">C. omega</taxonomicName>
|
||
, together with the populations from Bioko and Bimbia. The population from Mboro is regarded as a belonging to
|
||
<taxonomicName id="3AB92E433B6093D10CD52E8BD03BF91E" LSID-ZBK="urn:lsid:zoobank.org:act:58293C95-476D-42FC-B52D-62A8938702E3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion punctulatum Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="26" phylum="Chordata" rank="species" species="punctulatum">C. punctulatum</taxonomicName>
|
||
, according to nuclear DNA and diagnostic color patterns.
|
||
</paragraph>
|
||
<paragraph id="95A5473333F28941E65A7AF6DB71D979" pageNumber="27">
|
||
The results of the nuclear marker are congruent with the species assignment by diagnostic coloration characters in those cases where the mtDNA gives a different result. As also coloration pattern characters have to be regarded as nuclear encoded marker, the observation of incongruences with the results of the mtDNA study is in line with a most probable mitochondrial introgression in the studied populations. The problem of species assignment based on only one marker, especially a mitochondrial marker, is currently discussed in the frame of genetic 'barcoding' approaches (e.g.
|
||
<bibRefCitation id="7CC612EB02B583364CE3F94AFB5F2ACB" author="Moritz, C. & Cicero, C." pagination="354" refId="ref17894" refString="Moritz, C. & Cicero, C. (2004) DNA barcoding: promise and pitfalls. PLoS Biology, 2, e 354." type="journal article" year="2004">Moritz & Cicero 2004</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="85C00670C08B51B78AC202A9EB676428" author="Sonnenberg, R. & Nolte, A. W. & Tautz, D." refId="ref19191" refString="Sonnenberg, R., Nolte, A. W. & Tautz, D. (2007) An evaluation of LSU rDNA Dl-D 2 sequences for their use in species identification. Frontiers in Zoology, 4, 12 pp." type="book" year="2007">Sonnenberg et al. 2007</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="B7D03CE7A29583F1FFA8087D55A5E3CC" lastPageNumber="28" pageNumber="27">
|
||
This is important in so far as
|
||
<bibRefCitation id="FB7A6F9576B0B6E0BE05E795237AA56E" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="A77D27237E8996EC0DB91866DAC240F7" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. (2006)
|
||
</bibRefCitation>
|
||
use only the mitochondrial DNA cluster as argument for their inclusion of the populations of the southern clade (SC) into
|
||
<taxonomicName id="ACB5E90BA89116B362274485AAE2EBBE" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="27" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
, which then includes the
|
||
<taxonomicName id="561B33B05CC9D81B6DE24AC14EBB0E45" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="27" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
population from the Tiko area and the here described species
|
||
<taxonomicName id="8CC87FCB10D7259FA177024F5ED3F07E" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="27" phylum="Chordata" rank="species" species="koungueense">C. koungueense</taxonomicName>
|
||
and
|
||
<taxonomicName id="A44533B17B70F02A37F64F99CA8C9C86" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="27" phylum="Chordata" rank="species" species="omega">C. omega</taxonomicName>
|
||
in addition to the northern populations of
|
||
<taxonomicName id="93936A7AC229004DB9652BE43C089DFE" LSID-ZBK="urn:lsid:zoobank.org:act:58293C95-476D-42FC-B52D-62A8938702E3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion punctulatum Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="27" phylum="Chordata" rank="species" species="punctulatum">C. punctulatum</taxonomicName>
|
||
which can all be assign by coloration pattern of males to different groups. As the diagnosis by coloration and the results of the LSU sequences are compatible, this means that at least two independent marker (if coloration is counted as only one nuclear encoded marker) are congruent and differ in the same cases with the mitochondrial results, it seems most probable that the conflicting mitochondrial data are caused by mitochondrial introgression.
