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<mods:title id="A2113F24F36B0D951179B33867078B2E">Phylogenetic position of Dendropsophus gaucheri (Lescure and Marty 2000) highlights the need for an in-depth investigation of the phylogenetic relationships of Dendropsophus (Anura: Hylidae)</mods:title>
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<mods:namePart id="6DB754350A9E865C45C72E3CB0D64BF6">Noonan, Brice P.</mods:namePart>
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<mods:namePart id="EEB6E28C23AA929FDA3393E003DBDCE0">Blanc, Michel</mods:namePart>
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<mods:namePart id="ED0143E487AF17CDCEE189F71220AA9D">Orrico, Victor Goyannes Dill</mods:namePart>
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<heading id="D0C03D18EA32FFA8CEB05A29FC470616" bold="true" box="[151,954,1123,1150]" fontSize="11" level="1" pageId="6" pageNumber="65" reason="1">
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<taxonomicName id="4C37F1F7EA32FFA8CEB05A29FE350615" ID-CoL="34S5C" box="[151,456,1124,1150]" class="Amphibia" family="Hylidae" genus="Dendropsophus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="gaucheri">
<emphasis id="B9435666EA32FFA8CEB05A29FE350615" bold="true" box="[151,456,1124,1150]" italics="true" pageId="6" pageNumber="65">Dendropsophus gaucheri</emphasis>
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: Distribution and Evolutionary History
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</paragraph>
<paragraph id="8B888A74EA32FFA8CEB05AE7FF17071D" blockId="6.[151,1436,1194,2011]" pageId="6" pageNumber="65">
All specimens of 
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<emphasis id="B9435666EA32FFA8CF435AE1FE1106A8" box="[356,492,1195,1219]" italics="true" pageId="6" pageNumber="65">D. gaucheri</emphasis>
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analyzed herein are not only recovered as a monophyletic group but also display strikingly little genetic distance among them (Mean K2P distance = 0.009). Even if not included in the molecular analyses the conspecificity of the populations in isolated savannas in Pará (
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) is unambiguous. This confirms the occurrence of 
<taxonomicName id="4C37F1F7EA32FFA8CF795B55FE190745" box="[350,484,1302,1327]" class="Amphibia" family="Hylidae" genus="Dendropsophus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="gaucheri">
<emphasis id="B9435666EA32FFA8CF795B55FE190745" box="[350,484,1302,1327]" italics="true" pageId="6" pageNumber="65">D. gaucheri</emphasis>
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populations over the eastern part of the 
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shield. This species is co-occurring in this region with at least four other member of the 
<taxonomicName id="4C37F1F7EA32FFA8CCFF5B71FC5D0738" box="[728,928,1339,1363]" class="Amphibia" family="Hylidae" genus="Dendropsophus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="microcephalus">
<emphasis id="B9435666EA32FFA8CCFF5B71FC5D0738" box="[728,928,1339,1363]" italics="true" pageId="6" pageNumber="65">D. microcephalus</emphasis>
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group (
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<emphasis id="B9435666EA32FFA8CDDB5B71FBAE0738" box="[1020,1107,1339,1363]" italics="true" pageId="6" pageNumber="65">D. leali</emphasis>
</taxonomicName>
, 
<taxonomicName id="4C37F1F7EA32FFA8CA445B71FAF80738" box="[1123,1285,1339,1363]" class="Amphibia" family="Hylidae" genus="Dendropsophus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="minusculus">
<emphasis id="B9435666EA32FFA8CA445B71FAF80738" box="[1123,1285,1339,1363]" italics="true" pageId="6" pageNumber="65">D. minusculus</emphasis>
</taxonomicName>
, 
<emphasis id="B9435666EA32FFA8CB325B71FAAF0738" box="[1301,1362,1340,1363]" italics="true" pageId="6" pageNumber="65">D. sp</emphasis>
. &amp; 
<taxonomicName id="4C37F1F7EA32FFA8CBA65B71FF27071C" class="Amphibia" family="Hylidae" genus="Dendropsophus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="nanus">
<emphasis id="B9435666EA32FFA8CBA65B71FF27071C" italics="true" pageId="6" pageNumber="65">D. nanus</emphasis>
</taxonomicName>
).
