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<mods:title id="495A2B09476ACBDD41337DAE3072027F">Molecular systematics of terraranas (Anura: Brachycephaloidea) with an assessment of the effects of alignment and optimality criteria</mods:title>
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<mods:namePart id="A9AD73CB56225456D7BB27298C2E6BA8">Grant, Taran</mods:namePart>
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<mods:namePart id="0EF3A9F3C33F32643769E27818B19A1B">Frost, Darrel R.</mods:namePart>
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<subSubSection id="EFC616C4FF9AFFFACA9EFA6B9C6EBBF1" box="[151,376,1448,1474]" pageId="49" pageNumber="50" type="nomenclature">
<paragraph id="A763454FFF9AFFFACA9EFA6B9C6EBBF1" blockId="49.[151,376,1448,1474]" box="[151,376,1448,1474]" pageId="49" pageNumber="50">
<heading id="FC2BF223FF9AFFFACA9EFA6B9C6EBBF1" bold="true" box="[151,376,1448,1474]" fontSize="11" level="2" pageId="49" pageNumber="50" reason="3">
<taxonomicName id="60DC3ECCFF9AFFFACA9EFA6B9C6EBBF1" ID-CoL="7CL" authority="Fitzinger" box="[151,376,1448,1474]" class="Amphibia" family="Brachycephalidae" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="family">
<emphasis id="95A8995DFF9AFFFACA9EFA6B9C6EBBF1" bold="true" box="[151,376,1448,1474]" pageId="49" pageNumber="50">Brachycephalidae</emphasis>
</taxonomicName>
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<paragraph id="A763454FFF9AFFFACA9EFA2D9DEBB978" blockId="49.[151,1436,1518,2011]" pageId="49" pageNumber="50">
This taxon includes the sister taxa 
<taxonomicName id="60DC3ECCFF9AFFFAC813FA2D9FD9B834" authorityName="Fitzinger" authorityYear="1826" box="[538,719,1518,1543]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFAC813FA2D9FD9B834" box="[538,719,1518,1543]" italics="true" pageId="49" pageNumber="50">Brachycephalus</emphasis>
</taxonomicName>
and 
<taxonomicName id="60DC3ECCFF9AFFFAC90DFA2D9E8EB834" box="[772,920,1518,1543]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFAC90DFA2D9E8EB834" box="[772,920,1518,1543]" italics="true" pageId="49" pageNumber="50">Ischnocnema</emphasis>
</taxonomicName>
, a relationship first discovered by 
<bibRefCitation id="C34D38BEFF9AFFFACF15FA2D9C1EB818" author="Heinicke, M. P. &amp; Duellman, W. E. &amp; Hedges, S. B." pageId="49" pageNumber="50" pagination="10092 - 10097" refId="ref48952" refString="Heinicke, M. P., Duellman, W. E. &amp; Hedges, S. B. (2007) Major Caribbean and Central American frog faunas originated by oceanic dispersal. Proceedings of the National Academy of Sciences, USA, 104, 10092 - 10097. http: // dx. doi. org / 10.1073 / pnas. 0611051104" type="journal article" year="2007">
Heinicke 
<emphasis id="95A8995DFF9AFFFACF81FA2C9DA2B818" italics="true" pageId="49" pageNumber="50">et al.</emphasis>
(2007)
</bibRefCitation>
, who placed most southeastern Brazilian taxa then referred to “ 
<emphasis id="95A8995DFF9AFFFAC9C4F9D199B1B818" box="[973,1191,1554,1579]" italics="true" pageId="49" pageNumber="50">
<taxonomicName id="60DC3ECCFF9AFFFAC9C4F9D199B6B818" authority="(Frost et al., 2006)" baseAuthorityName="Frost" baseAuthorityYear="2006" box="[973,1184,1554,1579]" class="Amphibia" family="Eleutherodactylidae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">Eleutherodactylus</taxonomicName>
”
</emphasis>
(
<bibRefCitation id="C34D38BEFF9AFFFACEBFF9D1986EB818" author="Frost, D. R. &amp; Grant, T. &amp; Faivovich, J. &amp; Bain, R. H. &amp; Haas, A. &amp; Haddad, C. F. B. &amp; De Sa, R. O. &amp; Channing, A. &amp; Wilkinson, M. &amp; Donnellan, S. C. &amp; Raxworthy, C. J. &amp; Campbell, J. A. &amp; Blotto, B. L. &amp; Moler, P. &amp; Drewes, R. C. &amp; Nussbaum, R. A. &amp; Lynch, J. D. &amp; Green, D. M. &amp; Wheeler, W. C." box="[1206,1400,1554,1579]" pageId="49" pageNumber="50" pagination="1 - 370" refId="ref47074" refString="Frost, D. R., Grant, T., Faivovich, J., Bain, R. H., Haas, A., Haddad, C. F. B., De Sa, R. O., Channing, A., Wilkinson, M., Donnellan, S. C., Raxworthy, C. J., Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nussbaum, R. A., Lynch, J. D., Green, D. M. &amp; Wheeler, W. C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1 - 370. http: // dx. doi. org / 10.1206 / 0003 - 0090 (2006) 297 [0001: tatol] 2.0. co; 2" type="journal article" year="2006">
Frost 
<emphasis id="95A8995DFF9AFFFACEF1F9D09826B818" box="[1272,1328,1554,1579]" italics="true" pageId="49" pageNumber="50">et al.</emphasis>