|
||
</paragraph>
|
||
<paragraph id="C44611570AA92CF641C315A375DB8153" pageNumber="28">
|
||
Despite these results, the largest taxonomical problem within this genus was and in some part is the wastebasket taxon,
|
||
<taxonomicName id="24BE5028EC22DC8DD9A97E7BC799F7FA" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
. This species was described from a commercial import in 1929 (
|
||
<bibRefCitation id="D7E1F40CE77470BD25B5D1194F6FD2F4" author="Bruening, C." pagination="356" refId="ref16519" refString="Bruening, C. (1929 a) Der gruene Leuchtaugenfisch und der afrikanische Glanzkaerpfling. Wochenschrift fuer Aquarien- und Terrarienkunde, 26, 356." type="journal article" year="1929">
|
||
<normalizedToken id="6A6672367E28558F6D770C04F629C6A5" originalValue="Brüning">Bruening</normalizedToken>
|
||
1929a
|
||
</bibRefCitation>
|
||
), collected around Tiko at the foothills of Mount Cameroon. It was subsequently often synonymized with
|
||
<taxonomicName id="B110C42F01D3A396301AC0393501E265" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="bitaeniatum">C. bitaeniatum</taxonomicName>
|
||
or
|
||
<taxonomicName id="E4631C4F623F15DB76625E9A0D2361C3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="bivittatum">C. bivittatum</taxonomicName>
|
||
(
|
||
<bibRefCitation id="87FEDA3F409F27053F421F4715DDD2E6" author="Scheel, J. J." pagination="261 - 278" refId="ref18640" refString="Scheel, J. J. (1966) Notes on phenotypy, distribution and systematics of Aphyosemion bivittatum (Loennberg), with remarks on the chromosome number in the Rivulinae. Ichthyologica 37, 261 - 278." type="journal article" year="1966">Scheel 1966</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="7FC540DCF1171A83A9E8220C73D21236" author="Scheel, J. J." refId="ref18680" refString="Scheel, J. J. (1968) Rivulins of the Old World. TFH Publications, New Jersey, 480 pp." type="book" year="1968">1968</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="A2F0FC8C68B7794BF865D23A3BAB7301" author="Scheel, J. J." refId="ref18704" refString="Scheel, J. J. (1974) Rivuline Studies. Taxonomic Studies of Rivuline Cyprinodonts from Tropical Atlantic Africa (Rivulinae, Cyprinodontidae, Atheriniformes, Pisces). Annales du Musee royal de l’Afrique centrale, no. 211, Tervuren, Belgique, 150 pp." type="book" year="1974">1974</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="61093C3F324F6DD7CF4FF8F3F41DD10F" author="Scheel, J. J." refId="ref18754" refString="Scheel, J. J. (1990) Atlas of the Killifishes of the Old World. TFH Publication, Neptune City, New Jersey, 448 pp." type="book" year="1990">1990</bibRefCitation>
|
||
) or sometimes includes nearly all
|
||
<taxonomicName id="A05F01BD09599C2A57EEA3064EF926D2" LSID-ZBK="urn:lsid:zoobank.org:act:10DF0F9D-4060-4E5F-A156-DFE88550256D" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion Aphyosemion Radda 1971:118" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="genus">Chromaphyosemion</taxonomicName>
|
||
found on the Atlantic side of the coastal plains, except for
|
||
<taxonomicName id="75F0A81EC5FF8746A2AB1F456C01672A" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="bitaeniatum">C. bitaeniatum</taxonomicName>
|
||
(
|
||
<bibRefCitation id="4509E9DD23EBF64A74841CBA24C47462" author="Amiet, J. L." refId="ref16267" refString="Amiet, J. L. (1987) Le Genre Aphyosemion Myers (Pisces, Teleostei, Cyprinodontiformes). Faune du Cameroun, Volume 2. Sciences Nat, Compiegne, France, 262 pp." type="book" year="1987">Amiet 1987</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="A321FA46F5EC3069426E5CCFB5C91CEC" author="Huber, J. H." pagination="15 - 23" refId="ref17026" refString="Huber, J. H. (1998 a) Aphyosemion alpha n. sp. - Eine neue Art der Untergattung Chromaphyosemion mit einem ausgepraegtem Faerbungsmuster und einer besonders suedlichen Verbreitung. Das Aquarium, 350, 15 - 23." type="journal article" year="1998">Huber 1998a</bibRefCitation>
|
||
