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<document id="AE39B6114D125EAA0DC3BFA117BC6E0D" ID-DOI="10.7717/peerj.18265" IM.bibliography_approvedBy="diego" IM.illustrations_approvedBy="diego" IM.materialsCitations_approvedBy="diego" IM.metadata_approvedBy="diego" IM.tables_requiresApprovalFor="operationResults" IM.taxonomicNames_approvedBy="diego" IM.treatments_approvedBy="diego" checkinTime="1730107934756" checkinUser="diego" docAuthor="Toledo, Luís Felipe, Botelho, Lucas Machado, Carrasco-Medina, Andres Santiago, Gray, Jaimi A., Ernetti, Julia R., Gama, Joana Moura, Lyra, Mariana Lucio, Blackburn, David C., Nunes, Ivan &amp; Muscat, Edelcio" docDate="2024" docId="03C3F71A8B5F673E8086FCF2FDB2FDCF" docLanguage="en" docName="PeerJ.12.e18265.pdf" docOrigin="PeerJ 12" docTitle="Brachycephalus dacnis Toledo, Botelho, Carrasco-Medina, Gray, Ernetti, Gama, Lyra, Blackburn, Nunes &amp; Muscat, 2024, sp. nov." docType="treatment" docVersion="2" lastPageNumber="19" masterDocId="FFFA8F628B59672C8174FFFFFFFCFFC4" masterDocTitle="Among the world s smallest vertebrates: a new miniaturized flea-toad (Brachycephalidae) from the Atlantic rainforest" masterLastPageNumber="e 18265" masterPageNumber="e 18265" pageNumber="7" updateTime="1730108183273" updateUser="diego">
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<mods:title id="D061777ABE022A97B3BF3CDE65766B1E">Among the world s smallest vertebrates: a new miniaturized flea-toad (Brachycephalidae) from the Atlantic rainforest</mods:title>
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<mods:namePart id="0656965EF9A690A3A7D5C97CBF2BED09">Toledo, Luís Felipe</mods:namePart>
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<mods:namePart id="4E0D058BE9FC1FECBE03ED8DBE3FFC2D">Botelho, Lucas Machado</mods:namePart>
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<mods:namePart id="6400521652C8D0824814117C7E22157F">Carrasco-Medina, Andres Santiago</mods:namePart>
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<mods:namePart id="200E3CB4B029905B2F21F000D69CE62F">Gray, Jaimi A.</mods:namePart>
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<mods:namePart id="03842FFDDCAF58621841EE7909855FD7">Ernetti, Julia R.</mods:namePart>
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<mods:namePart id="942BFFADCC773290262F4E5BD57FCDB1">Gama, Joana Moura</mods:namePart>
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<mods:namePart id="045170E7DD49B0F5D7F6B7876DB06AC4">Lyra, Mariana Lucio</mods:namePart>
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<mods:namePart id="A7CF0B793BA855D1849A9BB4D67DF91A">Blackburn, David C.</mods:namePart>
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<mods:namePart id="B618B419DCF76CA4F3C0F2CC2744D9F3">Nunes, Ivan</mods:namePart>
</mods:name>
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<mods:namePart id="D9418FAC8E8BD43D935FD053311326F0">Muscat, Edelcio</mods:namePart>
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<mods:title id="0A09AF8499CB39D1CA5F51C30A674C4F">PeerJ</mods:title>
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<mods:date id="015DAB277535EF40C9D5E3F1A441CEE7">2024</mods:date>
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<mods:number id="38E706A77D444509E84F65F2FFE2A413">2024-10-25</mods:number>
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<mods:number id="B3E7C87EC40507AA3A4FC1490985F97E">12</mods:number>
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<mods:start id="4411280A3AC735173206ED756A06923F">e 18265</mods:start>
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<mods:classification id="3472FD77C9A846D1305B34DACF11D1D1">journal article</mods:classification>
<mods:identifier id="4736E9E757FA65635300CA365BB6680D" type="DOI">10.7717/peerj.18265</mods:identifier>
<mods:identifier id="BA884DD9BA856EC48B51B3FFE7F0A200" type="ZooBank">115413F3-9005-4006-863C-EA4BAD3C58AD</mods:identifier>
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<treatment id="03C3F71A8B5F673E8086FCF2FDB2FDCF" LSID="urn:lsid:plazi:treatment:03C3F71A8B5F673E8086FCF2FDB2FDCF" httpUri="http://treatment.plazi.org/id/03C3F71A8B5F673E8086FCF2FDB2FDCF" lastPageId="18" lastPageNumber="19" pageId="6" pageNumber="7">
<subSubSection id="C37015878B5F672A8086FCF2FC43FCEC" box="[498,959,781,808]" pageId="6" pageNumber="7" type="nomenclature">
<paragraph id="8BD5460C8B5F672A8086FCF2FC43FCEC" blockId="6.[498,1411,732,888]" box="[498,959,781,808]" pageId="6" pageNumber="7">
<heading id="D09DF1608B5F672A8086FCF2FC43FCEC" box="[498,959,781,808]" level="2" pageId="6" pageNumber="7">
<taxonomicName id="4C6A3D8F8B5F672A8086FCF2FD0AFCE3" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[498,758,781,807]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="dacnis" status="sp. nov.">
<emphasis id="B91E9A1E8B5F672A8086FCF2FD0AFCE3" box="[498,758,781,807]" italics="true" pageId="6" pageNumber="7">Brachycephalusdacnis</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A22D27658B5F672A838BFCF1FCA7FCEC" box="[767,859,782,808]" pageId="6" pageNumber="7" rank="species">sp. nov.</taxonomicNameLabel>
(
<figureCitation id="13515A898B5F672A8219FCF2FC52FCE3" box="[877,942,781,807]" captionStart="Figure 1" captionStartId="7.[524,589,673,695]" captionTargetBox="[498,1541,236,660]" captionTargetId="figure-144@7.[498,1299,234,645]" captionTargetPageId="7" captionText="Figure 1 Brachycephalus dacnis holotype (ZUEC-AMP 24982) adult male, SVL = 7.55 mm. (A) Dorsal and (B) ventral views. Full-size DOI: 10.7717/peerj.18265/fig-1" pageId="6" pageNumber="7">Fig. 1</figureCitation>
).
</heading>
</paragraph>
</subSubSection>
<subSubSection id="C37015878B5F672A8086FCCAFC04FC94" box="[498,1016,821,848]" pageId="6" pageNumber="7" type="reference_group">
<paragraph id="8BD5460C8B5F672A8086FCCAFC04FC94" blockId="6.[498,1411,732,888]" box="[498,1016,821,848]" pageId="6" pageNumber="7">
<taxonomicName id="4C6A3D8F8B5F672A8086FCCAFD32FC94" box="[498,718,821,848]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B91E9A1E8B5F672A8086FCCAFD58FC8B" box="[498,676,821,847]" italics="true" pageId="6" pageNumber="7">Brachycephalus</emphasis>
sp.
</taxonomicName>
(
<bibRefCitation id="EFFB3BFD8B5F672A8394FCCAFC14FC8B" author="Bornschein MR &amp; Ribeiro LF &amp; Teixeira L &amp; Belmonte-Lopes R &amp; Moraes LA &amp; de Correa L &amp; Mauricio GN &amp; Pie MR" box="[736,1000,821,847]" journalOrPublisher="PeerJ" pageId="6" pageNumber="7" part="9" publicationUrl="10.7717/peerj.10983" refId="ref12763" refString="Bornschein MR, Ribeiro LF, Teixeira L, Belmonte-Lopes R, Moraes LA, de Correa L, Mauricio GN, Nadaline Ι, Pie MR. 2021. A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae). PeerJ 9: e 10983 DOI 10.7717 / peerj. 10983." title="A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae)" type="journal volume" year="2021">
<emphasis id="B91E9A1E8B5F672A8394FCCAFC14FC8B" box="[736,1000,821,847]" italics="true" pageId="6" pageNumber="7">Bornschein et al., 2021</emphasis>
</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C37015878B5F672A8086FCA2FA7FFCBC" box="[498,1411,861,888]" pageId="6" pageNumber="7" type="description">
<paragraph id="8BD5460C8B5F672A8086FCA2FA7FFCBC" blockId="6.[498,1411,732,888]" box="[498,1411,861,888]" pageId="6" pageNumber="7">
LSID:
<uri id="FFFB4A0E8B5F672A834AFCA2FA83FCBC" box="[574,1407,861,888]" pageId="6" pageNumber="7">
urn:lsid:zoobank.org:act:
<uuid id="FFCC7CD98B5F672A8227FCA2FA83FCBC" box="[851,1407,861,888]" pageId="6" pageNumber="7">18F93070-2CAE-4817-8CCD-48C0C5DA5210</uuid>
</uri>
.
</paragraph>
</subSubSection>
<subSubSection id="C37015878B5F672A8086FC63FDC6FB91" pageId="6" pageNumber="7" type="etymology">
<paragraph id="8BD5460C8B5F672A8086FC63FDC6FB91" blockId="6.[498,1542,923,1109]" pageId="6" pageNumber="7">
<emphasis id="B91E9A1E8B5F672A8086FC63FD88FC72" bold="true" box="[498,628,924,950]" pageId="6" pageNumber="7">Etymology</emphasis>
The specific epithet name
<emphasis id="B91E9A1E8B5F672A82B5FC64FBF3FC71" box="[961,1039,923,949]" italics="true" pageId="6" pageNumber="7">dacnis</emphasis>
honors the Projeto Dacnis private reserve and NGO that has supported biodiversity research since
<quantity id="4C92EBE98B5F672A85FCFC3CFB22FC19" box="[1160,1246,963,989]" metricMagnitude="1" metricUnit="m" metricValue="5.1053999999999995" pageId="6" pageNumber="7" unit="in" value="2010.0">2010 in</quantity>
the municipalities of São José dos Campos, Miracatu and Ubatuba (where the new species was discovered), state of
<collectingRegion id="49AE88EE8B5F672A8086FBECFD94FBE9" box="[498,616,1043,1069]" country="Brazil" name="Sao Paulo" pageId="6" pageNumber="7">São Paulo</collectingRegion>
,
<collectingCountry id="F37D069C8B5F672A8302FBECFD41FBE9" box="[630,701,1043,1069]" name="Brazil" pageId="6" pageNumber="7">Brazil</collectingCountry>
. The name is used as an invariable noun in apposition to the generic name.
</paragraph>
</subSubSection>
<subSubSection id="C37015878B5F672A8086FB86FD0FF974" pageId="6" pageNumber="7" type="materials_examined">
<paragraph id="8BD5460C8B5F672A8086FB86FA63FB27" blockId="6.[498,1542,1145,1252]" pageId="6" pageNumber="7">
<materialsCitation id="3B024C518B5F672A8086FB86FA63FB27" collectingDate="2021-05-18" collectorName="A. Mariano" country="Brazil" location="Projeto Dacnis reserve" municipality="Ubatuba" pageId="6" pageNumber="7" specimenCode="ZUEC-AMP 24982" specimenCount="2" specimenCount-adult="1" specimenCount-male="1" stateProvince="Sao Paulo" typeStatus="holotype">
<emphasis id="B91E9A1E8B5F672A8086FB86FD9FFB57" bold="true" box="[498,611,1145,1171]" pageId="6" pageNumber="7">
<typeStatus id="54D1F8AE8B5F672A8086FB86FD9FFB57" box="[498,611,1145,1171]" pageId="6" pageNumber="7" type="holotype">Holotype</typeStatus>
</emphasis>
<specimenCount id="9D6C8D858B5F672A8306FB86FD49FB57" box="[626,693,1145,1171]" pageId="6" pageNumber="7" type="adult">Adult</specimenCount>
<specimenCount id="9D6C8D858B5F672A83C9FB86FD09FB57" box="[701,757,1145,1171]" pageId="6" pageNumber="7" type="male">male</specimenCount>
(
<specimenCode id="DBCCEE778B5F672A8271FB86FC17FB57" box="[773,1003,1145,1171]" pageId="6" pageNumber="7">ZUEC-AMP 24982</specimenCode>
) collected and recorded (FNJV 51341) at
<location id="8EB510D78B5F672A8086FB5EFD0BFB78" LSID="urn:lsid:plazi:treatment:03C3F71A8B5F673E8086FCF2FDB2FDCF:8EB510D78B5F672A8086FB5EFD0BFB78" box="[498,759,1185,1212]" country="Brazil" municipality="Ubatuba" name="Projeto Dacnis reserve" pageId="6" pageNumber="7" stateProvince="Sao Paulo">Projeto Dacnis reserve</location>
, municipality of
<collectingMunicipality id="6BB1DC768B5F672A82C3FB5EFBE1FB7F" box="[951,1053,1185,1211]" pageId="6" pageNumber="7">Ubatuba</collectingMunicipality>
, state of
<collectingRegion id="49AE88EE8B5F672A85F4FB5EFB0FFB7F" box="[1152,1267,1185,1211]" country="Brazil" name="Sao Paulo" pageId="6" pageNumber="7">São Paulo</collectingRegion>
,
<collectingCountry id="F37D069C8B5F672A858AFB5EFAB8FB7F" box="[1278,1348,1185,1211]" name="Brazil" pageId="6" pageNumber="7">Brazil</collectingCountry>
, on
<date id="FFD460CC8B5F672A8406FB5EF9FAFB7F" box="[1394,1542,1185,1212]" pageId="6" pageNumber="7" value="2021-05-18">
<collectingDate id="EF9099248B5F672A8406FB5EF9FAFB7F" box="[1394,1542,1185,1212]" pageId="6" pageNumber="7" value="2021-05-18">18 May 2021</collectingDate>
</date>
by
<collectorName id="269F23DA8B5F672A8363FB36FD62FB27" box="[535,670,1225,1251]" pageId="6" pageNumber="7">A. Mariano</collectorName>
(
<figureCitation id="13515A898B5F672A83C5FB36FD0EFB27" box="[689,754,1225,1251]" captionStart="Figure 1" captionStartId="7.[524,589,673,695]" captionTargetBox="[498,1541,236,660]" captionTargetId="figure-144@7.[498,1299,234,645]" captionTargetPageId="7" captionText="Figure 1 Brachycephalus dacnis holotype (ZUEC-AMP 24982) adult male, SVL = 7.55 mm. (A) Dorsal and (B) ventral views. Full-size DOI: 10.7717/peerj.18265/fig-1" pageId="6" pageNumber="7">Fig. 1</figureCitation>
;
<tableCitation id="C6E873B78B5F672A8276FB36FCA5FB27" box="[770,857,1225,1251]" captionStart="Table 1" captionStartId="7.[525,580,794,815]" captionTargetBox="[516,1524,894,1446]" captionTargetPageId="7" captionText="Table 1 Measurements of the holotype and adult paratypes (one adult that was dissected for internal anatomy was not measured) of Brachycephalus dacnis. Values presented as mean ± standard deviation (minimummaximum) in millimeters." pageId="6" pageNumber="7">Table 1</tableCitation>
). We also sequenced the DNA of this individual.
</materialsCitation>
</paragraph>
<paragraph id="8BD5460C8B5F672A8086FAF7FD0FF974" blockId="6.[498,1542,1288,1712]" pageId="6" pageNumber="7">
<emphasis id="B91E9A1E8B5F672A8086FAF7FD94FAE6" bold="true" box="[498,616,1288,1314]" pageId="6" pageNumber="7">
<typeStatus id="54D1F8AE8B5F672A8086FAF7FD94FAE6" box="[498,616,1288,1314]" pageId="6" pageNumber="7" type="paratype">Paratypes</typeStatus>
</emphasis>
<specimenCount id="9D6C8D858B5F672A8303FAF7FCE9FAE6" box="[631,789,1288,1314]" count="11" pageId="6" pageNumber="7" type="adult">Eleven adults</specimenCount>
, only considered those confirmed by DNA and/or call traits:
<materialsCitation id="3B024C518B5F672A8086FAD0FD95FAB6" collectingDate="2019-08-03" collectingDateMax="2019-08-09" collectingDateMin="2019-08-03" collectorName="R. Mitsuo &amp; R. C. Menegucc" pageId="6" pageNumber="7" specimenCode="HCLP-A.267, HCLP-A.268" specimenCount="2" specimenCount-adult="2" typeStatus="paratype">
<date id="FFD460CC8B5F672A8086FAD0FD2AFA8D" box="[498,726,1327,1354]" pageId="6" pageNumber="7" value="2019-08-03" valueMax="2019-08-09" valueMin="2019-08-03">
<collectingDate id="EF9099248B5F672A8086FAD0FD2AFA8D" box="[498,726,1327,1354]" pageId="6" pageNumber="7" value="2019-08-03" valueMax="2019-08-09" valueMin="2019-08-03">0309 August 2019</collectingDate>
</date>
by
<collectorName id="269F23DA8B5F672A8277FACFFC84FA8D" box="[771,888,1327,1354]" pageId="6" pageNumber="7">R. Mitsuo</collectorName>
and
<collectorName id="269F23DA8B5F672A82C1FAD0FB91FA8D" box="[949,1133,1327,1353]" pageId="6" pageNumber="7">R.C. Menegucci</collectorName>
(HCLP-A 26768; heard, but not recorded);
</materialsCitation>
<materialsCitation id="3B024C518B5F672A8306FAA8FD8BFA5E" collectingDate="2021-03-10" collectorName="E. Muscat &amp; L. M. Botelho" pageId="6" pageNumber="7" specimenCode="ZUEC-AMP 24978" specimenCount="1" typeStatus="paratype">
<date id="FFD460CC8B5F672A8306FAA8FCD9FAB5" box="[626,805,1367,1393]" pageId="6" pageNumber="7" value="2021-03-10">
<collectingDate id="EF9099248B5F672A8306FAA8FCD9FAB5" box="[626,805,1367,1393]" pageId="6" pageNumber="7" value="2021-03-10">10 March 2021</collectingDate>
</date>
by
<collectorName id="269F23DA8B5F672A8226FAA7FC3BFAB6" box="[850,967,1368,1394]" pageId="6" pageNumber="7">E. Muscat</collectorName>
and
<collectorName id="269F23DA8B5F672A8570FAA7FB61FAB5" box="[1028,1181,1367,1394]" pageId="6" pageNumber="7">L.M. Botelho</collectorName>
(
<specimenCode id="DBCCEE778B5F672A85DAFAA8FA68FAB5" box="[1198,1428,1367,1393]" pageId="6" pageNumber="7">ZUEC-AMP 24978</specimenCode>
; sequenced)
</materialsCitation>
;
<materialsCitation id="3B024C518B5F672A83F7FA80FA9AFA5E" box="[643,1382,1407,1434]" collectingDate="2021-05-05" collectorName="L. M. Botelho" pageId="6" pageNumber="7" specimenCode="ZUEC-AMP 24984" specimenCount="1" typeStatus="paratype">
<date id="FFD460CC8B5F672A83F7FA80FCE6FA5D" box="[643,794,1407,1434]" pageId="6" pageNumber="7" value="2021-05-05">
<collectingDate id="EF9099248B5F672A83F7FA80FCE6FA5D" box="[643,794,1407,1434]" pageId="6" pageNumber="7" value="2021-05-05">05 May 2021</collectingDate>
</date>
by
<collectorName id="269F23DA8B5F672A8232FA7FFC22FA5D" box="[838,990,1407,1434]" pageId="6" pageNumber="7">L.M. Botelho</collectorName>
(
<specimenCode id="DBCCEE778B5F672A8299FA80FB2EFA5D" box="[1005,1234,1407,1433]" pageId="6" pageNumber="7">ZUEC-AMP 24984</specimenCode>
; sequenced);
</materialsCitation>
<materialsCitation id="3B024C518B5F672A841AFA80FBE5FA06" collectingDate="2021-05-18" collectorName="A. Mariano" pageId="6" pageNumber="7" specimenCode="ZUEC-AMP 24979" specimenCount="1" typeStatus="paratype">
<date id="FFD460CC8B5F672A841AFA80F9FAFA5D" box="[1390,1542,1407,1434]" pageId="6" pageNumber="7" value="2021-05-18">
<collectingDate id="EF9099248B5F672A841AFA80F9FAFA5D" box="[1390,1542,1407,1434]" pageId="6" pageNumber="7" value="2021-05-18">18 May 2021</collectingDate>
</date>
by
<collectorName id="269F23DA8B5F672A8363FA58FD62FA05" box="[535,670,1447,1473]" pageId="6" pageNumber="7">A. Mariano</collectorName>
(
<specimenCode id="DBCCEE778B5F672A83DBFA58FC69FA05" box="[687,917,1447,1473]" pageId="6" pageNumber="7">ZUEC-AMP 24979</specimenCode>
; recorded);
</materialsCitation>
<materialsCitation id="3B024C518B5F672A8556FA58FB80FA2E" collectingDate="2021-05-24" collectorName="E. Muscat &amp; L. M. Botelho" pageId="6" pageNumber="7" specimenCode="ZUEC-AMP 24981" specimenCount="1" typeStatus="paratype">
<date id="FFD460CC8B5F672A8556FA58FB47FA05" box="[1058,1211,1447,1473]" pageId="6" pageNumber="7" value="2021-05-24">
<collectingDate id="EF9099248B5F672A8556FA58FB47FA05" box="[1058,1211,1447,1473]" pageId="6" pageNumber="7" value="2021-05-24">24 May 2021</collectingDate>
</date>
by
<collectorName id="269F23DA8B5F672A859DFA58FAA1FA05" box="[1257,1373,1447,1473]" pageId="6" pageNumber="7">E. Muscat</collectorName>
and
<collectorName id="269F23DA8B5F672A84EFFA58FDB0FA2D" pageId="6" pageNumber="7">L.M. Botelho</collectorName>
(
<specimenCode id="DBCCEE778B5F672A8329FA30FCBFFA2D" box="[605,835,1487,1513]" pageId="6" pageNumber="7">ZUEC-AMP 24981</specimenCode>
; recorded and sequenced);
</materialsCitation>
<materialsCitation id="3B024C518B5F672A85F1FA30FB5EF9D6" collectingDate="2021-02-22" collectingDateMax="2021-02-21" collectingDateMin="2021-02-21" collectorName="L. M. Botelho" pageId="6" pageNumber="7" specimenCode="ZUEC-AMP 25270" specimenCount="1" typeStatus="paratype">
<date id="FFD460CC8B5F672A85F1FA30FAC2FA2D" box="[1157,1342,1487,1513]" pageId="6" pageNumber="7" value="2021-02-22">
<collectingDate id="EF9099248B5F672A85F1FA30FAC2FA2D" box="[1157,1342,1487,1513]" pageId="6" pageNumber="7" value="2021-02-22">21, 22 February</collectingDate>
</date>
2022 by
<collectorName id="269F23DA8B5F672A84D8FA30FDB0F9D5" pageId="6" pageNumber="7">L.M. Botelho</collectorName>
(
<specimenCode id="DBCCEE778B5F672A832EFA08FCC2F9D5" box="[602,830,1526,1553]" pageId="6" pageNumber="7">ZUEC-AMP 25270</specimenCode>
; 25272; 2527475; sequenced);
</materialsCitation>
<materialsCitation id="3B024C518B5F672A85DDFA08FA6AF9FD" collectingDate="2022-04-14" collectorName="E. Muscat &amp; L. M. Botelho &amp; L. A. O. S Ferreira" pageId="6" pageNumber="7" specimenCode="ZUEC-AMP 25612" specimenCount="2" specimenCount-adult="1" specimenCount-male="1" typeStatus="paratype">
<date id="FFD460CC8B5F672A85DDFA08FABBF9D5" box="[1193,1351,1527,1553]" pageId="6" pageNumber="7" value="2022-04-14">
<collectingDate id="EF9099248B5F672A85DDFA08FABBF9D5" box="[1193,1351,1527,1553]" pageId="6" pageNumber="7" value="2022-04-14">14 April 2022</collectingDate>
</date>
by
<collectorName id="269F23DA8B5F672A8404FA08FA19F9D5" box="[1392,1509,1527,1553]" pageId="6" pageNumber="7">E. Muscat</collectorName>
,
<collectorName id="269F23DA8B5F672A8484FA08FD8BF9FC" pageId="6" pageNumber="7">L. M. Botelho</collectorName>
,
<collectorName id="269F23DA8B5F672A83F0F9E0FCBFF9FC" box="[644,835,1566,1593]" pageId="6" pageNumber="7">L.A.O.S Ferreira</collectorName>
(
<specimenCount id="9D6C8D858B5F672A8220F9E1FC6DF9FC" box="[852,913,1566,1592]" pageId="6" pageNumber="7" type="adult">adult</specimenCount>
<specimenCount id="9D6C8D858B5F672A82EEF9E1FC28F9FC" box="[922,980,1566,1592]" pageId="6" pageNumber="7" type="male">male</specimenCount>
;
<specimenCode id="DBCCEE778B5F672A8295F9E0FB39F9FD" box="[993,1221,1566,1593]" pageId="6" pageNumber="7">ZUEC-AMP 25612</specimenCode>
;
<figureCitation id="13515A898B5F672A85A6F9E1FAEFF9FD" box="[1234,1299,1566,1593]" captionStart="Figure 2" captionStartId="8.[524,589,1066,1088]" captionTargetBox="[498,1541,236,1053]" captionTargetId="figure-278@8.[498,1299,234,1038]" captionTargetPageId="8" captionText="Figure 2 Brachycephalus dacnis paratype (ZUEC-AMP 25612) adult male, SVL = 7.89 mm. (A) Dorsal view, (B) ventral view, (C) ventral view of hand, (D) ventral view of feet. Full-size DOI: 10.7717/peerj.18265/fig-2" pageId="6" pageNumber="7">Fig. 2</figureCitation>
; recorded)
</materialsCitation>
. All
<typeStatus id="54D1F8AE8B5F672A8086F9B8FD9EF9A5" box="[498,610,1607,1633]" pageId="6" pageNumber="7" type="paratype">paratypes</typeStatus>
were collected at the same locality as the
<typeStatus id="54D1F8AE8B5F672A8526F9B9FB47F9A4" box="[1106,1211,1606,1632]" pageId="6" pageNumber="7" type="holotype">holotype</typeStatus>
. Sex determination was not possible in most cases, except for the individual that was dissected (a female; see below) and the calling males.
</paragraph>
</subSubSection>
<subSubSection id="C37015878B5F672B8086F92BFBCAF92E" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7" type="diagnosis">
<paragraph id="8BD5460C8B5F672B8086F92BFBCAF92E" blockId="6.[498,1542,1748,1934]" lastBlockId="7.[498,1542,1544,1770]" lastPageId="7" lastPageNumber="8" pageId="6" pageNumber="7">
<emphasis id="B91E9A1E8B5F672A8086F92BFD96F92A" bold="true" box="[498,618,1748,1774]" pageId="6" pageNumber="7">Diagnosis</emphasis>
The new species is assigned to the genus
<taxonomicName id="4C6A3D8F8B5F672A8515F92BFAEFF92A" baseAuthorityName="Trueb &amp; Alberch" baseAuthorityYear="1985" box="[1121,1299,1748,1774]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="6" pageNumber="7" phylum="Chordata" rank="genus">
<emphasis id="B91E9A1E8B5F672A8515F92BFAEFF92A" box="[1121,1299,1748,1774]" italics="true" pageId="6" pageNumber="7">Brachycephalus</emphasis>
</taxonomicName>
because of its miniature body size, fewer phalanges and toes than a typical frog, fingers and toe tips not expanded but apically pointed, and toes lacking circumferential grooves. The new species can be diagnosed from its congeners by the following combination of characters: (1) “leptodactyliform” body shape; (2) adult body length (SVL) smaller than
<quantity id="4C92EBE98B5F672A8434F88BFA87F84A" box="[1344,1403,1908,1934]" metricMagnitude="-2" metricUnit="m" metricValue="1.0" pageId="6" pageNumber="7" unit="cm" value="1.0">1 cm</quantity>
; (3) distinct and functional toes II and V; (4) presence of vestigial fingers Iand IV; (5) distinct iris; (6) absence of dark markings on the skin over the pectoral region; (7) dark black or pale brown marbled venter with small white blotches in preserved specimens; (8) and advertisement call composed of one or two multi-pulsed (37 pulses) note with dominant frequency between 8.01 and 8.44 kHz, note duration between 0.03
<geoCoordinate id="EE5E20CB8B5E672B8509F957FB42F906" box="[1149,1214,1704,1730]" degrees="0.08" direction="south" orientation="latitude" pageId="7" pageNumber="8" precision="555" value="-0.08">0.08 s</geoCoordinate>
(when isolated), up to
<geoCoordinate id="EE5E20CB8B5E672B84B1F957F9FAF906" box="[1477,1542,1704,1730]" degrees="0.41" direction="south" orientation="latitude" pageId="7" pageNumber="8" precision="555" value="-0.41">0.41 s</geoCoordinate>
(when in pairs), and absence of attenuated notes.
