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<document id="72595708B17CB4467B0FEF786B73FB2D" ID-DOI="10.11646/zootaxa.4104.1.1" ID-GBIF-Dataset="9b3e7716-1099-48ca-803b-620c25d788f1" ID-ISSN="1175-5326" ID-Zenodo-Dep="265809" ID-ZooBank="D598E724-C9E4-4BBA-B25D-511300A47B1D" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.tables_requiresApprovalFor="existingObjects,plazi" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1461050759050" checkinUser="plazi" docAuthor="Duellman, William E., Marion, Angela B. &amp; Hedges, Blair" docDate="2016" docId="03EA87A5FF9C121FF3988FD036B4F243" docLanguage="en" docName="zootaxa.4104.pdf.imf" docOrigin="Zootaxa 4104 (1)" docStyle="DocumentStyle:5EBBA59367AD13919D70D935FA04F6A3.14:Zootaxa.2013-.monograph" docStyleId="5EBBA59367AD13919D70D935FA04F6A3" docStyleName="Zootaxa.2013-.monograph" docStyleVersion="14" docTitle="Hylidae" docType="treatment" docVersion="11" lastPageNumber="54" masterDocId="FFD3FFDDFFA91229F30F8D5E333EF35E" masterDocTitle="Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae)" masterLastPageNumber="109" masterPageNumber="1" pageNumber="54" updateTime="1698579563523" updateUser="plazi">
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<mods:title id="D41323072A4166198F080BAC190EE867">Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae)</mods:title>
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<mods:namePart id="984FDAFC1B6462ECC696160FCBACAA66">Duellman, William E.</mods:namePart>
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<treatment id="03EA87A5FF9C121FF3988FD036B4F243" ID-DOI="http://doi.org/10.5281/zenodo.5458573" ID-GBIF-Taxon="119611842" ID-Zenodo-Dep="5458573" LSID="urn:lsid:plazi:treatment:03EA87A5FF9C121FF3988FD036B4F243" httpUri="http://treatment.plazi.org/id/03EA87A5FF9C121FF3988FD036B4F243" lastPageId="54" lastPageNumber="54" pageId="53" pageNumber="54">
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<paragraph id="8BFC36B3FF9C121CF3988FD03255F1F6" blockId="53.[151,363,654,680]" box="[151,363,654,680]" pageId="53" pageNumber="54">
<heading id="D0B481DFFF9C121CF3988FD03255F1F6" bold="true" box="[151,363,654,680]" fontSize="11" level="1" pageId="53" pageNumber="54" reason="1">
<emphasis id="B937EAA1FF9C121CF3988FD03255F1F6" bold="true" box="[151,363,654,680]" pageId="53" pageNumber="54">
<taxonomicName id="4C434D30FF9C121CF3988FD033C3F1F6" ID-CoL="B9H" box="[151,253,654,680]" class="Amphibia" family="Hylidae" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="family">Hylidae</taxonomicName>
:
<taxonomicName id="4C434D30FF9C121CF2058FD03255F1F6" box="[266,363,654,680]" class="Amphibia" family="Hylidae" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="subFamily" subFamily="Hylinae">Hylinae</taxonomicName>
</emphasis>
</heading>
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<paragraph id="8BFC36B3FF9C121CF3988F8B33D1F0FC" blockId="53.[151,1437,725,2010]" pageId="53" pageNumber="54">
The divergence of North and Middle American hylid frogs from their relatives in South
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was in the early Oligocene, 32.9 (30.235.6) Mya. Thus, hyline frogs were in Middle
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in the early Miocene. Differentiation (crown node times) of the various genera occurred throughout the Miocene—as early as 26.1 Mya for the origin of
<taxonomicName id="4C434D30FF9C121CF3988E1F325EF004" box="[151,352,833,858]" class="Amphibia" family="Hylidae" genus="Megastomatohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF3988E1F325EF004" box="[151,352,833,858]" italics="true" pageId="53" pageNumber="54">Megastomatohyla</emphasis>
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to 13.1 Mya for the origin of
<emphasis id="B937EAA1FF9C121CF1B98E1F3001F004" box="[694,831,833,858]" italics="true" pageId="53" pageNumber="54">
<taxonomicName id="4C434D30FF9C121CF1B98E1F3005F004" box="[694,827,833,858]" class="Amphibia" family="Hylidae" genus="Exerodonta" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">Exerodonta</taxonomicName>
.