, b;
|
||
<bibRefCitation id="01A984E26D940E1948321CAE6873CDEA" author="Radda, A. C. & Puerzl, E." refId="ref18353" refString="Radda, A. C. & Puerzl, E. (1982) Killifische aus aller Welt. Feldfuehrer der Cyprinodontiformes der Laender der Bucht von Biafra (SO-Nigeria, West-Kamerun). Band 2. Verlag Otto Hofmann, Wien, 72 pp." type="book" year="1982">
|
||
Radda &
|
||
<normalizedToken id="C096345CC07B71C6EDEBD44DD54F76E1" originalValue="Pürzl">Puerzl</normalizedToken>
|
||
1982
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation id="D43288376BFC5559BDF502D396B02282" author="Radda, A. C. & Puerzl, E." refId="ref18442" refString="Radda, A. C. & Puerzl, E. (1987) Colour Atlas of Cyprinodonts of the Rain Forests of Tropical Africa. Verlag Otto Hofmann, Wien, 160 pp." type="book" year="1987">1987</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="2246B6469EA5661BDACC570957EAC090" author="Sonnenberg, R." pagination="79 - 94" refId="ref19032" refString="Sonnenberg, R. (2000) The Distribution of Chromaphyosemion Radda, 1971 (Teleostei: Cyprinodontiformes) on the Coastal Plains of West and Central Africa. In Rheinwald, G. (Ed.) Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium, Bonn. Bonner zoologische Monographien 46, pp. 79 - 94." type="journal article" year="2000">Sonnenberg 2000</bibRefCitation>
|
||
). In recent years several new species were described from this taxonomic wastebasket, namely
|
||
<taxonomicName id="7A29FF9EBA4B5666DA0ADC313BDF7B75" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="alpha">C. alpha</taxonomicName>
|
||
,
|
||
<taxonomicName id="B9A4CD7B164118C8C9ABB9CCCFC1EBA5" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="koungueense">C. koungueense</taxonomicName>
|
||
,
|
||
<taxonomicName id="CA112A5865F605ACA185231FDB140149" LSID-ZBK="urn:lsid:zoobank.org:act:494F6568-0395-45FD-B6C4-99E11F027244" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion melanogaster Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="melanogaster">C. melanogaster</taxonomicName>
|
||
,
|
||
<taxonomicName id="067D66FE79900F9B5BCE5B5E55D98210" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="omega">C. omega</taxonomicName>
|
||
,
|
||
<taxonomicName id="E80BE6692EDDCD077C2971B54F92843F" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
, and
|
||
<taxonomicName id="F312B2C17CC27CDFB13AF9C14BC4145C" LSID-ZBK="urn:lsid:zoobank.org:act:58293C95-476D-42FC-B52D-62A8938702E3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Chromaphyosemion punctulatum Legros, Zentz & Agnèse 2005" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="punctulatum">C. punctulatum</taxonomicName>
|
||
(
|
||
<bibRefCitation id="BD52809F431B7F7308083DF1E1C271CC" author="Amiet, J. L." pagination="83 - 95" refId="ref16305" refString="Amiet, J. L. (1991) Diagnoses de deux especes nouvelles d' Aphyosemion du Cameroun (Teleostei: Aplocheilidae). Ichthyological Exploration of Freshwaters, 2, 83 - 95." type="journal article" year="1991">Amiet 1991</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="BBBEE16D6B5424F263EFA8977B7686B5" author="Huber, J. H." pagination="15 - 23" refId="ref17026" refString="Huber, J. H. (1998 a) Aphyosemion alpha n. sp. - Eine neue Art der Untergattung Chromaphyosemion mit einem ausgepraegtem Faerbungsmuster und einer besonders suedlichen Verbreitung. Das Aquarium, 350, 15 - 23." type="journal article" year="1998">Huber 1998a</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="09310BF774C5D42B041D1025A30769EC" author="Legros, O. & Zentz, F. & Agnese, J. F." pagination="3 - 30" refId="ref17632" refString="Legros, O., Zentz, F. & Agnese, J. F. (2005) Description de deux nouveaux Chromaphyosemion (Teleostei: Aplocheilidae) du sud de la Plaine littorale camerounaise. AKFB Killi-Contact, 33, 3 - 30." type="journal article" year="2005">Legros et al. 2005</bibRefCitation>
|
||
; this paper). All these species are only more or less closely related and their inclusion in