</paragraph>
</subSubSection>
<caption id="DF1516848B5E672B8378FD5EFA17FD10" pageId="7" pageNumber="8" startId="7.[524,589,673,695]" targetBox="[498,1541,236,660]" targetPageId="7" targetType="figure">
<paragraph id="8BD5460C8B5E672B8378FD5EFA17FD10" blockId="7.[524,1515,673,725]" pageId="7" pageNumber="8">
<emphasis id="B91E9A1E8B5E672B8378FD5EFA17FD72" bold="true" box="[524,1515,673,695]" pageId="7" pageNumber="8">
Figure 1
<taxonomicName id="4C6A3D8F8B5E672B8306FD5EFCA4FD73" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[626,856,673,695]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5E672B8306FD5EFCA4FD73" bold="true" box="[626,856,673,695]" italics="true" pageId="7" pageNumber="8">Brachycephalusdacnis</emphasis>
</taxonomicName>
holotype (ZUEC-AMP 24982) adult male, SVL = 7.55 mm.
</emphasis>
(A) Dorsal and (B) ventral views. Full-size DOI: 10.7717/peerj.18265/fig-1
</paragraph>
</caption>
<caption id="DF1516848B5E672B8379FCE5FC7DFCAE" pageId="7" pageNumber="8" startId="7.[525,580,794,815]" targetBox="[516,1524,894,1446]" targetIsTable="true" targetPageId="7" targetType="table">
<paragraph id="8BD5460C8B5E672B8379FCE5FC7DFCAE" blockId="7.[525,1515,793,874]" pageId="7" pageNumber="8">
<emphasis id="B91E9A1E8B5E672B8379FCE5FBDAFC88" bold="true" pageId="7" pageNumber="8">
Table 1 Measurementsoftheholotypeandadultparatypes (oneadultthatwasdissectedforinternal anatomy was not measured) of
<taxonomicName id="4C6A3D8F8B5E672B8236FCC9FBE3FC88" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[834,1055,822,844]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="7" pageNumber="8" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5E672B8236FCC9FBE3FC88" bold="true" box="[834,1055,822,844]" italics="true" pageId="7" pageNumber="8">Brachycephalus dacnis</emphasis>
</taxonomicName>
.
</emphasis>
Values presented as mean ± standard deviation (minimum maximum) in millimeters.
</paragraph>
</caption>
<paragraph id="8BD5460C8B5E672B8370FC81FA26FA62" pageId="7" pageNumber="8">
<table id="F96AB4AC8B5E98D38370FC81FA08FA62" box="[516,1524,894,1446]" gridcols="3" gridrows="15" pageId="7" pageNumber="8">
<tr id="355A444E8B5E98D38370FC81FA08FC50" box="[516,1524,894,916]" gridrow="0" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FC81FDCAFC50" box="[516,566,894,916]" gridcol="0" gridrow="0" pageId="7" pageNumber="8">Trait</th>
<th id="768B2D328B5E98D3821FFC81FC3BFC50" box="[875,967,894,916]" gridcol="1" gridrow="0" pageId="7" pageNumber="8">Holotype</th>
<th id="768B2D328B5E98D38588FC81FA08FC50" box="[1276,1524,894,916]" gridcol="2" gridrow="0" pageId="7" pageNumber="8">
<emphasis id="B91E9A1E8B5E672B8588FC81FA08FC50" bold="true" box="[1276,1524,894,916]" pageId="7" pageNumber="8">
Adult paratypes (
<emphasis id="B91E9A1E8B5E672B84DFFC80FA44FC50" bold="true" box="[1451,1464,895,916]" italics="true" pageId="7" pageNumber="8">n</emphasis>
= 10)
</emphasis>
</th>
</tr>
<tr id="355A444E8B5E98D38370FC58FA08FC7A" box="[516,1524,935,958]" gridrow="1" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FC58FDCAFC7A" box="[516,566,935,958]" gridcol="0" gridrow="1" pageId="7" pageNumber="8">SVL</th>
<td id="768B2D328B5E98D3821FFC58FC3BFC7A" box="[875,967,935,958]" gridcol="1" gridrow="1" pageId="7" pageNumber="8">7.55</td>
<td id="768B2D328B5E98D38588FC58FA08FC7A" box="[1276,1524,935,958]" gridcol="2" gridrow="1" pageId="7" pageNumber="8">8.29 ± 1.03 (6.959.90)</td>
</tr>
<tr id="355A444E8B5E98D38370FC32FA08FC27" box="[516,1524,973,995]" gridrow="2" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FC32FDCAFC27" box="[516,566,973,995]" gridcol="0" gridrow="2" pageId="7" pageNumber="8">HL</th>
<td id="768B2D328B5E98D3821FFC32FC3BFC27" box="[875,967,973,995]" gridcol="1" gridrow="2" pageId="7" pageNumber="8">1.78</td>
<td id="768B2D328B5E98D38588FC32FA08FC27" box="[1276,1524,973,995]" gridcol="2" gridrow="2" pageId="7" pageNumber="8">2.59 ± 0.45 (1.873.18)</td>
</tr>
<tr id="355A444E8B5E98D38370FC0DFA08FBCD" box="[516,1524,1010,1033]" gridrow="3" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FC0DFDCAFBCD" box="[516,566,1010,1033]" gridcol="0" gridrow="3" pageId="7" pageNumber="8">HW</th>
<td id="768B2D328B5E98D3821FFC0DFC3BFBCD" box="[875,967,1010,1033]" gridcol="1" gridrow="3" pageId="7" pageNumber="8">1.78</td>
<td id="768B2D328B5E98D38588FC0DFA08FBCD" box="[1276,1524,1010,1033]" gridcol="2" gridrow="3" pageId="7" pageNumber="8">2.97 ± 0.55 (1.823.63)</td>
</tr>
<tr id="355A444E8B5E98D38370FBE7FA08FBEA" box="[516,1524,1048,1070]" gridrow="4" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FBE7FDCAFBEA" box="[516,566,1048,1070]" gridcol="0" gridrow="4" pageId="7" pageNumber="8">END</th>
<td id="768B2D328B5E98D3821FFBE7FC3BFBEA" box="[875,967,1048,1070]" gridcol="1" gridrow="4" pageId="7" pageNumber="8">0.37</td>
<td id="768B2D328B5E98D38588FBE7FA08FBEA" box="[1276,1524,1048,1070]" gridcol="2" gridrow="4" pageId="7" pageNumber="8">0.48 ± 0.23 (0.181.02)</td>
</tr>
<tr id="355A444E8B5E98D38370FBC1FA08FB90" box="[516,1524,1086,1108]" gridrow="5" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FBC1FDCAFB90" box="[516,566,1086,1108]" gridcol="0" gridrow="5" pageId="7" pageNumber="8">IOD</th>
<td id="768B2D328B5E98D3821FFBC1FC3BFB90" box="[875,967,1086,1108]" gridcol="1" gridrow="5" pageId="7" pageNumber="8">1.42</td>
<td id="768B2D328B5E98D38588FBC1FA08FB90" box="[1276,1524,1086,1108]" gridcol="2" gridrow="5" pageId="7" pageNumber="8">1.48 ± 0.32 (0.842.10)</td>
</tr>
<tr id="355A444E8B5E98D38370FB9CFA08FBBE" box="[516,1524,1123,1146]" gridrow="6" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FB9CFDCAFBBE" box="[516,566,1123,1146]" gridcol="0" gridrow="6" pageId="7" pageNumber="8">ED</th>
<td id="768B2D328B5E98D3821FFB9CFC3BFBBE" box="[875,967,1123,1146]" gridcol="1" gridrow="6" pageId="7" pageNumber="8">0.49</td>
<td id="768B2D328B5E98D38588FB9CFA08FBBE" box="[1276,1524,1123,1146]" gridcol="2" gridrow="6" pageId="7" pageNumber="8">0.94 ± 0.24 (0.501.30)</td>
</tr>
<tr id="355A444E8B5E98D38370FB76FA08FB5B" box="[516,1524,1161,1183]" gridrow="7" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FB76FDCAFB5B" box="[516,566,1161,1183]" gridcol="0" gridrow="7" pageId="7" pageNumber="8">ND</th>
<td id="768B2D328B5E98D3821FFB76FC3BFB5B" box="[875,967,1161,1183]" gridcol="1" gridrow="7" pageId="7" pageNumber="8">0.31</td>
<td id="768B2D328B5E98D38588FB76FA08FB5B" box="[1276,1524,1161,1183]" gridcol="2" gridrow="7" pageId="7" pageNumber="8">0.27 ± 0.08 (0.160.40)</td>
</tr>
<tr id="355A444E8B5E98D38370FB50FA08FB01" box="[516,1524,1199,1221]" gridrow="8" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FB50FDCAFB01" box="[516,566,1199,1221]" gridcol="0" gridrow="8" pageId="7" pageNumber="8">UAL</th>
<td id="768B2D328B5E98D3821FFB50FC3BFB01" box="[875,967,1199,1221]" gridcol="1" gridrow="8" pageId="7" pageNumber="8">1.30</td>
<td id="768B2D328B5E98D38588FB50FA08FB01" box="[1276,1524,1199,1221]" gridcol="2" gridrow="8" pageId="7" pageNumber="8">1.57 ± 0.22 (1.302.02)</td>
</tr>
<tr id="355A444E8B5E98D38370FB2BFA08FB2E" box="[516,1524,1236,1258]" gridrow="9" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FB2BFDCAFB2E" box="[516,566,1236,1258]" gridcol="0" gridrow="9" pageId="7" pageNumber="8">FAL</th>
<td id="768B2D328B5E98D3821FFB2BFC3BFB2E" box="[875,967,1236,1258]" gridcol="1" gridrow="9" pageId="7" pageNumber="8">2.95</td>
<td id="768B2D328B5E98D38588FB2BFA08FB2E" box="[1276,1524,1236,1258]" gridcol="2" gridrow="9" pageId="7" pageNumber="8">2.13 ± 0.30 (1.802.70)</td>
</tr>
<tr id="355A444E8B5E98D38370FB05FA08FAD4" box="[516,1524,1274,1296]" gridrow="10" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FB05FDCAFAD4" box="[516,566,1274,1296]" gridcol="0" gridrow="10" pageId="7" pageNumber="8">HAL</th>
<td id="768B2D328B5E98D3821FFB05FC3BFAD4" box="[875,967,1274,1296]" gridcol="1" gridrow="10" pageId="7" pageNumber="8">1.00</td>
<td id="768B2D328B5E98D38588FB05FA08FAD4" box="[1276,1524,1274,1296]" gridcol="2" gridrow="10" pageId="7" pageNumber="8">1.28 ± 0.23 (0.921.63)</td>
</tr>
<tr id="355A444E8B5E98D38370FAE0FA08FAF2" box="[516,1524,1311,1334]" gridrow="11" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FAE0FDCAFAF2" box="[516,566,1311,1334]" gridcol="0" gridrow="11" pageId="7" pageNumber="8">TBL</th>
<td id="768B2D328B5E98D3821FFAE0FC3BFAF2" box="[875,967,1311,1334]" gridcol="1" gridrow="11" pageId="7" pageNumber="8">3.59</td>
<td id="768B2D328B5E98D38588FAE0FA08FAF2" box="[1276,1524,1311,1334]" gridcol="2" gridrow="11" pageId="7" pageNumber="8">3.82 ± 0.48 (2.984.50)</td>
</tr>
<tr id="355A444E8B5E98D38370FABAFA08FA9F" box="[516,1524,1349,1371]" gridrow="12" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FABAFDCAFA9F" box="[516,566,1349,1371]" gridcol="0" gridrow="12" pageId="7" pageNumber="8">THL</th>
<td id="768B2D328B5E98D3821FFABAFC3BFA9F" box="[875,967,1349,1371]" gridcol="1" gridrow="12" pageId="7" pageNumber="8">4.00</td>
<td id="768B2D328B5E98D38588FABAFA08FA9F" box="[1276,1524,1349,1371]" gridcol="2" gridrow="12" pageId="7" pageNumber="8">3.77 ± 0.51 (3.255.00)</td>
</tr>
<tr id="355A444E8B5E98D38370FA95FA08FA45" box="[516,1524,1386,1409]" gridrow="13" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FA95FDCAFA45" box="[516,566,1386,1409]" gridcol="0" gridrow="13" pageId="7" pageNumber="8">FL</th>
<td id="768B2D328B5E98D3821FFA95FC3BFA45" box="[875,967,1386,1409]" gridcol="1" gridrow="13" pageId="7" pageNumber="8">4.50</td>
<td id="768B2D328B5E98D38588FA95FA08FA45" box="[1276,1524,1386,1409]" gridcol="2" gridrow="13" pageId="7" pageNumber="8">6.10 ± 0.41 (5.606.75)</td>
</tr>
<tr id="355A444E8B5E98D38370FA6FFA08FA62" box="[516,1524,1424,1446]" gridrow="14" pageId="7" pageNumber="8">
<th id="768B2D328B5E98D38370FA6FFDCAFA62" box="[516,566,1424,1446]" gridcol="0" gridrow="14" pageId="7" pageNumber="8">fL</th>
<td id="768B2D328B5E98D3821FFA6FFC3BFA62" box="[875,967,1424,1446]" gridcol="1" gridrow="14" pageId="7" pageNumber="8">2.75</td>
<td id="768B2D328B5E98D38588FA6FFA08FA62" box="[1276,1524,1424,1446]" gridcol="2" gridrow="14" pageId="7" pageNumber="8">3.16 ± 0.47 (2.354.00)</td>
</tr>
</table>
</paragraph>
<subSubSection id="C37015878B5E67218086F8F1FBABFA5B" lastPageId="13" lastPageNumber="14" pageId="7" pageNumber="8" type="description">
<paragraph id="8BD5460C8B5E67248086F8F1FA79F916" blockId="7.[498,1542,1806,1913]" lastBlockId="8.[498,1542,1202,1746]" lastPageId="8" lastPageNumber="9" pageId="7" pageNumber="8">
<emphasis id="B91E9A1E8B5E672B8086F8F1FD08F8EC" bold="true" box="[498,756,1806,1832]" pageId="7" pageNumber="8">
<typeStatus id="54D1F8AE8B5E672B8086F8F1FD9FF8EC" box="[498,611,1806,1832]" pageId="7" pageNumber="8" type="holotype">Holotype</typeStatus>
description
</emphasis>
Body slender and leptodactyliform (
<figureCitation id="13515A898B5E672B85DFF8F1FB10F8EC" box="[1195,1260,1806,1832]" captionStart="Figure 1" captionStartId="7.[524,589,673,695]" captionTargetBox="[498,1541,236,660]" captionTargetId="figure-144@7.[498,1299,234,645]" captionTargetPageId="7" captionText="Figure 1 Brachycephalus dacnis holotype (ZUEC-AMP 24982) adult male, SVL = 7.55 mm. (A) Dorsal and (B) ventral views. Full-size DOI: 10.7717/peerj.18265/fig-1" pageId="7" pageNumber="8">Fig. 1</figureCitation>
; SVL =
<quantity id="4C92EBE98B5E672B8439F8F1FA45F8ED" box="[1357,1465,1806,1833]" metricMagnitude="-3" metricUnit="m" metricValue="7.55" pageId="7" pageNumber="8" unit="mm" value="7.55">7.55 mm</quantity>
); head as wide as long (
<quantity id="4C92EBE98B5E672B83C6F8C9FCE1F894" box="[690,797,1846,1872]" metricMagnitude="-3" metricUnit="m" metricValue="1.78" pageId="7" pageNumber="8" unit="mm" value="1.78">1.78 mm</quantity>
;
<tableCitation id="C6E873B78B5E672B825EF8C9FC7CF894" box="[810,896,1846,1872]" captionStart="Table 1" captionStartId="7.[525,580,794,815]" captionTargetBox="[516,1524,894,1446]" captionTargetPageId="7" captionText="Table 1 Measurements of the holotype and adult paratypes (one adult that was dissected for internal anatomy was not measured) of Brachycephalus dacnis. Values presented as mean ± standard deviation (minimummaximum) in millimeters." pageId="7" pageNumber="8">Table 1</tableCitation>
), and one-fourth of total body length (HW/SVL = HL/ SVL = 23.6%); snout short and slightly mucronate in dorsal view and between rounded and vertical in lateral view; nostrils oval, not protuberant and directed anterolaterally;
<emphasis id="B91E9A1E8B51672484D8FB4DFDB2FB37" italics="true" pageId="8" pageNumber="9">canthus rostralis</emphasis>
distinct; loreal region slightly concave; lips nearly sigmoid; eyes slightly protruding in dorsal and lateral view, and directed anterolaterally, 27.5% ED/HW and ED/HL (as HW = HL); tympanum indistinct; vocal sac not externally expanded; vocal slits present; tongue longer than wide, with posterior half free; choanae small and rounded; vomerine teeth absent. Forearm more than twice (2.3 times) length of upper arm length; hand length shorter than forearm or upper arm lengths; hands with all fingers distinct; fingers Iand IV externally vestigial; fingers II and III robust; tips of fingers Iand IV (when visible) rounded, fingers II and III pointed; relative finger length I ≤ IV &lt;II &lt;III; subarticular tubercles absent; inner and outer metacarpal tubercles absent. Leg relatively long, total leg length (TBL + THL + FL =
<quantity id="4C92EBE98B516724839FF9BFFCA9F99F" box="[747,853,1600,1627]" metricMagnitude="-2" metricUnit="m" metricValue="1.21" pageId="8" pageNumber="9" unit="mm" value="12.1">12.1 mm</quantity>
) 160% of SVL (
<quantity id="4C92EBE98B5167248564F9BFFB8CF99F" box="[1040,1136,1600,1627]" metricMagnitude="-3" metricUnit="m" metricValue="7.6" pageId="8" pageNumber="9" unit="mm" value="7.6">7.6 mm</quantity>
); shank slightly shorter than thigh (SHL/SVL = 40.6% and THL/SVL = 47.5%); foot longer than tarsus and the shank; toe I externally absent; toes II, III, IV and V externally distinct and functional. Skin smooth without dermal ossifications. Measurements of
<typeStatus id="54D1F8AE8B5167248554F947FB7AF916" box="[1056,1158,1720,1746]" pageId="8" pageNumber="9" type="holotype">holotype</typeStatus>
presented in
<tableCitation id="C6E873B78B516724845CF947FA83F916" box="[1320,1407,1720,1746]" captionStart="Table 1" captionStartId="7.[525,580,794,815]" captionTargetBox="[516,1524,894,1446]" captionTargetPageId="7" captionText="Table 1 Measurements of the holotype and adult paratypes (one adult that was dissected for internal anatomy was not measured) of Brachycephalus dacnis. Values presented as mean ± standard deviation (minimummaximum) in millimeters." pageId="8" pageNumber="9">Table 1</tableCitation>
.
</paragraph>
<caption id="DF1516848B5167248378FBD5FA17FBBE" pageId="8" pageNumber="9" startId="8.[524,589,1066,1088]" targetBox="[498,1541,236,1053]" targetPageId="8" targetType="figure">
<paragraph id="8BD5460C8B5167248378FBD5FAC6FB99" blockId="8.[524,1515,1066,1146]" pageId="8" pageNumber="9">
<emphasis id="B91E9A1E8B5167248378FBD5FA17FBFB" bold="true" box="[524,1515,1066,1088]" pageId="8" pageNumber="9">
Figure 2
<taxonomicName id="4C6A3D8F8B5167248306FBD5FCA4FB84" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[626,856,1066,1088]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="8" pageNumber="9" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5167248306FBD5FCA4FB84" bold="true" box="[626,856,1066,1088]" italics="true" pageId="8" pageNumber="9">Brachycephalusdacnis</emphasis>
</taxonomicName>
paratype (ZUEC-AMP 25612) adult male, SVL = 7.89 mm.
</emphasis>
(A) Dorsal view, (B) ventral view, (C) ventral view of hand, (D) ventral view of feet.
</paragraph>
<paragraph id="8BD5460C8B516724853FFB9BFA17FBBE" blockId="8.[524,1515,1066,1146]" box="[1099,1515,1124,1146]" pageId="8" pageNumber="9">Full-size DOI: 10.7717/peerj.18265/fig-2</paragraph>
</caption>
<paragraph id="8BD5460C8B5167258086F909F9FAFB56" blockId="8.[498,1542,1782,1928]" lastBlockId="9.[498,1542,1065,1171]" lastPageId="9" lastPageNumber="10" pageId="8" pageNumber="9">
<emphasis id="B91E9A1E8B5167248086F909FCB6F8D4" bold="true" box="[498,842,1782,1808]" pageId="8" pageNumber="9">
Color in life of the
<typeStatus id="54D1F8AE8B5167248395F909FCB6F8D4" box="[737,842,1782,1808]" pageId="8" pageNumber="9" type="holotype">holotype</typeStatus>
</emphasis>
The general coloration of the dorsal body surface is yellowish-brown. Adark brown stripe extends laterally from the tip of the snout to the flanks. Dorsal surfaces of arms and legs interrupted by dark brown stripes. An x-shaped or dorsal mark is present. The lateral view of the body is dark brown; the region overlying the upper jaw is dark brown with distinct white spots. The ventral body surface is brown with small white blotches. The ventral surface of the hands and feet are dark brown with white blotches over each phalanx of the fingers and toes. The pupil is black, and the iris is bronze.
</paragraph>
<caption id="DF1516848B5067258378FC7BFA17FC35" pageId="9" pageNumber="10" startId="9.[524,589,900,922]" targetBox="[498,1299,234,868]" targetPageId="9" targetType="figure">
<paragraph id="8BD5460C8B5067258378FC7BFA17FC35" blockId="9.[524,1516,899,1010]" pageId="9" pageNumber="10">
<emphasis id="B91E9A1E8B5067258378FC7BFBEFFC73" bold="true" pageId="9" pageNumber="10">
Figure 3
<taxonomicName id="4C6A3D8F8B5067258319FC7CFCB1FC5D" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[621,845,899,921]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="9" pageNumber="10" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5067258319FC7CFCB1FC5D" bold="true" box="[621,845,899,921]" italics="true" pageId="9" pageNumber="10">Brachycephalus dacnis</emphasis>
</taxonomicName>
paratype individuals found at the Projeto Dacnis private reserve, municipality of Ubatuba, state of São Paulo, Brazil.
</emphasis>
(A) ZUEC-AMP 25272; (B) ZUEC-AMP 25274; (C and D) ZUEC-AMP 25275 displaying mouth-gaping behavior and the same individual on top of one herpetologist s fingertip. Full-size DOI: 10.7717/peerj.18265/fig-3
</paragraph>
</caption>
<paragraph id="8BD5460C8B5067258086FB48FCD8FA8C" blockId="9.[498,1542,1207,1353]" pageId="9" pageNumber="10">
<emphasis id="B91E9A1E8B5067258086FB48FD52FB15" bold="true" box="[498,686,1207,1233]" pageId="9" pageNumber="10">Color variation</emphasis>
Color and ornamentation patterns in this species vary considerably among individuals. Some frogs have a leaf-like dorsal coloration, while others have a rock-like dorsal coloration (
<figureCitation id="13515A898B506725824CFAF9FC84FAE5" box="[824,888,1286,1313]" captionStart="Figure 3" captionStartId="9.[524,589,900,922]" captionTargetBox="[498,1299,234,868]" captionTargetId="figure-274@9.[498,1299,234,872]" captionTargetPageId="9" captionText="Figure 3 Brachycephalus dacnis paratype individuals found at the Projeto Dacnis private reserve, municipality of Ubatuba, state of São Paulo, Brazil. (A) ZUEC-AMP 25272; (B) ZUEC-AMP 25274; (C and D) ZUEC-AMP 25275 displaying mouth-gaping behavior and the same individual on top of one herpetologists fingertip. Full-size DOI: 10.7717/peerj.18265/fig-3" pageId="9" pageNumber="10">Fig. 3</figureCitation>
). We did not observe significant variation in the ventral pattern of the individuals.
</paragraph>
<paragraph id="8BD5460C8B5067258086FA92FD27F9E3" blockId="9.[498,1541,1389,1575]" pageId="9" pageNumber="10">
<emphasis id="B91E9A1E8B5067258086FA92FC4AFA43" bold="true" box="[498,950,1389,1415]" pageId="9" pageNumber="10">
Color in preservative of the
<typeStatus id="54D1F8AE8B5067258239FA92FC4AFA43" box="[845,950,1389,1415]" pageId="9" pageNumber="10" type="holotype">holotype</typeStatus>
</emphasis>
(less than 1 year in preservative)The general coloration of the dorsal surface is reddish-brown (
<figureCitation id="13515A898B5067258530FA6AFB7AFA6B" box="[1092,1158,1429,1455]" captionStart="Figure 1" captionStartId="7.[524,589,673,695]" captionTargetBox="[498,1541,236,660]" captionTargetId="figure-144@7.[498,1299,234,645]" captionTargetPageId="7" captionText="Figure 1 Brachycephalus dacnis holotype (ZUEC-AMP 24982) adult male, SVL = 7.55 mm. (A) Dorsal and (B) ventral views. Full-size DOI: 10.7717/peerj.18265/fig-1" pageId="9" pageNumber="10">Fig. 1</figureCitation>
). The white spots overlying the upper jaw and ventral region can become pale cream; no
<emphasis id="B91E9A1E8B50672585EFFA42FAAEFA13" box="[1179,1362,1469,1495]" italics="true" pageId="9" pageNumber="10">linea masculina</emphasis>
is visible. The stripes on the arms and legs become more evident. The extremities of the fingers and toes become pale cream.