, 2006
</bibRefCitation>
) in a redelimited 
<taxonomicName id="60DC3ECCFF9AFFFACB3BF9F59CD0B87C" box="[306,454,1590,1615]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFACB3BF9F59CD0B87C" box="[306,454,1590,1615]" italics="true" pageId="49" pageNumber="50">Ischnocnema</emphasis>
</taxonomicName>
. Previously, 
<bibRefCitation id="C34D38BEFF9AFFFAC851F9F59EBFB87C" author="Caramaschi, U. &amp; Canedo, C." box="[600,937,1590,1615]" pageId="49" pageNumber="50" pagination="43 - 54" refId="ref43997" refString="Caramaschi, U. &amp; Canedo, C. (2006) Reassessment of the taxonomic status of the genera Ischnocnema Reinhardt and Lutken, 1862, and Oreobates Jimenez-de-la-Espada, 1872, with notes on the synonymy of Leiuperus verrucosus Reinhardt and Lutken, 1862 (Anura: Leptodactylidae). Zootaxa, 1116, 43 - 54." type="journal article" year="2006">Caramaschi &amp; Canedo (2006)</bibRefCitation>
had restricted 
<taxonomicName id="60DC3ECCFF9AFFFACE59F9F599F2B87C" box="[1104,1252,1590,1615]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFACE59F9F599F2B87C" box="[1104,1252,1590,1615]" italics="true" pageId="49" pageNumber="50">Ischnocnema</emphasis>
</taxonomicName>
to 
<taxonomicName id="60DC3ECCFF9AFFFACF01F9F59C11B841" baseAuthorityName="Reinhardt &amp; Lutken" baseAuthorityYear="1862" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="species" species="verrucosa">
<emphasis id="95A8995DFF9AFFFACF01F9F59C11B841" italics="true" pageId="49" pageNumber="50">Ischnocnema verrucosa</emphasis>
</taxonomicName>
, and placed it in the synonymy of 
<taxonomicName id="60DC3ECCFF9AFFFAC8DBF9999E8BB840" box="[722,925,1626,1651]" class="Amphibia" family="Eleutherodactylidae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFAC8DBF9999E8BB840" box="[722,925,1626,1651]" italics="true" pageId="49" pageNumber="50">Eleutherodactylus</emphasis>
</taxonomicName>
. A new test of the sister relationship of 
<taxonomicName id="60DC3ECCFF9AFFFACA9EF9BD9C3DB8A4" box="[151,299,1662,1687]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFACA9EF9BD9C3DB8A4" box="[151,299,1662,1687]" italics="true" pageId="49" pageNumber="50">Ischnocnema</emphasis>
</taxonomicName>
and 
<taxonomicName id="60DC3ECCFF9AFFFACB6DF9BD9F0EB8A4" authorityName="Fitzinger" authorityYear="1826" box="[356,536,1662,1687]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFACB6DF9BD9F0EB8A4" box="[356,536,1662,1687]" italics="true" pageId="49" pageNumber="50">Brachycephalus</emphasis>
</taxonomicName>
was recently provided by 
<bibRefCitation id="C34D38BEFF9AFFFAC941F9BD9964B8A4" author="Canedo, C. &amp; Haddad, C. F. B." box="[840,1138,1662,1687]" pageId="49" pageNumber="50" pagination="610 - 620" refId="ref43929" refString="Canedo, C. &amp; Haddad, C. F. B. (2012) Phylogenetic relationships within anuran clade Terrarana, with emphasis on the placement of Brazilian Atlantic rainforest frogs genus Ischnocnema (Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 65, 610 - 620. http: // dx. doi. org / 10.1016 / j. ympev. 2012.07.016" type="journal article" year="2012">Canedo &amp; Haddad (2012)</bibRefCitation>
, who found the taxa to be reciprocally monophyletic in both maximum likelihood and Bayesian analyses. Interestingly, their parsimony analysis of a similarity alignment placed 
<taxonomicName id="60DC3ECCFF9AFFFAC860F9059E08B8EC" authorityName="Fitzinger" authorityYear="1826" box="[617,798,1734,1759]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFAC860F9059E08B8EC" box="[617,798,1734,1759]" italics="true" pageId="49" pageNumber="50">Brachycephalus</emphasis>
</taxonomicName>
as the sister of 
<taxonomicName id="60DC3ECCFF9AFFFAC9D8F90599A2B8EC" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[977,1204,1734,1759]" class="Amphibia" family="Craugastoridae" genus="Haddadus" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="species" species="binotatus">
<emphasis id="95A8995DFF9AFFFAC9D8F90599A2B8EC" box="[977,1204,1734,1759]" italics="true" pageId="49" pageNumber="50">Haddadus binotatus</emphasis>
</taxonomicName>
, another taxon from southeastern 
<collectingCountry id="DFCB05DFFF9AFFFACB26F9299C61B930" box="[303,375,1770,1795]" name="Brazil" pageId="49" pageNumber="50">Brazil</collectingCountry>
. Such a relationship had never been proposed, and although it is not corroborated in our analyses, it surely deserves future attention, particularly with regard to the effects of alignment and optimality criterion.