</paragraph>
<paragraph id="8B888A74EA32FFA8CEE25BCFFB5304FD" blockId="6.[151,1436,1194,2011]" pageId="6" pageNumber="65">
The 
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locality (Sinnamary, 
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) is situated on the oriental limit of a formation of coastal savannas occurring almost continuously through 
<collectingCountry id="F320CAE4EA32FFA8CCA25BEBFD0D07D5" box="[645,752,1446,1470]" name="Suriname" pageId="6" pageNumber="65">Suriname</collectingCountry>
to the eastern coast of 
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. According to 
<bibRefCitation id="EFA6F785EA32FFA8CB275BEBFEF80789" pageId="6" pageNumber="65" refString="Avila-Pires, T. C. S., Hoogmoed, M. S. &amp; da Rocha, W. A. (2010) Notes on the Vertebrates of northern Para, Brazil: a forgotten part of the Guianan Region, I. Herpetofauna. Boletim do Museu Paraense Emilio Goeldi, 5 (1), 13 - 112." type="journal article">
Ávila-Pires 
<emphasis id="B9435666EA32FFA8CBAF5BE5FF500788" italics="true" pageId="6" pageNumber="65">et al</emphasis>
. (2010)
</bibRefCitation>
<taxonomicName id="4C37F1F7EA32FFA8CF2C5B81FE6C0788" box="[267,401,1483,1507]" class="Amphibia" family="Hylidae" genus="Dendropsophus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="gaucheri">
<emphasis id="B9435666EA32FFA8CF2C5B81FE6C0788" box="[267,401,1483,1507]" italics="true" pageId="6" pageNumber="65">D. gaucheri</emphasis>
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also occurs in coastal savannas of 
<collectingCountry id="F320CAE4EA32FFA8CD335B87FC7E0789" box="[788,899,1482,1506]" name="Suriname" pageId="6" pageNumber="65">Suriname</collectingCountry>
. Therefore the species probably occurs throughout this area, possibly as far as 
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and even 
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. On the east however, one hilly forested area on the 
<collectingCountry id="F320CAE4EA32FFA8CEB0585FFEC20441" box="[151,319,1554,1578]" name="French Guiana" pageId="6" pageNumber="65">French Guiana</collectingCountry>
coast apparently constitutes a barrier that limits the connection with coastal savannas in Amapá (
<collectingCountry id="F320CAE4EA32FFA8CEB8587BFF150425" box="[159,232,1590,1614]" name="Brazil" pageId="6" pageNumber="65">Brasil</collectingCountry>
). This region represents a distributional edge for many open habitat species (
<bibRefCitation id="EFA6F785EA32FFA8CA64587BFAB70425" author="Lescure" box="[1091,1354,1590,1615]" pageId="6" pageNumber="65" refString="Lescure, J. &amp; Marty, C. (2000) Atlas des amphibiens du Guyane. Patrimoine Naturel, 45, Paris, Museum National d'Histoire Naturelle, 388 pp." type="book" year="2000">Lescure &amp; Marty 2000</bibRefCitation>
). However, as the species is known to also occur in isolated savannas in the interior of the 
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shield close to the Amazon River, it seems likely that 
<taxonomicName id="4C37F1F7EA32FFA8CC0658CDFD5A04FD" box="[545,679,1662,1687]" class="Amphibia" family="Hylidae" genus="Dendropsophus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="gaucheri">
<emphasis id="B9435666EA32FFA8CC0658CDFD5A04FD" box="[545,679,1662,1687]" italics="true" pageId="6" pageNumber="65">D. gaucheri</emphasis>
</taxonomicName>
also occurs in the coastal savannas in Amapá.