</paragraph>
<paragraph id="8BD5460C8B5067278086F9B4FCB1FDAF" blockId="9.[498,1542,1611,1916]" lastBlockId="11.[498,1542,234,619]" lastPageId="11" lastPageNumber="12" pageId="9" pageNumber="10">
<emphasis id="B91E9A1E8B5067258086F9B4FCD6F9A1" bold="true" box="[498,810,1611,1637]" pageId="9" pageNumber="10">Internal anatomy: organs</emphasis>
(ZUEC-AMP 25273; female)After cutting the skin, the transparent abdominal muscles were observed to have a slight dark-brown coloration (melanocytes). Beneath the musculature, the liver and small intestine were also observed with similar dark-brown pigmentation. However, it was more prominent in the intestine than in the ventral musculature and liver (
<figureCitation id="13515A898B50672582AEF915FBD0F8C0" box="[986,1068,1770,1796]" captionStart="Figure 4" captionStartId="10.[524,589,1841,1863]" captionTargetBox="[502,1298,236,1809]" captionTargetId="figure-8@10.[498,1299,234,1813]" captionTargetPageId="10" captionText="Figure 4 Ventral dissection of Brachycephalus dacnis (ZUEC-AMP 25273) with the highlighted structures. (A) Abdominal muscles, (B) Liver, (C) Small intestine; (D) Atrium, (E) Ventricle, (F) Conus arteriosus, (G) Truncus arteriosus, (H) Carotid artery, (I) Systemic artery, (J) Parathyroid gland, (K) Eggs, (L) Oviduct. Full-size DOI: 10.7717/peerj.18265/fig-4" pageId="9" pageNumber="10">Fig. 4A</figureCitation>
). After removing the abdominal muscles and opening the pericardium, we observed the heart to have melanocytes scattered throughout the atria, ventricle,
<emphasis id="B91E9A1E8B5067258217F8C4FBE2F890" box="[867,1054,1851,1876]" italics="true" pageId="9" pageNumber="10">conus arteriosus</emphasis>
, and
<emphasis id="B91E9A1E8B5067258516F8C4FACDF890" box="[1122,1329,1851,1876]" italics="true" pageId="9" pageNumber="10">truncus arteriosus</emphasis>
. After cutting the pectoral girdle in the medial region of the weakly mineralized procoracoid cartilages, we observed that the dark pigmentation extends from the cranial region of the heart to the carotid and systemic arteries. We did not see this pigmentation in the pulmocutaneous artery (
<figureCitation id="13515A898B526727833EFEC5FD2BFE90" box="[586,727,314,340]" captionStart="Figure 4" captionStartId="10.[524,589,1841,1863]" captionTargetBox="[502,1298,236,1809]" captionTargetId="figure-8@10.[498,1299,234,1813]" captionTargetPageId="10" captionText="Figure 4 Ventral dissection of Brachycephalus dacnis (ZUEC-AMP 25273) with the highlighted structures. (A) Abdominal muscles, (B) Liver, (C) Small intestine; (D) Atrium, (E) Ventricle, (F) Conus arteriosus, (G) Truncus arteriosus, (H) Carotid artery, (I) Systemic artery, (J) Parathyroid gland, (K) Eggs, (L) Oviduct. Full-size DOI: 10.7717/peerj.18265/fig-4" pageId="11" pageNumber="12">Figs. 4B, 4C</figureCitation>
). Aslight extension of this dark pigmentation occurs in the musculature lateral to the systemic and carotid arteries (
<figureCitation id="13515A898B526727850FFE9DFAF8FEB8" box="[1147,1284,354,380]" captionStart="Figure 4" captionStartId="10.[524,589,1841,1863]" captionTargetBox="[502,1298,236,1809]" captionTargetId="figure-8@10.[498,1299,234,1813]" captionTargetPageId="10" captionText="Figure 4 Ventral dissection of Brachycephalus dacnis (ZUEC-AMP 25273) with the highlighted structures. (A) Abdominal muscles, (B) Liver, (C) Small intestine; (D) Atrium, (E) Ventricle, (F) Conus arteriosus, (G) Truncus arteriosus, (H) Carotid artery, (I) Systemic artery, (J) Parathyroid gland, (K) Eggs, (L) Oviduct. Full-size DOI: 10.7717/peerj.18265/fig-4" pageId="11" pageNumber="12">Figs. 4B, 4C</figureCitation>
). Near the bifurcation of the internal and external jugular arteries and between the systemic and carotid arteries, there were two black, ovoid parathyroid glands on each side (
<figureCitation id="13515A898B52672785BFFE4EFADCFE08" box="[1227,1312,433,460]" captionStart="Figure 4" captionStartId="10.[524,589,1841,1863]" captionTargetBox="[502,1298,236,1809]" captionTargetId="figure-8@10.[498,1299,234,1813]" captionTargetPageId="10" captionText="Figure 4 Ventral dissection of Brachycephalus dacnis (ZUEC-AMP 25273) with the highlighted structures. (A) Abdominal muscles, (B) Liver, (C) Small intestine; (D) Atrium, (E) Ventricle, (F) Conus arteriosus, (G) Truncus arteriosus, (H) Carotid artery, (I) Systemic artery, (J) Parathyroid gland, (K) Eggs, (L) Oviduct. Full-size DOI: 10.7717/peerj.18265/fig-4" pageId="11" pageNumber="12">Fig. 4C</figureCitation>
). Less black pigmentation was observed in the thoracic region; however, a few small spots were observed close to the white eggs and in the oviduct. The oviduct was transparent and between the left and right oviduct two mature eggs were observed, and the oviducts joined near the pubis (
<figureCitation id="13515A898B52672783DAFDAEFCC1FDAF" box="[686,829,593,619]" captionStart="Figure 4" captionStartId="10.[524,589,1841,1863]" captionTargetBox="[502,1298,236,1809]" captionTargetId="figure-8@10.[498,1299,234,1813]" captionTargetPageId="10" captionText="Figure 4 Ventral dissection of Brachycephalus dacnis (ZUEC-AMP 25273) with the highlighted structures. (A) Abdominal muscles, (B) Liver, (C) Small intestine; (D) Atrium, (E) Ventricle, (F) Conus arteriosus, (G) Truncus arteriosus, (H) Carotid artery, (I) Systemic artery, (J) Parathyroid gland, (K) Eggs, (L) Oviduct. Full-size DOI: 10.7717/peerj.18265/fig-4" pageId="11" pageNumber="12">Figs. 4D, 4E</figureCitation>
).
</paragraph>
<caption id="DF1516848B5367268378F8CEFA17F85A" pageId="10" pageNumber="11" startId="10.[524,589,1841,1863]" targetBox="[502,1298,236,1809]" targetPageId="10" targetType="figure">
<paragraph id="8BD5460C8B5367268378F8CEFA17F85A" blockId="10.[524,1516,1841,1952]" pageId="10" pageNumber="11">
<emphasis id="B91E9A1E8B5367268378F8CEFD8FF8A0" bold="true" pageId="10" pageNumber="11">
Figure 4 Ventral dissection of
<taxonomicName id="4C6A3D8F8B5367268227F8CEFBC8F883" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[851,1076,1841,1863]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="10" pageNumber="11" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5367268227F8CEFBC8F883" bold="true" box="[851,1076,1841,1863]" italics="true" pageId="10" pageNumber="11">Brachycephalus dacnis</emphasis>
</taxonomicName>
(ZUEC-AMP 25273) with the highlighted structures.
</emphasis>
(A) Abdominal muscles, (B) Liver, (C) Small intestine; (D) Atrium, (E) Ventricle, (F)
<emphasis id="B91E9A1E8B53672684C4F8B0FD9BF846" italics="true" pageId="10" pageNumber="11">Conus arteriosus</emphasis>
, (G)
<emphasis id="B91E9A1E8B53672683ECF892FCB9F846" box="[664,837,1901,1922]" italics="true" pageId="10" pageNumber="11">Truncus arteriosus</emphasis>
, (H) Carotid artery, (I) Systemic artery, (J) Parathyroid gland, (K) Eggs, (L) Oviduct. Full-size DOI: 10.7717/peerj.18265/fig-4
</paragraph>
</caption>
<paragraph id="8BD5460C8B5267278086FD70FCC9FCB5" blockId="11.[498,1542,655,1917]" pageId="11" pageNumber="12">
<emphasis id="B91E9A1E8B5267278086FD70FCC0FD6D" bold="true" box="[498,828,655,681]" pageId="11" pageNumber="12">Internal anatomy: skeleton</emphasis>
(ZUEC-AMP 24978; undetermined sex). The skeleton of
<taxonomicName id="4C6A3D8F8B526727849DFD6FFDC7FD15" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="11" pageNumber="12" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B526727849DFD6FFDC7FD15" italics="true" pageId="11" pageNumber="12">B. dacnis</emphasis>
</taxonomicName>
is broadly similar to that of other small species in the genus, including by lacking osteoderms and having unornamented dermal skull bones that are not co-ossified to the skin. This individual (
<figureCitation id="13515A898B5267278383FCF8FCC4FCE6" box="[759,824,775,802]" captionStart="Figure 5" captionStartId="12.[116,181,1269,1291]" captionTargetBox="[93,1510,238,1238]" captionTargetId="figure-137@12.[90,1512,234,1241]" captionTargetPageId="12" captionText="Figure 5 Skeletal anatomy of Brachycephalus dacnis (ZUEC-AMP 24978; undetermined sex) based on CT-scans. (A) The entire skeleton, with insets of (B) the skull (in dorsal, ventral, and right lateral views), (C) right hand and (D) left foot (both in dorsal view), (E) vertebral column (in dorsal view), (F) pectoral girdle (in ventral view), and (G) pelvic girdle (in right lateral view). Abbreviations: c, carpus; cl, clavicle; cor, coracoid; dt, distal tarsals; ex, exoccipital; fp, frontoparietal; il, ilium; isc, ischium; m, maxilla; mdI, manual digit I; mdV, manual digit V; min, mineralized cartilage; n, nasal; op, operculum; pd I, pedal digit I; pd V, pedal digit V; pm, premaxilla; pr, prootic; ps, parasphenoid; pt, pterygoid; pub, pubis; qj, quadratojugal; sac, sacrum; sc, scapula; ses, sesamoid; sm, septomaxilla; sp, sphenethmoid; sq, squamosal; ur, urostyle; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-5" pageId="11" pageNumber="12">Fig. 5</figureCitation>
) is osteologically mature with ossified mesopodials and complete, ossified distal long bones; the specimen is missing the right leg, which was removed for DNA analysis.
</paragraph>
<paragraph id="8BD5460C8B5267278366FC81FCD7F8C1" blockId="11.[498,1542,655,1917]" pageId="11" pageNumber="12">
The skull is compact, slightly longer than wide, and lacking ornamented dermal roofing bones. The frontoparietals, sphenethmoids, prootics, exoccipitals, and parasphenoid are incompletely synostosed with each other and often have clear separations between the bones. There are well-defined and bony margins to both the optic fenestrae and prootic foramina. The premaxillae are broad, clearly separated from one another, and have teeth; each has a robust
<emphasis id="B91E9A1E8B52672783B2FBB8FCA4FBA4" box="[710,856,1094,1120]" italics="true" pageId="11" pageNumber="12">pars dentalis</emphasis>
, and a robust alary process that is approximately as tall as wide and widely separated from the nasal. In ventral view, the maxillae are nearly straight and bear odontoids on the anterior third. The quadratojugals are thin and angled dorsally, and do not articulate with the maxillae. The pterygoids are slender, each with a long, straight, and forked anterior ramus that approaches but does not articulate with the adjacent maxilla, a short, subtriangular posterior ramus adjacent to but not articulating with the ventral ramus of the squamosal, and a short, broad medial ramus that is well separated from the prootic. Distinct neopalatines are not observable and likely not synostosed to the sphenethmoid. Distinct, small c-shaped vomers are present bordering the anterior margins of the choanae. Tall, thin, and curved septomaxillae are present at the anterior margin of the nasal capsule. The parasphenoid is triradiate with a long, rectangular anterior ramus. The squamosals are robust, and each has a prominent zygomatic ramus that is expanded ventrally as a sheet-like flange along its anterior margin and a long, slender posterior ramus that is adjacent but not fused to the prootic. The portions of the prootic are not completely synostosed and the broad fenestra ovalis contains a small, irregularly shaped operculum; a stapes (or columella) is not present. The posteromedial processes of the hyoid are ossified and slender. The arytenoid cartilages are not obviously mineralized.
</paragraph>
<paragraph id="8BD5460C8B5267208366F8ECFA5AF947" blockId="11.[498,1542,655,1917]" lastBlockId="12.[498,1542,1521,1906]" lastPageId="12" lastPageNumber="13" pageId="11" pageNumber="12">There are eight distinct, procoelous, non-imbricating presacral vertebrae that arenot fused to one another. The atlas lacks transverse processes and cotyles that are widely separated, a distance greater than the posterior diameter of the atlas centrum. Transverse processes of presacral II are distinctly shorter than those of IIIVIII. The sacrum is procoelous with stout transverse processes. Asesamoid at the distal transverse process of the sacrum is present at its articulation with the ilium. The urostyle bears a tall dorsal ridge that decreases in height posteriorly along the anterior two-thirds of the urostyle.</paragraph>
<caption id="DF1516848B5567208100FB0AFA17FA7E" pageId="12" pageNumber="13" startId="12.[116,181,1269,1291]" targetBox="[93,1510,238,1238]" targetPageId="12" targetType="figure">
<paragraph id="8BD5460C8B5567208100FB0AFAE4FA59" blockId="12.[116,1515,1268,1466]" pageId="12" pageNumber="13">
<emphasis id="B91E9A1E8B5567208100FB0AFB2DFACF" bold="true" box="[116,1233,1268,1291]" pageId="12" pageNumber="13">
Figure 5 Skeletal anatomy of
<taxonomicName id="4C6A3D8F8B55672080E8FB0BFD86FACE" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[412,634,1268,1290]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="12" pageNumber="13" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B55672080E8FB0BFD86FACE" bold="true" box="[412,634,1268,1290]" italics="true" pageId="12" pageNumber="13">Brachycephalus dacnis</emphasis>
</taxonomicName>
(ZUEC-AMP 24978; undetermined sex) based on CT-scans.
</emphasis>
(A) The entire skeleton, with insets of (B) the skull (in dorsal, ventral, and right lateral views), (C) right hand and (D) left foot (both in dorsal view), (E) vertebral column (in dorsal view), (F) pectoral girdle (in ventral view), and (G) pelvic girdle (in right lateral view). Abbreviations: c, carpus; cl, clavicle; cor, coracoid; dt, distal tarsals; ex, exoccipital; fp, frontoparietal; il, ilium; isc, ischium; m, maxilla; mdI, manual digit I; mdV, manual digit V; min, mineralized cartilage; n, nasal; op, operculum; pd I, pedal digit I; pd V, pedal digit V; pm, premaxilla; pr, prootic; ps, parasphenoid; pt, pterygoid; pub, pubis; qj, quadratojugal; sac, sacrum; sc, scapula; ses, sesamoid; sm, septomaxilla; sp, sphenethmoid; sq, squamosal; ur, urostyle; v, vomer.
</paragraph>
<paragraph id="8BD5460C8B556720853FFA5BFA17FA7E" blockId="12.[116,1515,1268,1466]" box="[1099,1515,1444,1466]" pageId="12" pageNumber="13">Full-size DOI: 10.7717/peerj.18265/fig-5</paragraph>
</caption>
<paragraph id="8BD5460C8B5567208366F96EFC38F8B6" blockId="12.[498,1542,1521,1906]" pageId="12" pageNumber="13">The pectoral girdle is not heavily ossified. The coracoids are expanded medially but do not meet at the midline. The coracoids are thin and articulate at the midline with a small, irregularly shaped element that may represent the omosternum. The scapula is short with a well-developed anterior process and a prominent supraglenoid foramen. The epicoracoid and procoracoid cartilages are weakly mineralized and do not form solid elements synostosed to the coracoid and clavicle.</paragraph>
<paragraph id="8BD5460C8B5467218366FF15FA0DFE0F" blockId="13.[498,1542,234,898]" pageId="13" pageNumber="14">The pelvic girdle is a robust element composed of synostosed ilium, pubis, and ischium; the publis and ischium appear to be incompletely ossified. The circular acetabulum is incompletely ossified but has well-defined margins. The shaft of the ilium is stout and mostly straight in both lateral and dorsal views, and has both a well-developed dorsal protuberance and a weakly developed dorsal crest. There is a broad ventral acetabular expansion comprising both the ilium and ossified pubis, but it is incompletely ossified.</paragraph>
<paragraph id="8BD5460C8B5467218366FE26FB61FC46" blockId="13.[498,1542,234,898]" pageId="13" pageNumber="14">The forearm is somewhat shorter than the humerus. The distal carpals (Element Yand IIV) are fused. The radiale and ulnare are large and subequal in size. The phalangeal formula for the manus is 1231. There appears to be a single, minute ossified prepollex. There is one small palmar sesamoid. The tips of the terminal manual phalanges are arrow-shaped in digits II and III but blunt and globular in Iand IV. The tibiofibula and femur are similar in length, and there is a small ossified fabella sesamoid near their articulation. There are two large distal tarsals and an ossified tarsal sesamoid near their articulation with the tibiofibula. The phalangeal formula for the pes is 12342 and there is a single small ossified prehallux. Medially, there are two small plantar sesamoids, whereas laterally there are two large plantar sesamoids. The tips of the terminal pedal phalanges are arrow-shaped in digits IIV but blunt in V.</paragraph>
<paragraph id="8BD5460C8B5467218086FC59FB51FB43" blockId="13.[498,1542,934,1439]" pageId="13" pageNumber="14">
<emphasis id="B91E9A1E8B5467218086FC59FCE3FC04" bold="true" box="[498,799,934,960]" pageId="13" pageNumber="14">Calls and calling activity</emphasis>
The new species is vocally active throughout the year, with peaks of activity when the humidity was higher than 90% (&gt;
<specimenCount id="9D6C8D858B54672185C4FC31FAE4FC2C" box="[1200,1304,974,1001]" count="50" pageId="13" pageNumber="14" type="male">50 males</specimenCount>
calling). The calling activity was mostly detected during daytime, with a higher number of individuals calling between 6:00 and 11:00 AM, from 3:00 to 6:00 PM, or anytime of the day after heavy rains. From August to April, we also heard calls during the night. Males frequently ceased calling for a few minutes after detecting the researcher s approach.
</paragraph>
<paragraph id="8BD5460C8B5467218366FB6AFBABFA5B" blockId="13.[498,1542,934,1439]" pageId="13" pageNumber="14">
The advertisement call of
<taxonomicName id="4C6A3D8F8B5467218231FB69FC53FB6B" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[837,943,1173,1199]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5467218231FB69FC53FB6B" box="[837,943,1173,1199]" italics="true" pageId="13" pageNumber="14">B. dacnis</emphasis>
</taxonomicName>
(
<figureCitation id="13515A898B54672182B5FB6AFBE8FB6B" box="[961,1044,1173,1199]" captionStart="Figure 6" captionStartId="14.[116,181,1171,1193]" captionTargetBox="[90,1541,236,1158]" captionTargetId="figure-189@14.[90,1512,234,1143]" captionTargetPageId="14" captionText="Figure 6 Geographic distribution, advertisement calls and phylogeny. (A) Type localities of Brachycephalus dacnis (red star) and B. hermogenesi (black star), and additional distribution records of B. dacnis (red triangles, based on call recordings published in Bornschein et al., 2021). This map was created using the free and open-source software QGIS, utilizing publicly available base data sources. (B) Brachycephalus dacnis holotypes advertisement call (FNJV 51341), recorded at Projeto Dacnis private reserve (type locality), Ubatuba, São Paulo (left) and B. hermogenesi topotypes advertisement call (FNJV 51343), recorded at Picinguaba (type locality), Ubatuba, São Paulo (right). (C) Schematic representation of the Bayesian phylogenetic inference of Brachycephalus based on the mitochondrially encoded 16S rDNA fragment. Black dots indicate fully supported clades (posterior probability, PP =&gt; 99). White dots indicate PP&gt; 0.95. Node supports below pp = 0.95 are not shown. Complete tree in the supplement (Fig. S1). Full-size DOI: 10.7717/peerj.18265/fig-6" pageId="13" pageNumber="14">Fig. 6B</figureCitation>
) can be simple or complex, respectively containing one or two multipulsed (three to seven pulses) notes, emitted in mean intervals of 2.1 ±
<geoCoordinate id="EE5E20CB8B5467218322FB1AFD66FAC4" box="[598,666,1253,1280]" degrees="0.32" direction="south" orientation="latitude" pageId="13" pageNumber="14" precision="555" value="-0.32">0.32 s</geoCoordinate>
(1.672.69). Out of the 17 recordings made, only two presented calls with two notes. The mean note duration was 0.05 ±
<geoCoordinate id="EE5E20CB8B5467218557FAF2FB9BFAEC" box="[1059,1127,1293,1320]" degrees="0.01" direction="south" orientation="latitude" pageId="13" pageNumber="14" precision="555" value="-0.01">0.01 s</geoCoordinate>
(0.030.08), and the call with two notes could reach up to
<geoCoordinate id="EE5E20CB8B5467218264FACAFCA8FA8B" box="[784,852,1333,1359]" degrees="0.41" direction="south" orientation="latitude" pageId="13" pageNumber="14" precision="555" value="-0.41">0.41 s</geoCoordinate>
. The call frequency varied from 6.89 (minimum measured frequency) to 8.96 (maximum measured frequency) kHz, while the mean dominant frequency was 8.22 ± 0.1 kHz (8.018.44) (
<tableCitation id="C6E873B78B5467218284FA7AFBBBFA5B" box="[1008,1095,1413,1439]" captionStart="Table 2" captionStartId="15.[117,172,248,269]" captionTargetBox="[108,1523,319,1447]" captionTargetPageId="15" captionText="Table 2 Advertisement call traits described for three flea-toads species from Brachycephalus genus. Call traits present as mean ± standard deviation (range)." pageId="13" pageNumber="14">Table 2</tableCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C37015878B54673D8086FA3CFA8AFB8E" lastPageId="17" lastPageNumber="18" pageId="13" pageNumber="14" type="discussion">
<paragraph id="8BD5460C8B5467228086FA3CFA55F94A" blockId="13.[498,1542,1475,1900]" lastBlockId="14.[498,1539,1452,1678]" lastPageId="14" lastPageNumber="15" pageId="13" pageNumber="14">
<emphasis id="B91E9A1E8B5467218086FA3CFBEFFA19" bold="true" box="[498,1043,1475,1501]" pageId="13" pageNumber="14">Phylogenetic inference and genetic distances</emphasis>
Our phylogenetic analysis supported the distinction of
<taxonomicName id="4C6A3D8F8B54672183ECFA13FCFDF9C1" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[664,769,1515,1541]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B54672183ECFA13FCFDF9C1" box="[664,769,1515,1541]" italics="true" pageId="13" pageNumber="14">B. dacnis</emphasis>
</taxonomicName>
from all the other members of the genus. All the specimens are clustered within a strongly supported clade (posterior probability, PP = 1.0). The new species is recovered as the sister species of
<taxonomicName id="4C6A3D8F8B5467218298F9C4FB67F991" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[1004,1179,1595,1621]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B5467218298F9C4FB67F991" box="[1004,1179,1595,1621]" italics="true" pageId="13" pageNumber="14">B. hermogenesi</emphasis>
</taxonomicName>
, and this clade sister of
<taxonomicName id="4C6A3D8F8B54672184CFF9C4FDA7F9B8" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="sulfuratu">
<emphasis id="B91E9A1E8B54672184CFF9C4FDA7F9B8" italics="true" pageId="13" pageNumber="14">B. sulfuratu</emphasis>
</taxonomicName>
s +
<taxonomicName id="4C6A3D8F8B54672183FCF99CFD0DF9B8" authorityName="Pombal, Wistuba &amp; Bornschein" authorityYear="1998" box="[648,753,1635,1660]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="pernix">
<emphasis id="B91E9A1E8B54672183FCF99CFD0DF9B8" box="[648,753,1635,1660]" italics="true" pageId="13" pageNumber="14">B. pernix</emphasis>
</taxonomicName>
species group (
<figureCitation id="13515A898B54672182DFF99DFBD7F9B9" box="[939,1067,1634,1661]" captionStart="Figure 6" captionStartId="14.[116,181,1171,1193]" captionTargetBox="[90,1541,236,1158]" captionTargetId="figure-189@14.[90,1512,234,1143]" captionTargetPageId="14" captionText="Figure 6 Geographic distribution, advertisement calls and phylogeny. (A) Type localities of Brachycephalus dacnis (red star) and B. hermogenesi (black star), and additional distribution records of B. dacnis (red triangles, based on call recordings published in Bornschein et al., 2021). This map was created using the free and open-source software QGIS, utilizing publicly available base data sources. (B) Brachycephalus dacnis holotypes advertisement call (FNJV 51341), recorded at Projeto Dacnis private reserve (type locality), Ubatuba, São Paulo (left) and B. hermogenesi topotypes advertisement call (FNJV 51343), recorded at Picinguaba (type locality), Ubatuba, São Paulo (right). (C) Schematic representation of the Bayesian phylogenetic inference of Brachycephalus based on the mitochondrially encoded 16S rDNA fragment. Black dots indicate fully supported clades (posterior probability, PP =&gt; 99). White dots indicate PP&gt; 0.95. Node supports below pp = 0.95 are not shown. Complete tree in the supplement (Fig. S1). Full-size DOI: 10.7717/peerj.18265/fig-6" pageId="13" pageNumber="14">Figs. 6C; S</figureCitation>
<figureCitation id="13515A898B546721855FF99CFBCAF9B9" box="[1067,1078,1635,1661]" captionStart="Figure 1" captionStartId="7.[524,589,673,695]" captionTargetBox="[498,1541,236,660]" captionTargetId="figure-144@7.[498,1299,234,645]" captionTargetPageId="7" captionText="Figure 1 Brachycephalus dacnis holotype (ZUEC-AMP 24982) adult male, SVL = 7.55 mm. (A) Dorsal and (B) ventral views. Full-size DOI: 10.7717/peerj.18265/fig-1" pageId="13" pageNumber="14">1</figureCitation>
, Table S4). The phylogenetic inference recovered
<taxonomicName id="4C6A3D8F8B5467218318F974FD36F960" authorityName="Napoli, Caramaschi, Cruz &amp; Dias" authorityYear="2011" box="[620,714,1674,1700]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="pulex">
<emphasis id="B91E9A1E8B5467218318F974FD36F960" box="[620,714,1674,1700]" italics="true" pageId="13" pageNumber="14">B. pulex</emphasis>
</taxonomicName>
sister to two diverse clades, one containing ((
<taxonomicName id="4C6A3D8F8B546721859DF974FAC5F960" authorityName="Almeida-Silva, Silva-Soares, Rodrigues &amp; Verdade" authorityYear="2021" box="[1257,1337,1675,1700]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="puri">
<emphasis id="B91E9A1E8B546721859DF974FAC5F960" box="[1257,1337,1675,1700]" italics="true" pageId="13" pageNumber="14">B. puri</emphasis>
</taxonomicName>
+ two putative distinct species) + (
<taxonomicName id="4C6A3D8F8B54672183ADF94CFC91F908" baseAuthorityName="Izecksohn" baseAuthorityYear="1971" box="[729,877,1714,1740]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="didactylus">
<emphasis id="B91E9A1E8B54672183ADF94CFC91F908" box="[729,877,1714,1740]" italics="true" pageId="13" pageNumber="14">B. didactylus</emphasis>
</taxonomicName>
+ (
<taxonomicName id="4C6A3D8F8B54672182EDF94CFBE5F908" box="[921,1049,1714,1740]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="clarissae">
<emphasis id="B91E9A1E8B54672182EDF94CFBE5F908" box="[921,1049,1714,1740]" italics="true" pageId="13" pageNumber="14">B. clarissae</emphasis>
</taxonomicName>
+ (
<taxonomicName id="4C6A3D8F8B5467218532F94CFB21F908" authorityName="Pombal" authorityYear="2001" box="[1094,1245,1714,1740]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="vertebralis">
<emphasis id="B91E9A1E8B5467218532F94CFB21F908" box="[1094,1245,1714,1740]" italics="true" pageId="13" pageNumber="14">B. vertebralis</emphasis>
</taxonomicName>
species group +
<taxonomicName id="4C6A3D8F8B54672184D2F94CFD91F930" baseAuthorityName="Spix" baseAuthorityYear="1824" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="B91E9A1E8B54672184D2F94CFD91F930" italics="true" pageId="13" pageNumber="14">B. ephippium</emphasis>
</taxonomicName>
species group)) and the other containing ((
<taxonomicName id="4C6A3D8F8B5467218506F924FADCF930" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[1138,1312,1754,1780]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B5467218506F924FADCF930" box="[1138,1312,1754,1780]" italics="true" pageId="13" pageNumber="14">B. hermogenesi</emphasis>
</taxonomicName>
+
<taxonomicName id="4C6A3D8F8B5467218437F924FA50F930" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[1347,1452,1754,1780]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5467218437F924FA50F930" box="[1347,1452,1754,1780]" italics="true" pageId="13" pageNumber="14">B. dacnis</emphasis>
</taxonomicName>
) + (
<taxonomicName id="4C6A3D8F8B5467218496F924FD99F8D8" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B5467218496F924FD99F8D8" italics="true" pageId="13" pageNumber="14">B. sulfuratus</emphasis>
</taxonomicName>
+
<taxonomicName id="4C6A3D8F8B54672183F0F8FCFD10F8D8" authorityName="Pombal, Wistuba &amp; Bornschein" authorityYear="1998" box="[644,748,1795,1820]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="pernix">
<emphasis id="B91E9A1E8B54672183F0F8FCFD10F8D8" box="[644,748,1795,1820]" italics="true" pageId="13" pageNumber="14">B. pernix</emphasis>
</taxonomicName>
species group) (
<figureCitation id="13515A898B54672182DFF8FDFBDBF8D8" box="[939,1063,1794,1820]" captionStart="Figure 6" captionStartId="14.[116,181,1171,1193]" captionTargetBox="[90,1541,236,1158]" captionTargetId="figure-189@14.[90,1512,234,1143]" captionTargetPageId="14" captionText="Figure 6 Geographic distribution, advertisement calls and phylogeny. (A) Type localities of Brachycephalus dacnis (red star) and B. hermogenesi (black star), and additional distribution records of B. dacnis (red triangles, based on call recordings published in Bornschein et al., 2021). This map was created using the free and open-source software QGIS, utilizing publicly available base data sources. (B) Brachycephalus dacnis holotypes advertisement call (FNJV 51341), recorded at Projeto Dacnis private reserve (type locality), Ubatuba, São Paulo (left) and B. hermogenesi topotypes advertisement call (FNJV 51343), recorded at Picinguaba (type locality), Ubatuba, São Paulo (right). (C) Schematic representation of the Bayesian phylogenetic inference of Brachycephalus based on the mitochondrially encoded 16S rDNA fragment. Black dots indicate fully supported clades (posterior probability, PP =&gt; 99). White dots indicate PP&gt; 0.95. Node supports below pp = 0.95 are not shown. Complete tree in the supplement (Fig. S1). Full-size DOI: 10.7717/peerj.18265/fig-6" pageId="13" pageNumber="14">Figs. 6C, S</figureCitation>