|
||
<taxonomicName id="70558E8F77FE969430C112BEEE3CE0CF" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
was never based on diagnostic characters. The type locality is only approximately known and within this area several differently colored forms can be found. Currently a form with very red to orange coloration on fins and body can be distinguished (Figs. 22- 23, as SPP in Figs. 15-18, Map 1), which is found around Tiko, Moliwe and also in the area of Kumba (which was described as
|
||
<taxonomicName id="1F4BF7254E41AA959B51D26494315077" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="volcanum">C. volcanum</taxonomicName>
|
||
) and shares many characters with
|
||
<taxonomicName id="4E7ECD33749698417494687EB73DA936" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
as given in the description by
|
||
<bibRefCitation id="FE8E4444F05771B56E0D3F1C1381B5E3" author="Bruening, C." pagination="356" refId="ref16519" refString="Bruening, C. (1929 a) Der gruene Leuchtaugenfisch und der afrikanische Glanzkaerpfling. Wochenschrift fuer Aquarien- und Terrarienkunde, 26, 356." type="journal article" year="1929">
|
||
<normalizedToken id="06FE989C86235E487D2B902F37232B6C" originalValue="Brüning">Bruening</normalizedToken>
|
||
(1929a)
|
||
</bibRefCitation>
|
||
and later
|
||
<bibRefCitation id="1E1E9AE60DB2B43C4A8F7AA233DAB20F" author="Meinken, H." pagination="17 - 20" refId="ref17837" refString="Meinken, H. (1930 a) Zwei neue Fundulopanchaxarten aus Kamerun. Wochenschrift fuer Aquarien- und Terrarienkunde, 27, 17 - 20." type="journal article" year="1930">Meinken (1930a)</bibRefCitation>
|
||
; a second form with bluish to pink sides and yellow to greenish fins (Fig. 25, as cf SPP in Figs. 15-18, Map 1), which is known from the eastern flanks of Mount Cameroon at Owe, Lykoko and Muyuka and further north with one population known from Oron (Cross river system) in Nigeria (
|
||
<bibRefCitation id="42F02F29B8975B4C6F2E57C71657D806" author="Radda, A. C. & Puerzl, E." refId="ref18353" refString="Radda, A. C. & Puerzl, E. (1982) Killifische aus aller Welt. Feldfuehrer der Cyprinodontiformes der Laender der Bucht von Biafra (SO-Nigeria, West-Kamerun). Band 2. Verlag Otto Hofmann, Wien, 72 pp." type="book" year="1982">
|
||
Radda &
|
||
<normalizedToken id="37E59C7190390A807B1C23F4E76B4766" originalValue="Pürzl">Puerzl</normalizedToken>
|
||
1982
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="CC76E70CFB2B2F0CF6C569D19872E438" author="Sonnenberg, R." pagination="79 - 94" refId="ref19032" refString="Sonnenberg, R. (2000) The Distribution of Chromaphyosemion Radda, 1971 (Teleostei: Cyprinodontiformes) on the Coastal Plains of West and Central Africa. In Rheinwald, G. (Ed.) Isolated Vertebrate Communities in the Tropics. Proceedings of the 4 th International Symposium, Bonn. Bonner zoologische Monographien 46, pp. 79 - 94." type="journal article" year="2000">Sonnenberg 2000</bibRefCitation>
|
||
); and a group of populations called
|
||
<taxonomicName id="0EF0302EC8BF8E373DD5E94DFDD351CA" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="sp. 08">C. sp. 08</taxonomicName>
|
||
which were first mentioned by
|
||
<bibRefCitation id="91D441E2E4749D0848CABFF863A70119" author="Amiet, J. L." refId="ref16267" refString="Amiet, J. L. (1987) Le Genre Aphyosemion Myers (Pisces, Teleostei, Cyprinodontiformes). Faune du Cameroun, Volume 2. Sciences Nat, Compiegne, France, 262 pp." type="book" year="1987">Amiet (1987)</bibRefCitation>
|
||
and considered to be
|
||
<taxonomicName id="EB301321EBEA7BEAEAC4E9293E975BEA" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="loennbergii">C. loennbergii</taxonomicName>
|
||
populations north of the Sanaga (Fig. 26, as sp8 in Figs. 15-18; Map 1). Additionally there are several populations found southeast of a line Tiko-Kumba, which are difficult to add to one of the former three forms based on color patterns. All these populations, together with