</paragraph>
<paragraph id="A763454FFF9AFFF9CACCF8959FCEBCB6" blockId="49.[151,1436,1518,2011]" lastBlockId="50.[150,1438,151,1187]" lastPageId="50" lastPageNumber="51" pageId="49" pageNumber="50">
<taxonomicName id="60DC3ECCFF9AFFFACACCF8959C6FB95C" authorityName="Fitzinger" authorityYear="1826" box="[197,377,1878,1903]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFACACCF8959C6FB95C" box="[197,377,1878,1903]" italics="true" pageId="49" pageNumber="50">Brachycephalus</emphasis>
</taxonomicName>
.—This genus of small, diurnal frogs that usually are brightly colored and contain tetrodotoxin (
<bibRefCitation id="C34D38BEFF9AFFFACA97F8B99C72B9A0" author="Pires, Jr. O. R. &amp; Sebben, A. &amp; Schwartz, E. N. F. &amp; Morales, R. A. V. &amp; Bloch Jr, C. &amp; Schwartz, C. A." box="[158,356,1914,1939]" pageId="49" pageNumber="50" pagination="73 - 79" refId="ref54713" refString="Pires, Jr. O. R., Sebben, A., Schwartz, E. N. F., Morales, R. A. V., Bloch Jr, C. &amp; Schwartz, C. A. 2005. Further report of the occurrence of tetrodotoxin and new analogues in the anuran family Brachycephalidae. Toxicon, 45, 73 - 79. http: // dx. doi. org / 10.1016 / j. toxicon. 2004.09.016" type="journal article" year="2005">
Pires 
<emphasis id="95A8995DFF9AFFFACAEAF8B89C09B9A0" box="[227,287,1914,1939]" italics="true" pageId="49" pageNumber="50">et al.</emphasis>
2005
</bibRefCitation>
) is endemic to southeastern 
<collectingCountry id="DFCB05DFFF9AFFFAC8BCF8B99FEBB9A0" box="[693,765,1914,1939]" name="Brazil" pageId="49" pageNumber="50">Brazil</collectingCountry>
. Its unique pectoral girdle has long been recognized as a synapomorphy of the genus (
<bibRefCitation id="C34D38BEFF9AFFFACBF6F85D9FD2B984" author="Izecksohn, E." box="[511,708,1950,1975]" pageId="49" pageNumber="50" pagination="1 - 12" refId="ref50002" refString="Izecksohn, E. (1971) Novo genero e nova especie de Brachycephalidae do Estado do Rio do Janeiro, Brazil. (Amphibia, Anura). Boletim do Museu Nacional, Nova Serie, Zoologia, 280, 1 - 12." type="journal article" year="1971">Izecksohn, 1971</bibRefCitation>
; 
<bibRefCitation id="C34D38BEFF9AFFFAC8D0F85D9937B984" author="Pombal, J. P. Jr. &amp; Gasparini, J. L." box="[729,1057,1950,1975]" pageId="49" pageNumber="50" pagination="87 - 93" refId="ref54914" refString="Pombal, J. P. Jr. &amp; Gasparini, J. L. (2006) A new Brachycephalus (Anura, Brachycephalidae) from the Atlantic Rainforest of Espirito Santo, southeastern Brazil. South American Journal of Herpetology 1, 87 - 93. http: // dx. doi. org / 10.2994 / 1808 - 9798 (2006) 1 [87: anbabf] 2.0. co; 2" type="journal article" year="2006">Pombal &amp; Gasparini, 2006</bibRefCitation>
) and was used to synonymize 
<taxonomicName id="60DC3ECCFF9AFFFACA9EF8019C3DB9E8" authorityName="Izecksohn" authorityYear="1971" box="[151,299,1986,2011]" class="Amphibia" family="Brachycephalidae" genus="Psyllophryne" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFACA9EF8019C3DB9E8" box="[151,299,1986,2011]" italics="true" pageId="49" pageNumber="50">Psyllophryne</emphasis>
</taxonomicName>
(
<emphasis id="95A8995DFF9AFFFACB4AF8009C70B9E9" box="[323,358,1987,2010]" italics="true" pageId="49" pageNumber="50">i.e.</emphasis>
, 
<taxonomicName id="60DC3ECCFF9AFFFACB75F8009F11B9E8" authorityName="Izecksohn" authorityYear="1971" box="[380,519,1986,2011]" class="Amphibia" family="Brachycephalidae" genus="Psyllophryne" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="species" species="didactyla">
<emphasis id="95A8995DFF9AFFFACB75F8009F11B9E8" box="[380,519,1986,2011]" italics="true" pageId="49" pageNumber="50">P. didactyla</emphasis>
</taxonomicName>
and 
<taxonomicName id="60DC3ECCFF9AFFFAC846F8009E14B9E8" box="[591,770,1986,2011]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="95A8995DFF9AFFFAC846F8009E14B9E8" box="[591,770,1986,2011]" italics="true" pageId="49" pageNumber="50">P. hermogenesi</emphasis>
</taxonomicName>
) with 
<taxonomicName id="60DC3ECCFF9AFFFAC952F8019906B9E8" authorityName="Fitzinger" authorityYear="1826" box="[859,1040,1986,2011]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="49" pageNumber="50" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF9AFFFAC952F8019906B9E8" box="[859,1040,1986,2011]" italics="true" pageId="49" pageNumber="50">Brachycephalus</emphasis>
</taxonomicName>
by 
<bibRefCitation id="C34D38BEFF9AFFFACE43F80199EEB9E8" author="Kaplan, M." box="[1098,1272,1986,2011]" pageId="49" pageNumber="50" pagination="225 - 227" refId="ref50138" refString="Kaplan, M. (2002) Histology of the anteroventral part of the breast-shoulder apparatus of Brachycephalus ephippium (Brachycephalidae) with comments on the validity of the genus Psyllophryne (Brachycephalidae). Amphibia-Reptilia, 23, 225 - 227." type="journal article" year="2002">Kaplan (2002)</bibRefCitation>
, a synonymy subsequently validated by analyses of DNA sequences that found the two species to be nested among other species of 
<taxonomicName id="60DC3ECCFF99FFF9CAB1FF7F9C7BBEE6" authorityName="Fitzinger" authorityYear="1826" box="[184,365,188,213]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9CAB1FF7F9C7BBEE6" box="[184,365,188,213]" italics="true" pageId="50" pageNumber="51">Brachycephalus</emphasis>
</taxonomicName>
(
<bibRefCitation id="C34D38BEFF99FFF9CB74FF7F9FFCBEE6" author="Clemente-Carvalho, R. B. G. &amp; Klaczko, J. &amp; Perez, I. S. &amp; Alves, A. C. R. &amp; Haddad, C. F. B. &amp; Reis, S. F." box="[381,746,188,213]" pageId="50" pageNumber="51" pagination="79 - 89" refId="ref44269" refString="Clemente-Carvalho, R. B. G., Klaczko, J., Perez, I. S., Alves, A. C. R., Haddad, C. F. B. &amp; Reis, S. F. (2011) Molecular phylogenetic relationships and phenotypic diversity in miniaturized toadlets, genus Brachycephalus (Amphibia: Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 61, 79 - 89. http: // dx. doi. org / 10.1016 / j. ympev. 2011.05.017" type="journal article" year="2011">
Clemente-Carvalho 
<emphasis id="95A8995DFF99FFF9C86DFF7E9F89BEE6" box="[612,671,188,213]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
, 2011
</bibRefCitation>
; current results). 