<figureCitation id="13515A898B5467218553F8FDFBCEF8D8" box="[1063,1074,1794,1820]" captionStart="Figure 1" captionStartId="7.[524,589,673,695]" captionTargetBox="[498,1541,236,660]" captionTargetId="figure-144@7.[498,1299,234,645]" captionTargetPageId="7" captionText="Figure 1 Brachycephalus dacnis holotype (ZUEC-AMP 24982) adult male, SVL = 7.55 mm. (A) Dorsal and (B) ventral views. Full-size DOI: 10.7717/peerj.18265/fig-1" pageId="13" pageNumber="14">1</figureCitation>
, Table S4). The 16S uncorrected genetic distance between the new species and
<taxonomicName id="4C6A3D8F8B54672182C2F8D5FB98F880" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[950,1124,1834,1860]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="13" pageNumber="14" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B54672182C2F8D5FB98F880" box="[950,1124,1834,1860]" italics="true" pageId="13" pageNumber="14">B. hermogenesi</emphasis>
</taxonomicName>
ranged from 6.8% to 7.8% (mean 7.1%;
<tableCitation id="C6E873B78B546721834EF8AEFD63F8A8" box="[570,671,1873,1900]" captionStart="Table 1" captionStartId="7.[525,580,794,815]" captionTargetBox="[516,1524,894,1446]" captionTargetPageId="7" captionText="Table 1 Measurements of the holotype and adult paratypes (one adult that was dissected for internal anatomy was not measured) of Brachycephalus dacnis. Values presented as mean ± standard deviation (minimummaximum) in millimeters." pageId="13" pageNumber="14">Table S1</tableCitation>
). These distances are higher than distances found between species of
<emphasis id="B91E9A1E8B54672184A3F8ADFA13F8AF" box="[1495,1519,1874,1899]" italics="true" pageId="13" pageNumber="14">B.</emphasis>
<taxonomicName id="4C6A3D8F8B5767228086FA52FD91FA03" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[498,621,1453,1479]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="B91E9A1E8B5767228086FA52FD91FA03" box="[498,621,1453,1479]" italics="true" pageId="14" pageNumber="15">ephippium</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6A3D8F8B57672283DEFA52FCBDFA03" authorityName="Pombal" authorityYear="2001" box="[682,833,1453,1479]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="vertebralis">
<emphasis id="B91E9A1E8B57672283DEFA52FCBDFA03" box="[682,833,1453,1479]" italics="true" pageId="14" pageNumber="15">B. vertebralis</emphasis>
</taxonomicName>
species groups (3.2% to 5.9%,
<tableCitation id="C6E873B78B57672285DAFA52FAE8FA03" box="[1198,1300,1453,1479]" captionStart="Table 1" captionStartId="7.[525,580,794,815]" captionTargetBox="[516,1524,894,1446]" captionTargetPageId="7" captionText="Table 1 Measurements of the holotype and adult paratypes (one adult that was dissected for internal anatomy was not measured) of Brachycephalus dacnis. Values presented as mean ± standard deviation (minimummaximum) in millimeters." pageId="14" pageNumber="15">Table S1</tableCitation>
). The relationships between species within each of the species groups containing the bright colored and bufoniform species (
<taxonomicName id="4C6A3D8F8B5767228390FA02FC83F9D2" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[740,895,1532,1558]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="B91E9A1E8B5767228390FA02FC83F9D2" box="[740,895,1532,1558]" italics="true" pageId="14" pageNumber="15">B. ephippium</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B57672282FAFA02FBD9F9D2" authorityName="Pombal" authorityYear="2001" box="[910,1061,1532,1558]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="vertebralis">
<emphasis id="B91E9A1E8B57672282FAFA02FBD9F9D2" box="[910,1061,1532,1558]" italics="true" pageId="14" pageNumber="15">B. vertebralis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6A3D8F8B5767228517FA02FB30F9D2" authorityName="Pombal, Wistuba &amp; Bornschein" authorityYear="1998" box="[1123,1228,1533,1558]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="pernix">
<emphasis id="B91E9A1E8B5767228517FA02FB30F9D2" box="[1123,1228,1533,1558]" italics="true" pageId="14" pageNumber="15">B. pernix</emphasis>
</taxonomicName>
species groups) are in general low supported (
<figureCitation id="13515A898B576722827EF9DBFCA6F9FA" box="[778,858,1572,1598]" captionStart="Figure 1" captionStartId="7.[524,589,673,695]" captionTargetBox="[498,1541,236,660]" captionTargetId="figure-144@7.[498,1299,234,645]" captionTargetPageId="7" captionText="Figure 1 Brachycephalus dacnis holotype (ZUEC-AMP 24982) adult male, SVL = 7.55 mm. (A) Dorsal and (B) ventral views. Full-size DOI: 10.7717/peerj.18265/fig-1" pageId="14" pageNumber="15">Fig. S1</figureCitation>
). Furthermore, some species were not recovered as monophyletic (
<emphasis id="B91E9A1E8B57672283D2F9B2FD3CF9A2" box="[678,704,1613,1638]" italics="true" pageId="14" pageNumber="15">i.e</emphasis>
.,
<taxonomicName id="4C6A3D8F8B57672283A2F9B2FCB1F9A2" authorityName="Alves, Sawaya, Reis &amp; Haddad" authorityYear="2009" box="[726,845,1613,1638]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="pitanga">
<emphasis id="B91E9A1E8B57672283A2F9B2FCB1F9A2" box="[726,845,1613,1638]" italics="true" pageId="14" pageNumber="15">B. pitanga</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6A3D8F8B57672282FEF9B2FBF8F9A2" authorityName="Clemente-Carvalho, Giaretta, Condez, Haddad &amp; Reis" authorityYear="2012" box="[906,1028,1613,1638]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="guarani">
<emphasis id="B91E9A1E8B57672282FEF9B2FBF8F9A2" box="[906,1028,1613,1638]" italics="true" pageId="14" pageNumber="15">B. guarani</emphasis>
</taxonomicName>
;
<taxonomicName id="4C6A3D8F8B5767228560F9B2FB2AF9A2" authorityName="Bornschein, Morato, Firkowski, Ribeiro &amp; Pie" authorityYear="2015" box="[1044,1238,1613,1638]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="mariaeterezae">
<emphasis id="B91E9A1E8B5767228560F9B2FB2AF9A2" box="[1044,1238,1613,1638]" italics="true" pageId="14" pageNumber="15">B. mariaeterezae</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6A3D8F8B5767228467F9B2FA66F9A2" authorityName="Bornschein, Morato, Firkowski, Ribeiro &amp; Pie" authorityYear="2015" box="[1299,1434,1612,1638]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="olivaceus">
<emphasis id="B91E9A1E8B5767228467F9B2FA66F9A2" box="[1299,1434,1612,1638]" italics="true" pageId="14" pageNumber="15">B. olivaceus</emphasis>
</taxonomicName>
;
<figureCitation id="13515A898B57672284DEF9B3FA05F9A2" box="[1450,1529,1612,1638]" captionStart="Figure 1" captionStartId="7.[524,589,673,695]" captionTargetBox="[498,1541,236,660]" captionTargetId="figure-144@7.[498,1299,234,645]" captionTargetPageId="7" captionText="Figure 1 Brachycephalus dacnis holotype (ZUEC-AMP 24982) adult male, SVL = 7.55 mm. (A) Dorsal and (B) ventral views. Full-size DOI: 10.7717/peerj.18265/fig-1" pageId="14" pageNumber="15">Fig. S1</figureCitation>
) and the genetic distances estimated were extremely low, less than 1% (
<tableCitation id="C6E873B78B5767228447F98BFA65F94A" box="[1331,1433,1652,1678]" captionStart="Table 1" captionStartId="7.[525,580,794,815]" captionTargetBox="[516,1524,894,1446]" captionTargetPageId="7" captionText="Table 1 Measurements of the holotype and adult paratypes (one adult that was dissected for internal anatomy was not measured) of Brachycephalus dacnis. Values presented as mean ± standard deviation (minimummaximum) in millimeters." pageId="14" pageNumber="15">Table S1</tableCitation>
).
</paragraph>
<caption id="DF1516848B5767228100FB6CFA17FAB2" pageId="14" pageNumber="15" startId="14.[116,181,1171,1193]" targetBox="[90,1541,236,1158]" targetPageId="14" targetType="figure">
<paragraph id="8BD5460C8B5767228100FB6CFA17FAB2" blockId="14.[116,1515,1171,1398]" pageId="14" pageNumber="15">
<emphasis id="B91E9A1E8B5767228100FB6CFCD4FB6C" bold="true" box="[116,808,1171,1193]" pageId="14" pageNumber="15">Figure 6 Geographicdistribution, advertisementcallsandphylogeny.</emphasis>
(A) Typelocalitiesof
<taxonomicName id="4C6A3D8F8B5767228288FB6CFB2CFB6C" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[1020,1232,1171,1192]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5767228288FB6CFB2CFB6C" box="[1020,1232,1171,1192]" italics="true" pageId="14" pageNumber="15">Brachycephalusdacnis</emphasis>
</taxonomicName>
(redstar) and
<taxonomicName id="4C6A3D8F8B576722842AFB6BFA17FB6C" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[1374,1515,1171,1193]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B576722842AFB6BFA17FB6C" box="[1374,1515,1171,1193]" italics="true" pageId="14" pageNumber="15">B. hermogenesi</emphasis>
</taxonomicName>
(black star), and additional distribution records of
<taxonomicName id="4C6A3D8F8B576722832CFB4EFD52FB01" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[600,686,1200,1222]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B576722832CFB4EFD52FB01" box="[600,686,1200,1222]" italics="true" pageId="14" pageNumber="15">B. dacnis</emphasis>
</taxonomicName>
(red triangles, based on call recordings published in
<bibRefCitation id="EFFB3BFD8B57672285D3FB4FFA81FB01" author="Bornschein MR &amp; Ribeiro LF &amp; Teixeira L &amp; Belmonte-Lopes R &amp; Moraes LA &amp; de Correa L &amp; Mauricio GN &amp; Pie MR" box="[1191,1405,1200,1222]" journalOrPublisher="PeerJ" pageId="14" pageNumber="15" part="9" publicationUrl="10.7717/peerj.10983" refId="ref12763" refString="Bornschein MR, Ribeiro LF, Teixeira L, Belmonte-Lopes R, Moraes LA, de Correa L, Mauricio GN, Nadaline Ι, Pie MR. 2021. A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae). PeerJ 9: e 10983 DOI 10.7717 / peerj. 10983." title="A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae)" type="journal volume" year="2021">
<emphasis id="B91E9A1E8B57672285D3FB4FFA81FB01" box="[1191,1405,1200,1222]" italics="true" pageId="14" pageNumber="15">Bornschein et al., 2021</emphasis>
</bibRefCitation>
). This map was created using the free and open-source software QGIS, utilizing publicly available base data sources. (B)
<taxonomicName id="4C6A3D8F8B57672285DFFB31FA7DFB27" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[1195,1409,1230,1251]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B57672285DFFB31FA7DFB27" box="[1195,1409,1230,1251]" italics="true" pageId="14" pageNumber="15">Brachycephalus dacnis</emphasis>
</taxonomicName>
holotype s advertisement call (FNJV 51341), recorded at Projeto Dacnis private reserve (type locality), Ubatuba, São Paulo (left) and
<taxonomicName id="4C6A3D8F8B5767228582FB14FA7FFAC4" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[1270,1411,1259,1280]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B5767228582FB14FA7FFAC4" box="[1270,1411,1259,1280]" italics="true" pageId="14" pageNumber="15">B. hermogenesi</emphasis>
</taxonomicName>
topotype s advertisement call (FNJV 51343), recorded at Picinguaba (type locality), Ubatuba, São Paulo (right). (C) Schematic representation of the Bayesian phylogenetic inference of
<taxonomicName id="4C6A3D8F8B576722801BFADAFDFDFAFE" baseAuthorityName="Trueb &amp; Alberch" baseAuthorityYear="1985" box="[367,513,1317,1338]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="genus">
<emphasis id="B91E9A1E8B576722801BFADAFDFDFAFE" box="[367,513,1317,1338]" italics="true" pageId="14" pageNumber="15">Brachycephalus</emphasis>
</taxonomicName>
based on the mitochondrially encoded 16S rDNA fragment. Black dots indicate fully supported clades (posterior probability, PP =&gt; 99). Whitedotsindicate PP&gt; 0.95. Nodesupportsbelowpp = 0.95 arenot shown. Completetreeinthesupplement (Fig. S1). Full-size DOI: 10.7717/peerj.18265/fig-6
</paragraph>
</caption>
<paragraph id="8BD5460C8B57673C8086F94DFA09F918" blockId="14.[498,1542,1714,1900]" lastBlockId="16.[498,1542,932,1915]" lastPageId="16" lastPageNumber="17" pageId="14" pageNumber="15">
<emphasis id="B91E9A1E8B5767228086F94DFC99F908" bold="true" box="[498,869,1714,1741]" pageId="14" pageNumber="15">Comparison with other species</emphasis>
<taxonomicName id="4C6A3D8F8B5767228201F94DFB8AF908" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[885,1142,1714,1740]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5767228201F94DFB8AF908" box="[885,1142,1714,1740]" italics="true" pageId="14" pageNumber="15">Brachycephalus dacnis</emphasis>
</taxonomicName>
is distinguished from all species of the
<taxonomicName id="4C6A3D8F8B5767228354F924FD47F930" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[544,699,1754,1780]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="B91E9A1E8B5767228354F924FD47F930" box="[544,699,1754,1780]" italics="true" pageId="14" pageNumber="15">B. ephippium</emphasis>
</taxonomicName>
(
<taxonomicName id="4C6A3D8F8B57672283B9F924FC9EF930" authorityName="Miranda-Ribeiro" authorityYear="1920" box="[717,866,1754,1780]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="atelopoide">
<emphasis id="B91E9A1E8B57672283B9F924FC9EF930" box="[717,866,1754,1780]" italics="true" pageId="14" pageNumber="15">B. atelopoide</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5767228206F924FC0FF930" authorityName="Guimaraes, Luz, Rocha &amp; Feio" authorityYear="2017" box="[882,1011,1754,1780]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="darkside">
<emphasis id="B91E9A1E8B5767228206F924FC0FF930" box="[882,1011,1754,1780]" italics="true" pageId="14" pageNumber="15">B. darkside</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5767228576F924FB60F930" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[1026,1180,1754,1780]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="B91E9A1E8B5767228576F924FB60F930" box="[1026,1180,1754,1780]" italics="true" pageId="14" pageNumber="15">B. ephippium</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B57672285D8F924FABEF930" authorityName="Miranda-Ribeiro" authorityYear="1920" box="[1196,1346,1754,1780]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="garbeanus">
<emphasis id="B91E9A1E8B57672285D8F924FABEF930" box="[1196,1346,1754,1780]" italics="true" pageId="14" pageNumber="15">B. garbeanus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5767228425F924FA35F930" box="[1361,1481,1754,1780]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="ibitinga">
<emphasis id="B91E9A1E8B5767228425F924FA35F930" box="[1361,1481,1754,1780]" italics="true" pageId="14" pageNumber="15">B. ibitinga</emphasis>
</taxonomicName>
,
<emphasis id="B91E9A1E8B57672284ACF924FA0CF930" box="[1496,1520,1755,1780]" italics="true" pageId="14" pageNumber="15">B.</emphasis>
<emphasis id="B91E9A1E8B5767228086F8FCFD74F8D8" box="[498,648,1795,1820]" italics="true" pageId="14" pageNumber="15">margaritatus</emphasis>
, and
<taxonomicName id="4C6A3D8F8B57672283B8F8FCFC8EF8D8" box="[716,882,1794,1820]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="rotenbergae">
<emphasis id="B91E9A1E8B57672283B8F8FCFC8EF8D8" box="[716,882,1794,1820]" italics="true" pageId="14" pageNumber="15">B. rotenbergae</emphasis>
</taxonomicName>
),
<taxonomicName id="4C6A3D8F8B57672282F8F8FCFBDFF8D8" authorityName="Pombal" authorityYear="2001" box="[908,1059,1794,1820]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="vertebralis">
<emphasis id="B91E9A1E8B57672282F8F8FCFBDFF8D8" box="[908,1059,1794,1820]" italics="true" pageId="14" pageNumber="15">B. vertebralis</emphasis>
</taxonomicName>
(
<taxonomicName id="4C6A3D8F8B5767228541F8FCFB63F8D8" authorityName="Pombal &amp; Gasparini" authorityYear="2006" box="[1077,1183,1794,1820]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="alipioi">
<emphasis id="B91E9A1E8B5767228541F8FCFB63F8D8" box="[1077,1183,1794,1820]" italics="true" pageId="14" pageNumber="15">B. alipioi</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B57672285DAF8FCFAB6F8D8" authorityName="Miranda-Ribeiro" authorityYear="1920" box="[1198,1354,1794,1820]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="bufonoides">
<emphasis id="B91E9A1E8B57672285DAF8FCFAB6F8D8" box="[1198,1354,1794,1820]" italics="true" pageId="14" pageNumber="15">B. bufonoides</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B576722842DF8FCFA3BF8D8" authorityName="Condez, Clemente-Carvalho, Haddad &amp; Reis" authorityYear="2014" box="[1369,1479,1795,1820]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="crispus">
<emphasis id="B91E9A1E8B576722842DF8FCFA3BF8D8" box="[1369,1479,1795,1820]" italics="true" pageId="14" pageNumber="15">B. crispus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B57672284A3F8FCFA12F8D8" box="[1495,1518,1795,1820]" pageId="14" pageNumber="15">
<emphasis id="B91E9A1E8B57672284A3F8FCFA12F8D8" box="[1495,1518,1795,1820]" italics="true" pageId="14" pageNumber="15">B.</emphasis>
</taxonomicName>
<emphasis id="B91E9A1E8B5767228086F8D4FDB0F880" box="[498,588,1835,1860]" italics="true" pageId="14" pageNumber="15">guarani</emphasis>
,
<emphasis id="B91E9A1E8B576722832DF8D4FC7CF880" box="[601,896,1834,1860]" italics="true" pageId="14" pageNumber="15">
<taxonomicName id="4C6A3D8F8B576722832DF8D4FD19F880" box="[601,741,1834,1860]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="herculeus">B. herculeus</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B576722839BF8D4FC7CF880" authorityName="Miranda-Ribeiro" authorityYear="1920" box="[751,896,1834,1860]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="nodoterga">B. nodoterga</taxonomicName>
</emphasis>
,
<taxonomicName id="4C6A3D8F8B57672282F8F8D4FBFCF880" authorityName="Alves, Sawaya, Reis &amp; Haddad" authorityYear="2009" box="[908,1024,1835,1860]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="pitanga">
<emphasis id="B91E9A1E8B57672282F8F8D4FBFCF880" box="[908,1024,1835,1860]" italics="true" pageId="14" pageNumber="15">B. pitanga</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5767228579F8D4FBA0F880" authorityName="Haddad, Alves, Clemente-Carvalho &amp; Reis" authorityYear="2010" box="[1037,1116,1834,1860]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="toby">
<emphasis id="B91E9A1E8B5767228579F8D4FBA0F880" box="[1037,1116,1834,1860]" italics="true" pageId="14" pageNumber="15">B. toby</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C6A3D8F8B57672285EEF8D4FAD3F880" authorityName="Pombal" authorityYear="2001" box="[1178,1327,1834,1860]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="vertebralis">
<emphasis id="B91E9A1E8B57672285EEF8D4FAD3F880" box="[1178,1327,1834,1860]" italics="true" pageId="14" pageNumber="15">B. vertebralis</emphasis>
</taxonomicName>
), and
<taxonomicName id="4C6A3D8F8B5767228403F8D4FA22F880" authorityName="Pombal, Wistuba &amp; Bornschein" authorityYear="1998" box="[1399,1502,1835,1860]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="pernix">
<emphasis id="B91E9A1E8B5767228403F8D4FA22F880" box="[1399,1502,1835,1860]" italics="true" pageId="14" pageNumber="15">B. pernix</emphasis>
</taxonomicName>
(
<taxonomicName id="4C6A3D8F8B576722849AF8D4FDBBF8A8" authorityName="Monteiro, Condez, Garcia, Comitti, Amaral &amp; Haddad" authorityYear="2018" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="actaeus">
<emphasis id="B91E9A1E8B576722849AF8D4FDBBF8A8" italics="true" pageId="14" pageNumber="15">B. actaeus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5767228320F8ACFD00F8A8" authorityName="Bornschein, Ribeiro, Blackburn, Stanley &amp; Pie" authorityYear="2016" box="[596,764,1874,1900]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="albolineatus">
<emphasis id="B91E9A1E8B5767228320F8ACFD00F8A8" box="[596,764,1874,1900]" italics="true" pageId="14" pageNumber="15">B. albolineatus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B576722827DF8ACFC47F8A8" authorityName="Ribeiro, Firkowski, Bornschein &amp; Pie" authorityYear="2015" box="[777,955,1875,1900]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="auroguttatus">
<emphasis id="B91E9A1E8B576722827DF8ACFC47F8A8" box="[777,955,1875,1900]" italics="true" pageId="14" pageNumber="15">B. auroguttatus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B57672282B3F8ACFBB6F8A8" authorityName="Pie, Bornschein, Firkowski, Belmonte-Lopes &amp; Ribeiro" authorityYear="2015" box="[967,1098,1874,1900]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="boticario">
<emphasis id="B91E9A1E8B57672282B3F8ACFBB6F8A8" box="[967,1098,1874,1900]" italics="true" pageId="14" pageNumber="15">B. boticario</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5767228522F8ACFB1CF8A8" authorityName="Ribeiro, Alves, Haddad &amp; Reis" authorityYear="2005" box="[1110,1248,1874,1900]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="brunneus">
<emphasis id="B91E9A1E8B5767228522F8ACFB1CF8A8" box="[1110,1248,1874,1900]" italics="true" pageId="14" pageNumber="15">B. brunneus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5767228598F8ACFA8EF8A8" authorityName="Ribeiro, Blackburn, Stanley, Pie &amp; Bornschein" authorityYear="2017" box="[1260,1394,1874,1900]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="coloratus">
<emphasis id="B91E9A1E8B5767228598F8ACFA8EF8A8" box="[1260,1394,1874,1900]" italics="true" pageId="14" pageNumber="15">B. coloratus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B576722840AF8ACFA03F8A8" authorityName="Ribeiro, Blackburn, Stanley, Pie &amp; Bornschein" authorityYear="2017" box="[1406,1535,1875,1900]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="14" pageNumber="15" phylum="Chordata" rank="species" species="curupira">
<emphasis id="B91E9A1E8B576722840AF8ACFA03F8A8" box="[1406,1535,1875,1900]" italics="true" pageId="14" pageNumber="15">B. curupira</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5667238086F9F4FD7BF9E0" authorityName="Alves, Ribeiro, Haddad &amp; Reis" authorityYear="2006" box="[498,647,1546,1572]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="ferruginus">
<emphasis id="B91E9A1E8B5667238086F9F4FD7BF9E0" box="[498,647,1546,1572]" italics="true" pageId="15" pageNumber="16">B. ferruginus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B56672383E1F9F4FCB4F9E0" authorityName="Pie, Bornschein, Firkowski, Belmonte-Lopes &amp; Ribeiro" authorityYear="2015" box="[661,840,1546,1572]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="fuscolineatus">
<emphasis id="B91E9A1E8B56672383E1F9F4FCB4F9E0" box="[661,840,1546,1572]" italics="true" pageId="15" pageNumber="16">B. fuscolineatus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5667238222F9F4FC17F9E0" authorityName="Ribeiro, Alves, Haddad &amp; Reis" authorityYear="2005" box="[854,1003,1546,1572]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="izecksohni">
<emphasis id="B91E9A1E8B5667238222F9F4FC17F9E0" box="[854,1003,1546,1572]" italics="true" pageId="15" pageNumber="16">B. izecksohni</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B566723828EF9F4FB7AF9E0" authorityName="Ribeiro, Firkowski &amp; Pie" authorityYear="2015" box="[1018,1158,1546,1572]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="leopardos">
<emphasis id="B91E9A1E8B566723828EF9F4FB7AF9E0" box="[1018,1158,1546,1572]" italics="true" pageId="15" pageNumber="16">B. leopardos</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B56672385E0F9F4FAA9F9E0" authorityName="Bornschein, Morato, Firkowski, Ribeiro &amp; Pie" authorityYear="2015" box="[1172,1365,1547,1572]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="mariaeterezae">
<emphasis id="B91E9A1E8B56672385E0F9F4FAA9F9E0" box="[1172,1365,1547,1572]" italics="true" pageId="15" pageNumber="16">B. mariaeterezae</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5667238417F9F4F9FCF9E0" authorityName="Pie, Ribeiro, Confetti, Nadaline &amp; Bornschein" authorityYear="2018" box="[1379,1536,1547,1572]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="mirissimus">
<emphasis id="B91E9A1E8B5667238417F9F4F9FCF9E0" box="[1379,1536,1547,1572]" italics="true" pageId="15" pageNumber="16">B. mirissimus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5667238086F9CCFD85F988" authorityName="Bornschein, Morato, Firkowski, Ribeiro &amp; Pie" authorityYear="2015" box="[498,633,1586,1612]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="olivaceus">
<emphasis id="B91E9A1E8B5667238086F9CCFD85F988" box="[498,633,1586,1612]" italics="true" pageId="15" pageNumber="16">B. olivaceus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B56672383FDF9CCFD0EF988" authorityName="Pombal, Wistuba &amp; Bornschein" authorityYear="1998" box="[649,754,1587,1612]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="pernix">
<emphasis id="B91E9A1E8B56672383FDF9CCFD0EF988" box="[649,754,1587,1612]" italics="true" pageId="15" pageNumber="16">B. pernix</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B5667238276F9CCFC7CF988" authorityName="Alves, Ribeiro, Haddad &amp; Reis" authorityYear="2006" box="[770,896,1586,1612]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="pombali">
<emphasis id="B91E9A1E8B5667238276F9CCFC7CF988" box="[770,896,1586,1612]" italics="true" pageId="15" pageNumber="16">B. pombali</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B56672382FBF9CCFBCCF988" authorityName="Pie &amp; Ribeiro" authorityYear="2015" box="[911,1072,1587,1612]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="quiririensis">
<emphasis id="B91E9A1E8B56672382FBF9CCFBCCF988" box="[911,1072,1587,1612]" italics="true" pageId="15" pageNumber="16">B. quiririensis</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B566723854BF9CCFB3AF988" authorityName="Mângia, Santana, de Oliveira Drummond, Sabagh, Ugioni, Costa &amp; Wachlevski" authorityYear="2023" box="[1087,1222,1586,1612]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="tabuleiro">
<emphasis id="B91E9A1E8B566723854BF9CCFB3AF988" box="[1087,1222,1586,1612]" italics="true" pageId="15" pageNumber="16">B. tabuleiro</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B56672385A2F9CCFA93F988" authorityName="Garey, Lima, Hartmann &amp; Haddad" authorityYear="2012" box="[1238,1391,1586,1612]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="tridactylus">
<emphasis id="B91E9A1E8B56672385A2F9CCFA93F988" box="[1238,1391,1586,1612]" italics="true" pageId="15" pageNumber="16">B. tridactylus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C6A3D8F8B56672384C6F9CCFD97F9B0" authorityName="Ribeiro, Firkowski, Bornschein &amp; Pie" authorityYear="2015" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="verrucosus">
<emphasis id="B91E9A1E8B56672384C6F9CCFD97F9B0" italics="true" pageId="15" pageNumber="16">B. verrucosus</emphasis>
</taxonomicName>
) species groups by the combination of its small adult body size, a leptodactyliform body form (
<emphasis id="B91E9A1E8B566723823FF97CFC9EF958" box="[843,866,1667,1692]" italics="true" pageId="15" pageNumber="16">vs</emphasis>
. bufoniform), and a dark-brown coloration (
<emphasis id="B91E9A1E8B5667238402F97CFA71F958" box="[1398,1421,1667,1692]" italics="true" pageId="15" pageNumber="16">vs</emphasis>
. vibrant coloration such as greenish, yellowish, orange or reddish) (
<bibRefCitation id="EFFB3BFD8B56672385D7F956FA80F900" author="Ribeiro LF &amp; Bornschein MR &amp; Belmonte-Lopes R &amp; Firkowski CR &amp; Morato SAA &amp; Pie MR" box="[1187,1404,1705,1732]" pageId="15" pageNumber="16" refId="ref14995" refString="Ribeiro LF, Bornschein MR, Belmonte-Lopes R, Firkowski CR, Morato SAA, Pie MR. 2015." type="book" year="2015">
<emphasis id="B91E9A1E8B56672385D7F956FA80F900" box="[1187,1404,1705,1732]" italics="true" pageId="15" pageNumber="16">Ribeiro et al., 2015</emphasis>
</bibRefCitation>
;
<bibRefCitation id="EFFB3BFD8B56672384FEF955FDD5F928" author="Lyra ML &amp; Rancilhac L &amp; Irisarri I &amp; Kunzel S &amp; Sanchez E &amp; Condez TH &amp; Padilla O &amp; Sole M &amp; Toledo LF &amp; Haddad CFB &amp; Vences M." journalOrPublisher="Toxins" pageId="15" pageNumber="16" part="13" publicationUrl="10.3390/toxins13110816" refId="ref14139" refString="Lyra ML, Monteiro ΙPC, Rancilhac L, Irisarri I, Kunzel S, Sanchez E, Condez TH, Rojas- Padilla O, Sole M, Toledo LF, Haddad CFB, Vences M. 2021. Initial phylotranscriptomic confirmation of homoplastic evolution of the conspicuous coloration and bufoniform morphology of pumpkin-toadlets in the genus Brachycephalus. Toxins 13 (11): 816 DOI 10.3390 / toxins 13110816." title="Initial phylotranscriptomic confirmation of homoplastic evolution of the conspicuous coloration and bufoniform morphology of pumpkin-toadlets in the genus Brachycephalus" type="journal volume" year="2021">
<emphasis id="B91E9A1E8B56672384FEF955FDD5F928" italics="true" pageId="15" pageNumber="16">Lyra et al., 2021</emphasis>
</bibRefCitation>
).