</paragraph>
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Interestingly, the genetic distance between the sample from 
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and samples from interior savannas is low. Given the fragmented distribution of the habitat of this species, this isolation is probably recent. Since the data are very limited, any further interpretation is subject to caution. Nevertheless, we argue that these data provide interesting support for hypothesized forest openings during Quaternary, a highly controversial subject (
<bibRefCitation id="EFA6F785EA32FFA8CB765943FF2E0521" author="Haffer" pageId="6" pageNumber="65" refString="Haffer, J. (2008) Hypotheses to explain the origin of species in Amazonia. Brazilian Journal of Biology, 68, 917 - 947." type="journal article" year="2008">Haffer 2008</bibRefCitation>
; 
<bibRefCitation id="EFA6F785EA32FFA8CEFA597FFE2F0521" author="Colinvaux" box="[221,466,1842,1867]" pageId="6" pageNumber="65" refString="Colinvaux, P. A., De Oliveira, P. E. &amp; Bush, M. B. (2000) Amazonian and Neotropical plant communities on glacial time-scales: The failure of the aridity and refuge hypotheses. Quaternary Science Reviews, 19, 141 - 169." type="journal article" year="2000">
Colinvaux 
<emphasis id="B9435666EA32FFA8CF7F5979FE750520" box="[344,392,1843,1867]" italics="true" pageId="6" pageNumber="65">et al</emphasis>
. 2000
</bibRefCitation>
; 
<bibRefCitation id="EFA6F785EA32FFA8CFFA597FFDA40521" author="Bush" box="[477,601,1842,1866]" pageId="6" pageNumber="65" refString="Bush, M. B. (1994) Amazonian speciation - a necessarily complex model. Journal of Biogeography, 21, 5 - 17." type="journal article" year="1994">Bush 1994</bibRefCitation>
). A similar pattern of isolated populations with low genetic divergence has been observed in 
<taxonomicName id="4C37F1F7EA32FFA8CF78591BFDD80505" box="[351,549,1878,1902]" class="Reptilia" family="Viperidae" genus="Crotalus" kingdom="Animalia" order="Squamata" pageId="6" pageNumber="65" phylum="Chordata" rank="species" species="durissus">
<emphasis id="B9435666EA32FFA8CF78591BFDD80505" box="[351,549,1878,1902]" italics="true" pageId="6" pageNumber="65">Crotalus durissus</emphasis>
</taxonomicName>
by 
<bibRefCitation id="EFA6F785EA32FFA8CC76591AFCC40505" box="[593,825,1878,1903]" pageId="6" pageNumber="65" refString="Wuster, W., Ferguson, J. E., Quijada-Mascarenas, J. A., Pook, C. E., Salomao, M. D. &amp; Thorpe, R. S. (2005) Tracing an invasion: landbridges, refugia, and the phylogeography of the Neotropical rattlesnake (Serpentes: Viperidae: Crotalus durissus). Molecular Ecology, 14, 1095 - 1108." type="journal article">
Wüster 
<emphasis id="B9435666EA32FFA8CC8C5915FD200505" box="[683,733,1878,1903]" italics="true" pageId="6" pageNumber="65">et al</emphasis>
. (2005)
</bibRefCitation>
. They interpreted such pattern in an open habitat species as the testimony of dispersal during periods of forest opening in Amazonia. Initially, the observed biogeographic patterns in Amazonia led 
<bibRefCitation id="EFA6F785EA32FFA8CC0F59D3FD3B05DD" author="Haffer" box="[552,710,1950,1974]" pageId="6" pageNumber="65" refString="Haffer, J. (1969) Speciation in amazonian forest birds. Science, 165, 131 - 137." type="journal article" year="1969">Haffer (1969)</bibRefCitation>
to suggest isolation of moist forest refugia during periods of increased aridity coincident with temperate glacial maxima. However, palynology, geomorphology (
<bibRefCitation id="EFA6F785EA32FFA9CB2E598FFF0502C4" author="Colinvaux" lastPageId="7" lastPageNumber="66" pageId="6" pageNumber="65" refString="Colinvaux, P. A., De Oliveira, P. E. &amp; Bush, M. B. (2000) Amazonian and Neotropical plant communities on glacial time-scales: The failure of the aridity and refuge hypotheses. Quaternary Science Reviews, 19, 141 - 169." type="journal article" year="2000">
Colinvaux 
<emphasis id="B9435666EA32FFA9CBAF5989FF5002DB" italics="true" lastPageId="7" lastPageNumber="66" pageId="6" pageNumber="65">et al</emphasis>
. 2000
</bibRefCitation>
) and climate modeling (