|
||
<taxonomicName id="96C8AF09BE4DF37FDF84E9F803851513" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
(Fig. 21), form a weakly differentiated group according to the mitochondrial data (
|
||
<bibRefCitation id="B2740767267145B89D08DA1D87A7CF0A" author="Agnese, J. - F. & Zentz, F. & Legros, O. & Sellos, D." pagination="332 - 346" refId="ref16129" refString="Agnese, J. - F., Zentz, F., Legros, O. & Sellos, D. (2006) Phylogenetic relationships and phylogeography of the killifish species of the subgenus Chromaphyosemion (Radda, 1971) in West Africa, inferred from mitochondrial DNA sequences. Molecular Phylogenetics and Evolution, 40, 332 - 346." type="journal article" year="2006">
|
||
<normalizedToken id="B4C67EE7703A41EAB69CE07A7AEDD9D3" originalValue="Agnèse">Agnese</normalizedToken>
|
||
et al. 2006
|
||
</bibRefCitation>
|
||
, Figs. 15-16). The exclusion and description of
|
||
<taxonomicName id="900C0F06DF7BF9473BD36C3FBB635625" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="koungueense">C. koungueense</taxonomicName>
|
||
and
|
||
<taxonomicName id="B84D4B36F45D866B5D18B4DC894F5FA3" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="omega">C. omega</taxonomicName>
|
||
in this publication restricts the still problematic taxon
|
||
<taxonomicName id="1A1D1036970B88E664FD3CD4EB5AEC9F" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
to a smaller and phylogenetically closely related groups of populations, which form the northern clade. Still it is currently difficult, with the exception of
|
||
<taxonomicName id="7F1D355DFA46727C0BC97F30EAA94D35" LSID-ZBK="urn:lsid:zoobank.org:act:FD680552-D332-415D-8B46-60937B2486FC" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" lsidName-ZBK="Aphyosemion Chromaphyosemion poliaki Amiet 1991:89" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="poliaki">C. poliaki</taxonomicName>
|
||
, to give unambiguous combinations of diagnostic color characters for these forms. In addition, it remains problematic to assign the specimens, on which the description of
|
||
<taxonomicName id="A32D836A3869330F2C176CD51BEF0B51" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
is based, to one of two biogeographically possible forms. As the description of
|
||
<taxonomicName id="4BC04D927FC3F60028141E84CE5116A7" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
was based on live specimens, a lot of information on male color pattern is known (
|
||
<bibRefCitation id="ABCA1AB0DB82F570BFD3555C26D26642" author="Bruening, C." pagination="356" refId="ref16519" refString="Bruening, C. (1929 a) Der gruene Leuchtaugenfisch und der afrikanische Glanzkaerpfling. Wochenschrift fuer Aquarien- und Terrarienkunde, 26, 356." type="journal article" year="1929">
|
||
<normalizedToken id="0E3AA1BE4B4835FE4C485094FD1D21A3" originalValue="Brüning">Bruening</normalizedToken>
|
||
1929a
|
||
</bibRefCitation>
|
||
, b;
|
||
<bibRefCitation id="7669A54B671852DC98F6129A3601F382" author="Meinken, H." pagination="17 - 20" refId="ref17837" refString="Meinken, H. (1930 a) Zwei neue Fundulopanchaxarten aus Kamerun. Wochenschrift fuer Aquarien- und Terrarienkunde, 27, 17 - 20." type="journal article" year="1930">Meinken1930a</bibRefCitation>
|
||
) but actually it is not possible to decide on which form the description is based. I currently find better evidence for the more reddish form with orange-yellow anal fin, based on
|
||
<bibRefCitation id="CEC5246A31D0043D3DD80259A0F83B68" author="Bruening, C." pagination="356" refId="ref16519" refString="Bruening, C. (1929 a) Der gruene Leuchtaugenfisch und der afrikanische Glanzkaerpfling. Wochenschrift fuer Aquarien- und Terrarienkunde, 26, 356." type="journal article" year="1929">
|
||
<normalizedToken id="B26944FC6493AC54BCF27D60615D5447" originalValue="Brünings">Bruenings</normalizedToken>
|
||
(1929a)
|
||
</bibRefCitation>
|
||
and
|
||