<taxonomicName id="4C6A3D8F8B5667238335F92EFCBFF92F" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[577,835,1745,1771]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B5667238335F92EFCBFF92F" box="[577,835,1745,1771]" italics="true" pageId="15" pageNumber="16">Brachycephalus dacnis</emphasis>
</taxonomicName>
is also distinguished from species of the
<taxonomicName id="4C6A3D8F8B566723846FF92DFA49F92F" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[1307,1461,1745,1771]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="B91E9A1E8B566723846FF92DFA49F92F" box="[1307,1461,1745,1771]" italics="true" pageId="15" pageNumber="16">B. ephippium</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6A3D8F8B566723849AF92DFD95F8D7" authorityName="Pombal" authorityYear="2001" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="vertebralis">
<emphasis id="B91E9A1E8B566723849AF92DFD95F8D7" italics="true" pageId="15" pageNumber="16">B. vertebralis</emphasis>
</taxonomicName>
species groups by its smooth skin, distinct frontoparietals, nasals, vomers, and parasphenoid that arenot synostosed
<emphasis id="B91E9A1E8B56672382C4F8DDFC2FF8FF" box="[944,979,1826,1851]" italics="true" pageId="15" pageNumber="16">via</emphasis>
a bony sphenethmoid (
<figureCitation id="13515A898B5667238598F8DEFACBF8F8" box="[1260,1335,1825,1852]" captionStart="Figure 5" captionStartId="12.[116,181,1269,1291]" captionTargetBox="[93,1510,238,1238]" captionTargetId="figure-137@12.[90,1512,234,1241]" captionTargetPageId="12" captionText="Figure 5 Skeletal anatomy of Brachycephalus dacnis (ZUEC-AMP 24978; undetermined sex) based on CT-scans. (A) The entire skeleton, with insets of (B) the skull (in dorsal, ventral, and right lateral views), (C) right hand and (D) left foot (both in dorsal view), (E) vertebral column (in dorsal view), (F) pectoral girdle (in ventral view), and (G) pelvic girdle (in right lateral view). Abbreviations: c, carpus; cl, clavicle; cor, coracoid; dt, distal tarsals; ex, exoccipital; fp, frontoparietal; il, ilium; isc, ischium; m, maxilla; mdI, manual digit I; mdV, manual digit V; min, mineralized cartilage; n, nasal; op, operculum; pd I, pedal digit I; pd V, pedal digit V; pm, premaxilla; pr, prootic; ps, parasphenoid; pt, pterygoid; pub, pubis; qj, quadratojugal; sac, sacrum; sc, scapula; ses, sesamoid; sm, septomaxilla; sp, sphenethmoid; sq, squamosal; ur, urostyle; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-5" pageId="15" pageNumber="16">Figs. 5</figureCitation>
,
<figureCitation id="13515A898B5667238432F8DDFAA8F8F8" box="[1350,1364,1826,1852]" captionStart="Figure 7" captionStartId="16.[116,181,738,760]" captionTargetBox="[90,1541,234,725]" captionTargetId="figure-414@16.[90,1512,234,710]" captionTargetPageId="16" captionText="Figure 7 Selected skulls of other Brachycephalus species. Cranial elements labeled are discussed in comparisons of B. dacnis to other species. Top row is the leptodactyliform “flea toads”, including B. hermogenesi (sister species of B. dacnis), B. clarissae, B. sulfuratus, and B. didactylus. Bottom row contains representatives of the three named species groups: pernix group (B.albolineatus, B. pernix), vertebralis group (B. vertebralis), and ephippium group (B. ephippium). Museum catalog numbers are listed below each species name. Abbreviations: fp, frontoparietal; n, nasal; ps, parasphenoid; pt, pterygoid; qj, quadratojugal; sp, sphenethmoid; sq, squamosal; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-7" pageId="15" pageNumber="16">7</figureCitation>
), presence of a quadratojugal, robust anterior process of the pterygoid, lack of hyperossification in the skull, distinct coracoid and clavicle and lack of mineralized epicoracoid cartilages, lack of vertebral fusions, lack of bony plates in the axial skeleton, and presence of phalanges on all digits (
<bibRefCitation id="EFFB3BFD8B49673C8330FC32FCFEFC23" author="Lyra ML &amp; Rancilhac L &amp; Irisarri I &amp; Kunzel S &amp; Sanchez E &amp; Condez TH &amp; Padilla O &amp; Sole M &amp; Toledo LF &amp; Haddad CFB &amp; Vences M." box="[580,770,972,999]" journalOrPublisher="Toxins" pageId="16" pageNumber="17" part="13" publicationUrl="10.3390/toxins13110816" refId="ref14139" refString="Lyra ML, Monteiro ΙPC, Rancilhac L, Irisarri I, Kunzel S, Sanchez E, Condez TH, Rojas- Padilla O, Sole M, Toledo LF, Haddad CFB, Vences M. 2021. Initial phylotranscriptomic confirmation of homoplastic evolution of the conspicuous coloration and bufoniform morphology of pumpkin-toadlets in the genus Brachycephalus. Toxins 13 (11): 816 DOI 10.3390 / toxins 13110816." title="Initial phylotranscriptomic confirmation of homoplastic evolution of the conspicuous coloration and bufoniform morphology of pumpkin-toadlets in the genus Brachycephalus" type="journal volume" year="2021">
<emphasis id="B91E9A1E8B49673C8330FC32FCFEFC23" box="[580,770,972,999]" italics="true" pageId="16" pageNumber="17">Lyra et al., 2021</emphasis>
</bibRefCitation>
;
<bibRefCitation id="EFFB3BFD8B49673C8265FC33FB84FC23" author="Clemente-Carvalho RBG &amp; Antoniazzi MM &amp; Haddad CFB &amp; Alves ACR &amp; Rocha HS &amp; Pereira GR &amp; Oliveira DF &amp; Lopes RT &amp; dos Reis SF" box="[785,1144,972,999]" journalOrPublisher="Journal of Morphology" pageId="16" pageNumber="17" pagination="1285 - 1295" part="270" publicationUrl="10.1002/jmor.10755" refId="ref12993" refString="Clemente-Carvalho RBG, Antoniazzi MM, Ιared C, Haddad CFB, Alves ACR, Rocha HS, Pereira GR, Oliveira DF, Lopes RT, dos Reis SF. 2009. Hyperossification in miniaturized toadlets of the genus Brachycephalus (Amphibia: Anura: Brachycephalidae): microscopic structure and macroscopic patterns of variation. Journal of Morphology 270: 1285 - 1295 DOI 10.1002 / jmor. 10755." title="Hyperossification in miniaturized toadlets of the genus Brachycephalus (Amphibia: Anura: Brachycephalidae): microscopic structure and macroscopic patterns of variation" type="journal article" year="2009">
<emphasis id="B91E9A1E8B49673C8265FC33FB84FC23" box="[785,1144,972,999]" italics="true" pageId="16" pageNumber="17">Clemente-Carvalho et al., 2009</emphasis>
</bibRefCitation>
,
<bibRefCitation id="EFFB3BFD8B49673C85F2FC32FB41FC23" author="Clemente-Carvalho RBG &amp; Giaretta AA &amp; Condez TH &amp; Haddad CFB &amp; dos Reis SF" box="[1158,1213,973,999]" journalOrPublisher="Herpetologic" pageId="16" pageNumber="17" pagination="365 - 374" part="68" publicationUrl="10.2307/23255788" refId="ref13065" refString="Clemente-Carvalho RBG, Giaretta AA, Condez TH, Haddad CFB, dos Reis SF. 2012. A new species of miniaturized toadlet, genus Brachycephalus (Anura: Brachycephalidae), from the Atlantic Forest of southeastern Brazil. Herpetologic 68: 365 - 374 DOI 10.2307 / 23255788." title="A new species of miniaturized toadlet, genus Brachycephalus (Anura: Brachycephalidae), from the Atlantic Forest of southeastern Brazil" type="journal article" year="2012">
<emphasis id="B91E9A1E8B49673C85F2FC32FB41FC23" box="[1158,1213,973,999]" italics="true" pageId="16" pageNumber="17">2012</emphasis>
</bibRefCitation>
;
<bibRefCitation id="EFFB3BFD8B49673C85B8FC33FA56FC23" author="Condez TH &amp; Clemente-Carvalho RBG &amp; Haddad CFB &amp; dos Reis SF" box="[1228,1450,972,999]" journalOrPublisher="Herpetologica" pageId="16" pageNumber="17" pagination="89 - 99" part="70" publicationUrl="10.1655/HERPETOLOGICA-D-13-00044" refId="ref13117" refString="Condez TH, Clemente-Carvalho RBG, Haddad CFB, dos Reis SF. 2014. A new species of Brachycephalus (Anura: Brachycephalidae) from the highlands of the Atlantic Forest, Southeastern Brazil. Herpetologica 70: 89 - 99 DOI 10.1655 / HERPETOLOGICA-D- 13 - 00044." title="A new species of Brachycephalus (Anura: Brachycephalidae) from the highlands of the Atlantic Forest, Southeastern Brazil" type="journal article" year="2014">
<emphasis id="B91E9A1E8B49673C85B8FC33FA56FC23" box="[1228,1450,972,999]" italics="true" pageId="16" pageNumber="17">Condez et al., 2014</emphasis>
</bibRefCitation>
,
<bibRefCitation id="EFFB3BFD8B49673C84CDFC33FA0CFC22" author="Condez TH &amp; Garcia PCA &amp; Amaral IB &amp; Haddad CFB" box="[1465,1520,972,998]" journalOrPublisher="Zootaxa" pageId="16" pageNumber="17" pagination="40 - 56" part="4083" publicationUrl="10.11646/zootaxa.4083.1.2" refId="ref13218" refString="Condez TH, Monteiro ΙPC, Comitti EΙ, Garcia PCA, Amaral IB, Haddad CFB. 2016. A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil. Zootaxa 4083: 40 - 56 DOI 10.11646 / zootaxa. 4083.1.2." title="A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil" type="journal article" year="2016">
<emphasis id="B91E9A1E8B49673C84CDFC33FA0CFC22" box="[1465,1520,972,998]" italics="true" pageId="16" pageNumber="17">2016</emphasis>
</bibRefCitation>
;
<bibRefCitation id="EFFB3BFD8B49673C8086FC0BFD25FBCA" author="Haddad CFB &amp; Alves ACR &amp; Clemente-Carvalho RBG &amp; dos Reis SF" box="[498,729,1012,1038]" journalOrPublisher="Copeia" pageId="16" pageNumber="17" pagination="410 - 420" part="3" publicationUrl="10.1643/CH-09-102" refId="ref13663" refString="Haddad CFB, Alves ACR, Clemente-Carvalho RBG, dos Reis SF. 2010. A new species of Brachycephalus from the Atlantic Rain Forest in Sao Paulo state, southeastern Brazil (Amphibia: Anura: Brachycephalidae). Copeia 3: 410 - 420 DOI 10.1643 / CH- 09 - 102." title="A new species of Brachycephalus from the Atlantic Rain Forest in Sao Paulo state, southeastern Brazil (Amphibia: Anura: Brachycephalidae)" type="journal article" year="2010">
<emphasis id="B91E9A1E8B49673C8086FC0BFD25FBCA" box="[498,729,1012,1038]" italics="true" pageId="16" pageNumber="17">Haddad et al., 2010</emphasis>
</bibRefCitation>
;
<bibRefCitation id="EFFB3BFD8B49673C839CFC0BFC7AFBCA" author="Pombal" box="[744,902,1012,1038]" journalOrPublisher="Boletin del Museo Nacional" pageId="16" pageNumber="17" pagination="1 - 12" part="526" refId="ref14732" refString="Pombal ΙP Ιr. 2010. A posicao taxonomica das &quot; variedades &quot; de Brachycephalus ephippium (Spix, 1824) descritas por Miranda-Ribeiro, 1920 (Amphibia, Anura, Brachycephalidae). Boletin del Museo Nacional 526: 1 - 12." title="A posicao taxonomica das &quot; variedades &quot; de Brachycephalus ephippium (Spix, 1824) descritas por Miranda-Ribeiro, 1920 (Amphibia, Anura, Brachycephalidae)" type="journal article" year="2010">
<emphasis id="B91E9A1E8B49673C839CFC0BFC7AFBCA" box="[744,902,1012,1038]" italics="true" pageId="16" pageNumber="17">Pombal, 2010</emphasis>
</bibRefCitation>
). From the other six leptodactyliform species, a.k.a. flea-toads (
<taxonomicName id="4C6A3D8F8B49673C8307FBE2FCF9FBF2" baseAuthorityName="Izecksohn" baseAuthorityYear="1971" box="[627,773,1052,1078]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="didactylus">
<emphasis id="B91E9A1E8B49673C8307FBE2FCF9FBF2" box="[627,773,1052,1078]" italics="true" pageId="16" pageNumber="17">B. didactylus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C8266FBE2FC73FBF2" box="[786,911,1052,1078]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="clarissae">
<emphasis id="B91E9A1E8B49673C8266FBE2FC73FBF2" box="[786,911,1052,1078]" italics="true" pageId="16" pageNumber="17">B. clarissae</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C82E8FBE2FBBBFBF2" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[924,1095,1052,1078]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B49673C82E8FBE2FBBBFBF2" box="[924,1095,1052,1078]" italics="true" pageId="16" pageNumber="17">B. hermogenesi</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C8520FBE2FB53FBF2" authorityName="Napoli, Caramaschi, Cruz &amp; Dias" authorityYear="2011" box="[1108,1199,1052,1078]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="pulex">
<emphasis id="B91E9A1E8B49673C8520FBE2FB53FBF2" box="[1108,1199,1052,1078]" italics="true" pageId="16" pageNumber="17">B. pulex</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C85C8FBE2FAF5FBF2" authorityName="Almeida-Silva, Silva-Soares, Rodrigues &amp; Verdade" authorityYear="2021" box="[1212,1289,1053,1078]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="puri">
<emphasis id="B91E9A1E8B49673C85C8FBE2FAF5FBF2" box="[1212,1289,1053,1078]" italics="true" pageId="16" pageNumber="17">B. puri</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C6A3D8F8B49673C843CFBE2FA24FBF2" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[1352,1496,1052,1078]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B49673C843CFBE2FA24FBF2" box="[1352,1496,1052,1078]" italics="true" pageId="16" pageNumber="17">B. sulfuratus</emphasis>
</taxonomicName>
),
<taxonomicName id="4C6A3D8F8B49673C849AFBE2FDC7FB9A" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C849AFBE2FDC7FB9A" italics="true" pageId="16" pageNumber="17">B. dacnis</emphasis>
</taxonomicName>
is distinguished by skin texture, toe morphology, coloration, and some skeletal traits. From
<taxonomicName id="4C6A3D8F8B49673C83F2FB92FCFAFB42" box="[646,774,1132,1158]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="clarissae">
<emphasis id="B91E9A1E8B49673C83F2FB92FCFAFB42" box="[646,774,1132,1158]" italics="true" pageId="16" pageNumber="17">B. clarissae</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C8261FB92FC82FB42" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[789,894,1132,1158]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C8261FB92FC82FB42" box="[789,894,1132,1158]" italics="true" pageId="16" pageNumber="17">B. dacnis</emphasis>
</taxonomicName>
is distinguished by having a smooth dorsal texture (granular in
<taxonomicName id="4C6A3D8F8B49673C83F2FB6BFCFAFB6A" box="[646,774,1172,1198]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="clarissae">
<emphasis id="B91E9A1E8B49673C83F2FB6BFCFAFB6A" box="[646,774,1172,1198]" italics="true" pageId="16" pageNumber="17">B. clarissae</emphasis>
</taxonomicName>
), a distinct and functional toe V (vestigial in
<taxonomicName id="4C6A3D8F8B49673C846FFB6BFA66FB6A" box="[1307,1434,1172,1198]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="clarissae">
<emphasis id="B91E9A1E8B49673C846FFB6BFA66FB6A" box="[1307,1434,1172,1198]" italics="true" pageId="16" pageNumber="17">B. clarissae</emphasis>
</taxonomicName>
), lacking synostosis among cranial bones (sphenthmoids, parasphenoid, and frontoparietals synostosed in
<taxonomicName id="4C6A3D8F8B49673C83EEFB1BFCE6FB39" box="[666,794,1251,1277]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="clarissae">
<emphasis id="B91E9A1E8B49673C83EEFB1BFCE6FB39" box="[666,794,1251,1277]" italics="true" pageId="16" pageNumber="17">B. clarissae</emphasis>
</taxonomicName>
;
<figureCitation id="13515A898B49673C825DFB1CFC89FB3A" box="[809,885,1251,1278]" captionStart="Figure 5" captionStartId="12.[116,181,1269,1291]" captionTargetBox="[93,1510,238,1238]" captionTargetId="figure-137@12.[90,1512,234,1241]" captionTargetPageId="12" captionText="Figure 5 Skeletal anatomy of Brachycephalus dacnis (ZUEC-AMP 24978; undetermined sex) based on CT-scans. (A) The entire skeleton, with insets of (B) the skull (in dorsal, ventral, and right lateral views), (C) right hand and (D) left foot (both in dorsal view), (E) vertebral column (in dorsal view), (F) pectoral girdle (in ventral view), and (G) pelvic girdle (in right lateral view). Abbreviations: c, carpus; cl, clavicle; cor, coracoid; dt, distal tarsals; ex, exoccipital; fp, frontoparietal; il, ilium; isc, ischium; m, maxilla; mdI, manual digit I; mdV, manual digit V; min, mineralized cartilage; n, nasal; op, operculum; pd I, pedal digit I; pd V, pedal digit V; pm, premaxilla; pr, prootic; ps, parasphenoid; pt, pterygoid; pub, pubis; qj, quadratojugal; sac, sacrum; sc, scapula; ses, sesamoid; sm, septomaxilla; sp, sphenethmoid; sq, squamosal; ur, urostyle; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-5" pageId="16" pageNumber="17">Figs. 5</figureCitation>
,
<figureCitation id="13515A898B49673C82F0FB1BFC6EFB3A" box="[900,914,1252,1278]" captionStart="Figure 7" captionStartId="16.[116,181,738,760]" captionTargetBox="[90,1541,234,725]" captionTargetId="figure-414@16.[90,1512,234,710]" captionTargetPageId="16" captionText="Figure 7 Selected skulls of other Brachycephalus species. Cranial elements labeled are discussed in comparisons of B. dacnis to other species. Top row is the leptodactyliform “flea toads”, including B. hermogenesi (sister species of B. dacnis), B. clarissae, B. sulfuratus, and B. didactylus. Bottom row contains representatives of the three named species groups: pernix group (B.albolineatus, B. pernix), vertebralis group (B. vertebralis), and ephippium group (B. ephippium). Museum catalog numbers are listed below each species name. Abbreviations: fp, frontoparietal; n, nasal; ps, parasphenoid; pt, pterygoid; qj, quadratojugal; sp, sphenethmoid; sq, squamosal; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-7" pageId="16" pageNumber="17">7</figureCitation>
), phalanges on pedal digits Iand V (lacking in
<taxonomicName id="4C6A3D8F8B49673C84BDFB1BFA1DFB39" box="[1481,1505,1252,1277]" pageId="16" pageNumber="17">
<emphasis id="B91E9A1E8B49673C84BDFB1BFA1DFB39" box="[1481,1505,1252,1277]" italics="true" pageId="16" pageNumber="17">B.</emphasis>
</taxonomicName>
<emphasis id="B91E9A1E8B49673C8086FAF4FDAEFAE1" box="[498,594,1291,1317]" italics="true" pageId="16" pageNumber="17">clarissae</emphasis>
), lacking mineralized epicoracoid cartilages (heavily mineralized and fused with pectoral bones in
<taxonomicName id="4C6A3D8F8B49673C83CBFACBFCC1FA89" box="[703,829,1331,1357]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="clarissae">
<emphasis id="B91E9A1E8B49673C83CBFACBFCC1FA89" box="[703,829,1331,1357]" italics="true" pageId="16" pageNumber="17">B. clarissae</emphasis>
</taxonomicName>
), and by having a distinct iris (indistinct in
<taxonomicName id="4C6A3D8F8B49673C8441FACBFA4FFA89" box="[1333,1459,1331,1357]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="clarissae">
<emphasis id="B91E9A1E8B49673C8441FACBFA4FFA89" box="[1333,1459,1331,1357]" italics="true" pageId="16" pageNumber="17">B. clarissae</emphasis>
</taxonomicName>
) (
<bibRefCitation id="EFFB3BFD8B49673C84BAFACCFD8DFAB1" author="Folly M &amp; Vrcibradic D &amp; Siqueira CC &amp; da Rocha CFD &amp; Machado AS &amp; Lopes RT" journalOrPublisher="Ichthyology &amp; Herpetology" pageId="16" pageNumber="17" pagination="585 - 601" part="110" publicationUrl="10.1643/h2020144" refId="ref13413" refString="Folly M, Vrcibradic D, Siqueira CC, da Rocha CFD, Machado AS, Lopes RT, Pombal ΙP Ιr. 2022. A new species of Brachycephalus (Anura, Brachycephalidae) from a montane Atlantic rainforest of southeastern Brazil, with a reappraisal of the species groups in the genus. Ichthyology &amp; Herpetology 110: 585 - 601 DOI 10.1643 / h 2020144." title="A new species of Brachycephalus (Anura, Brachycephalidae) from a montane Atlantic rainforest of southeastern Brazil, with a reappraisal of the species groups in the genus" type="journal article" year="2022">
<emphasis id="B91E9A1E8B49673C84BAFACCFD8DFAB1" italics="true" pageId="16" pageNumber="17">Folly et al., 2022</emphasis>
</bibRefCitation>
).
