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<mods:title id="5C6825014F16B10C02535FF0115D8606">A taxonomic revision of the Neotropical poison frog genus Ranitomeya (Amphibia: Dendrobatidae) 3083</mods:title>
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<emphasis id="A7EEEA8C950D1516C8FC9EDB3C98FB93" bold="true" box="[151,415,1013,1039]" italics="true" pageId="77" pageNumber="78">Ranitomeya variabilis</emphasis>
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species group
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<figureCitation id="0DA12A1B950D1516C8FC99393DE3FBB3" box="[151,228,1047,1071]" captionStart="FIGURE 3" captionStartId="23.[151,244,1397,1418]" captionTargetBox="[151,1436,167,1383]" captionTargetId="figure-307@23.[151,1436,167,1383]" captionTargetPageId="23" captionText="FIGURE 3. A consensus Bayesian phylogeny based on 1011 base pairs of aligned mitochondrial DNA sequences of the 12S (12s rRNA), 16S (16s rRNA) and cytb (cytochrome-b gene) regions. Thickened branches represent nodes with posterior probabilities 90 and greater, other values are shown on nodes. Taxon labels depict current specific epithet, number in tree, the epithet being used prior to this revision (contained in parentheses), and the collection locality. A. Top segment. B. Middle segment. C. Bottom segment of phylogeny." figureDoi="http://doi.org/10.5281/zenodo.5244220" httpUri="https://zenodo.org/record/5244220/files/figure.png" pageId="77" pageNumber="78">Figs. 3</figureCitation>
, 
<figureCitation id="0DA12A1B950D1516C89A99393C06FBB3" box="[241,257,1047,1071]" captionStart="FIGURE 4" captionStartId="24.[151,244,1452,1473]" captionTargetBox="[168,1416,206,1417]" captionTargetId="figure-261@24.[151,1436,195,1431]" captionTargetPageId="24" captionText="FIGURE 4. Putative species tree for Andinobates, Excidobates, and Ranitomeya. Placement of species where molecular data were lacking (A. altobueyensis, A. viridis, A. abditus, A. daleswansoni and R. opisthomelas) was based on morphology. Andinobates altobueyensis and A. viridis were placed as sister taxa due to the absence of dark pigmentation on dorsal body and limbs and overall similar dorsal coloration and patterning. These species were placed as sister to A. fulguritus (sequenced) on the basis of similar dorsal coloration (bright green to greenish-yellow). Andinobates opisthomelas was placed in the bombetes group in a polytomy with A. bombetes and A. virolinensis (both sequenced) due to their similar advertisement calls and morphology, particularly their red dorsal pattern and marbled venter. Andinobates daleswansoni was placed as sister to A. dorisswansonae due to the absence of a well-defined first toe in both species. Andinobates abditus was placed in the bombetes group based on a larval synapomorphy which appears to be diagnostic of that group (wide medial gap in the papillae on the posterior labium). However, A. abditus was placed as the sister species to all other members of the bombetes group due to the absence of bright dorsal coloration and isolated geographic distribution. Andinobates abditus is currently the only species of its genus known to occur in the east-Andean versant, thus its placement remains speculative until molecular data become available. Photo credits: Thomas Ostrowski, Karl-Heinz Jungfer, Victor Luna-Mora, Giovanni Chaves-Portilla." figureDoi="http://doi.org/10.5281/zenodo.5244223" httpUri="https://zenodo.org/record/5244223/files/figure.png" pageId="77" pageNumber="78">4</figureCitation>