<bibRefCitation id="C34D38BEFF99FFF9C9BDFF7F999FBEE6" author="Frost, D. R. &amp; Grant, T. &amp; Faivovich, J. &amp; Bain, R. H. &amp; Haas, A. &amp; Haddad, C. F. B. &amp; De Sa, R. O. &amp; Channing, A. &amp; Wilkinson, M. &amp; Donnellan, S. C. &amp; Raxworthy, C. J. &amp; Campbell, J. A. &amp; Blotto, B. L. &amp; Moler, P. &amp; Drewes, R. C. &amp; Nussbaum, R. A. &amp; Lynch, J. D. &amp; Green, D. M. &amp; Wheeler, W. C." box="[948,1161,188,213]" pageId="50" pageNumber="51" pagination="1 - 370" refId="ref47074" refString="Frost, D. R., Grant, T., Faivovich, J., Bain, R. H., Haas, A., Haddad, C. F. B., De Sa, R. O., Channing, A., Wilkinson, M., Donnellan, S. C., Raxworthy, C. J., Campbell, J. A., Blotto, B. L., Moler, P., Drewes, R. C., Nussbaum, R. A., Lynch, J. D., Green, D. M. &amp; Wheeler, W. C. (2006) The amphibian tree of life. Bulletin of the American Museum of Natural History, 297, 1 - 370. http: // dx. doi. org / 10.1206 / 0003 - 0090 (2006) 297 [0001: tatol] 2.0. co; 2" type="journal article" year="2006">
Frost 
<emphasis id="95A8995DFF99FFF9C9FEFF7E9925BEE6" box="[1015,1075,188,213]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
(2006)
</bibRefCitation>
, 
<bibRefCitation id="C34D38BEFF99FFF9CE9EFF7F9881BEE6" author="Heinicke, M. P. &amp; Duellman, W. E. &amp; Hedges, S. B." box="[1175,1431,188,213]" pageId="50" pageNumber="51" pagination="10092 - 10097" refId="ref48952" refString="Heinicke, M. P., Duellman, W. E. &amp; Hedges, S. B. (2007) Major Caribbean and Central American frog faunas originated by oceanic dispersal. Proceedings of the National Academy of Sciences, USA, 104, 10092 - 10097. http: // dx. doi. org / 10.1073 / pnas. 0611051104" type="journal article" year="2007">
Heinicke 
<emphasis id="95A8995DFF99FFF9CF0FFF7E9857BEE6" box="[1286,1345,188,213]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
(2007)
</bibRefCitation>
, 
<bibRefCitation id="C34D38BEFF99FFF9CA9EFF1C9C84BECB" author="Hedges, S. B. &amp; Duellman, W. E. &amp; Heinicke, M. P." box="[151,402,223,248]" pageId="50" pageNumber="51" pagination="1 - 182" refId="ref48854" refString="Hedges, S. B., Duellman, W. E. &amp; Heinicke, M. P. (2008 a) New World direct-developing frogs (Anura, Terrarana), molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182." type="journal article" year="2008">
Hedges 
<emphasis id="95A8995DFF99FFF9CAFDFF229C39BECB" box="[244,303,223,248]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
(2008a)
</bibRefCitation>
, and 
<bibRefCitation id="C34D38BEFF99FFF9CBDAFF239FCEBECB" author="Pyron, R. A. &amp; Wiens, J. J." box="[467,728,223,248]" pageId="50" pageNumber="51" pagination="543 - 583" refId="ref55319" refString="Pyron, R. A. &amp; Wiens, J. J. (2011) A large-scale phylogeny of Amphibia including over 2,800 species, and a revised classification of extant frogs, salamanders, and caecilians. Molecular Phylogenetics and Evolution, 61, 543 - 583. http: // dx. doi. org / 10.1016 / j. ympev. 2011.06.012" type="journal article" year="2011">Pyron &amp; Wiens (2011)</bibRefCitation>
only included 
<taxonomicName id="60DC3ECCFF99FFF9C98EFF229909BECB" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[903,1055,223,248]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="95A8995DFF99FFF9C98EFF229909BECB" box="[903,1055,223,248]" italics="true" pageId="50" pageNumber="51">B. ephippium</emphasis>
</taxonomicName>
in their analyses, and 
<bibRefCitation id="C34D38BEFF99FFF9CF2EFF1C9C55BF2E" author="Canedo, C. &amp; Haddad, C. F. B." pageId="50" pageNumber="51" pagination="610 - 620" refId="ref43929" refString="Canedo, C. &amp; Haddad, C. F. B. (2012) Phylogenetic relationships within anuran clade Terrarana, with emphasis on the placement of Brazilian Atlantic rainforest frogs genus Ischnocnema (Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 65, 610 - 620. http: // dx. doi. org / 10.1016 / j. ympev. 2012.07.016" type="journal article" year="2012">Canedo &amp; Haddad (2012)</bibRefCitation>
added 
<taxonomicName id="60DC3ECCFF99FFF9CB93FEC69F40BF2E" baseAuthorityName="Izecksohn" baseAuthorityYear="1971" box="[410,598,260,285]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="didactylus">
<emphasis id="95A8995DFF99FFF9CB93FEC69CA4BF2F" box="[410,434,261,284]" italics="true" pageId="50" pageNumber="51">B.</emphasis>
cf. 
<emphasis id="95A8995DFF99FFF9CBEAFEC79F40BF2E" box="[483,598,260,285]" italics="true" pageId="50" pageNumber="51">didactylus</emphasis>
</taxonomicName>
. 
<bibRefCitation id="C34D38BEFF99FFF9C861FEC79ECABF2E" author="Clemente-Carvalho, R. B. G. &amp; Klaczko, J. &amp; Perez, I. S. &amp; Alves, A. C. R. &amp; Haddad, C. F. B. &amp; Reis, S. F." box="[616,988,260,285]" pageId="50" pageNumber="51" pagination="79 - 89" refId="ref44269" refString="Clemente-Carvalho, R. B. G., Klaczko, J., Perez, I. S., Alves, A. C. R., Haddad, C. F. B. &amp; Reis, S. F. (2011) Molecular phylogenetic relationships and phenotypic diversity in miniaturized toadlets, genus Brachycephalus (Amphibia: Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 61, 79 - 89. http: // dx. doi. org / 10.1016 / j. ympev. 2011.05.017" type="journal article" year="2011">
Clemente-Carvalho 
<emphasis id="95A8995DFF99FFF9C944FEC69E9FBF2E" box="[845,905,260,285]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
(2011)
</bibRefCitation>
provided the first extensive molecular phylogenetic analysis of the genus by including 14 of the 18 currently recognized species (
<bibRefCitation id="C34D38BEFF99FFF9CEACFEE49DC5BF56" author="Pombal, J. P. Jr. &amp; Izecksohn, E." pageId="50" pageNumber="51" pagination="443 - 451" refId="ref54990" refString="Pombal, J. P. Jr. &amp; Izecksohn, E. (2011) Uma nova especie de Brachycephalus (Anura, Brachycephalidae). Papeis Avulsos de Zoologia, Sao Paulo, 51, 443 - 451. http: // dx. doi. org / 10.1590 / s 0031 - 10492007001300001" type="journal article" year="2011">Pombal &amp; Izecksohn, 2011</bibRefCitation>
). Inadequate outgroup sampling (the tree was rooted on a single terminal of 
<taxonomicName id="60DC3ECCFF99FFF9CE50FE8F99FBBF56" box="[1113,1261,332,357]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9CE50FE8F99FBBF56" box="[1113,1261,332,357]" italics="true" pageId="50" pageNumber="51">Ischnocnema</emphasis>
</taxonomicName>
) prevented the monophyly of 
<taxonomicName id="60DC3ECCFF99FFF9CB37FEAC9CE4BFBB" authorityName="Fitzinger" authorityYear="1826" box="[318,498,367,392]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9CB37FEAC9CE4BFBB" box="[318,498,367,392]" italics="true" pageId="50" pageNumber="51">Brachycephalus</emphasis>
</taxonomicName>
, the position of 
<taxonomicName id="60DC3ECCFF99FFF9C8A7FEAC9E54BFBB" authorityName="Izecksohn" authorityYear="1971" box="[686,834,367,392]" class="Amphibia" family="Brachycephalidae" genus="Psyllophryne" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9C8A7FEAC9E54BFBB" box="[686,834,367,392]" italics="true" pageId="50" pageNumber="51">Psyllophryne</emphasis>
</taxonomicName>
within Brachycephaloidea, or the sister relationships with 
<taxonomicName id="60DC3ECCFF99FFF9CAD9FE579C72BF9E" box="[208,356,404,429]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9CAD9FE579C72BF9E" box="[208,356,404,429]" italics="true" pageId="50" pageNumber="51">Ischnocnema</emphasis>
</taxonomicName>
from being tested in that study. Nevertheless, our results agree with Clemente-Carvalho 
<emphasis id="95A8995DFF99FFF9CF59FE56989CBF9E" box="[1360,1418,404,429]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
’s (2011) Bayesian analysis of the concatenated nuclear and mitochondrial data in recovering two major sister clades, a southern clade formed by locally endemic species restricted to the high and humid Atlantic Forest of 
<collectingRegion id="65188BADFF99FFF9CF2EFE1F9865BFC6" box="[1319,1395,476,501]" country="Brazil" name="Parana" pageId="50" pageNumber="51">Paraná</collectingRegion>
(
<taxonomicName id="60DC3ECCFF99FFF9CF8DFE1E9DE9BC2B" authorityName="Ribeiro, Alves, Haddad &amp; Reis" authorityYear="2005" class="Amphibia" family="Strabomantidae" genus="Hypodactylus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="brunneus">
<emphasis id="95A8995DFF99FFF9CF8DFE1E9DE9BC2B" italics="true" pageId="50" pageNumber="51">B. brunneus</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9CB06FDC29CB0BC2B" authorityName="Alves, Ribeiro, Haddad &amp; Reis" authorityYear="2006" box="[271,422,511,536]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="ferruginus">
<emphasis id="95A8995DFF99FFF9CB06FDC29CB0BC2B" box="[271,422,511,536]" italics="true" pageId="50" pageNumber="51">B. ferruginus</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9CBBFFDC29F5ABC2B" authorityName="Ribeiro, Alves, Haddad &amp; Reis" authorityYear="2005" box="[438,588,511,536]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="izecksohni">
<emphasis id="95A8995DFF99FFF9CBBFFDC29F5ABC2B" box="[438,588,511,536]" italics="true" pageId="50" pageNumber="51">B. izecksohni</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9C854FDC29FD3BC24" authorityName="Pombal, Wistuba &amp; Bornschein" authorityYear="1998" box="[605,709,512,536]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="pernix">
<emphasis id="95A8995DFF99FFF9C854FDC29FD3BC24" box="[605,709,512,536]" italics="true" pageId="50" pageNumber="51">B. pernix</emphasis>
</taxonomicName>
, and 
<taxonomicName id="60DC3ECCFF99FFF9C901FDC29E93BC2B" authorityName="Alves, Ribeiro, Haddad &amp; Reis" authorityYear="2006" box="[776,901,511,536]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="pombali">
<emphasis id="95A8995DFF99FFF9C901FDC29E93BC2B" box="[776,901,511,536]" italics="true" pageId="50" pageNumber="51">B. pombali</emphasis>
</taxonomicName>
) and the northern clade, allopatric to the first, distributed from farther north in 
<collectingRegion id="65188BADFF99FFF9C80CFDE79F47BC0E" box="[517,593,548,573]" country="Brazil" name="Parana" pageId="50" pageNumber="51">Paraná</collectingRegion>
to 
<collectingRegion id="65188BADFF99FFF9C87CFDE79E03BC0E" box="[629,789,548,573]" country="Brazil" name="Espirito Santo" pageId="50" pageNumber="51">Espírito Santo</collectingRegion>
(
<taxonomicName id="60DC3ECCFF99FFF9C92CFDE69E98BC0E" authorityName="Pombal &amp; Gasparini" authorityYear="2006" box="[805,910,548,573]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="alipioi">
<emphasis id="95A8995DFF99FFF9C92CFDE69E98BC0E" box="[805,910,548,573]" italics="true" pageId="50" pageNumber="51">B. alipioi</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9C995FDE69938BC0E" baseAuthorityName="Izecksohn" baseAuthorityYear="1971" box="[924,1070,548,573]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="didactylus">
<emphasis id="95A8995DFF99FFF9C995FDE69938BC0E" box="[924,1070,548,573]" italics="true" pageId="50" pageNumber="51">B. didactylus</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9CE35FDE699C7BC0E" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[1084,1233,548,573]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="95A8995DFF99FFF9CE35FDE699C7BC0E" box="[1084,1233,548,573]" italics="true" pageId="50" pageNumber="51">B. ephippium</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9CEE9FDE69863BC0E" authorityName="Miranda-Ribeiro" authorityYear="1920" box="[1248,1397,548,573]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="garbeanus">
<emphasis id="95A8995DFF99FFF9CEE9FDE69863BC0E" box="[1248,1397,548,573]" italics="true" pageId="50" pageNumber="51">B. garbeanus</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9CF8DFDE69C30BC53" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="95A8995DFF99FFF9CF8DFDE69C30BC53" italics="true" pageId="50" pageNumber="51">B. hermogenesi</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9CB34FD8A9CC0BC53" authorityName="Miranda-Ribeiro" authorityYear="1920" box="[317,470,583,608]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="nodoterga">
<emphasis id="95A8995DFF99FFF9CB34FD8A9CC0BC53" box="[317,470,583,608]" italics="true" pageId="50" pageNumber="51">B. nodoterga</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9CBE4FD8A9F7CBC6C" authorityName="Alves, Sawaya, Reis &amp; Haddad" authorityYear="2009" box="[493,618,584,608]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="pitanga">
<emphasis id="95A8995DFF99FFF9CBE4FD8A9F7CBC6C" box="[493,618,584,608]" italics="true" pageId="50" pageNumber="51">B. pitanga</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9C88BFD8A9FCEBC53" authorityName="Haddad, Alves, Clemente-Carvalho &amp; Reis" authorityYear="2010" box="[642,728,583,608]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="toby">
<emphasis id="95A8995DFF99FFF9C88BFD8A9FCEBC53" box="[642,728,583,608]" italics="true" pageId="50" pageNumber="51">B. toby</emphasis>
</taxonomicName>
, and 
<taxonomicName id="60DC3ECCFF99FFF9C92EFD8A9EDEBC53" authorityName="Pombal" authorityYear="2001" box="[807,968,583,608]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="vertebralis">
<emphasis id="95A8995DFF99FFF9C92EFD8A9EDEBC53" box="[807,968,583,608]" italics="true" pageId="50" pageNumber="51">B. vertebralis</emphasis>
</taxonomicName>
), with the two species of the former 
<taxonomicName id="60DC3ECCFF99FFF9CA9EFDAF9C3DBCB6" authorityName="Izecksohn" authorityYear="1971" box="[151,299,620,645]" class="Amphibia" family="Brachycephalidae" genus="Psyllophryne" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9CA9EFDAF9C3DBCB6" box="[151,299,620,645]" italics="true" pageId="50" pageNumber="51">Psyllophryne</emphasis>
</taxonomicName>
placed separately in the second clade.
</paragraph>
<paragraph id="A763454FFF99FFF9CACCFD4C9EDDBA90" blockId="50.[150,1438,151,1187]" pageId="50" pageNumber="51">
<emphasis id="95A8995DFF99FFF9CACCFD4C9C6EBC9B" box="[197,376,655,680]" italics="true" pageId="50" pageNumber="51">
<taxonomicName id="60DC3ECCFF99FFF9CACCFD4C9C4EBC9B" box="[197,344,655,680]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">Ischnocnema</taxonomicName>
.—
</emphasis>
This genus is distributed within the Atlantic Forest of south, southeastern, and northeastern 
<collectingCountry id="DFCB05DFFF99FFF9CA9EFD779DCDBCFE" box="[151,219,692,717]" name="Brazil" pageId="50" pageNumber="51">Brazil</collectingCountry>
and 
<collectingRegion id="65188BADFF99FFF9CB16FD779C9FBCFE" box="[287,393,692,717]" country="Argentina" name="Misiones" pageId="50" pageNumber="51">Misiones</collectingRegion>
, 
<collectingCountry id="DFCB05DFFF99FFF9CB92FD779F1CBCFE" box="[411,522,692,717]" name="Argentina" pageId="50" pageNumber="51">Argentina</collectingCountry>
(
<bibRefCitation id="C34D38BEFF99FFF9C816FD779E59BCFE" author="Canedo, C. &amp; Haddad, C. F. B." box="[543,847,692,717]" pageId="50" pageNumber="51" pagination="610 - 620" refId="ref43929" refString="Canedo, C. &amp; Haddad, C. F. B. (2012) Phylogenetic relationships within anuran clade Terrarana, with emphasis on the placement of Brazilian Atlantic rainforest frogs genus Ischnocnema (Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 65, 610 - 620. http: // dx. doi. org / 10.1016 / j. ympev. 2012.07.016" type="journal article" year="2012">Canedo &amp; Haddad, 2012</bibRefCitation>
; 
<bibRefCitation id="C34D38BEFF99FFF9C968FD779EFBBCFE" author="Frost, D. R." box="[865,1005,692,717]" pageId="50" pageNumber="51" refId="ref47018" refString="Frost, D. R. (2014) Amphibian Species of the World, an Online Reference. Version 6.0 American Museum of Natural History, New York, USA. Available at http: // research. amnh. org / herpetology / amphibia / index. html (accessed February 04 2014)" type="url" year="2014">Frost, 2014</bibRefCitation>
). The monophyly of the genus was hypothesized by 
<bibRefCitation id="C34D38BEFF99FFF9CB55FD149F4BBCC3" author="Heinicke, M. P. &amp; Duellman, W. E. &amp; Hedges, S. B." box="[348,605,727,752]" pageId="50" pageNumber="51" pagination="10092 - 10097" refId="ref48952" refString="Heinicke, M. P., Duellman, W. E. &amp; Hedges, S. B. (2007) Major Caribbean and Central American frog faunas originated by oceanic dispersal. Proceedings of the National Academy of Sciences, USA, 104, 10092 - 10097. http: // dx. doi. org / 10.1073 / pnas. 0611051104" type="journal article" year="2007">
Heinicke 
<emphasis id="95A8995DFF99FFF9CBC5FD1A9F1EBCC3" box="[460,520,727,752]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
(2007)
</bibRefCitation>
and 
<bibRefCitation id="C34D38BEFF99FFF9C892FD149E8CBCC3" author="Hedges, S. B. &amp; Duellman, W. E. &amp; Heinicke, M. P." box="[667,922,727,752]" pageId="50" pageNumber="51" pagination="1 - 182" refId="ref48854" refString="Hedges, S. B., Duellman, W. E. &amp; Heinicke, M. P. (2008 a) New World direct-developing frogs (Anura, Terrarana), molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182." type="journal article" year="2008">
Hedges 
<emphasis id="95A8995DFF99FFF9C8F3FD1A9E20BCC3" box="[762,822,727,752]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
(2008a)
</bibRefCitation>
, who analyzed 5 of the 33 species. A more complete test of generic monophyly was recently provided by 
<bibRefCitation id="C34D38BEFF99FFF9C974FD3F99BBBD26" author="Canedo, C. &amp; Haddad, C. F. B." box="[893,1197,764,789]" pageId="50" pageNumber="51" pagination="610 - 620" refId="ref43929" refString="Canedo, C. &amp; Haddad, C. F. B. (2012) Phylogenetic relationships within anuran clade Terrarana, with emphasis on the placement of Brazilian Atlantic rainforest frogs genus Ischnocnema (Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 65, 610 - 620. http: // dx. doi. org / 10.1016 / j. ympev. 2012.07.016" type="journal article" year="2012">Canedo &amp; Haddad (2012)</bibRefCitation>
in a study that was published while the present manuscript was being written, which unfortunately precluded the use of their data in our analyses. Nevertheless, their taxon sampling of other brachycephalids was large enough (214 species) to provide a strong test of the monophyly of 
<taxonomicName id="60DC3ECCFF99FFF9C860FCA49FEBBDB3" box="[617,765,871,896]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9C860FCA49FEBBDB3" box="[617,765,871,896]" italics="true" pageId="50" pageNumber="51">Ischnocnema</emphasis>
</taxonomicName>
. Their major finding was a polyphyletic 
<taxonomicName id="60DC3ECCFF99FFF9CECDFCA49C9EBD96" authority="sensu Hedges et al. (2008 a)" authorityName="Hedges" authorityYear="2008" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9CECDFCA4984EBDB3" box="[1220,1368,871,896]" italics="true" pageId="50" pageNumber="51">Ischnocnema</emphasis>
sensu 
<bibRefCitation id="C34D38BEFF99FFF9CA9EFC4F9C9EBD96" author="Hedges, S. B. &amp; Duellman, W. E. &amp; Heinicke, M. P." box="[151,392,908,933]" pageId="50" pageNumber="51" pagination="1 - 182" refId="ref48854" refString="Hedges, S. B., Duellman, W. E. &amp; Heinicke, M. P. (2008 a) New World direct-developing frogs (Anura, Terrarana), molecular phylogeny, classification, biogeography, and conservation. Zootaxa, 1737, 1 - 182." type="journal article" year="2008">
Hedges 
<emphasis id="95A8995DFF99FFF9CAFBFC4E9C3CBD96" box="[242,298,908,933]" italics="true" pageId="50" pageNumber="51">et al.</emphasis>
(2008a)
</bibRefCitation>
</taxonomicName>
with three named species (
<taxonomicName id="60DC3ECCFF99FFF9C8B0FC4E9E42BD96" baseAuthorityName="Bokermann" baseAuthorityYear="1975" box="[697,852,908,933]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="paulodutrai">
<emphasis id="95A8995DFF99FFF9C8B0FC4E9E42BD96" box="[697,852,908,933]" italics="true" pageId="50" pageNumber="51">I. paulodutrai</emphasis>
</taxonomicName>
, 
<taxonomicName id="60DC3ECCFF99FFF9C96BFC4E9EC5BD97" baseAuthorityName="Boulenger" baseAuthorityYear="1888" box="[866,979,909,932]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="ramagii">
<emphasis id="95A8995DFF99FFF9C96BFC4E9EC5BD97" box="[866,979,909,932]" italics="true" pageId="50" pageNumber="51">I. ramagii</emphasis>
</taxonomicName>
, and 
<taxonomicName id="60DC3ECCFF99FFF9CE19FC4E997BBD96" baseAuthorityName="Bokermann" baseAuthorityYear="1975" box="[1040,1133,908,933]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="vinhai">
<emphasis id="95A8995DFF99FFF9CE19FC4E997BBD96" box="[1040,1133,908,933]" italics="true" pageId="50" pageNumber="51">I. vinhai</emphasis>
</taxonomicName>
) and two unnamed species recovered as part of the 
<taxonomicName id="60DC3ECCFF99FFF9CBAAFC739FD8BDFB" baseAuthorityName="Gunther" baseAuthorityYear="1858" box="[419,718,943,968]" class="Amphibia" family="Strabomantidae" genus="Pristimantis" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="conspicillatus">