<taxonomicName id="4C6A3D8F8B49673C83FDFAA4FC70FAB1" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[649,908,1371,1397]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C83FDFAA4FC70FAB1" box="[649,908,1371,1397]" italics="true" pageId="16" pageNumber="17">Brachycephalus dacnis</emphasis>
</taxonomicName>
has a distinct, and functional toe V (like
<taxonomicName id="4C6A3D8F8B49673C8400FAA3F9FAFAB1" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[1396,1542,1371,1397]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B49673C8400FAA3F9FAFAB1" box="[1396,1542,1371,1397]" italics="true" pageId="16" pageNumber="17">B. sulfuratus</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6A3D8F8B49673C8350FA7CFD2CFA59" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[548,720,1411,1437]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B49673C8350FA7CFD2CFA59" box="[548,720,1411,1437]" italics="true" pageId="16" pageNumber="17">B. hermogenesi</emphasis>
</taxonomicName>
), whereas it is vestigial in
<taxonomicName id="4C6A3D8F8B49673C8288FA7CFB71FA59" baseAuthorityName="Izecksohn" baseAuthorityYear="1971" box="[1020,1165,1411,1437]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="didactylus">
<emphasis id="B91E9A1E8B49673C8288FA7CFB71FA59" box="[1020,1165,1411,1437]" italics="true" pageId="16" pageNumber="17">B. didactylus</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFFB3BFD8B49673C85EAFA7CFA81FA59" author="Haddad CFB &amp; Alves ACR &amp; Clemente-Carvalho RBG &amp; dos Reis SF" box="[1182,1405,1411,1437]" journalOrPublisher="Copeia" pageId="16" pageNumber="17" pagination="410 - 420" part="3" publicationUrl="10.1643/CH-09-102" refId="ref13663" refString="Haddad CFB, Alves ACR, Clemente-Carvalho RBG, dos Reis SF. 2010. A new species of Brachycephalus from the Atlantic Rain Forest in Sao Paulo state, southeastern Brazil (Amphibia: Anura: Brachycephalidae). Copeia 3: 410 - 420 DOI 10.1643 / CH- 09 - 102." title="A new species of Brachycephalus from the Atlantic Rain Forest in Sao Paulo state, southeastern Brazil (Amphibia: Anura: Brachycephalidae)" type="journal article" year="2010">
<emphasis id="B91E9A1E8B49673C85EAFA7CFA81FA59" box="[1182,1405,1411,1437]" italics="true" pageId="16" pageNumber="17">Haddad et al., 2010</emphasis>
</bibRefCitation>
;
<emphasis id="B91E9A1E8B49673C84FDFA7CFD6DFA01" italics="true" pageId="16" pageNumber="17">Giaretta &amp; Sawaya, 1998</emphasis>
). Neither
<taxonomicName id="4C6A3D8F8B49673C827FFA54FC88FA00" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[779,884,1450,1476]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C827FFA54FC88FA00" box="[779,884,1450,1476]" italics="true" pageId="16" pageNumber="17">B. dacnis</emphasis>
</taxonomicName>
nor
<taxonomicName id="4C6A3D8F8B49673C82DAFA54FBA7FA00" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[942,1115,1450,1476]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B49673C82DAFA54FBA7FA00" box="[942,1115,1450,1476]" italics="true" pageId="16" pageNumber="17">B. hermogenesi</emphasis>
</taxonomicName>
lack phalanges on the toes, whereas both
<taxonomicName id="4C6A3D8F8B49673C8345FA2CFD39FA28" baseAuthorityName="Izecksohn" baseAuthorityYear="1971" box="[561,709,1490,1516]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="didactylus">
<emphasis id="B91E9A1E8B49673C8345FA2CFD39FA28" box="[561,709,1490,1516]" italics="true" pageId="16" pageNumber="17">B. didactylus</emphasis>
</taxonomicName>
(pedal digit count: 0, 2, 3, 4, 0) and
<taxonomicName id="4C6A3D8F8B49673C850CFA2CFAF7FA28" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[1144,1291,1490,1516]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B49673C850CFA2CFAF7FA28" box="[1144,1291,1490,1516]" italics="true" pageId="16" pageNumber="17">B. sulfuratus</emphasis>
</taxonomicName>
(0, 2, 3, 4, 2) lack phalanges on one or more digit. In
<taxonomicName id="4C6A3D8F8B49673C82F3FA04FBE4F9D0" baseAuthorityName="Izecksohn" baseAuthorityYear="1971" box="[903,1048,1530,1556]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="didactylus">
<emphasis id="B91E9A1E8B49673C82F3FA04FBE4F9D0" box="[903,1048,1530,1556]" italics="true" pageId="16" pageNumber="17">B. didactylus</emphasis>
</taxonomicName>
, there is a trapezoidal chest mark, which is absent in
<taxonomicName id="4C6A3D8F8B49673C8312F9DCFBEBF9F9" authority="(Almeida-Silva et al., 2021)" baseAuthorityName="Almeida-Silva" baseAuthorityYear="2021" box="[614,1047,1570,1597]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C8312F9DCFD33F9F8" box="[614,719,1570,1596]" italics="true" pageId="16" pageNumber="17">B. dacnis</emphasis>
(
<bibRefCitation id="EFFB3BFD8B49673C8396F9DDFBF1F9F8" author="Almeida-Silva D &amp; Silva-Soares T &amp; Rodrigues MT &amp; Verdade VK" box="[738,1037,1570,1596]" journalOrPublisher="Zootaxa" pageId="16" pageNumber="17" pagination="517 - 532" part="5068" publicationUrl="10.11646/zootaxa.5068.4.3" refId="ref12432" refString="Almeida-Silva D, Silva-Soares T, Rodrigues MT, Verdade VK. 2021. New species of flea-toad, genus Brachycephalus (Anura: Brachycephalidae) from the Atlantic Forest of Espirito Santo, Brazil. Zootaxa 5068: 517 - 532 DOI 10.11646 / zootaxa. 5068.4.3." title="New species of flea-toad, genus Brachycephalus (Anura: Brachycephalidae) from the Atlantic Forest of Espirito Santo, Brazil" type="journal article" year="2021">
<emphasis id="B91E9A1E8B49673C8396F9DDFBF1F9F8" box="[738,1037,1570,1596]" italics="true" pageId="16" pageNumber="17">Almeida-Silva et al., 2021</emphasis>
</bibRefCitation>
)
</taxonomicName>
, as well as a fully synostosed braincase (
<figureCitation id="13515A898B49673C8088F9B5FDC7F9A0" box="[508,571,1610,1636]" captionStart="Figure 7" captionStartId="16.[116,181,738,760]" captionTargetBox="[90,1541,234,725]" captionTargetId="figure-414@16.[90,1512,234,710]" captionTargetPageId="16" captionText="Figure 7 Selected skulls of other Brachycephalus species. Cranial elements labeled are discussed in comparisons of B. dacnis to other species. Top row is the leptodactyliform “flea toads”, including B. hermogenesi (sister species of B. dacnis), B. clarissae, B. sulfuratus, and B. didactylus. Bottom row contains representatives of the three named species groups: pernix group (B.albolineatus, B. pernix), vertebralis group (B. vertebralis), and ephippium group (B. ephippium). Museum catalog numbers are listed below each species name. Abbreviations: fp, frontoparietal; n, nasal; ps, parasphenoid; pt, pterygoid; qj, quadratojugal; sp, sphenethmoid; sq, squamosal; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-7" pageId="16" pageNumber="17">Fig. 7</figureCitation>
), a wire-like anterior process of the pterygoid, no quadratojugal, heavily mineralized epicoracoid cartilages fused with the pectoral bones, fused presacral vertebrae VI, VII, and VIII, and no phalanges on manual digit Ior IV. From
<taxonomicName id="4C6A3D8F8B49673C850FF965FAF2F977" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[1147,1294,1689,1715]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B49673C850FF965FAF2F977" box="[1147,1294,1689,1715]" italics="true" pageId="16" pageNumber="17">B. sulfuratus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C8468F965FA7AF977" box="[1308,1414,1689,1715]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C8468F965FA7AF977" box="[1308,1414,1689,1715]" italics="true" pageId="16" pageNumber="17">B. dacnis</emphasis>
</taxonomicName>
is distinguished by the absence of an inverted V-shaped chest mark (
<bibRefCitation id="EFFB3BFD8B49673C8473F93EFA19F91F" author="Condez TH &amp; Garcia PCA &amp; Amaral IB &amp; Haddad CFB" box="[1287,1509,1729,1755]" journalOrPublisher="Zootaxa" pageId="16" pageNumber="17" pagination="40 - 56" part="4083" publicationUrl="10.11646/zootaxa.4083.1.2" refId="ref13218" refString="Condez TH, Monteiro ΙPC, Comitti EΙ, Garcia PCA, Amaral IB, Haddad CFB. 2016. A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil. Zootaxa 4083: 40 - 56 DOI 10.11646 / zootaxa. 4083.1.2." title="A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil" type="journal article" year="2016">
<emphasis id="B91E9A1E8B49673C8473F93EFA19F91F" box="[1287,1509,1729,1755]" italics="true" pageId="16" pageNumber="17">Condez et al., 2016</emphasis>
</bibRefCitation>
).
</paragraph>
<caption id="DF1516848B5667238101FF07FEDCFEE8" pageId="15" pageNumber="16" startId="15.[117,172,248,269]" targetBox="[108,1523,319,1447]" targetIsTable="true" targetPageId="15" targetType="table">
<paragraph id="8BD5460C8B5667238101FF07FEDCFEE8" blockId="15.[117,1515,248,300]" pageId="15" pageNumber="16">
<emphasis id="B91E9A1E8B5667238101FF07FB95FEC9" bold="true" box="[117,1129,248,270]" pageId="15" pageNumber="16">
Table 2 Advertisement call traits described for three flea-toads species from
<taxonomicName id="4C6A3D8F8B56672382F3FF07FBDCFECA" baseAuthorityName="Trueb &amp; Alberch" baseAuthorityYear="1985" box="[903,1056,248,270]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B91E9A1E8B56672382F3FF07FBDCFECA" bold="true" box="[903,1056,248,270]" italics="true" pageId="15" pageNumber="16">Brachycephalus</emphasis>
</taxonomicName>
genus.
</emphasis>
Call traits present as mean ± standard deviation (range).
</paragraph>
</caption>
<paragraph id="8BD5460C8B5667238039FEC0FA5CFA63" pageId="15" pageNumber="16">
<table id="F96AB4AC8B5698D38118FEC0FA0FFA63" box="[108,1523,319,1447]" gridcols="7" gridrows="23" pageId="15" pageNumber="16">
<tr id="355A444E8B5698D38118FEC0FA0FFE0F" box="[108,1523,319,459]" gridrow="0" pageId="15" pageNumber="16" rowspan-0="1">
<th id="768B2D328B5698D38039FEC0FDD0FE0F" box="[333,556,319,459]" gridcol="1" gridrow="0" pageId="15" pageNumber="16">
<taxonomicName id="4C6A3D8F8B5667238039FEC0FDD0FE91" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[333,556,319,341]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="dacnis">Brachycephalus dacnis</taxonomicName>
six individuals (including the holotype) Dacnis, Ubatuba
</th>
<th id="768B2D328B5698D38336FEC0FCD8FE0F" box="[578,804,319,459]" gridcol="2" gridrow="0" pageId="15" pageNumber="16">
<taxonomicName id="4C6A3D8F8B5667238336FEC0FCDDFE91" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[578,801,319,341]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="dacnis">Brachycephalus dacnis</taxonomicName>
Corcovado, SP and Trilha do Corisco, RƖ
</th>
<th id="768B2D328B5698D3824FFEC0FC20FE0F" box="[827,988,319,459]" gridcol="3" gridrow="0" pageId="15" pageNumber="16">
<taxonomicName id="4C6A3D8F8B566723824FFEC0FC28FE91" baseAuthorityName="Trueb &amp; Alberch" baseAuthorityYear="1985" box="[827,980,319,341]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">Brachycephalus</taxonomicName>
<taxonomicName id="4C6A3D8F8B566723824FFEA2FC48FEB7" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[827,948,349,371]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B566723824FFEA2FC48FEB7" bold="true" box="[827,948,349,371]" italics="true" pageId="15" pageNumber="16">hermogenesi</emphasis>
</taxonomicName>
five individuals Ubatuba
</th>
<th id="768B2D328B5698D38298FEC0FB6FFE0F" box="[1004,1171,319,459]" gridcol="4" gridrow="0" pageId="15" pageNumber="16">
<taxonomicName id="4C6A3D8F8B5667238298FEC0FB79FE91" baseAuthorityName="Trueb &amp; Alberch" baseAuthorityYear="1985" box="[1004,1157,319,341]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">Brachycephalus</taxonomicName>
<taxonomicName id="4C6A3D8F8B5667238298FEA2FB99FEB7" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[1004,1125,349,371]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="hermogenesi">hermogenesi</taxonomicName>
Several locations
</th>
<th id="768B2D328B5698D385D7FEC0FAC0FE0F" box="[1187,1340,319,459]" gridcol="5" gridrow="0" pageId="15" pageNumber="16">
<taxonomicName id="4C6A3D8F8B56672385D7FEC0FAC0FE91" baseAuthorityName="Trueb &amp; Alberch" baseAuthorityYear="1985" box="[1187,1340,319,341]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">Brachycephalus</taxonomicName>
<taxonomicName id="4C6A3D8F8B56672385D7FEA2FAF8FEB7" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[1187,1284,349,371]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B56672385D7FEA2FAF8FEB7" bold="true" box="[1187,1284,349,371]" italics="true" pageId="15" pageNumber="16">sulfuratus</emphasis>
</taxonomicName>
four topotypes São Francisco do Sul
</th>
<th id="768B2D328B5698D38420FEC0FA0FFE0F" box="[1364,1523,319,459]" gridcol="6" gridrow="0" pageId="15" pageNumber="16">
<emphasis id="B91E9A1E8B5667238420FEC0FA52FE69" bold="true" box="[1364,1517,319,429]" pageId="15" pageNumber="16">
<taxonomicName id="4C6A3D8F8B5667238420FEC0FA11FE91" baseAuthorityName="Trueb &amp; Alberch" baseAuthorityYear="1985" box="[1364,1517,319,341]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="genus">
<emphasis id="B91E9A1E8B5667238420FEC0FA11FE91" bold="true" box="[1364,1517,319,341]" italics="true" pageId="15" pageNumber="16">Brachycephalus</emphasis>
</taxonomicName>
<taxonomicName id="4C6A3D8F8B5667238420FEA2FA49FEB7" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[1364,1461,349,371]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="15" pageNumber="16" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B5667238420FEA2FA49FEB7" bold="true" box="[1364,1461,349,371]" italics="true" pageId="15" pageNumber="16">sulfuratus</emphasis>
</taxonomicName>
Several locations
</emphasis>
</th>
</tr>
<tr id="355A444E8B5698D38118FE21FA0FFE30" box="[108,1523,478,500]" gridrow="1" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FE21FEC1FE30" box="[108,317,478,500]" gridcol="0" gridrow="1" pageId="15" pageNumber="16">Call duration (s)</th>
<td id="768B2D328B5698D38039FE21FDD0FE30" box="[333,556,478,500]" gridcol="1" gridrow="1" pageId="15" pageNumber="16">0.05 ± 0.01</td>
<td id="768B2D328B5698D38336FE21FCD8FE30" box="[578,804,478,500]" gridcol="2" gridrow="1" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D3824FFE21FC20FE30" box="[827,988,478,500]" gridcol="3" gridrow="1" pageId="15" pageNumber="16">0.56 ± 0.14</td>
<td id="768B2D328B5698D38298FE21FB6FFE30" box="[1004,1171,478,500]" gridcol="4" gridrow="1" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D385D7FE21FAC0FE30" box="[1187,1340,478,500]" gridcol="5" gridrow="1" pageId="15" pageNumber="16">1.70 ± 0.10</td>
<td id="768B2D328B5698D38420FE21FA0FFE30" box="[1364,1523,478,500]" gridcol="6" gridrow="1" pageId="15" pageNumber="16">-</td>
</tr>
<tr id="355A444E8B5698D38118FDFCFA0FFDDE" box="[108,1523,515,538]" gridrow="2" pageId="15" pageNumber="16" rowspan-0="1" rowspan-2="1" rowspan-4="1" rowspan-6="1">
<td id="768B2D328B5698D38039FDFCFDD0FDDE" box="[333,556,515,538]" gridcol="1" gridrow="2" pageId="15" pageNumber="16">(0.030.41)</td>
<td id="768B2D328B5698D3824FFDFCFC20FDDE" box="[827,988,515,538]" gridcol="3" gridrow="2" pageId="15" pageNumber="16">(0.410.78)</td>
<td id="768B2D328B5698D385D7FDFCFAC0FDDE" box="[1187,1340,515,538]" gridcol="5" gridrow="2" pageId="15" pageNumber="16">(1.502.30)</td>
</tr>
<tr id="355A444E8B5698D38118FDD6FA0FFDFB" box="[108,1523,553,575]" gridrow="3" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FDD6FEC1FDFB" box="[108,317,553,575]" gridcol="0" gridrow="3" pageId="15" pageNumber="16">Inter-call interval (s)</th>
<td id="768B2D328B5698D38039FDD6FDD0FDFB" box="[333,556,553,575]" gridcol="1" gridrow="3" pageId="15" pageNumber="16">2.1 ± 0.06</td>
<td id="768B2D328B5698D38336FDD6FCD8FDFB" box="[578,804,553,575]" gridcol="2" gridrow="3" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D3824FFDD6FC20FDFB" box="[827,988,553,575]" gridcol="3" gridrow="3" pageId="15" pageNumber="16">2.70 ± 0.15</td>
<td id="768B2D328B5698D38298FDD6FB6FFDFB" box="[1004,1171,553,575]" gridcol="4" gridrow="3" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D385D7FDD6FAC0FDFB" box="[1187,1340,553,575]" gridcol="5" gridrow="3" pageId="15" pageNumber="16">4.5 ± 1.7</td>
<td id="768B2D328B5698D38420FDD6FA0FFDFB" box="[1364,1523,553,575]" gridcol="6" gridrow="3" pageId="15" pageNumber="16">-</td>
</tr>
<tr id="355A444E8B5698D38118FDB0FA0FFDA1" box="[108,1523,591,613]" gridrow="4" pageId="15" pageNumber="16" rowspan-0="1" rowspan-2="1" rowspan-4="1" rowspan-6="1">
<td id="768B2D328B5698D38039FDB0FDD0FDA1" box="[333,556,591,613]" gridcol="1" gridrow="4" pageId="15" pageNumber="16">(1.672.69)</td>
<td id="768B2D328B5698D3824FFDB0FC20FDA1" box="[827,988,591,613]" gridcol="3" gridrow="4" pageId="15" pageNumber="16">(2.522.91)</td>
<td id="768B2D328B5698D385D7FDB0FAC0FDA1" box="[1187,1340,591,613]" gridcol="5" gridrow="4" pageId="15" pageNumber="16">(3.17.4)</td>
</tr>
<tr id="355A444E8B5698D38118FD8BFA0FFD6C" box="[108,1523,628,680]" gridrow="5" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FD8BFEC1FD6C" box="[108,317,628,680]" gridcol="0" gridrow="5" pageId="15" pageNumber="16">Number of notes per call</th>
<td id="768B2D328B5698D38039FD8BFDD0FD6C" box="[333,556,628,680]" gridcol="1" gridrow="5" pageId="15" pageNumber="16">12</td>
<td id="768B2D328B5698D38336FD8BFCD8FD6C" box="[578,804,628,680]" gridcol="2" gridrow="5" pageId="15" pageNumber="16"></td>
<td id="768B2D328B5698D3824FFD8BFC20FD6C" box="[827,988,628,680]" gridcol="3" gridrow="5" pageId="15" pageNumber="16">26</td>
<td id="768B2D328B5698D38298FD8BFB6FFD6C" box="[1004,1171,628,680]" gridcol="4" gridrow="5" pageId="15" pageNumber="16"></td>
<td id="768B2D328B5698D385D7FD8BFAC0FD6C" box="[1187,1340,628,680]" gridcol="5" gridrow="5" pageId="15" pageNumber="16">46</td>
<td id="768B2D328B5698D38420FD8BFA0FFD6C" box="[1364,1523,628,680]" gridcol="6" gridrow="5" pageId="15" pageNumber="16"></td>
</tr>
<tr id="355A444E8B5698D38118FD48FA0FFD2E" box="[108,1523,695,746]" gridrow="6" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FD48FEC1FD2E" box="[108,317,695,746]" gridcol="0" gridrow="6" pageId="15" pageNumber="16">
Presence of attenuated notes
<emphasis id="B91E9A1E8B566723806EFD2BFEDFFD21" box="[282,291,724,741]" italics="true" pageId="15" pageNumber="16">*</emphasis>
</th>
<td id="768B2D328B5698D38039FD48FDD0FD2E" box="[333,556,695,746]" gridcol="1" gridrow="6" pageId="15" pageNumber="16">No</td>
<td id="768B2D328B5698D38336FD48FCD8FD2E" box="[578,804,695,746]" gridcol="2" gridrow="6" pageId="15" pageNumber="16">No</td>
<td id="768B2D328B5698D3824FFD48FC20FD2E" box="[827,988,695,746]" gridcol="3" gridrow="6" pageId="15" pageNumber="16">Yes</td>
<td id="768B2D328B5698D38298FD48FB6FFD2E" box="[1004,1171,695,746]" gridcol="4" gridrow="6" pageId="15" pageNumber="16">Yes</td>
<td id="768B2D328B5698D385D7FD48FAC0FD2E" box="[1187,1340,695,746]" gridcol="5" gridrow="6" pageId="15" pageNumber="16">No</td>
<td id="768B2D328B5698D38420FD48FA0FFD2E" box="[1364,1523,695,746]" gridcol="6" gridrow="6" pageId="15" pageNumber="16">No</td>
</tr>
<tr id="355A444E8B5698D38118FD05FA0FFCD4" box="[108,1523,762,784]" gridrow="7" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FD05FEC1FCD4" box="[108,317,762,784]" gridcol="0" gridrow="7" pageId="15" pageNumber="16">Note duration (s)</th>
<td id="768B2D328B5698D38039FD05FDD0FCD4" box="[333,556,762,784]" gridcol="1" gridrow="7" pageId="15" pageNumber="16">The same as call</td>
<td id="768B2D328B5698D38336FD05FCD8FCD4" box="[578,804,762,784]" gridcol="2" gridrow="7" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D3824FFD05FC20FCD4" box="[827,988,762,784]" gridcol="3" gridrow="7" pageId="15" pageNumber="16">0.25 ± 0.04</td>
<td id="768B2D328B5698D38298FD05FB6FFCD4" box="[1004,1171,762,784]" gridcol="4" gridrow="7" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D385D7FD05FAC0FCD4" box="[1187,1340,762,784]" gridcol="5" gridrow="7" pageId="15" pageNumber="16">0.17 ± 0.01</td>
<td id="768B2D328B5698D38420FD05FA0FFCD4" box="[1364,1523,762,784]" gridcol="6" gridrow="7" pageId="15" pageNumber="16">-</td>
</tr>
<tr id="355A444E8B5698D38118FCE8FA0FFCF2" box="[108,1523,791,822]" gridrow="8" pageId="15" pageNumber="16" rowspan-0="1" rowspan-2="1" rowspan-4="1" rowspan-6="1">
<td id="768B2D328B5698D38039FCE8FDD0FCF2" box="[333,556,791,822]" gridcol="1" gridrow="8" pageId="15" pageNumber="16">duration</td>
<td id="768B2D328B5698D3824FFCE8FC20FCF2" box="[827,988,791,822]" gridcol="3" gridrow="8" pageId="15" pageNumber="16">(0.170.27)</td>
<td id="768B2D328B5698D385D7FCE8FAC0FCF2" box="[1187,1340,791,822]" gridcol="5" gridrow="8" pageId="15" pageNumber="16">(0.130.21)</td>
</tr>
<tr id="355A444E8B5698D38118FCBAFA0FFC9F" box="[108,1523,837,859]" gridrow="9" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FCBAFEC1FC9F" box="[108,317,837,859]" gridcol="0" gridrow="9" pageId="15" pageNumber="16">Inter-note interval (s)</th>
<td id="768B2D328B5698D38039FCBAFDD0FC9F" box="[333,556,837,859]" gridcol="1" gridrow="9" pageId="15" pageNumber="16">n/a</td>
<td id="768B2D328B5698D38336FCBAFCD8FC9F" box="[578,804,837,859]" gridcol="2" gridrow="9" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D3824FFCBAFC20FC9F" box="[827,988,837,859]" gridcol="3" gridrow="9" pageId="15" pageNumber="16">2.70 ± 0.15</td>
<td id="768B2D328B5698D38298FCBAFB6FFC9F" box="[1004,1171,837,859]" gridcol="4" gridrow="9" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D385D7FCBAFAC0FC9F" box="[1187,1340,837,859]" gridcol="5" gridrow="9" pageId="15" pageNumber="16">n/a</td>
<td id="768B2D328B5698D38420FCBAFA0FFC9F" box="[1364,1523,837,859]" gridcol="6" gridrow="9" pageId="15" pageNumber="16">-</td>
</tr>
<tr id="355A444E8B5698D38118FC94FA0FFC45" box="[108,1523,875,897]" gridrow="10" pageId="15" pageNumber="16" rowspan-0="1" rowspan-1="1" rowspan-2="1" rowspan-4="1" rowspan-5="1" rowspan-6="1">
<td id="768B2D328B5698D3824FFC94FC20FC45" box="[827,988,875,897]" gridcol="3" gridrow="10" pageId="15" pageNumber="16">(2.522.91)</td>
</tr>
<tr id="355A444E8B5698D38118FC6FFA0FFC62" box="[108,1523,912,934]" gridrow="11" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FC6FFEC1FC62" box="[108,317,912,934]" gridcol="0" gridrow="11" pageId="15" pageNumber="16">Note repetition rate</th>
<td id="768B2D328B5698D38039FC6FFDD0FC62" box="[333,556,912,934]" gridcol="1" gridrow="11" pageId="15" pageNumber="16">19.58 ± 1.84</td>
<td id="768B2D328B5698D38336FC6FFCD8FC62" box="[578,804,912,934]" gridcol="2" gridrow="11" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D3824FFC6FFC20FC62" box="[827,988,912,934]" gridcol="3" gridrow="11" pageId="15" pageNumber="16">4.12 ± 0.48</td>
<td id="768B2D328B5698D38298FC6FFB6FFC62" box="[1004,1171,912,934]" gridcol="4" gridrow="11" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D385D7FC6FFAC0FC62" box="[1187,1340,912,934]" gridcol="5" gridrow="11" pageId="15" pageNumber="16">0.2 ± 0.1</td>
<td id="768B2D328B5698D38420FC6FFA0FFC62" box="[1364,1523,912,934]" gridcol="6" gridrow="11" pageId="15" pageNumber="16">-</td>
</tr>
<tr id="355A444E8B5698D38118FC51FA0FFC08" box="[108,1523,942,972]" gridrow="12" pageId="15" pageNumber="16" rowspan-2="1" rowspan-4="1" rowspan-6="1">
<th id="768B2D328B5698D38118FC51FEC1FC08" box="[108,317,942,972]" gridcol="0" gridrow="12" pageId="15" pageNumber="16">(notes/s)</th>
<td id="768B2D328B5698D38039FC51FDD0FC08" box="[333,556,942,972]" gridcol="1" gridrow="12" pageId="15" pageNumber="16">(12.5529.41)</td>
<td id="768B2D328B5698D3824FFC51FC20FC08" box="[827,988,942,972]" gridcol="3" gridrow="12" pageId="15" pageNumber="16">(3.584.88)</td>
<td id="768B2D328B5698D385D7FC51FAC0FC08" box="[1187,1340,942,972]" gridcol="5" gridrow="12" pageId="15" pageNumber="16">(0.20.3)</td>
</tr>
<tr id="355A444E8B5698D38118FC24FA0FFBCA" box="[108,1523,987,1038]" gridrow="13" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FC24FEC1FBCA" box="[108,317,987,1038]" gridcol="0" gridrow="13" pageId="15" pageNumber="16">Number of pulses per note</th>
<td id="768B2D328B5698D38039FC24FDD0FBCA" box="[333,556,987,1038]" gridcol="1" gridrow="13" pageId="15" pageNumber="16">37</td>
<td id="768B2D328B5698D38336FC24FCD8FBCA" box="[578,804,987,1038]" gridcol="2" gridrow="13" pageId="15" pageNumber="16">116</td>
<td id="768B2D328B5698D3824FFC24FC20FBCA" box="[827,988,987,1038]" gridcol="3" gridrow="13" pageId="15" pageNumber="16">12</td>
<td id="768B2D328B5698D38298FC24FB6FFBCA" box="[1004,1171,987,1038]" gridcol="4" gridrow="13" pageId="15" pageNumber="16">23</td>
<td id="768B2D328B5698D385D7FC24FAC0FBCA" box="[1187,1340,987,1038]" gridcol="5" gridrow="13" pageId="15" pageNumber="16">79</td>
<td id="768B2D328B5698D38420FC24FA0FFBCA" box="[1364,1523,987,1038]" gridcol="6" gridrow="13" pageId="15" pageNumber="16">214</td>
</tr>
<tr id="355A444E8B5698D38118FBE1FA0FFBF0" box="[108,1523,1054,1076]" gridrow="14" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FBE1FEC1FBF0" box="[108,317,1054,1076]" gridcol="0" gridrow="14" pageId="15" pageNumber="16">Minimum frequency</th>
<td id="768B2D328B5698D38039FBE1FDD0FBF0" box="[333,556,1054,1076]" gridcol="1" gridrow="14" pageId="15" pageNumber="16">7.27 ± 0.12</td>
<td id="768B2D328B5698D38336FBE1FCD8FBF0" box="[578,804,1054,1076]" gridcol="2" gridrow="14" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D3824FFBE1FC20FBF0" box="[827,988,1054,1076]" gridcol="3" gridrow="14" pageId="15" pageNumber="16">7.94 ± 0.08</td>
<td id="768B2D328B5698D38298FBE1FB6FFBF0" box="[1004,1171,1054,1076]" gridcol="4" gridrow="14" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D385D7FBE1FAC0FBF0" box="[1187,1340,1054,1076]" gridcol="5" gridrow="14" pageId="15" pageNumber="16">4.9 ± 0.3</td>
<td id="768B2D328B5698D38420FBE1FA0FFBF0" box="[1364,1523,1054,1076]" gridcol="6" gridrow="14" pageId="15" pageNumber="16">-</td>
</tr>
<tr id="355A444E8B5698D38118FBC4FA0FFBAB" box="[108,1523,1083,1135]" gridrow="15" pageId="15" pageNumber="16" rowspan-2="1" rowspan-4="1" rowspan-6="1">
<th id="768B2D328B5698D38118FBC4FEC1FBAB" box="[108,317,1083,1135]" gridcol="0" gridrow="15" pageId="15" pageNumber="16">(Frequency 5%) (kHz)</th>