</emphasis>
A major geographic divergence occurred in the early Miocene, 23.9 (22.325.6) Mya, when the clade now known as the Holarctic hylines diverged from the tropical hylines.
</paragraph>
<paragraph id="8BFC36B3FF9C121CF3C88EF331FDF7B8" blockId="53.[151,1437,725,2010]" pageId="53" pageNumber="54">
The geological uplifts and volcanism in
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and Central
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began in the late Cretaceous and continues to the present. Only three clades of lowland inhabitants exist in the Middle American tropics; these are in a major clade that also includes
<emphasis id="B937EAA1FF9C121CF10F8EAB31BDF750" box="[512,643,1013,1038]" italics="true" pageId="53" pageNumber="54">
<taxonomicName id="4C434D30FF9C121CF10F8EAB3141F750" box="[512,639,1013,1038]" class="Amphibia" family="Hylidae" genus="Isthmohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">Isthmohyla</taxonomicName>
,
</emphasis>
which has stream-adapted tadpoles. The three lowland clades radiated (crown node times) in the mid-Miocene—
<taxonomicName id="4C434D30FF9C121CF162894731F2F76C" box="[621,716,1049,1074]" class="Amphibia" family="Hylidae" genus="Smilisca" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF162894731F2F76C" box="[621,716,1049,1074]" italics="true" pageId="53" pageNumber="54">Smilisca</emphasis>
</taxonomicName>
13.2 (10.615.9) Mya,
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<emphasis id="B937EAA1FF9C121CF0DB89473764F76C" box="[980,1114,1049,1074]" italics="true" pageId="53" pageNumber="54">Tlalocohyla</emphasis>
</taxonomicName>
15.5 (13.018.1) Mya, and a casque-headed clade (
<emphasis id="B937EAA1FF9C121CF294896331B7F708" box="[411,649,1085,1110]" italics="true" pageId="53" pageNumber="54">
<taxonomicName id="4C434D30FF9C121CF29489633136F708" box="[411,520,1085,1110]" class="Amphibia" family="Hylidae" genus="Anotheca" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">Anotheca</taxonomicName>
,
<taxonomicName id="4C434D30FF9C121CF115896331BBF708" box="[538,645,1085,1110]" class="Amphibia" family="Hylidae" genus="Diaglena" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">Diaglena</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C434D30FF9C121CF1C389603016F70B" box="[716,808,1086,1109]" class="Amphibia" family="Hylidae" genus="Triprion" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF1C389603016F70B" box="[716,808,1086,1109]" italics="true" pageId="53" pageNumber="54">Triprion</emphasis>
</taxonomicName>
) 14.3 (11.217.4) Mya. Slightly later the last clade differentiated into
<taxonomicName id="4C434D30FF9C121CF265893F32ECF724" box="[362,466,1121,1146]" class="Amphibia" family="Hylidae" genus="Diaglena" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF265893F32ECF724" box="[362,466,1121,1146]" italics="true" pageId="53" pageNumber="54">Diaglena</emphasis>
</taxonomicName>
and
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<emphasis id="B937EAA1FF9C121CF103893C3157F727" box="[524,617,1122,1145]" italics="true" pageId="53" pageNumber="54">Triprion</emphasis>
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in xeric lowland habitats with tadpoles developing in temporary ponds, and the cloud forest-inhabitant,
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<taxonomicName id="4C434D30FF9C121CF2F289DB3154F7C0" box="[509,618,1157,1182]" class="Amphibia" family="Hylidae" genus="Anotheca" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">Anotheca</taxonomicName>
,
</emphasis>
with tadpoles developing in tree holes. All species of
<taxonomicName id="4C434D30FF9C121CF7DE89DB3669F7C0" box="[1233,1367,1157,1182]" class="Amphibia" family="Hylidae" genus="Tlalocohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF7DE89DB3669F7C0" box="[1233,1367,1157,1182]" italics="true" pageId="53" pageNumber="54">Tlalocohyla</emphasis>
</taxonomicName>
range in the lowlands of
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; one species extends as far south as
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.