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<figureCitation id="0DA12A1B950D1516C96699393C1AFBB3" box="[269,285,1047,1071]" captionStart="FIGURE 9" captionStartId="39.[151,244,1424,1445]" captionTargetBox="[161,1436,606,1398]" captionTargetId="figure-326@39.[151,1436,553,1404]" captionTargetPageId="39" captionText="FIGURE 9. Known elevation distributions of Ranitomeya. Dotted line is mean for all samples. Dark boxes display the total elevation range of each species, within each contains a corresponding box plot." figureDoi="http://doi.org/10.5281/zenodo.5244239" httpUri="https://zenodo.org/record/5244239/files/figure.png" pageId="77" pageNumber="78">9</figureCitation>
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<figureCitation id="0DA12A1B950D1516C94299393C40FBB3" box="[297,327,1047,1071]" captionStart="FIGURE 14" captionStartId="46.[151,244,1819,1840]" captionTargetBox="[156,1424,1117,1789]" captionTargetId="figure-447@46.[151,1436,1116,1801]" captionTargetPageId="46" captionText="FIGURE 14. Advertisement calls of Ranitomeya species in the variabilis group and defleri group. A. Ranitomeya amazonica from 23 km S Iquitos, Loreto, Peru (type locality), recorded at 26° C; B. Ranitomeya amazonica from French Guiana, unknown temperature (call courtesy Erik Poelman); C. Ranitomeya variabilis from Cainarachi valley, San Martín, Peru, recorded at 22° C. D. Ranitomeya variabilis from Cerro Yupatí, Amazonas, Colombia, recorded at 27° C; E. Ranitomeya variabilis from Saposoa, San Martín, Peru, recorded at 24.5 C; F. Ranitomeya defleri from Rio Apaporis, Vaupés, Colombia, recorded at 26° C." figureDoi="http://doi.org/10.5281/zenodo.5244255" httpUri="https://zenodo.org/record/5244255/files/figure.png" pageId="77" pageNumber="78">14</figureCitation>
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<figureCitation id="0DA12A1B950D1516C93899393C9EFBB3" box="[339,409,1047,1072]" captionStart-0="FIGURE 32" captionStart-1="FIGURE 33" captionStart-2="FIGURE 34" captionStart-3="FIGURE 35" captionStart-4="FIGURE 36" captionStart-5="FIGURE 37" captionStartId-0="79.[151,250,1856,1879]" captionStartId-1="80.[151,250,1861,1884]" captionStartId-2="81.[151,248,1847,1870]" captionStartId-3="82.[151,250,1868,1891]" captionStartId-4="84.[151,244,998,1019]" captionStartId-5="84.[151,244,1923,1944]" captionTargetBox-0="[163,1422,154,1830]" captionTargetBox-1="[159,1425,156,1841]" captionTargetBox-2="[161,1423,152,1832]" captionTargetBox-3="[161,1422,155,1835]" captionTargetBox-4="[315,1278,203,971]" captionTargetBox-5="[330,1262,1142,1889]" captionTargetId-0="figure-142@79.[151,1436,141,1848]" captionTargetId-1="figure-146@80.[151,1436,141,1850]" captionTargetId-2="figure-146@81.[151,1436,137,1845]" captionTargetId-3="figure-91@82.[151,1436,141,1849]" captionTargetId-4="figure-63@84.[315,1279,193,979]" captionTargetId-5="figure-66@84.[315,1279,1127,1900]" captionTargetPageId-0="79" captionTargetPageId-1="80" captionTargetPageId-2="81" captionTargetPageId-3="82" captionTargetPageId-4="84" captionTargetPageId-5="84" captionText-0="FIGURE 32. Ranitomeya Plate 11. variabilis group. A–P: Ranitomeya variabilis (Highland morph): A: Chazuta, San Martin, Peru; B–G: Upper Cainarachi Valley, San Martin, Peru (Ω); H &amp; I: Borja, Loreto, Peru (24, 46Φ); J: Parque Nacional Ichigkat Muja, Amazonas (D. Rodriquez-Mercado); K: Saposoa, San Martin, Peru (Ω); L: Comparision between Saposoa and upper Cainarachi Valley populations (respectively); M: Xanthosoma sp. that was being used by R. variabilis for tadpole and egg deposition near Saposoa, San Martin, Peru; N: Saposoa, San Martin, Peru (Ω); O: Tocache, San Martin, Peru (C. Torres); P: Macas, Morona Santiago, Ecuador (J. Verkade, Ω). (nΦ = number of individual in phylogeny, Ω = population sampled in phylogeny)." captionText-1="FIGURE 33. Ranitomeya Plate 12. variabilis group: A–P: Ranitomeya variabilis (Lowland morph): A: Contamana, Loreto, Peru (42Φ); B: Shamboyacu, San Martin, Peru (41Φ); C: Lower Huallaga Canyon, San Martin, Peru (‡); D: Callanayacu, San Martin, Peru(‡); E: Barranquita, San Martin, Peru (‡); F: Pongo de Cainarachi, San Martin, Peru (7,8,11Φ); G &amp; H: Bonilla, San Martin, Peru (1,3, 9 Φ); I &amp; J: Varadero, Loreto, Peru (27Φ); K: Quebrada Blanco, Loreto, Peru (Ω); L: Rio Boncuya, Loreto, Peru (G. Gagliardi); M: Upper Rio Nanay, Loreto, Peru; N: Middle Rio Nanay, Loreto, Peru; O &amp; P: Archidona, Napo, Ecuador (EHP and J. Verkade). (nΦ = number of individual in phylogeny, Ω = population sampled in phylogeny)." captionText-2="FIGURE 34. Ranitomeya Plate 13. variabilis group: A–K: Ranitomeya variabilis: A: Archidona, Napo, Ecuador (J. Verkade); B– D: Macuma, Morona-Santiago, Ecuador (J. Verkade and EHP); E &amp;F: Puyo, Pastaza, Ecuador (j. Verkade) G: Yupati, Vaupés, Colombia (14 Φ); H: Embryos, Archidona, Napo, Ecuador (J. Verkade); I: Breeding pair of R. variabilis, Archidona, Napo, Ecuador (J. Verkade); J &amp; K: Yupati, Vaupés, Colombia (14-16 Φ). L–Q: Ranitomeya amazonica: L: Km 41 Iquitos, Loreto, Peru (ET, 26Φ); M: Km 31 Iquitos, Loreto, Peru (PPP); N &amp; O; Km 26 Iquitos, Loreto, Peru (ET, 25, 27Φ); P: Upper Rio Mazan-Pintuyacu, Loreto, Peru (J. J. Lopez-Rojas); Q: Lower Rio Mazan, Loreto, Peru (J. J. Lopez-Rojas). (nΦ = number of individual in phylogeny)." captionText-3="FIGURE 35. Ranitomeya Plate 14. variabilis group: A–Q: Ranitomeya amazonica: A–C: Iquitos, Loreto, Peru; D–K: ‘Arena Blanca’, Loreto, Peru (16–18, 20–24Φ); L: 30 km west of Pevas, Loreto Peru; M: Rio Sucusari, Loreto, Peru (B. Pieper); N &amp; O: Nouragues French Guiana (EHP); P: French Guiana (B.P. Noonan, Ω); Q: Estação Científica Ferreira Penna-Caxiuanã, Para, Brazil.(nΦ = number of individual in phylogeny, Ω=population sampled in phylogeny)." captionText-4="FIGURE 36. Known distribution of Ranitomeya amazonica. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya)." captionText-5="FIGURE 37. Known distribution of Ranitomeya variabilis. The inset map displays the geographic extent of distributions (black circles = all other Ranitomeya)." figureDoi-0="http://doi.org/10.5281/zenodo.5244315" figureDoi-1="http://doi.org/10.5281/zenodo.5244319" figureDoi-2="http://doi.org/10.5281/zenodo.5244321" figureDoi-3="http://doi.org/10.5281/zenodo.5244325" figureDoi-4="http://doi.org/10.5281/zenodo.5244329" figureDoi-5="http://doi.org/10.5281/zenodo.5244331" httpUri-0="https://zenodo.org/record/5244315/files/figure.png" httpUri-1="https://zenodo.org/record/5244319/files/figure.png" httpUri-2="https://zenodo.org/record/5244321/files/figure.png" httpUri-3="https://zenodo.org/record/5244325/files/figure.png" httpUri-4="https://zenodo.org/record/5244329/files/figure.png" httpUri-5="https://zenodo.org/record/5244331/files/figure.png" pageId="77" pageNumber="78">