<td id="768B2D328B5698D38039FBC4FDD0FBAB" box="[333,556,1083,1135]" gridcol="1" gridrow="15" pageId="15" pageNumber="16">(6.897.50)</td>
<td id="768B2D328B5698D3824FFBC4FC20FBAB" box="[827,988,1083,1135]" gridcol="3" gridrow="15" pageId="15" pageNumber="16">(7.848.01)</td>
<td id="768B2D328B5698D385D7FBC4FAC0FBAB" box="[1187,1340,1083,1135]" gridcol="5" gridrow="15" pageId="15" pageNumber="16">(4.55.2)</td>
</tr>
<tr id="355A444E8B5698D38118FB81FA0FFB50" box="[108,1523,1150,1172]" gridrow="16" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FB81FEC1FB50" box="[108,317,1150,1172]" gridcol="0" gridrow="16" pageId="15" pageNumber="16">Dominant frequency</th>
<td id="768B2D328B5698D38039FB81FDD0FB50" box="[333,556,1150,1172]" gridcol="1" gridrow="16" pageId="15" pageNumber="16">8.22 ± 0.10</td>
<td id="768B2D328B5698D38336FB81FCD8FB50" box="[578,804,1150,1172]" gridcol="2" gridrow="16" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D3824FFB81FC20FB50" box="[827,988,1150,1172]" gridcol="3" gridrow="16" pageId="15" pageNumber="16">8.38 ± 0.18</td>
<td id="768B2D328B5698D38298FB81FB6FFB50" box="[1004,1171,1150,1172]" gridcol="4" gridrow="16" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D385D7FB81FAC0FB50" box="[1187,1340,1150,1172]" gridcol="5" gridrow="16" pageId="15" pageNumber="16">6.6 ± 0.1</td>
<td id="768B2D328B5698D38420FB81FA0FFB50" box="[1364,1523,1150,1172]" gridcol="6" gridrow="16" pageId="15" pageNumber="16">-</td>
</tr>
<tr id="355A444E8B5698D38118FB63FA0FFB7E" box="[108,1523,1180,1210]" gridrow="17" pageId="15" pageNumber="16" rowspan-2="1" rowspan-4="1" rowspan-6="1">
<th id="768B2D328B5698D38118FB63FEC1FB7E" box="[108,317,1180,1210]" gridcol="0" gridrow="17" pageId="15" pageNumber="16">(kHz)</th>
<td id="768B2D328B5698D38039FB63FDD0FB7E" box="[333,556,1180,1210]" gridcol="1" gridrow="17" pageId="15" pageNumber="16">(8.018.44)</td>
<td id="768B2D328B5698D3824FFB63FC20FB7E" box="[827,988,1180,1210]" gridcol="3" gridrow="17" pageId="15" pageNumber="16">(8.188.61)</td>
<td id="768B2D328B5698D385D7FB63FAC0FB7E" box="[1187,1340,1180,1210]" gridcol="5" gridrow="17" pageId="15" pageNumber="16">(6.56.7)</td>
</tr>
<tr id="355A444E8B5698D38118FB35FA0FFB24" box="[108,1523,1226,1248]" gridrow="18" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FB35FEC1FB24" box="[108,317,1226,1248]" gridcol="0" gridrow="18" pageId="15" pageNumber="16">Maximum frequency</th>
<td id="768B2D328B5698D38039FB35FDD0FB24" box="[333,556,1226,1248]" gridcol="1" gridrow="18" pageId="15" pageNumber="16">8.62 ± 0.03</td>
<td id="768B2D328B5698D38336FB35FCD8FB24" box="[578,804,1226,1248]" gridcol="2" gridrow="18" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D3824FFB35FC20FB24" box="[827,988,1226,1248]" gridcol="3" gridrow="18" pageId="15" pageNumber="16">8.91 ± 0.19</td>
<td id="768B2D328B5698D38298FB35FB6FFB24" box="[1004,1171,1226,1248]" gridcol="4" gridrow="18" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D385D7FB35FAC0FB24" box="[1187,1340,1226,1248]" gridcol="5" gridrow="18" pageId="15" pageNumber="16">9.1 ± 0.2</td>
<td id="768B2D328B5698D38420FB35FA0FFB24" box="[1364,1523,1226,1248]" gridcol="6" gridrow="18" pageId="15" pageNumber="16">-</td>
</tr>
<tr id="355A444E8B5698D38118FB18FA0FFADE" box="[108,1523,1255,1306]" gridrow="19" pageId="15" pageNumber="16" rowspan-2="1" rowspan-4="1" rowspan-6="1">
<th id="768B2D328B5698D38118FB18FEC1FADE" box="[108,317,1255,1306]" gridcol="0" gridrow="19" pageId="15" pageNumber="16">(Frequency 95%) (kHz)</th>
<td id="768B2D328B5698D38039FB18FDD0FADE" box="[333,556,1255,1306]" gridcol="1" gridrow="19" pageId="15" pageNumber="16">(8.448.96)</td>
<td id="768B2D328B5698D3824FFB18FC20FADE" box="[827,988,1255,1306]" gridcol="3" gridrow="19" pageId="15" pageNumber="16">(8.709.13)</td>
<td id="768B2D328B5698D385D7FB18FAC0FADE" box="[1187,1340,1255,1306]" gridcol="5" gridrow="19" pageId="15" pageNumber="16">(9.010.7)</td>
</tr>
<tr id="355A444E8B5698D38118FAD6FA0FFA84" box="[108,1523,1321,1344]" gridrow="20" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FAD6FEC1FA84" box="[108,317,1321,1344]" gridcol="0" gridrow="20" pageId="15" pageNumber="16">Frequency bandwidth</th>
<td id="768B2D328B5698D38039FAD6FDD0FA84" box="[333,556,1321,1344]" gridcol="1" gridrow="20" pageId="15" pageNumber="16">1.35 ± 0.12</td>
<td id="768B2D328B5698D38336FAD6FCD8FA84" box="[578,804,1321,1344]" gridcol="2" gridrow="20" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D3824FFAD6FC20FA84" box="[827,988,1321,1344]" gridcol="3" gridrow="20" pageId="15" pageNumber="16">0.97 ± 0.18</td>
<td id="768B2D328B5698D38298FAD6FB6FFA84" box="[1004,1171,1321,1344]" gridcol="4" gridrow="20" pageId="15" pageNumber="16">-</td>
<td id="768B2D328B5698D385D7FAD6FAC0FA84" box="[1187,1340,1321,1344]" gridcol="5" gridrow="20" pageId="15" pageNumber="16">n/a</td>
<td id="768B2D328B5698D38420FAD6FA0FFA84" box="[1364,1523,1321,1344]" gridcol="6" gridrow="20" pageId="15" pageNumber="16">-</td>
</tr>
<tr id="355A444E8B5698D38118FAB8FA0FFAA1" box="[108,1523,1351,1381]" gridrow="21" pageId="15" pageNumber="16" rowspan-2="1" rowspan-4="1" rowspan-5="1" rowspan-6="1">
<th id="768B2D328B5698D38118FAB8FEC1FAA1" box="[108,317,1351,1381]" gridcol="0" gridrow="21" pageId="15" pageNumber="16">(90%) (kHz)</th>
<td id="768B2D328B5698D38039FAB8FDD0FAA1" box="[333,556,1351,1381]" gridcol="1" gridrow="21" pageId="15" pageNumber="16">(0.941.89)</td>
<td id="768B2D328B5698D3824FFAB8FC20FAA1" box="[827,988,1351,1381]" gridcol="3" gridrow="21" pageId="15" pageNumber="16">(0.781.29)</td>
</tr>
<tr id="355A444E8B5698D38118FA8AFA0FFA63" box="[108,1523,1397,1447]" gridrow="22" pageId="15" pageNumber="16">
<th id="768B2D328B5698D38118FA8AFEC1FA63" box="[108,317,1397,1447]" gridcol="0" gridrow="22" pageId="15" pageNumber="16">References</th>
<td id="768B2D328B5698D38039FA8AFDD0FA63" box="[333,556,1397,1447]" gridcol="1" gridrow="22" pageId="15" pageNumber="16">Present study</td>
<td id="768B2D328B5698D38336FA8AFCD8FA63" box="[578,804,1397,1447]" gridcol="2" gridrow="22" pageId="15" pageNumber="16">
<bibRefCitation id="EFFB3BFD8B5667238336FA8AFCD8FA4E" author="Bornschein MR &amp; Ribeiro LF &amp; Teixeira L &amp; Belmonte-Lopes R &amp; Moraes LA &amp; de Correa L &amp; Mauricio GN &amp; Pie MR" box="[578,804,1397,1418]" journalOrPublisher="PeerJ" pageId="15" pageNumber="16" part="9" publicationUrl="10.7717/peerj.10983" refId="ref12763" refString="Bornschein MR, Ribeiro LF, Teixeira L, Belmonte-Lopes R, Moraes LA, de Correa L, Mauricio GN, Nadaline Ι, Pie MR. 2021. A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae). PeerJ 9: e 10983 DOI 10.7717 / peerj. 10983." title="A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae)" type="journal volume" year="2021">
<emphasis id="B91E9A1E8B5667238336FA8AFCD8FA4E" box="[578,804,1397,1418]" italics="true" pageId="15" pageNumber="16">Bornschein et al. (2021)</emphasis>
</bibRefCitation>
</td>
<td id="768B2D328B5698D3824FFA8AFC20FA63" box="[827,988,1397,1447]" gridcol="3" gridrow="22" pageId="15" pageNumber="16">Present study</td>
<td id="768B2D328B5698D38298FA8AFB6FFA63" box="[1004,1171,1397,1447]" gridcol="4" gridrow="22" pageId="15" pageNumber="16">Bornschein et al. (2021)</td>
<td id="768B2D328B5698D385D7FA8AFAC0FA63" box="[1187,1340,1397,1447]" gridcol="5" gridrow="22" pageId="15" pageNumber="16">Condez et al. (2016)</td>
<td id="768B2D328B5698D38420FA8AFA0FFA63" box="[1364,1523,1397,1447]" gridcol="6" gridrow="22" pageId="15" pageNumber="16">
<emphasis id="B91E9A1E8B5667238420FA8AFA5CFA63" italics="true" pageId="15" pageNumber="16">
<bibRefCitation id="EFFB3BFD8B5667238420FA8AFA0FFA4E" author="Bornschein MR &amp; Ribeiro LF &amp; Teixeira L &amp; Belmonte-Lopes R &amp; Moraes LA &amp; de Correa L &amp; Mauricio GN &amp; Pie MR" box="[1364,1523,1397,1418]" journalOrPublisher="PeerJ" pageId="15" pageNumber="16" part="9" publicationUrl="10.7717/peerj.10983" refId="ref12763" refString="Bornschein MR, Ribeiro LF, Teixeira L, Belmonte-Lopes R, Moraes LA, de Correa L, Mauricio GN, Nadaline Ι, Pie MR. 2021. A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae). PeerJ 9: e 10983 DOI 10.7717 / peerj. 10983." title="A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae)" type="journal volume" year="2021">Bornschein et al.</bibRefCitation>
(2021)
</emphasis>
</td>
</tr>
</table>
</paragraph>
<caption id="DF1516848B49673C8100FD1DFA17FCA9" pageId="16" pageNumber="17" startId="16.[116,181,738,760]" targetBox="[90,1541,234,725]" targetPageId="16" targetType="figure">
<paragraph id="8BD5460C8B49673C8100FD1DFA17FCA9" blockId="16.[116,1515,738,877]" pageId="16" pageNumber="17">
<emphasis id="B91E9A1E8B49673C8100FD1DFD58FD3C" bold="true" box="[116,676,738,760]" pageId="16" pageNumber="17">
Figure 7 Selected skulls of other
<taxonomicName id="4C6A3D8F8B49673C80CEFD1DFD58FD3C" box="[442,676,738,760]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="undetermined">
<emphasis id="B91E9A1E8B49673C80CEFD1DFDAFFD3C" bold="true" box="[442,595,738,760]" italics="true" pageId="16" pageNumber="17">Brachycephalus</emphasis>
species.
</taxonomicName>
</emphasis>
Cranial elements labeled are discussed in comparisons of
<taxonomicName id="4C6A3D8F8B49673C85BFFD1CFADCFD33" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[1227,1312,738,760]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C85BFFD1CFADCFD33" box="[1227,1312,738,760]" italics="true" pageId="16" pageNumber="17">B. dacnis</emphasis>
</taxonomicName>
to other species. Top row is the leptodactyliform “flea toads”, including
<taxonomicName id="4C6A3D8F8B49673C833FFCFFFD2BFCD1" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[587,727,768,789]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B49673C833FFCFFFD2BFCD1" box="[587,727,768,789]" italics="true" pageId="16" pageNumber="17">B. hermogenesi</emphasis>
</taxonomicName>
(sister species of
<taxonomicName id="4C6A3D8F8B49673C820DFCFFFC31FCD1" authorityName="Toledo &amp; Botelho &amp; Carrasco-Medina &amp; Gray &amp; Ernetti &amp; Gama &amp; Lyra &amp; Blackburn &amp; Nunes &amp; Muscat" authorityYear="2024" box="[889,973,768,789]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C820DFCFFFC31FCD1" box="[889,973,768,789]" italics="true" pageId="16" pageNumber="17">B. dacnis</emphasis>
</taxonomicName>
),
<taxonomicName id="4C6A3D8F8B49673C82ABFCFFFBBAFCD1" box="[991,1094,768,789]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="clarissae">
<emphasis id="B91E9A1E8B49673C82ABFCFFFBBAFCD1" box="[991,1094,768,789]" italics="true" pageId="16" pageNumber="17">B. clarissae</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C8524FCFFFB3AFCD1" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[1104,1222,768,789]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B49673C8524FCFFFB3AFCD1" box="[1104,1222,768,789]" italics="true" pageId="16" pageNumber="17">B. sulfuratus</emphasis>
</taxonomicName>
, and
<taxonomicName id="4C6A3D8F8B49673C858CFCFFFA8CFCD1" baseAuthorityName="Izecksohn" baseAuthorityYear="1971" box="[1272,1392,768,789]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="didactylus">
<emphasis id="B91E9A1E8B49673C858CFCFFFA8CFCD1" box="[1272,1392,768,789]" italics="true" pageId="16" pageNumber="17">B. didactylus</emphasis>
</taxonomicName>
. Bottom row contains representatives of the three named species groups:
<taxonomicName id="4C6A3D8F8B49673C83D1FCE2FD1DFCF6" authorityName="Pombal, Wistuba &amp; Bornschein" authorityYear="1998" box="[677,737,797,818]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="pernix">
<emphasis id="B91E9A1E8B49673C83D1FCE2FD1DFCF6" box="[677,737,797,818]" italics="true" pageId="16" pageNumber="17">pernix</emphasis>
</taxonomicName>
group (
<taxonomicName id="4C6A3D8F8B49673C825FFCE1FC49FCF6" authorityName="Bornschein, Ribeiro, Blackburn, Stanley &amp; Pie" authorityYear="2016" box="[811,949,797,819]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="albolineatus">
<emphasis id="B91E9A1E8B49673C825FFCE1FC49FCF6" box="[811,949,797,819]" italics="true" pageId="16" pageNumber="17">B. albolineatus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C82B4FCE1FBE8FCF6" authorityName="Pombal, Wistuba &amp; Bornschein" authorityYear="1998" box="[960,1044,797,819]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="pernix">
<emphasis id="B91E9A1E8B49673C82B4FCE1FBE8FCF6" box="[960,1044,797,819]" italics="true" pageId="16" pageNumber="17">B. pernix</emphasis>
</taxonomicName>
),
<taxonomicName id="4C6A3D8F8B49673C8552FCE2FB7BFCF6" authorityName="Pombal" authorityYear="2001" box="[1062,1159,797,818]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="vertebralis">
<emphasis id="B91E9A1E8B49673C8552FCE2FB7BFCF6" box="[1062,1159,797,818]" italics="true" pageId="16" pageNumber="17">vertebralis</emphasis>
</taxonomicName>
group (
<taxonomicName id="4C6A3D8F8B49673C85A6FCE1FAB7FCF6" authorityName="Pombal" authorityYear="2001" box="[1234,1355,797,819]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="vertebralis">
<emphasis id="B91E9A1E8B49673C85A6FCE1FAB7FCF6" box="[1234,1355,797,819]" italics="true" pageId="16" pageNumber="17">B. vertebralis</emphasis>
</taxonomicName>
), and
<taxonomicName id="4C6A3D8F8B49673C84F3FCE2FA17FCF6" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[1415,1515,797,818]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="B91E9A1E8B49673C84F3FCE2FA17FCF6" box="[1415,1515,797,818]" italics="true" pageId="16" pageNumber="17">ephippium</emphasis>
</taxonomicName>
group (
<taxonomicName id="4C6A3D8F8B49673C81CEFCC4FECBFC8B" baseAuthorityName="Spix" baseAuthorityYear="1824" box="[186,311,826,848]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="ephippium">
<emphasis id="B91E9A1E8B49673C81CEFCC4FECBFC8B" box="[186,311,826,848]" italics="true" pageId="16" pageNumber="17">B. ephippium</emphasis>
</taxonomicName>
). Museum catalog numbers are listed below each species name. Abbreviations: fp, frontoparietal; n, nasal; ps, parasphenoid; pt, pterygoid; qj, quadratojugal; sp, sphenethmoid; sq, squamosal; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-7
</paragraph>
</caption>
<paragraph id="8BD5460C8B49673D8086F916FC0CFDDF" blockId="16.[498,1542,932,1915]" lastBlockId="17.[498,1542,234,1098]" lastPageId="17" lastPageNumber="18" pageId="16" pageNumber="17">
Although this trait varies in
<taxonomicName id="4C6A3D8F8B49673C8240F915FC38F8C7" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[820,964,1769,1795]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B49673C8240F915FC38F8C7" box="[820,964,1769,1795]" italics="true" pageId="16" pageNumber="17">B. sulfuratus</emphasis>
</taxonomicName>
(see
<bibRefCitation id="EFFB3BFD8B49673C8288F916FB07F8C0" author="Bornschein MR &amp; Ribeiro LF &amp; Teixeira L &amp; Belmonte-Lopes R &amp; Moraes LA &amp; de Correa L &amp; Mauricio GN &amp; Pie MR" box="[1020,1275,1769,1796]" journalOrPublisher="PeerJ" pageId="16" pageNumber="17" part="9" publicationUrl="10.7717/peerj.10983" refId="ref12763" refString="Bornschein MR, Ribeiro LF, Teixeira L, Belmonte-Lopes R, Moraes LA, de Correa L, Mauricio GN, Nadaline Ι, Pie MR. 2021. A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae). PeerJ 9: e 10983 DOI 10.7717 / peerj. 10983." title="A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae)" type="journal volume" year="2021">
<emphasis id="B91E9A1E8B49673C8288F916FB07F8C0" box="[1020,1275,1769,1796]" italics="true" pageId="16" pageNumber="17">Bornschein et al., 2021</emphasis>
</bibRefCitation>
), it still can distinguish some individuals. From
<taxonomicName id="4C6A3D8F8B49673C8265F8EDFC58F8EF" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[785,932,1809,1835]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B49673C8265F8EDFC58F8EF" box="[785,932,1809,1835]" italics="true" pageId="16" pageNumber="17">B. sulfuratus</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C82C7F8EDFBE0F8EF" box="[947,1052,1809,1835]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C82C7F8EDFBE0F8EF" box="[947,1052,1809,1835]" italics="true" pageId="16" pageNumber="17">B. dacnis</emphasis>
</taxonomicName>
is distinguished by its thin quadratojugal (
<figureCitation id="13515A898B49673C8088F8C6FDC2F890" box="[508,574,1849,1876]" captionStart="Figure 7" captionStartId="16.[116,181,738,760]" captionTargetBox="[90,1541,234,725]" captionTargetId="figure-414@16.[90,1512,234,710]" captionTargetPageId="16" captionText="Figure 7 Selected skulls of other Brachycephalus species. Cranial elements labeled are discussed in comparisons of B. dacnis to other species. Top row is the leptodactyliform “flea toads”, including B. hermogenesi (sister species of B. dacnis), B. clarissae, B. sulfuratus, and B. didactylus. Bottom row contains representatives of the three named species groups: pernix group (B.albolineatus, B. pernix), vertebralis group (B. vertebralis), and ephippium group (B. ephippium). Museum catalog numbers are listed below each species name. Abbreviations: fp, frontoparietal; n, nasal; ps, parasphenoid; pt, pterygoid; qj, quadratojugal; sp, sphenethmoid; sq, squamosal; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-7" pageId="16" pageNumber="17">Fig. 7</figureCitation>
), and medially expanded coracoid. From
<taxonomicName id="4C6A3D8F8B49673C8558F8C5FB81F897" authorityName="Almeida-Silva, Silva-Soares, Rodrigues &amp; Verdade" authorityYear="2021" box="[1068,1149,1850,1875]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="puri">
<emphasis id="B91E9A1E8B49673C8558F8C5FB81F897" box="[1068,1149,1850,1875]" italics="true" pageId="16" pageNumber="17">B. puri</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B49673C85F8F8C5FB09F897" box="[1164,1269,1849,1875]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B49673C85F8F8C5FB09F897" box="[1164,1269,1849,1875]" italics="true" pageId="16" pageNumber="17">B. dacnis</emphasis>
</taxonomicName>
is distinguished by having a distinct toe II (reduced in
<taxonomicName id="4C6A3D8F8B49673C82F3F89DFC29F8BF" authorityName="Almeida-Silva, Silva-Soares, Rodrigues &amp; Verdade" authorityYear="2021" box="[903,981,1890,1915]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="16" pageNumber="17" phylum="Chordata" rank="species" species="puri">
<emphasis id="B91E9A1E8B49673C82F3F89DFC29F8BF" box="[903,981,1890,1915]" italics="true" pageId="16" pageNumber="17">B. puri</emphasis>
</taxonomicName>
), by its dark black or pale brown marbled venter with small white blotches (uniformly brown in
<taxonomicName id="4C6A3D8F8B48673D8556FF14FB8EFEC0" authorityName="Almeida-Silva, Silva-Soares, Rodrigues &amp; Verdade" authorityYear="2021" box="[1058,1138,235,260]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="puri">
<emphasis id="B91E9A1E8B48673D8556FF14FB8EFEC0" box="[1058,1138,235,260]" italics="true" pageId="17" pageNumber="18">B. puri</emphasis>
</taxonomicName>
), and by sometimes having an X-shaped dorsal mark (inverted V-shaped in
<taxonomicName id="4C6A3D8F8B48673D8575FEECFBB3FEE8" authorityName="Almeida-Silva, Silva-Soares, Rodrigues &amp; Verdade" authorityYear="2021" box="[1025,1103,275,300]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="puri">
<emphasis id="B91E9A1E8B48673D8575FEECFBB3FEE8" box="[1025,1103,275,300]" italics="true" pageId="17" pageNumber="18">B. puri</emphasis>
</taxonomicName>
) (
<bibRefCitation id="EFFB3BFD8B48673D851FFEEDFA6CFEE8" author="Almeida-Silva D &amp; Silva-Soares T &amp; Rodrigues MT &amp; Verdade VK" box="[1131,1424,274,300]" journalOrPublisher="Zootaxa" pageId="17" pageNumber="18" pagination="517 - 532" part="5068" publicationUrl="10.11646/zootaxa.5068.4.3" refId="ref12432" refString="Almeida-Silva D, Silva-Soares T, Rodrigues MT, Verdade VK. 2021. New species of flea-toad, genus Brachycephalus (Anura: Brachycephalidae) from the Atlantic Forest of Espirito Santo, Brazil. Zootaxa 5068: 517 - 532 DOI 10.11646 / zootaxa. 5068.4.3." title="New species of flea-toad, genus Brachycephalus (Anura: Brachycephalidae) from the Atlantic Forest of Espirito Santo, Brazil" type="journal article" year="2021">
<emphasis id="B91E9A1E8B48673D851FFEEDFA6CFEE8" box="[1131,1424,274,300]" italics="true" pageId="17" pageNumber="18">Almeida-Silva et al., 2021</emphasis>
</bibRefCitation>
). From
<taxonomicName id="4C6A3D8F8B48673D849AFEECFDCCFE90" authorityName="Napoli, Caramaschi, Cruz &amp; Dias" authorityYear="2011" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="pulex">
<emphasis id="B91E9A1E8B48673D849AFEECFDCCFE90" italics="true" pageId="17" pageNumber="18">B. pulex</emphasis>
</taxonomicName>
,
<taxonomicName id="4C6A3D8F8B48673D834BFEC4FD5BFE90" box="[575,679,314,340]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D834BFEC4FD5BFE90" box="[575,679,314,340]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
is distinguished by the presence of vestigial fingers Iand IV (absent in
<taxonomicName id="4C6A3D8F8B48673D849AFEC4FDCCFEB8" authorityName="Napoli, Caramaschi, Cruz &amp; Dias" authorityYear="2011" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="pulex">
<emphasis id="B91E9A1E8B48673D849AFEC4FDCCFEB8" italics="true" pageId="17" pageNumber="18">B. pulex</emphasis>
</taxonomicName>
), the presence of distinct and functional toe II and V (absent or vestigial, respectively, in
<taxonomicName id="4C6A3D8F8B48673D8365FE75FD92FE60" authorityName="Napoli, Caramaschi, Cruz &amp; Dias" authorityYear="2011" box="[529,622,394,420]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="pulex">
<emphasis id="B91E9A1E8B48673D8365FE75FD92FE60" box="[529,622,394,420]" italics="true" pageId="17" pageNumber="18">B. pulex</emphasis>
</taxonomicName>
), and lacking a chest mark (presence of an inverted V-shaped chest mark in
<taxonomicName id="4C6A3D8F8B48673D849AFE75FDCCFE0F" authorityName="Napoli, Caramaschi, Cruz &amp; Dias" authorityYear="2011" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="pulex">
<emphasis id="B91E9A1E8B48673D849AFE75FDCCFE0F" italics="true" pageId="17" pageNumber="18">B. pulex</emphasis>
</taxonomicName>
) (
<bibRefCitation id="EFFB3BFD8B48673D8339FE4EFC84FE08" author="Almeida-Silva D &amp; Silva-Soares T &amp; Rodrigues MT &amp; Verdade VK" box="[589,888,433,460]" journalOrPublisher="Zootaxa" pageId="17" pageNumber="18" pagination="517 - 532" part="5068" publicationUrl="10.11646/zootaxa.5068.4.3" refId="ref12432" refString="Almeida-Silva D, Silva-Soares T, Rodrigues MT, Verdade VK. 2021. New species of flea-toad, genus Brachycephalus (Anura: Brachycephalidae) from the Atlantic Forest of Espirito Santo, Brazil. Zootaxa 5068: 517 - 532 DOI 10.11646 / zootaxa. 5068.4.3." title="New species of flea-toad, genus Brachycephalus (Anura: Brachycephalidae) from the Atlantic Forest of Espirito Santo, Brazil" type="journal article" year="2021">
<emphasis id="B91E9A1E8B48673D8339FE4EFC84FE08" box="[589,888,433,460]" italics="true" pageId="17" pageNumber="18">Almeida-Silva et al., 2021</emphasis>
</bibRefCitation>
).
<taxonomicName id="4C6A3D8F8B48673D82E5FE4EFB68FE0F" box="[913,1172,433,459]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D82E5FE4EFB68FE0F" box="[913,1172,433,459]" italics="true" pageId="17" pageNumber="18">Brachycephalusdacnis</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6A3D8F8B48673D85A6FE4DFA7CFE0F" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[1234,1408,433,459]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B48673D85A6FE4DFA7CFE0F" box="[1234,1408,433,459]" italics="true" pageId="17" pageNumber="18">B. hermogenesi</emphasis>
</taxonomicName>
are morphologically similar, including in their skeletons (
<figureCitation id="13515A898B48673D8513FE26FB4EFE30" box="[1127,1202,473,500]" captionStart="Figure 5" captionStartId="12.[116,181,1269,1291]" captionTargetBox="[93,1510,238,1238]" captionTargetId="figure-137@12.[90,1512,234,1241]" captionTargetPageId="12" captionText="Figure 5 Skeletal anatomy of Brachycephalus dacnis (ZUEC-AMP 24978; undetermined sex) based on CT-scans. (A) The entire skeleton, with insets of (B) the skull (in dorsal, ventral, and right lateral views), (C) right hand and (D) left foot (both in dorsal view), (E) vertebral column (in dorsal view), (F) pectoral girdle (in ventral view), and (G) pelvic girdle (in right lateral view). Abbreviations: c, carpus; cl, clavicle; cor, coracoid; dt, distal tarsals; ex, exoccipital; fp, frontoparietal; il, ilium; isc, ischium; m, maxilla; mdI, manual digit I; mdV, manual digit V; min, mineralized cartilage; n, nasal; op, operculum; pd I, pedal digit I; pd V, pedal digit V; pm, premaxilla; pr, prootic; ps, parasphenoid; pt, pterygoid; pub, pubis; qj, quadratojugal; sac, sacrum; sc, scapula; ses, sesamoid; sm, septomaxilla; sp, sphenethmoid; sq, squamosal; ur, urostyle; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-5" pageId="17" pageNumber="18">Figs. 5</figureCitation>
,
<figureCitation id="13515A898B48673D85B4FE25FB32FE30" box="[1216,1230,474,500]" captionStart="Figure 7" captionStartId="16.[116,181,738,760]" captionTargetBox="[90,1541,234,725]" captionTargetId="figure-414@16.[90,1512,234,710]" captionTargetPageId="16" captionText="Figure 7 Selected skulls of other Brachycephalus species. Cranial elements labeled are discussed in comparisons of B. dacnis to other species. Top row is the leptodactyliform “flea toads”, including B. hermogenesi (sister species of B. dacnis), B. clarissae, B. sulfuratus, and B. didactylus. Bottom row contains representatives of the three named species groups: pernix group (B.albolineatus, B. pernix), vertebralis group (B. vertebralis), and ephippium group (B. ephippium). Museum catalog numbers are listed below each species name. Abbreviations: fp, frontoparietal; n, nasal; ps, parasphenoid; pt, pterygoid; qj, quadratojugal; sp, sphenethmoid; sq, squamosal; v, vomer. Full-size DOI: 10.7717/peerj.18265/fig-7" pageId="17" pageNumber="18">7</figureCitation>
), but can be recognized as distinct through acoustic and genetic traits.