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<emphasis id="B937EAA1FF9C121CF0ED89F7377FF79C" box="[994,1089,1193,1218]" italics="true" pageId="53" pageNumber="54">Smilisca</emphasis>
</taxonomicName>
consists of six species ranging throughout Central
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and tropical
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.
</paragraph>
<paragraph id="8BFC36B3FF9C121CF3C889AF3262F580" blockId="53.[151,1437,725,2010]" pageId="53" pageNumber="54">
Currently there are 108 species of stream-breeding hylids in nine genera endemic to Middle
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. The small stream-breeding
<taxonomicName id="4C434D30FF9C121CF291884B3136F670" box="[414,520,1301,1326]" class="Amphibia" family="Hylidae" genus="Rheohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="19" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="B937EAA1FF9C121CF291884B3136F670" box="[414,520,1301,1326]" italics="true" pageId="53" pageNumber="54">Rheohyla</emphasis>
</taxonomicName>
diverged from the clade that became the large, arboreal hylines in the earliest Miocene, 23.0 (17.828.2) Mya. The fringe-limbed tree frogs of the genus
<taxonomicName id="4C434D30FF9C121CF0D588673750F60C" box="[986,1134,1337,1362]" class="Amphibia" family="Hylidae" genus="Ecnomiohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF0D588673750F60C" box="[986,1134,1337,1362]" italics="true" pageId="53" pageNumber="54">Ecnomiohyla</emphasis>
</taxonomicName>
have disjunct distributions from Oaxaca,
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, through
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, whereas
<taxonomicName id="4C434D30FF9C121CF1E288033069F628" box="[749,855,1373,1398]" class="Amphibia" family="Hylidae" genus="Rheohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="19" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="B937EAA1FF9C121CF1E288033069F628" box="[749,855,1373,1398]" italics="true" pageId="53" pageNumber="54">Rheohyla</emphasis>
</taxonomicName>
occurs only in
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west of the Isthmus of Tehuantepec. Nested within the clade of inhabitants of Middle American lowlands is the stream-breading
<emphasis id="B937EAA1FF9C121CF39888FB3224F6E0" box="[151,282,1445,1470]" italics="true" pageId="53" pageNumber="54">
<taxonomicName id="4C434D30FF9C121CF39888FB3228F6E0" box="[151,278,1445,1470]" class="Amphibia" family="Hylidae" genus="Isthmohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">Isthmohyla</taxonomicName>
,
</emphasis>
which consists of 15 species ranging discontinuously in highlands from
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to central
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.
<taxonomicName id="4C434D30FF9C121CF3988897322DF6BC" box="[151,275,1481,1506]" class="Amphibia" family="Hylidae" genus="Isthmohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF3988897322DF6BC" box="[151,275,1481,1506]" italics="true" pageId="53" pageNumber="54">Isthmohyla</emphasis>
</taxonomicName>
diverged from ancestral
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<emphasis id="B937EAA1FF9C121CF14C8897319CF6BC" box="[579,674,1481,1506]" italics="true" pageId="53" pageNumber="54">Smilisca</emphasis>
</taxonomicName>
in the early Miocene, 21.4 (18.724.1) Mya. All other streambreeding hylids belong to one major clade. Two of these clades possibly diverged in the late Oligocene.
<taxonomicName id="4C434D30FF9C121CF3988B4F325EF574" box="[151,352,1553,1578]" class="Amphibia" family="Hylidae" genus="Megastomatohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF3988B4F325EF574" box="[151,352,1553,1578]" italics="true" pageId="53" pageNumber="54">Megastomatohyla</emphasis>
</taxonomicName>
split with
<taxonomicName id="4C434D30FF9C121CF2D98B4F314BF574" box="[470,629,1553,1578]" class="Amphibia" family="Hylidae" genus="Charadrahyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF2D98B4F314BF574" box="[470,629,1553,1578]" italics="true" pageId="53" pageNumber="54">Charadrahyla</emphasis>
</taxonomicName>
about 26.1 (20.531.7) Mya in the Mexican highlands. The crown node time of the clade containing the genera
<emphasis id="B937EAA1FF9C121CF16B8B6B30FEF510" box="[612,960,1589,1614]" italics="true" pageId="53" pageNumber="54">
<taxonomicName id="4C434D30FF9C121CF16B8B6B31C0F510" box="[612,766,1589,1614]" class="Amphibia" family="Hylidae" genus="Bromeliohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">Bromeliohyla</taxonomicName>
,
<taxonomicName id="4C434D30FF9C121CF0028B6B3083F510" box="[781,957,1589,1614]" class="Amphibia" family="Hylidae" genus="Duellmanohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">Duellmanohyla</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C434D30FF9C121CF0F38B6B3744F510" box="[1020,1146,1589,1614]" class="Amphibia" family="Hylidae" genus="Ptychohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF0F38B6B3744F510" box="[1020,1146,1589,1614]" italics="true" pageId="53" pageNumber="54">Ptychohyla</emphasis>
</taxonomicName>
, species occurring in the Central American highlands, is about 17.4 (14.919.9) Mya. These genera of yet undetermined affinities contain two bromeliad-breeding species in the genus
<taxonomicName id="4C434D30FF9C121CF1AF8B233009F5C8" box="[672,823,1661,1686]" class="Amphibia" family="Hylidae" genus="Bromeliohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF1AF8B233009F5C8" box="[672,823,1661,1686]" italics="true" pageId="53" pageNumber="54">Bromeliohyla</emphasis>
</taxonomicName>
and
<taxonomicName id="4C434D30FF9C121CF07D8B233721F5C8" box="[882,1055,1661,1686]" class="Amphibia" family="Hylidae" genus="Duellmanohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF07D8B233721F5C8" box="[882,1055,1661,1686]" italics="true" pageId="53" pageNumber="54">Duellmanohyla</emphasis>
</taxonomicName>
with eight species inhabiting the Central American highlands. That region is also inhabited by
<taxonomicName id="4C434D30FF9C121CF05F8BFF30F0F5E4" box="[848,974,1697,1722]" class="Amphibia" family="Hylidae" genus="Ptychohyla" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF05F8BFF30F0F5E4" box="[848,974,1697,1722]" italics="true" pageId="53" pageNumber="54">Ptychohyla</emphasis>
</taxonomicName>
with 13 species, some of which occur in southern
<collectingCountry id="F3547623FF9C121CF3F18B9B3266F580" box="[254,344,1733,1758]" name="Mexico" pageId="53" pageNumber="54">Mexico</collectingCountry>
.
</paragraph>
<paragraph id="8BFC36B3FF9C121FF3C88BB736B4F243" blockId="53.[151,1437,725,2010]" lastBlockId="54.[151,1436,151,285]" lastPageId="54" lastPageNumber="55" pageId="53" pageNumber="54">
The Isthmus of Tehuantepec in southern
<collectingCountry id="F3547623FF9C121CF1998BB731D5F45C" box="[662,747,1769,1794]" name="Mexico" pageId="53" pageNumber="54">Mexico</collectingCountry>
seems to have played a significant role in the differentiation and distribution of genera of stream-breeding hylines.