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37
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</paragraph>
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<tableCitation id="D8180325950D1516C8FC99153DF3FBCF" box="[151,244,1083,1107]" captionStart="TABLE 1" captionStartId="12.[151,241,151,175]" captionText="TABLE 1A (continued on next page). Currently accepted taxa (based on Frost et al. 2011). History of alpha-taxonomic and species group (in parentheses) classification of taxa in the genera Ranitomeya and Andinobates. Dendrobates (minutus)* = by implication, &quot;+&quot; species which were not described at the time of classification, &quot;*&quot; species which were not acknowledged by the authors." pageId="77" pageNumber="78">Tables 1</tableCitation>
, 
<tableCitation id="D8180325950D1516C96899153C16FBCF" box="[259,273,1083,1107]" captionStart="TABLE 4" captionStartId="27.[151,241,157,181]" captionText="TABLE 4A (continued on next page). Basic Tadpole Morphology." pageId="77" pageNumber="78">4</tableCitation>
<emphasis id="A7EEEA8C950D1516C97B99123C19FBC8" bold="true" box="[272,286,1084,1108]" pageId="77" pageNumber="78">–</emphasis>
6
</paragraph>
<paragraph id="9525369E950D1516C8FC99533CB6FB25" blockId="77.[151,1437,1149,1508]" pageId="77" pageNumber="78">
A monophyletic assemblage of two species: 
<taxonomicName id="529A4D1D950D1516CACB99503841FB09" authority="Zimmermann &amp; Zimmermann 1988" authorityName="sensu Zimmermann &amp; Zimmermann" authorityYear="1988" box="[672,1350,1149,1173]" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="77" pageNumber="78" phylum="Chordata" rank="species" species="variabilis">
<emphasis id="A7EEEA8C950D1516CACB99503E9BFB09" box="[672,924,1149,1173]" italics="true" pageId="77" pageNumber="78">Ranitomeya variabilis</emphasis>
<bibRefCitation id="F10B4B6F950D1516CBCC99533841FB09" author="Zimmermann, H. &amp; Zimmermann, E." box="[935,1350,1149,1173]" pageId="77" pageNumber="78" pagination="125 - 160" refId="ref80585" refString="Zimmermann, H., &amp; Zimmermann, E. (1988) Etho-Taxonomie und zoogeographische Artengruppenbildung bei Pfeilgiftfroschen (Anura: Dendrobatidae). Salamandra, 24, 125 - 160." type="journal article" year="1988">Zimmermann &amp; Zimmermann 1988</bibRefCitation>
</taxonomicName>
and 
<taxonomicName id="529A4D1D950D1516CDEF99503CAAFB25" authority="Schulte 1999" authorityName="Schulte" authorityYear="1999" class="Amphibia" family="Dendrobatidae" genus="Ranitomeya" kingdom="Animalia" order="Anura" pageId="77" pageNumber="78" phylum="Chordata" rank="species" species="amazonica">
<emphasis id="A7EEEA8C950D1516CDEF99503C17FB25" italics="true" pageId="77" pageNumber="78">R. amazonica</emphasis>
<bibRefCitation id="F10B4B6F950D1516C97C998F3CAAFB25" author="Schulte, R." box="[279,429,1185,1209]" pageId="77" pageNumber="78" refId="ref78797" refString="Schulte, R. (1999) Pfeilgiftfrosche &quot; Artenteil - Peru &quot;. INBICO, Wailblingen, Germany, 294 pp." type="book" year="1999">Schulte 1999</bibRefCitation>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection id="DD806515950D1516C8AC99EA3892FA78" pageId="77" pageNumber="78" type="diagnosis">
<paragraph id="9525369E950D1516C8AC99EA3892FA78" blockId="77.[151,1437,1149,1508]" pageId="77" pageNumber="78">