<bibRefCitation id="EFA6F785EA33FFA9CC325EDAFD1A02C4" author="Mayle" box="[533,743,151,176]" pageId="7" pageNumber="66" refString="Mayle, F. E., Beerling, D. J., Gosling, W. D. &amp; Bush, M. B. (2004) Responses of Amazonian ecosystems to climatic and atmospheric carbon dioxide changes since the last glacial maximum. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences, 359, 499 - 514." type="journal article" year="2004">
Mayle 
<emphasis id="B9435666EA33FFA9CC4F5ED4FD6602DB" box="[616,667,152,176]" italics="true" pageId="7" pageNumber="66">et al</emphasis>
. 2004
</bibRefCitation>
) suggests that Quaternary climatic changes were not strong enough to produce broad fragmentation of Amazonian forest. Though there is little empirical data supporting broad scale turnover of biomes (forest vs. savannas) within Amazonia, there is evidence that peripheral and ecotonal areas, like in the GS, have suffered reduced precipitation and forest fragmentation (
<bibRefCitation id="EFA6F785EA33FFA9CA6C5F49FAE70377" author="Mayle" box="[1099,1306,260,284]" pageId="7" pageNumber="66" refString="Mayle, F. E., Beerling, D. J., Gosling, W. D. &amp; Bush, M. B. (2004) Responses of Amazonian ecosystems to climatic and atmospheric carbon dioxide changes since the last glacial maximum. Philosophical Transactions of the Royal Society of London Series B-Biological Sciences, 359, 499 - 514." type="journal article" year="2004">
Mayle 
<emphasis id="B9435666EA33FFA9CABB5F48FB330377" box="[1180,1230,260,284]" italics="true" pageId="7" pageNumber="66">et al</emphasis>
. 2004
</bibRefCitation>
). In 
<collectingCountry id="F320CAE4EA33FFA9CB695F49FF160354" name="French Guiana" pageId="7" pageNumber="66">French Guiana</collectingCountry>
, palynological evidence suggests two phases of forest retreat at least during the Holocene (
<bibRefCitation id="EFA6F785EA33FFA9CB355F6AFF2C030F" author="Ledru" pageId="7" pageNumber="66" refString="Ledru, M. - P., Blanc, P., Charles-Dominique, P., Fournier, M., Martin, L., Riera, B. &amp; Tardy, C. (2000) Reconstitution de l'ecosysteme forestier guyanais au cours de l'Holocene superieur: apport de la palynologie. In Servant, M. &amp; Servant Vildary, S. (Eds), Dynamique a long terme des ecosystemes forestiers intertropicaux, UNESCO, pp. 199 - 204." type="book chapter" year="2000">
Ledru 
<emphasis id="B9435666EA33FFA9CB465F64FA68032B" box="[1377,1429,296,320]" italics="true" pageId="7" pageNumber="66">et al</emphasis>
. 2000
</bibRefCitation>
) and charcoal deposits indicate widespread burning during this time (
<bibRefCitation id="EFA6F785EA33FFA9CDC75F01FABF030F" author="Charles-Dominique" box="[992,1346,332,356]" pageId="7" pageNumber="66" refString="Charles-Dominique, P., Blanc, P., Larpin, D., Ledru, M. - P., Riera, B., Sarthou, C., Servant, M. &amp; Tardy, C. (1998) Forest perturbations and biodiversity during the last ten thousand years in French Guiana. Acta Oecologica-International Journal of Ecology, 19, 295 - 302." type="journal article" year="1998">
Charles-Dominique 
<emphasis id="B9435666EA33FFA9CAE15F00FB05030F" box="[1222,1272,332,356]" italics="true" pageId="7" pageNumber="66">et al</emphasis>
. 1998
</bibRefCitation>
). Moreover, patches of savanna and different 
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of forests peppered through the eastern GS are the testimony of these recent changes in vegetation distribution (
<bibRefCitation id="EFA6F785EA33FFA9CC525FD9FCEF03C7" author="Servant" box="[629,786,404,428]" pageId="7" pageNumber="66" refString="Servant, M. (2000) Diversite actuelle de la foret tropicale et changements passes du climat: Le programme ecosystemes et paleoecosystemes des forets intertropicales (ECOFIT). Bilan et perspectives. In Servant, M. &amp; Servant Vildary, S. (Eds), Dynamique a long terme des ecosystemes forestiers intertropicaux, UNESCO, pp. 13 - 18." type="book chapter" year="2000">Servant 2000</bibRefCitation>
). Further investigation focusing on comparing phylogeographic patterns across open habitat adapted species vs. forest adapted species should provide important insights into how these changes have impacted biodiversity in Amazonia.
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