<taxonomicName id="4C434D30FF9C121CF1F08A5330BFF478" box="[767,897,1805,1830]" class="Amphibia" family="Hylidae" genus="Exerodonta" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9C121CF1F08A5330BFF478" box="[767,897,1805,1830]" italics="true" pageId="53" pageNumber="54">Exerodonta</emphasis>
</taxonomicName>
split from the
<emphasis id="B937EAA1FF9C121CF7258A53361BF478" box="[1066,1317,1805,1830]" italics="true" pageId="53" pageNumber="54">Plectrohyla-Sarcohyla</emphasis>
lineage in the late Oligocene 27.5 (23.731.3) Mya.
<emphasis id="B937EAA1FF9C121CF17F8A6F31C6F414" box="[624,760,1841,1866]" italics="true" pageId="53" pageNumber="54">
<taxonomicName id="4C434D30FF9C121CF17F8A6F31CAF414" box="[624,756,1841,1866]" class="Amphibia" family="Hylidae" genus="Exerodonta" kingdom="Animalia" order="Anura" pageId="53" pageNumber="54" phylum="Chordata" rank="genus">Exerodonta</taxonomicName>
,
</emphasis>
which radiated (crown node time) in the mid-Miocene 13.1 (10.415.8) Mya, inhabits cloud forest and pine-oak forests at elevations of
<quantity id="4CBB9B56FF9C121CF0FB8A0B37BBF433" box="[1012,1157,1877,1902]" metricMagnitude="3" metricUnit="m" metricValue="1.305" metricValueMax="2.16" metricValueMin="0.45" pageId="53" pageNumber="54" unit="m" value="1305.0" valueMax="2160.0" valueMin="450.0">4502160 m</quantity>
; nine species occur west of the Isthmus of Tehuantepec and two live east of the isthmus. Likewise in the mid-Miocene, 15.7 (11.519.9) Mya,
<emphasis id="B937EAA1FF9C121CF3D58AC33249F4E8" box="[218,375,1949,1974]" italics="true" pageId="53" pageNumber="54">Charadrohyla</emphasis>
was diversifying in the streams in southern
<collectingCountry id="F3547623FF9C121CF07E8AC330F4F4E8" box="[881,970,1949,1974]" name="Mexico" pageId="53" pageNumber="54">Mexico</collectingCountry>
, where five species live today; one other species occurs in Chiapas to the east of the Isthmus of Tehuantepec. The most striking example of diversification on opposite sides of the isthmus is
<taxonomicName id="4C434D30FF9F121FF12E8DC9319DF3EE" box="[545,675,151,176]" class="Amphibia" family="Hylidae" genus="Plectrohyla" kingdom="Animalia" order="Anura" pageId="54" pageNumber="55" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9F121FF12E8DC9319DF3EE" box="[545,675,151,176]" italics="true" pageId="54" pageNumber="55">Plectrohyla</emphasis>
</taxonomicName>
(18 species) in the northern Central American highlands to the east of the isthmus and
<taxonomicName id="4C434D30FF9F121FF27C8DE232DDF38B" box="[371,483,188,213]" class="Amphibia" family="Hylidae" genus="Sarcohyla" kingdom="Animalia" order="Anura" pageId="54" pageNumber="18" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="B937EAA1FF9F121FF27C8DE232DDF38B" box="[371,483,188,213]" italics="true" pageId="54" pageNumber="55">Sarcohyla</emphasis>
</taxonomicName>
(24 species) in the Mexican highlands to the west of the isthmus. They split in the early mid-Miocene 18.6 (18.615.9) Mya. The times of diversification of ancestral stocks on either side of the isthmus coincide with the volcanism in the Miocene that elevated nuclear Central
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and southern
<collectingCountry id="F3547623FF9F121FF6228C5A36B8F243" box="[1325,1414,260,285]" name="Mexico" pageId="54" pageNumber="55">Mexico</collectingCountry>
.
</paragraph>
</subSubSection>
</treatment>
</document>