<emphasis id="95A8995DFF99FFF9CBAAFC739FD8BDFB" box="[419,718,943,968]" italics="true" pageId="50" pageNumber="51">Pristimantis conspicillatus</emphasis>
</taxonomicName>
species group. Also, 
<taxonomicName id="60DC3ECCFF99FFF9C9B4FC6C99AABDFB" baseAuthorityName="Bokermann" baseAuthorityYear="1975" box="[957,1212,943,968]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="bilineata">
<emphasis id="95A8995DFF99FFF9C9B4FC6C99AABDFB" box="[957,1212,943,968]" italics="true" pageId="50" pageNumber="51">Ischnocnema bilineata</emphasis>
</taxonomicName>
was found to be the sister of a clade composed of the genera 
<taxonomicName id="60DC3ECCFF99FFF9C880FC179FFCBDDE" box="[649,746,980,1005]" class="Amphibia" family="Strabomantidae" genus="Noblella" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9C880FC179FFCBDDE" box="[649,746,980,1005]" italics="true" pageId="50" pageNumber="51">Noblella</emphasis>
</taxonomicName>
and 
<taxonomicName id="60DC3ECCFF99FFF9C923FC179EBFBDDE" authorityName="Heyer" authorityYear="1969" box="[810,937,980,1005]" class="Amphibia" family="Strabomantidae" genus="Barycholos" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9C923FC179EBFBDDE" box="[810,937,980,1005]" italics="true" pageId="50" pageNumber="51">Barycholos</emphasis>
</taxonomicName>
in 
<taxonomicName id="60DC3ECCFF99FFF9C9D1FC179961BDDE" authorityName="Hedges, Duellman &amp; Heinicke" authorityYear="2008" box="[984,1143,980,1005]" class="Amphibia" family="Craugastoridae" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="subFamily" subFamily="Holoadeninae">Holoadeninae</taxonomicName>
. Accordingly, 
<bibRefCitation id="C34D38BEFF99FFF9CF2DFC179C5EBA23" author="Canedo, C. &amp; Haddad, C. F. B." pageId="50" pageNumber="51" pagination="610 - 620" refId="ref43929" refString="Canedo, C. &amp; Haddad, C. F. B. (2012) Phylogenetic relationships within anuran clade Terrarana, with emphasis on the placement of Brazilian Atlantic rainforest frogs genus Ischnocnema (Anura: Brachycephalidae). Molecular Phylogenetics and Evolution, 65, 610 - 620. http: // dx. doi. org / 10.1016 / j. ympev. 2012.07.016" type="journal article" year="2012">Canedo &amp; Haddad (2012)</bibRefCitation>
placed three nominal species of 
<taxonomicName id="60DC3ECCFF99FFF9C8E3FC349E68BA23" box="[746,894,1015,1040]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9C8E3FC349E68BA23" box="[746,894,1015,1040]" italics="true" pageId="50" pageNumber="51">Ischnocnema</emphasis>
</taxonomicName>
in 
<taxonomicName id="60DC3ECCFF99FFF9C9BAFC3B992ABA3C" box="[947,1084,1016,1039]" class="Amphibia" family="Strabomantidae" genus="Pristimantis" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9C9BAFC3B992ABA3C" box="[947,1084,1016,1039]" italics="true" pageId="50" pageNumber="51">Pristimantis</emphasis>
</taxonomicName>
, but left &quot; 
<emphasis id="95A8995DFF99FFF9CECCFC34988ABA23" box="[1221,1436,1015,1040]" italics="true" pageId="50" pageNumber="51">
<taxonomicName id="60DC3ECCFF99FFF9CECCFC349885BA23" box="[1221,1427,1015,1040]" class="Amphibia" family="Eleutherodactylidae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">Eleutherodactylus</taxonomicName>
&quot;
</emphasis>
<taxonomicName id="60DC3ECCFF99FFF9CA9EFBDD9CCDBA04" authority="Bokermann, 1975" authorityName="Bokermann" authorityYear="1975" box="[151,475,1054,1079]" class="Amphibia" family="Eleutherodactylidae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="species" species="bilineatus">
<emphasis id="95A8995DFF99FFF9CA9EFBDD9C13BA04" box="[151,261,1054,1079]" italics="true" pageId="50" pageNumber="51">bilineatus</emphasis>
Bokermann, 1975
</taxonomicName>
&quot;1974&quot; as 
<taxonomicName id="60DC3ECCFF99FFF9C856FBE39FEBBA04" box="[607,765,1054,1079]" class="Diplopoda" higherTaxonomySource="CoL" kingdom="Animalia" order="Sedis" pageId="50" pageNumber="124" phylum="Arthropoda" rank="order">
<emphasis id="95A8995DFF99FFF9C856FBE39FEBBA04" box="[607,765,1054,1079]" italics="true" pageId="50" pageNumber="51">incertae sedis</emphasis>
</taxonomicName>
within 
<taxonomicName id="60DC3ECCFF99FFF9C951FBDD9EE5BA04" authorityName="Hedges, Duellman &amp; Heinicke" authorityYear="2008" box="[856,1011,1054,1079]" class="Amphibia" family="Craugastoridae" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="subFamily" subFamily="Holoadeninae">Holoadeninae</taxonomicName>
<superScript id="50A9E807FF99FFF9C9FDFBD89EEABA1A" attach="left" box="[1012,1020,1051,1065]" fontSize="6" pageId="50" pageNumber="51">6</superScript>
(of 
<taxonomicName id="60DC3ECCFF99FFF9CE39FBDD99CBBA04" box="[1072,1245,1054,1079]" class="Amphibia" family="Craugastoridae" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="family">Craugastoridae</taxonomicName>
, below). For the remaining species of 
<taxonomicName id="60DC3ECCFF99FFF9CB97FB819F24BA68" box="[414,562,1090,1115]" class="Amphibia" family="Brachycephalidae" genus="Ischnocnema" kingdom="Animalia" order="Anura" pageId="50" pageNumber="51" phylum="Chordata" rank="genus">
<emphasis id="95A8995DFF99FFF9CB97FB819F24BA68" box="[414,562,1090,1115]" italics="true" pageId="50" pageNumber="51">Ischnocnema</emphasis>
</taxonomicName>
, they recognized four monophyletic species series and left two species unassigned to any group, although five species were only tentatively assigned to groups based on their morphology. We follow their assignment of species to species series (see Appendix 2).
</paragraph>
</subSubSection>
</treatment>
</document>