<bibRefCitation id="38519EEF15792C05259F127F3D09239E" author="Meinken, H." pagination="17 - 20" refId="ref17837" refString="Meinken, H. (1930 a) Zwei neue Fundulopanchaxarten aus Kamerun. Wochenschrift fuer Aquarien- und Terrarienkunde, 27, 17 - 20." type="journal article" year="1930">Meinkens (1930a)</bibRefCitation>
|
||
color description (sides metallic golden or greenish-blue, ventral and on caudal peduncle golden yellow, color of dorsal fin red-brown in center, anal fin greenish-yellow at base and golden-yellow in distal part (
|
||
<bibRefCitation id="BBC770B8A9B2E48A6C54F3D5D3C95C39" author="Bruening, C." pagination="356" refId="ref16519" refString="Bruening, C. (1929 a) Der gruene Leuchtaugenfisch und der afrikanische Glanzkaerpfling. Wochenschrift fuer Aquarien- und Terrarienkunde, 26, 356." type="journal article" year="1929">
|
||
<normalizedToken id="5E0BF124AC3CE09DBB03AB64402EBA02" originalValue="Brüning">Bruening</normalizedToken>
|
||
1929a
|
||
</bibRefCitation>
|
||
;
|
||
<bibRefCitation id="DBD2A4894613D1E4F0428E7137DA6706" author="Meinken, H." pagination="17 - 20" refId="ref17837" refString="Meinken, H. (1930 a) Zwei neue Fundulopanchaxarten aus Kamerun. Wochenschrift fuer Aquarien- und Terrarienkunde, 27, 17 - 20." type="journal article" year="1930">Meinken 1930a</bibRefCitation>
|
||
)). This opinion is expressed in the application of the name
|
||
<taxonomicName id="F67088B8E5845DFA4DA24413D2FF0F78" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" species="splendopleure">C. splendopleure</taxonomicName>
|
||
for this form in the present paper. It does not anticipate any other decision in the future and a final solution has to be found before the taxonomy of this group can be solved.
|
||
</paragraph>
|
||
<paragraph id="652735EC0252D2B09D47885F6693D543" pageNumber="28">
|
||
As a first step toward a stable taxonomy, which takes phylogenetic information into consideration, it is now possible to remove all the taxa included in
|
||
<taxonomicName id="58BB3DDFEEE960999F06C374C385C0B8" authority="Huber (2000)" authorityName="Huber" authorityYear="2000" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" phylum="Chordata" rank="species" species="splendopleure">
|
||
C. splendopleure sensu
|
||
<bibRefCitation id="BD6ECB5B7B0DF8407B1E0549C62085A5" author="Huber, J. H." refId="ref17097" refString="Huber, J. H. (2000) Killi - Data 2000. Societe Francaise d'Ichthyologie, Paris (german edition), 450 pp." type="book" year="2000">Huber (2000)</bibRefCitation>
|
||
</taxonomicName>
|
||
or
|
||
<taxonomicName id="43DAE636963DFDB1D375B09402173153" authority="Scheel (1990)" authorityName="Scheel" authorityYear="1990" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" phylum="Chordata" rank="species" species="bitaeniatum">
|
||
C. bitaeniatum sensu
|
||
<bibRefCitation id="0B9E9D07F701C42C16AE1AA5EDFE8A9C" author="Scheel, J. J." refId="ref18754" refString="Scheel, J. J. (1990) Atlas of the Killifishes of the Old World. TFH Publication, Neptune City, New Jersey, 448 pp." type="book" year="1990">Scheel (1990)</bibRefCitation>
|
||
</taxonomicName>
|
||
, which are not related to the clade containing
|
||
<taxonomicName id="238192C8F5D70BFB7C5087C42676D211" class="Actinopterygii" family="Nothobranchiidae" genus="Chromaphyosemion" kingdom="Animalia" order="Cyprinodontiformes" pageNumber="28" phylum="Chordata" rank="species" sensu="stricto" species="splendopleure" status="s.s.">C. splendopleure</taxonomicName>
|
||
<taxonomicNameLabel id="A90BC71A2CD36B25D512646F685D5BE9" sensu="stricto">s.s.</taxonomicNameLabel>
|
||
(the northern clade NC) and are distinguishable by stable characters of the male color pattern. This allows the reduction of a polymorphic or variable species into diagnosable units with the additional argument of support of phylogenetic information. I want to stress that the diagnoses of the new species are also possible without the molecular phylogeny.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |