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<document id="006718DBBA17BAD34F5141F27E2D559E" 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="03EA87A5FF9A121AF3988EF331CBF45E" 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="52" masterDocId="FFD3FFDDFFA91229F30F8D5E333EF35E" masterDocTitle="Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae)" masterLastPageNumber="109" masterPageNumber="1" pageNumber="52" updateTime="1698579563523" updateUser="plazi">
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<mods:title id="629EBE5445A3B71944875B8BF8F50C43">Phylogenetics, classification, and biogeography of the treefrogs (Amphibia: Anura: Arboranae)</mods:title>
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<paragraph id="8BFC36B3FF9A121AF3988EF332ECF099" blockId="51.[151,466,941,967]" box="[151,466,941,967]" pageId="51" pageNumber="52">
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<taxonomicName id="4C434D30FF9A121AF3988EF333C3F099" ID-CoL="B9H" box="[151,253,941,967]" class="Amphibia" family="Hylidae" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="family">Hylidae</taxonomicName>
: Cophomantinae
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<paragraph id="8BFC36B3FF9A121AF3988EAA300AF673" blockId="51.[151,1437,1012,1792]" pageId="51" pageNumber="52">
Shortly after the origin of the clade herein referred to as cophomantines, an ancestral stock inhabited the earlyuplifted western part of the continent (the proto Andes). This clade representing current
<emphasis id="B937EAA1FF9A121AF7FD89493221F70B" italics="true" pageId="51" pageNumber="52">
<taxonomicName id="4C434D30FF9A121AF7FD894936A6F76E" box="[1266,1432,1047,1072]" class="Amphibia" family="Hylidae" genus="Colomascirtus" kingdom="Animalia" order="Anura" pageId="51" pageNumber="30" phylum="Chordata" rank="genus" status="gen. nov.">Colomascirtus</taxonomicName>
,
<taxonomicName id="4C434D30FF9A121AF39889623225F70B" box="[151,283,1084,1109]" class="Amphibia" family="Hylidae" genus="Hyloscirtus" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="genus">Hyloscirtus</taxonomicName>
,
</emphasis>
and
<taxonomicName id="4C434D30FF9A121AF26D896232D0F70B" box="[354,494,1084,1109]" class="Amphibia" family="Hylidae" genus="Myersiohyla" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9A121AF26D896232D0F70B" box="[354,494,1084,1109]" italics="true" pageId="51" pageNumber="52">Myersiohyla</emphasis>
</taxonomicName>
diverged (crown node) in the mid-Eocene 47.5 (38.057.0) Mya. These frogs inhabited streams, and their tadpoles developed large suctorial mouths with many rows of labial teeth as they dispersed through the ever-rising Andes. An early divergence (the crown node) in northern South
<collectingCountry id="F3547623FF9A121AF63389DA36A2F7C3" box="[1340,1436,1156,1181]" name="United States of America" pageId="51" pageNumber="52">America</collectingCountry>
restricted
<taxonomicName id="4C434D30FF9A121AF20289F932BBF79E" box="[269,389,1191,1216]" class="Amphibia" family="Microhylidae" genus="Myersiella" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9A121AF20289F932BBF79E" box="[269,389,1191,1216]" italics="true" pageId="51" pageNumber="52">Myersiella</emphasis>
</taxonomicName>
to the
<collectingCountry id="F3547623FF9A121AF2E989F93109F79E" box="[486,567,1191,1216]" name="Guyana" pageId="51" pageNumber="52">Guiana</collectingCountry>
Highlands, whereas in the Andes in the early Oligocene
<taxonomicName id="4C434D30FF9A121AF7F689F936A2F79E" box="[1273,1436,1191,1216]" class="Amphibia" family="Hylidae" genus="Colomascirtus" kingdom="Animalia" order="Anura" pageId="51" pageNumber="30" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="B937EAA1FF9A121AF7F689F936A2F79E" box="[1273,1436,1191,1216]" italics="true" pageId="51" pageNumber="52">Colomascirtus</emphasis>
</taxonomicName>
differentiated from
<taxonomicName id="4C434D30FF9A121AF278899232C6F7BB" box="[375,504,1228,1253]" class="Amphibia" family="Hylidae" genus="Hyloscirtus" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9A121AF278899232C6F7BB" box="[375,504,1228,1253]" italics="true" pageId="51" pageNumber="52">Hyloscirtus</emphasis>
</taxonomicName>
33.3 (29.337.3) Mya.
<bibRefCitation id="EFD24B42FF9A121AF0018992373EF7BA" box="[782,1024,1228,1253]" pageId="51" pageNumber="52" refString="Coloma, L. A., Carvajal-Endara, S., Duenas, J. F., Paredes-Recalde, A., Morales-Mite, M. A., Almeida-Reinoso, D., Tapia, E. E., Hutter, C. R., Toral-Contreras, E. &amp; Guayasamin, J. M. (2012) Molecular phylogenetics of stream treefrogs of the Hyloscirtus larinopygion group (Anura: Hylidae), and description of two new species from Ecuador. Zootaxa, 3364 (4), 1 - 78. http: // dx. doi. org / 10.11646 / zootaxa. 3686.4.3" type="journal article">
Coloma
<emphasis id="B937EAA1FF9A121AF07F89933092F7BB" box="[880,940,1228,1253]" italics="true" pageId="51" pageNumber="52">et al.</emphasis>
(2012)
</bibRefCitation>
estimated the minimum divergence time of these events as 61.2 and 40.9 Mya, respectively. The latter inhabited low to moderate elevations, whereas
<taxonomicName id="4C434D30FF9A121AF398884A3205F673" box="[151,315,1300,1325]" class="Amphibia" family="Hylidae" genus="Colomascirtus" kingdom="Animalia" order="Anura" pageId="51" pageNumber="30" phylum="Chordata" rank="genus" status="gen. nov.">
<emphasis id="B937EAA1FF9A121AF398884A3205F673" box="[151,315,1300,1325]" italics="true" pageId="51" pageNumber="52">Colomascirtus</emphasis>
</taxonomicName>
came to inhabit streams at higher elevations.
</paragraph>
<paragraph id="8BFC36B3FF9A121AF3C888693610F6E3" blockId="51.[151,1437,1012,1792]" pageId="51" pageNumber="52">
The other major clade within Cophomantinae subsequently diverged into two lineages in the late Eocene, 36.8 (32.840.8) Mya. One clade consisted solely of
<emphasis id="B937EAA1FF9A121AF1B4880230BCF62B" box="[699,898,1372,1397]" italics="true" pageId="51" pageNumber="52">
<taxonomicName id="4C434D30FF9A121AF1B488023043F62B" box="[699,893,1372,1397]" class="Amphibia" family="Hylidae" genus="Bokermannohyla" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="genus">Bokermannohyla</taxonomicName>
,
</emphasis>
which inhabited the mountains of southeastern
<collectingCountry id="F3547623FF9A121AF398882133E5F6C6" box="[151,219,1407,1432]" name="Brazil" pageId="51" pageNumber="52">Brazil</collectingCountry>
and developed stream-adapted tadpoles. In the other clade, a lowland lineage,
<emphasis id="B937EAA1FF9A121AF74A882137FCF6C6" box="[1093,1218,1407,1432]" italics="true" pageId="51" pageNumber="52">
<taxonomicName id="4C434D30FF9A121AF74A88213780F6C6" box="[1093,1214,1407,1432]" class="Amphibia" family="Hylidae" genus="Hypsiboas" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="genus">Hypsiboas</taxonomicName>
,
</emphasis>
differentiated from a highland group,
<taxonomicName id="4C434D30FF9A121AF26C88FA32C1F6E3" box="[355,511,1444,1469]" class="Amphibia" family="Hylidae" genus="Aplastodiscus" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9A121AF26C88FA32C1F6E3" box="[355,511,1444,1469]" italics="true" pageId="51" pageNumber="52">Aplastodiscus</emphasis>
</taxonomicName>
with stream-adapted tadpoles, in the late Eocene, 34.2 (30.937.5) Mya.
</paragraph>
<paragraph id="8BFC36B3FF9A121AF3C8889931CBF45E" blockId="51.[151,1437,1012,1792]" pageId="51" pageNumber="52">
The earliest divergence within
<taxonomicName id="4C434D30FF9A121AF128889931A3F6BE" box="[551,669,1479,1504]" class="Amphibia" family="Hylidae" genus="Hypsiboas" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9A121AF128889931A3F6BE" box="[551,669,1479,1504]" italics="true" pageId="51" pageNumber="52">Hypsiboas</emphasis>
</taxonomicName>
occurred in the mid-Oligocene 31.9 (28.435.5) Mya when a large group of species (
<taxonomicName id="4C434D30FF9A121AF26B88B332E1F55B" box="[356,479,1516,1541]" class="Amphibia" family="Hylidae" genus="Hypsiboas" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="species" species="benitezi">
<emphasis id="B937EAA1FF9A121AF26B88B33246F55A" box="[356,376,1517,1540]" italics="true" pageId="51" pageNumber="52">H</emphasis>
.
<emphasis id="B937EAA1FF9A121AF28788B232E1F55B" box="[392,479,1516,1541]" italics="true" pageId="51" pageNumber="52">benitezi</emphasis>
</taxonomicName>
<taxonomicName id="4C434D30FF9A121AF2E288B3315DF55B" box="[493,611,1516,1541]" class="Amphibia" family="Hylidae" genus="Hypsiboas" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="species" species="sibleszi">
H.
<emphasis id="B937EAA1FF9A121AF11E88B2315DF55B" box="[529,611,1516,1541]" italics="true" pageId="51" pageNumber="52">sibleszi</emphasis>
</taxonomicName>
) occupied northeastern South
<collectingCountry id="F3547623FF9A121AF0CE88B2371FF55B" box="[961,1057,1516,1541]" name="United States of America" pageId="51" pageNumber="52">America</collectingCountry>
including the
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Highlands. Differentiation of most clades of
<taxonomicName id="4C434D30FF9A121AF11C8B5131B7F576" box="[531,649,1551,1576]" class="Amphibia" family="Hylidae" genus="Hypsiboas" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="genus">
<emphasis id="B937EAA1FF9A121AF11C8B5131B7F576" box="[531,649,1551,1576]" italics="true" pageId="51" pageNumber="52">Hypsiboas</emphasis>
</taxonomicName>
occurred in the Miocene. For example, the long-legged tree frogs of the
<taxonomicName id="4C434D30FF9A121AF3CF8B6A32E2F513" box="[192,476,1588,1613]" class="Amphibia" family="Hylidae" genus="Hypsiboas" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="species" species="albopunctatus">
<emphasis id="B937EAA1FF9A121AF3CF8B6A32E2F513" box="[192,476,1588,1613]" italics="true" pageId="51" pageNumber="52">Hypsiboas albopunctatus</emphasis>
</taxonomicName>
Group differentiated about 17.4 (14.020.8) Mya in the Amazon Basin and dispersed into coastal
<collectingCountry id="F3547623FF9A121AF2288B093251F52E" box="[295,367,1623,1648]" name="Brazil" pageId="51" pageNumber="52">Brazil</collectingCountry>
. Likewise, the ancestral large, arboreal frogs in
<taxonomicName id="4C434D30FF9A121AF0A58B0937E2F52E" box="[938,1244,1623,1648]" class="Amphibia" family="Hylidae" genus="Hypsiboas" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="species" species="albomarginatus">
<emphasis id="B937EAA1FF9A121AF0A58B0937E2F52E" box="[938,1244,1623,1648]" italics="true" pageId="51" pageNumber="52">Hypsiboas albomarginatus</emphasis>
</taxonomicName>
Group diverged about 19.2 (15.422.9) Mya. Other species groups are more recent. For example, the clade of Andean species in the
<taxonomicName id="4C434D30FF9A121AF3988BC13259F5E6" box="[151,359,1695,1720]" class="Amphibia" family="Hylidae" genus="Hypsiboas" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="species" species="balzani">
<emphasis id="B937EAA1FF9A121AF3988BC13259F5E6" box="[151,359,1695,1720]" italics="true" pageId="51" pageNumber="52">Hypsiboas balzani</emphasis>
</taxonomicName>
Group differentiated in the Pliocene about 5.0 (13.86.1) Mya. Similarly, the Pliocene was the time of divergence of the ancestral stock of the
<taxonomicName id="4C434D30FF9A121AF1AA8B9A30ABF583" box="[677,917,1732,1757]" class="Amphibia" family="Hylidae" genus="Hypsiboas" kingdom="Animalia" order="Anura" pageId="51" pageNumber="52" phylum="Chordata" rank="species" species="pulchellus">
<emphasis id="B937EAA1FF9A121AF1AA8B9A30ABF583" box="[677,917,1732,1757]" italics="true" pageId="51" pageNumber="52">Hypsiboas pulchellus</emphasis>
</taxonomicName>
Group, 5.7 (4.86.5) Mya that came to inhabit subtropical highlands from
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to southern
<collectingCountry id="F3547623FF9A121AF1A68BB931CFF45E" box="[681,753,1767,1792]" name="Brazil" pageId="51" pageNumber="52">Brazil</collectingCountry>
.
</paragraph>
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