<emphasis id="A7EEEA8C950D1516C8AC99EA3F06FB42" bold="true" box="[199,513,1220,1246]" pageId="77" pageNumber="78">Definition and diagnosis.</emphasis>
<materialsCitation id="25F23CC3950D1516CA6199EB3896FA78" collectionCode="SVL" pageId="77" pageNumber="78" specimenCode="LTRF 2" specimenCount="6" specimenCount-egg="6">
Medium to large adult size (
<collectionCode id="F38BAE5B950D1516CB3699EB3E91FB43" box="[861,918,1221,1247]" pageId="77" pageNumber="78">SVL</collectionCode>
<quantity id="52629B7B950D1516CBCB99EB3923FB43" box="[928,1060,1221,1247]" metricMagnitude="-2" metricUnit="m" metricValue="1.75" metricValueMax="2.1" metricValueMin="1.4" pageId="77" pageNumber="78" unit="mm" value="17.5" valueMax="21.0" valueMin="14.0">
14 
<emphasis id="A7EEEA8C950D1516CBD699E93ECCFB42" box="[957,971,1223,1246]" italics="true" pageId="77" pageNumber="78">–</emphasis>
21 mm
</quantity>
); two prevalent dorsal patterns: large ovoid black spots on greenish dorsum or complete yellow to red dorsolateral stripes, middorsal stripe and oblique lateral stripes (note that occasionally these species possess ventrolateral stripes, or an intermediate between oblique lateral and ventrolateral stripes, suggesting this trait is a continuous character); large intestine of larvae entirely pigmented; 
<specimenCode id="C53C9EE5950D1516CA3398743FB2FAE8" box="[600,693,1370,1396]" collectionCode="LTRF" pageId="77" pageNumber="78">LTRF 2</specimenCode>
(2)/3(1), oral disc emarginated; larvae gray, tadpoles cannibalistic on embryos and other tadpoles; eggs gray; neither males nor females territorial; promiscuous mating system with male parental care; females produce between 2 
<emphasis id="A7EEEA8C950D1516CB5698893E4CFA22" box="[829,843,1447,1470]" italics="true" pageId="77" pageNumber="78">–</emphasis>
<specimenCount id="839CFD17950D1516CB20988B3E9EFA23" box="[843,921,1445,1471]" pageId="77" pageNumber="78" type="egg">6 eggs</specimenCount>
per mating. Advertisement calls consist of short, regularly spaced buzz calls, notes 0.16 
<emphasis id="A7EEEA8C950D1516CACB98E23FA9FA7F" box="[672,686,1484,1507]" italics="true" pageId="77" pageNumber="78">–</emphasis>
0.44 sec in length, repeated at 24 
<emphasis id="A7EEEA8C950D1516CC4698E2393CFA7F" box="[1069,1083,1484,1507]" italics="true" pageId="77" pageNumber="78">–</emphasis>
70 notes per minute (
<figureCitation id="0DA12A1B950D1516CD5B98E43880FA78" box="[1328,1415,1482,1508]" captionStart="FIGURE 14" captionStartId="46.[151,244,1819,1840]" captionTargetBox="[156,1424,1117,1789]" captionTargetId="figure-447@46.[151,1436,1116,1801]" captionTargetPageId="46" captionText="FIGURE 14. Advertisement calls of Ranitomeya species in the variabilis group and defleri group. A. Ranitomeya amazonica from 23 km S Iquitos, Loreto, Peru (type locality), recorded at 26° C; B. Ranitomeya amazonica from French Guiana, unknown temperature (call courtesy Erik Poelman); C. Ranitomeya variabilis from Cainarachi valley, San Martín, Peru, recorded at 22° C. D. Ranitomeya variabilis from Cerro Yupatí, Amazonas, Colombia, recorded at 27° C; E. Ranitomeya variabilis from Saposoa, San Martín, Peru, recorded at 24.5 C; F. Ranitomeya defleri from Rio Apaporis, Vaupés, Colombia, recorded at 26° C." figureDoi="http://doi.org/10.5281/zenodo.5244255" httpUri="https://zenodo.org/record/5244255/files/figure.png" pageId="77" pageNumber="78">Fig. 14</figureCitation>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
</treatment>
</document>