</paragraph>
<paragraph id="8BD5460C8B48673D8366FDD6FA8AFB8E" blockId="17.[498,1542,234,1098]" pageId="17" pageNumber="18">
The advertisement call of
<taxonomicName id="4C6A3D8F8B48673D8231FDD5FC53FD87" box="[837,943,553,579]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D8231FDD5FC53FD87" box="[837,943,553,579]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
is distinguished from
<taxonomicName id="4C6A3D8F8B48673D85CDFDD5FA9BFD87" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[1209,1383,553,579]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B48673D85CDFDD5FA9BFD87" box="[1209,1383,553,579]" italics="true" pageId="17" pageNumber="18">B. hermogenesi</emphasis>
</taxonomicName>
and
<taxonomicName id="4C6A3D8F8B48673D84D0FDD5FA40FD87" box="[1444,1468,554,579]" pageId="17" pageNumber="18">
<emphasis id="B91E9A1E8B48673D84D0FDD5FA40FD87" box="[1444,1468,554,579]" italics="true" pageId="17" pageNumber="18">B.</emphasis>
</taxonomicName>
<emphasis id="B91E9A1E8B48673D8086FDAEFD99FDAF" box="[498,613,593,619]" italics="true" pageId="17" pageNumber="18">sulfuratus</emphasis>
by generally having fewer notes per call (one or two notes per call in
<taxonomicName id="4C6A3D8F8B48673D84E3FDADFA03FDAF" box="[1431,1535,593,619]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D84E3FDADFA03FDAF" box="[1431,1535,593,619]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
,
<quantity id="4C92EBE98B48673D8086FD86FDC1FD57" box="[498,573,633,659]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" metricValueMax="1.524" metricValueMin="0.508" pageId="17" pageNumber="18" unit="in" value="4.0" valueMax="6.0" valueMin="2.0">26 in</quantity>
<taxonomicName id="4C6A3D8F8B48673D8332FD86FD08FD57" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[582,756,633,659]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B48673D8332FD86FD08FD57" box="[582,756,633,659]" italics="true" pageId="17" pageNumber="18">B. hermogenesi</emphasis>
</taxonomicName>
,
<quantity id="4C92EBE98B48673D8277FD86FCB2FD57" box="[771,846,633,659]" metricMagnitude="-1" metricUnit="m" metricValue="1.27" metricValueMax="1.524" metricValueMin="1.016" pageId="17" pageNumber="18" unit="in" value="5.0" valueMax="6.0" valueMin="4.0">46 in</quantity>
<taxonomicName id="4C6A3D8F8B48673D8223FD86FC16FD57" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[855,1002,633,659]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B48673D8223FD86FC16FD57" box="[855,1002,633,659]" italics="true" pageId="17" pageNumber="18">B. sulfuratus</emphasis>
</taxonomicName>
), shorter note duration (≤
<geoCoordinate id="EE5E20CB8B48673D8457FD86FA9BFD57" box="[1315,1383,633,659]" degrees="0.08" direction="south" orientation="latitude" pageId="17" pageNumber="18" precision="555" value="-0.08">0.08 s</geoCoordinate>
in
<taxonomicName id="4C6A3D8F8B48673D84E4FD86FA05FD57" box="[1424,1529,633,659]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D84E4FD86FA05FD57" box="[1424,1529,633,659]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
; ≥
<geoCoordinate id="EE5E20CB8B48673D8377FD5EFDB9FD7F" box="[515,581,673,699]" degrees="0.17" direction="south" orientation="latitude" pageId="17" pageNumber="18" precision="555" value="-0.17">0.17 s</geoCoordinate>
in
<taxonomicName id="4C6A3D8F8B48673D831DFD5EFCE9FD7F" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[617,789,673,699]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B48673D831DFD5EFCE9FD7F" box="[617,789,673,699]" italics="true" pageId="17" pageNumber="18">B. hermogenesi</emphasis>
</taxonomicName>
, and ≥
<geoCoordinate id="EE5E20CB8B48673D8212FD5FFC66FD7F" box="[870,922,672,699]" degrees="1.5" direction="south" orientation="latitude" pageId="17" pageNumber="18" precision="5555" value="-1.5">1.5 s</geoCoordinate>
in
<taxonomicName id="4C6A3D8F8B48673D82CAFD5EFBB3FD7F" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[958,1103,673,699]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B48673D82CAFD5EFBB3FD7F" box="[958,1103,673,699]" italics="true" pageId="17" pageNumber="18">B. sulfuratus</emphasis>
</taxonomicName>
), and shorter call duration (≤
<geoCoordinate id="EE5E20CB8B48673D84D2FD5EFA14FD7F" box="[1446,1512,673,699]" degrees="0.08" direction="south" orientation="latitude" pageId="17" pageNumber="18" precision="555" value="-0.08">0.08 s</geoCoordinate>
in
<taxonomicName id="4C6A3D8F8B48673D8086FD36FDA7FD26" box="[498,603,712,738]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D8086FD36FDA7FD26" box="[498,603,712,738]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
in calls with one note and ≤
<geoCoordinate id="EE5E20CB8B48673D82C5FD37FC09FD27" box="[945,1013,712,739]" degrees="0.41" direction="south" orientation="latitude" pageId="17" pageNumber="18" precision="555" value="-0.41">0.41 s</geoCoordinate>
in calls with two notes; ≥
<geoCoordinate id="EE5E20CB8B48673D8452FD37FA97FD27" box="[1318,1387,712,739]" degrees="0.41" direction="south" orientation="latitude" pageId="17" pageNumber="18" precision="555" value="-0.41">0.41 s</geoCoordinate>
in
<taxonomicName id="4C6A3D8F8B48673D84E7FD36FA57FD26" box="[1427,1451,713,738]" pageId="17" pageNumber="18">
<emphasis id="B91E9A1E8B48673D84E7FD36FA57FD26" box="[1427,1451,713,738]" italics="true" pageId="17" pageNumber="18">B.</emphasis>
</taxonomicName>
<emphasis id="B91E9A1E8B48673D8086FD0FFD7CFCCE" box="[498,640,752,778]" italics="true" pageId="17" pageNumber="18">hermogenesi</emphasis>
, and ≥
<geoCoordinate id="EE5E20CB8B48673D83A6FD0FFCE9FCCF" box="[722,789,752,779]" degrees="0.13" direction="south" orientation="latitude" pageId="17" pageNumber="18" precision="555" value="-0.13">0.13 s</geoCoordinate>
in
<taxonomicName id="4C6A3D8F8B48673D824FFD0EFC31FCCE" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[827,973,752,778]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B48673D824FFD0EFC31FCCE" box="[827,973,752,778]" italics="true" pageId="17" pageNumber="18">B. sulfuratus</emphasis>
</taxonomicName>
).
<taxonomicName id="4C6A3D8F8B48673D8290FD0FFB1BFCCE" box="[996,1255,752,778]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D8290FD0FFB1BFCCE" box="[996,1255,752,778]" italics="true" pageId="17" pageNumber="18">Brachycephalus dacnis</emphasis>
</taxonomicName>
generally presents fewer pulses per note (37) than
<taxonomicName id="4C6A3D8F8B48673D8246FCE6FC3AFCF6" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[818,966,792,818]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B48673D8246FCE6FC3AFCF6" box="[818,966,792,818]" italics="true" pageId="17" pageNumber="18">B. sulfuratus</emphasis>
</taxonomicName>
(711;
<bibRefCitation id="EFFB3BFD8B48673D8554FCE7FB02FCF6" author="Condez TH &amp; Garcia PCA &amp; Amaral IB &amp; Haddad CFB" box="[1056,1278,792,818]" journalOrPublisher="Zootaxa" pageId="17" pageNumber="18" pagination="40 - 56" part="4083" publicationUrl="10.11646/zootaxa.4083.1.2" refId="ref13218" refString="Condez TH, Monteiro ΙPC, Comitti EΙ, Garcia PCA, Amaral IB, Haddad CFB. 2016. A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil. Zootaxa 4083: 40 - 56 DOI 10.11646 / zootaxa. 4083.1.2." title="A new species of flea-toad (Anura: Brachycephalidae) from southern Atlantic Forest, Brazil" type="journal article" year="2016">
<emphasis id="B91E9A1E8B48673D8554FCE7FB02FCF6" box="[1056,1278,792,818]" italics="true" pageId="17" pageNumber="18">Condez et al., 2016</emphasis>
</bibRefCitation>
), and higher than
<taxonomicName id="4C6A3D8F8B48673D84A9FCE6FA09FCF6" box="[1501,1525,793,818]" pageId="17" pageNumber="18">
<emphasis id="B91E9A1E8B48673D84A9FCE6FA09FCF6" box="[1501,1525,793,818]" italics="true" pageId="17" pageNumber="18">B.</emphasis>
</taxonomicName>
<emphasis id="B91E9A1E8B48673D8086FCBFFD7CFC9E" box="[498,640,832,858]" italics="true" pageId="17" pageNumber="18">hermogenesi</emphasis>
(13) (see also
<bibRefCitation id="EFFB3BFD8B48673D8246FCBFFBCDFC9E" author="Bornschein MR &amp; Ribeiro LF &amp; Teixeira L &amp; Belmonte-Lopes R &amp; Moraes LA &amp; de Correa L &amp; Mauricio GN &amp; Pie MR" box="[818,1073,832,858]" journalOrPublisher="PeerJ" pageId="17" pageNumber="18" part="9" publicationUrl="10.7717/peerj.10983" refId="ref12763" refString="Bornschein MR, Ribeiro LF, Teixeira L, Belmonte-Lopes R, Moraes LA, de Correa L, Mauricio GN, Nadaline Ι, Pie MR. 2021. A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae). PeerJ 9: e 10983 DOI 10.7717 / peerj. 10983." title="A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae)" type="journal volume" year="2021">
<emphasis id="B91E9A1E8B48673D8246FCBFFBCDFC9E" box="[818,1073,832,858]" italics="true" pageId="17" pageNumber="18">Bornschein et al., 2021</emphasis>
</bibRefCitation>
). The absence of attenuated notes (
<emphasis id="B91E9A1E8B48673D84BCFCBEFD0DFC46" italics="true" pageId="17" pageNumber="18">
sensu
<bibRefCitation id="EFFB3BFD8B48673D8086FC97FD0DFC46" author="Bornschein MR &amp; Ribeiro LF &amp; Teixeira L &amp; Belmonte-Lopes R &amp; Moraes LA &amp; de Correa L &amp; Mauricio GN &amp; Pie MR" box="[498,753,872,898]" journalOrPublisher="PeerJ" pageId="17" pageNumber="18" part="9" publicationUrl="10.7717/peerj.10983" refId="ref12763" refString="Bornschein MR, Ribeiro LF, Teixeira L, Belmonte-Lopes R, Moraes LA, de Correa L, Mauricio GN, Nadaline Ι, Pie MR. 2021. A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae). PeerJ 9: e 10983 DOI 10.7717 / peerj. 10983." title="A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae)" type="journal volume" year="2021">Bornschein et al., 2021</bibRefCitation>
</emphasis>
) in the calls of
<taxonomicName id="4C6A3D8F8B48673D82EAFC96FBF9FC46" box="[926,1029,872,898]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D82EAFC96FBF9FC46" box="[926,1029,872,898]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
, distinguishes it from
<taxonomicName id="4C6A3D8F8B48673D8470FC96FA53FC46" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[1284,1455,872,898]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B48673D8470FC96FA53FC46" box="[1284,1455,872,898]" italics="true" pageId="17" pageNumber="18">B. hermogenesi</emphasis>
</taxonomicName>
(which can present attenuated advertisement call notes) (see
<bibRefCitation id="EFFB3BFD8B48673D851DFC6FFA8CFC6E" author="Bornschein MR &amp; Ribeiro LF &amp; Teixeira L &amp; Belmonte-Lopes R &amp; Moraes LA &amp; de Correa L &amp; Mauricio GN &amp; Pie MR" box="[1129,1392,912,938]" journalOrPublisher="PeerJ" pageId="17" pageNumber="18" part="9" publicationUrl="10.7717/peerj.10983" refId="ref12763" refString="Bornschein MR, Ribeiro LF, Teixeira L, Belmonte-Lopes R, Moraes LA, de Correa L, Mauricio GN, Nadaline Ι, Pie MR. 2021. A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae). PeerJ 9: e 10983 DOI 10.7717 / peerj. 10983." title="A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae)" type="journal volume" year="2021">
<emphasis id="B91E9A1E8B48673D851DFC6FFA8CFC6E" box="[1129,1392,912,938]" italics="true" pageId="17" pageNumber="18">Bornschein et al., 2021</emphasis>
</bibRefCitation>
). The dominant frequency of the advertisement call of
<taxonomicName id="4C6A3D8F8B48673D8547FC47FB60FC15" box="[1075,1180,951,977]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D8547FC47FB60FC15" box="[1075,1180,951,977]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
overlaps with that of
<taxonomicName id="4C6A3D8F8B48673D84D5FC47FD7CFC3D" pageId="17" pageNumber="18">
<emphasis id="B91E9A1E8B48673D84D5FC47FA45FC15" box="[1441,1465,952,977]" italics="true" pageId="17" pageNumber="18">B.</emphasis>
<emphasis id="B91E9A1E8B48673D8086FC20FD7CFC3D" box="[498,640,991,1017]" italics="true" pageId="17" pageNumber="18">hermogenesi</emphasis>
</taxonomicName>
, but it is higher than that of
<taxonomicName id="4C6A3D8F8B48673D82AFFC1FFB92FC3D" authorityName="Condez, Monteiro, Comitti, Garcia, Amaral &amp; Haddad" authorityYear="2016" box="[987,1134,991,1017]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="sulfuratus">
<emphasis id="B91E9A1E8B48673D82AFFC1FFB92FC3D" box="[987,1134,991,1017]" italics="true" pageId="17" pageNumber="18">B. sulfuratus</emphasis>
</taxonomicName>
. However, the minimum and maximum frequencies of the call of
<taxonomicName id="4C6A3D8F8B48673D82E5FBF7FC04FBE5" box="[913,1016,1031,1057]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D82E5FBF7FC04FBE5" box="[913,1016,1031,1057]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
are lower and do not overlap with those of
<taxonomicName id="4C6A3D8F8B48673D849AFBF7FD7CFB8D" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B48673D849AFBF7FD7CFB8D" italics="true" pageId="17" pageNumber="18">B. hermogenesi</emphasis>
</taxonomicName>
. Call parameters of these three species are presented in
<tableCitation id="C6E873B78B48673D846CFBD0FA8CFB8D" box="[1304,1392,1071,1097]" captionStart="Table 2" captionStartId="15.[117,172,248,269]" captionTargetBox="[108,1523,319,1447]" captionTargetPageId="15" captionText="Table 2 Advertisement call traits described for three flea-toads species from Brachycephalus genus. Call traits present as mean ± standard deviation (range)." pageId="17" pageNumber="18">Table 2</tableCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="C37015878B48673E8086FB91FA2CFEE9" lastPageId="18" lastPageNumber="19" pageId="17" pageNumber="18" type="biology_ecology">
<paragraph id="8BD5460C8B48673D8086FB91FA2AF9A2" blockId="17.[498,1542,1133,1878]" pageId="17" pageNumber="18">
<emphasis id="B91E9A1E8B48673D8086FB91FD52FB43" bold="true" box="[498,686,1133,1160]" pageId="17" pageNumber="18">Natural history</emphasis>
We observed
<taxonomicName id="4C6A3D8F8B48673D822AFB92FB9EFB43" box="[862,1122,1133,1159]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D822AFB92FB9EFB43" box="[862,1122,1133,1159]" italics="true" pageId="17" pageNumber="18">Brachycephalus dacnis</emphasis>
</taxonomicName>
occupying areas with dense vegetation, tall canopy, reduced sunlight incidence, a thick layer of litter, and moist soil. These areas lie between small hills, forming a valley. The leaf litter along the slopes lies in small clusters at the base of trees and creeping plants. These areas have a small stream nearby with a substrate composed of rock and sand. Water flow intensity varies with precipitation and can soak the adjacent soil. The new species also occurs in an area characterized by thinner vegetation, mainly shrubs, with more solar incidence, and a thick litter layer, and consequently, less moisture. In these sites, finding frogs was difficult because of their small size, cryptic coloration, cryptozoic habits, and the behavior of ceasing calling with any disturbance in their surroundings. After thorough searches, we found the individuals either between juçara palm (
<emphasis id="B91E9A1E8B48673D853CFA02FB09F9D2" box="[1096,1269,1532,1558]" italics="true" pageId="17" pageNumber="18">Euterpes edulis</emphasis>
) roots or under rotten trunks. In these areas,
<taxonomicName id="4C6A3D8F8B48673D8389F9DAFC9BF9FA" box="[765,871,1572,1598]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D8389F9DAFC9BF9FA" box="[765,871,1572,1598]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
was syntopic with
<taxonomicName id="4C6A3D8F8B48673D853FF9DAFB05F9FA" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" box="[1099,1273,1572,1598]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B48673D853FF9DAFB05F9FA" box="[1099,1273,1572,1598]" italics="true" pageId="17" pageNumber="18">B. hermogenesi</emphasis>
</taxonomicName>
. When approached, some individuals occasionally exhibited mouth-gaping (
<figureCitation id="13515A898B48673D85F0F9B3FB25F9A2" box="[1156,1241,1612,1638]" captionStart="Figure 3" captionStartId="9.[524,589,900,922]" captionTargetBox="[498,1299,234,868]" captionTargetId="figure-274@9.[498,1299,234,872]" captionTargetPageId="9" captionText="Figure 3 Brachycephalus dacnis paratype individuals found at the Projeto Dacnis private reserve, municipality of Ubatuba, state of São Paulo, Brazil. (A) ZUEC-AMP 25272; (B) ZUEC-AMP 25274; (C and D) ZUEC-AMP 25275 displaying mouth-gaping behavior and the same individual on top of one herpetologists fingertip. Full-size DOI: 10.7717/peerj.18265/fig-3" pageId="17" pageNumber="18">Fig. 3C</figureCitation>
). We also observed a
</paragraph>
<paragraph id="8BD5460C8B48673D8086F98BFDD1F91A" blockId="17.[498,1542,1133,1878]" pageId="17" pageNumber="18">
<quantity id="4C92EBE98B48673D8086F98BFDB7F94A" box="[498,587,1652,1678]" metricMagnitude="-4" metricUnit="m" metricValue="7.0" pageId="17" pageNumber="18" unit="mm" value="0.7">0.7 mm</quantity>
<taxonomicName id="4C6A3D8F8B48673D8327F98BFC85F94A" box="[595,889,1652,1678]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" isUncertain="true" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D8327F98BFCF9F94A" box="[595,773,1652,1678]" italics="true" pageId="17" pageNumber="18">Brachycephalus</emphasis>
cf.
<emphasis id="B91E9A1E8B48673D8244F98BFC85F94A" box="[816,889,1652,1678]" italics="true" pageId="17" pageNumber="18">dacnis</emphasis>
</taxonomicName>
individual (not collectedbut likely a
<taxonomicName id="4C6A3D8F8B48673D8447F98BFA67F94A" box="[1331,1435,1652,1678]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B48673D8447F98BFA67F94A" box="[1331,1435,1652,1678]" italics="true" pageId="17" pageNumber="18">B. dacnis</emphasis>
</taxonomicName>
, as no
<taxonomicName id="4C6A3D8F8B48673D849AF98BFD7CF971" baseAuthorityName="Giaretta &amp; Sawaya" baseAuthorityYear="1998" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="17" pageNumber="18" phylum="Chordata" rank="species" species="hermogenesi">
<emphasis id="B91E9A1E8B48673D849AF98BFD7CF971" italics="true" pageId="17" pageNumber="18">B. hermogenesi</emphasis>
</taxonomicName>
was found in the same area) jumping
<quantity id="4C92EBE98B48673D854FF963FB6AF972" box="[1083,1174,1692,1718]" metricMagnitude="-1" metricUnit="m" metricValue="2.18" pageId="17" pageNumber="18" unit="cm" value="21.8">21.8 cm</quantity>
(31 times its SVL) (ZUEC-VID 796).
</paragraph>
<paragraph id="8BD5460C8B48673E8366F913FA2CFEE9" blockId="17.[498,1542,1133,1878]" lastBlockId="18.[498,1488,234,301]" lastPageId="18" lastPageNumber="19" pageId="17" pageNumber="18">
We identified the stomach contents of two individuals. During dissection, we found one individual (ZUEC-AMP 25273) with four items in its stomach, identified to order: one Collembola, one
<taxonomicName id="4C6A3D8F8B48673D83CEF8C4FCC0F891" box="[698,828,1851,1877]" class="Insecta" kingdom="Animalia" order="Coleoptera" pageId="17" pageNumber="18" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
, one Hymenoptera (Hexapoda), and one Acari (Sarcoptiformes) (
<figureCitation id="13515A898B4B673E83BDFF15FCE4FEC0" box="[713,792,234,260]" captionStart="Figure 2" captionStartId="8.[524,589,1066,1088]" captionTargetBox="[498,1541,236,1053]" captionTargetId="figure-278@8.[498,1299,234,1038]" captionTargetPageId="8" captionText="Figure 2 Brachycephalus dacnis paratype (ZUEC-AMP 25612) adult male, SVL = 7.89 mm. (A) Dorsal view, (B) ventral view, (C) ventral view of hand, (D) ventral view of feet. Full-size DOI: 10.7717/peerj.18265/fig-2" pageId="18" pageNumber="19">Fig. S2</figureCitation>
). In the microCT scan of another individual (ZUEC-AMP 24978), we identified one item, an Anobiinae beetle (
<taxonomicName id="4C6A3D8F8B4B673E8517FEEDFB14FEE8" box="[1123,1256,274,300]" class="Insecta" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="order">Coleoptera</taxonomicName>
:
<taxonomicName id="4C6A3D8F8B4B673E8587FEEDFAAAFEE9" box="[1267,1366,274,301]" class="Insecta" family="Ptinidae" kingdom="Animalia" order="Coleoptera" pageId="18" pageNumber="19" phylum="Arthropoda" rank="family">Ptinidae</taxonomicName>
) (
<figureCitation id="13515A898B4B673E8405FEEDFA3CFEE8" box="[1393,1472,274,300]" captionStart="Figure 2" captionStartId="8.[524,589,1066,1088]" captionTargetBox="[498,1541,236,1053]" captionTargetId="figure-278@8.[498,1299,234,1038]" captionTargetPageId="8" captionText="Figure 2 Brachycephalus dacnis paratype (ZUEC-AMP 25612) adult male, SVL = 7.89 mm. (A) Dorsal view, (B) ventral view, (C) ventral view of hand, (D) ventral view of feet. Full-size DOI: 10.7717/peerj.18265/fig-2" pageId="18" pageNumber="19">Fig. S2</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C37015878B4B673E8086FEAFFDB2FDCF" pageId="18" pageNumber="19" type="distribution">
<paragraph id="8BD5460C8B4B673E8086FEAFFDB2FDCF" blockId="18.[498,1542,336,523]" pageId="18" pageNumber="19">
<emphasis id="B91E9A1E8B4B673E8086FEAFFD77FEAE" bold="true" box="[498,651,336,362]" pageId="18" pageNumber="19">Distribution</emphasis>
<taxonomicName id="4C6A3D8F8B4B673E83EEFEAFFC62FEAE" box="[666,926,336,362]" class="Amphibia" family="Brachycephalidae" genus="Brachycephalus" kingdom="Animalia" order="Anura" pageId="18" pageNumber="19" phylum="Chordata" rank="species" species="dacnis">
<emphasis id="B91E9A1E8B4B673E83EEFEAFFC62FEAE" box="[666,926,336,362]" italics="true" pageId="18" pageNumber="19">Brachycephalus dacnis</emphasis>
</taxonomicName>
is known from its
<typeStatus id="54D1F8AE8B4B673E85F5FEAEFB4FFEAF" box="[1153,1203,337,363]" pageId="18" pageNumber="19">type</typeStatus>
locality, at Projeto Dacnis private reserve in Ubatuba, state of
<collectingRegion id="49AE88EE8B4B673E82E3FE86FBF7FE56" box="[919,1035,376,403]" country="Brazil" name="Sao Paulo" pageId="18" pageNumber="19">São Paulo</collectingRegion>
(SP),
<collectingCountry id="F37D069C8B4B673E8522FE87FB60FE56" box="[1110,1180,376,402]" name="Brazil" pageId="18" pageNumber="19">Brazil</collectingCountry>
, and two additional localities, based on acoustic records previously published in
<bibRefCitation id="EFFB3BFD8B4B673E8530FE5FFAA4FE7E" author="Bornschein MR &amp; Ribeiro LF &amp; Teixeira L &amp; Belmonte-Lopes R &amp; Moraes LA &amp; de Correa L &amp; Mauricio GN &amp; Pie MR" box="[1092,1368,416,442]" journalOrPublisher="PeerJ" pageId="18" pageNumber="19" part="9" publicationUrl="10.7717/peerj.10983" refId="ref12763" refString="Bornschein MR, Ribeiro LF, Teixeira L, Belmonte-Lopes R, Moraes LA, de Correa L, Mauricio GN, Nadaline Ι, Pie MR. 2021. A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae). PeerJ 9: e 10983 DOI 10.7717 / peerj. 10983." title="A review of the diagnosis and geographical distribution of the recently described flea toad Brachycephalus sulfuratus in relation to B. hermogenesi (Anura: Brachycephalidae)" type="journal volume" year="2021">
<emphasis id="B91E9A1E8B4B673E8530FE5FFAA4FE7E" box="[1092,1368,416,442]" italics="true" pageId="18" pageNumber="19">Bornschein et al. (2021)</emphasis>
</bibRefCitation>
: Corcovado, municipality of Ubatuba, SP, and Trilha do Corisco, municipality of Paraty,
<collectingRegion id="49AE88EE8B4B673E8416FE37F9FAFE26" box="[1378,1542,456,482]" country="Brazil" name="Rio de Janeiro" pageId="18" pageNumber="19">Rio de Janeiro</collectingRegion>
(
<figureCitation id="13515A898B4B673E8088FE0FFDC2FDCE" box="[508,574,496,522]" captionStart="Figure 6" captionStartId="14.[116,181,1171,1193]" captionTargetBox="[90,1541,236,1158]" captionTargetId="figure-189@14.[90,1512,234,1143]" captionTargetPageId="14" captionText="Figure 6 Geographic distribution, advertisement calls and phylogeny. (A) Type localities of Brachycephalus dacnis (red star) and B. hermogenesi (black star), and additional distribution records of B. dacnis (red triangles, based on call recordings published in Bornschein et al., 2021). This map was created using the free and open-source software QGIS, utilizing publicly available base data sources. (B) Brachycephalus dacnis holotypes advertisement call (FNJV 51341), recorded at Projeto Dacnis private reserve (type locality), Ubatuba, São Paulo (left) and B. hermogenesi topotypes advertisement call (FNJV 51343), recorded at Picinguaba (type locality), Ubatuba, São Paulo (right). (C) Schematic representation of the Bayesian phylogenetic inference of Brachycephalus based on the mitochondrially encoded 16S rDNA fragment. Black dots indicate fully supported clades (posterior probability, PP =&gt; 99). White dots indicate PP&gt; 0.95. Node supports below pp = 0.95 are not shown. Complete tree in the supplement (Fig. S1). Full-size DOI: 10.7717/peerj.18265/fig-6" pageId="18" pageNumber="19">Fig. 6</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>