treatments-xml/data/04/21/27/0421277A8B3BB5419DE962FEFB5852F5.xml

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<mods:title id="78353D25C22BFA0C8B6A1E6C8494A754">A new moss salamander, genus Nototriton (Caudata: Plethodontidae), from the Cordillera de Talamanca, in the Costa Rica-Panama border region</mods:title>
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<mods:namePart id="AC56C497EB490225292D1FB86ECC6A67">Arias, Erick</mods:namePart>
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<emphasis id="BEFC4A7E8B3BB54C9DE962FEFDE75787" bold="true" box="[151,532,151,178]" pageId="6" pageNumber="493">
<taxonomicName id="4B88EDEF8B3BB54C9DE962FEFE435784" ID-CoL="47Z9R" authority="Arias &amp; Kubicki, 2018" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[151,432,151,177]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="costaricense" status="sp. nov.">
<emphasis id="BEFC4A7E8B3BB54C9DE962FEFE435784" bold="true" box="[151,432,151,177]" italics="true" pageId="6" pageNumber="493">Nototriton costaricense</emphasis>
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<heading id="D77F21008B3BB54C9DE962B7FE0257CD" bold="true" box="[151,497,222,248]" fontSize="11" level="1" pageId="6" pageNumber="493" reason="1">
<emphasis id="BEFC4A7E8B3BB54C9DE962B7FE0257CD" bold="true" box="[151,497,222,248]" pageId="6" pageNumber="493">Southern Moss Salamander</emphasis>
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<paragraph id="8C37966C8B3BB54C9DE96368FEDD562F" blockId="6.[151,497,222,282]" box="[151,302,257,282]" pageId="6" pageNumber="493">
(
<figureCitation id="14B38AE98B3BB54C9DE06368FED5562F" box="[158,294,257,282]" captionStart-0="FIGURE 3" captionStart-1="FIGURE 4" captionStart-2="FIGURE 5" captionStartId-0="6.[151,250,1339,1361]" captionStartId-1="7.[151,250,1584,1606]" captionStartId-2="9.[151,250,1002,1024]" captionTargetBox-0="[166,1424,830,1298]" captionTargetBox-1="[150,1437,555,1559]" captionTargetBox-2="[151,1435,447,977]" captionTargetId-0="figure@6.[151,1436,805,1315]" captionTargetId-1="figure@7.[150,1437,555,1559]" captionTargetId-2="figure@9.[151,1435,447,977]" captionTargetPageId-0="6" captionTargetPageId-1="7" captionTargetPageId-2="9" captionText-0="FIGURE 3. Nototriton costaricense sp. nov. Photograph taken byBK." captionText-1="FIGURE 4. Images demonstrating the dorsal, lateral, and ventral aspects of the holotype of Nototriton costaricense sp. nov. Photographs taken by BK." captionText-2="FIGURE 5. Views of the nostrils of Nototriton costaricense sp. nov. Photograph taken by BK." httpUri-0="https://zenodo.org/record/1136408/files/figure.png" httpUri-1="https://zenodo.org/record/1136410/files/figure.png" httpUri-2="https://zenodo.org/record/1136412/files/figure.png" pageId="6" pageNumber="493">Figures 3–5</figureCitation>
)
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<paragraph id="8C37966C8B3BB54C9DE96320FAC856B3" blockId="6.[151,1437,329,714]" pageId="6" pageNumber="493">
<materialsCitation id="3CE09C318B3BB54C9DE96320FAC856B3" ID-GBIF-Occurrence="1802522134" collectingDate="2015-03-11" collectionCode="UCR" collectorName="Erick Arias" country="Costa Rica" elevation="1500" location="Distrito de Telire" municipality="Cantón de Talamanca" pageId="6" pageNumber="493" specimenCode="UCR 22900" specimenCount="1" specimenCount-subadult="1" stateProvince="Provincia de Limón" typeStatus="holotype">
<emphasis id="BEFC4A7E8B3BB54C9DE96320FEF85657" bold="true" box="[151,267,329,354]" pageId="6" pageNumber="493">
<typeStatus id="533328CE8B3BB54C9DE96320FEF55657" box="[151,262,329,354]" pageId="6" pageNumber="493" type="holotype">Holotype</typeStatus>
.
</emphasis>
<specimenCode id="DC2E3E178B3BB54C9C6C6320FE685654" box="[274,411,329,353]" pageId="6" pageNumber="493">UCR 22900</specimenCode>
, a 
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from 
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: 
<collectingRegion id="4E4C588E8B3BB54C9F996320FC3A5657" box="[743,969,329,354]" country="Costa Rica" name="Limon" pageId="6" pageNumber="493">Provincia de Limón</collectingRegion>
: 
<collectingMunicipality id="6C530C168B3BB54C9EAA6320FB385657" box="[980,1227,329,354]" pageId="6" pageNumber="493">Cantón de Talamanca</collectingMunicipality>
: 
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: 
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, 
<location id="8957C0B78B3BB54C9C716307FD7656B3" LSID="urn:lsid:plazi:treatment:0421277A8B3BB5419DE962FEFB5852F5:8957C0B78B3BB54C9C716307FD7656B3" box="[271,645,365,390]" country="Costa Rica" municipality="Cantón de Talamanca" name="Parque Internacional La Amistad" pageId="6" pageNumber="493" stateProvince="Provincia de Limón">Parque Internacional La Amistad</location>
, ca. 
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<quantity id="4B703B898B3BB54C9FC96304FCFF56B0" box="[695,780,365,390]" metricMagnitude="3" metricUnit="m" metricValue="1.5" pageId="6" pageNumber="493" unit="m" value="1500.0">1500 m</quantity>
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a.s.l., obtained by 
<collectorName id="217DF3BA8B3BB54C9E9E6304FB9356B3" box="[992,1120,365,390]" pageId="6" pageNumber="493">Erick Arias</collectorName>
on 
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<date id="F836B0AC8B3BB54C99F46307FACB56B3" box="[1162,1336,365,390]" pageId="6" pageNumber="493" value="2015-03-11">11 March 2015</date>
</collectingDate>
.
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</paragraph>
</subSubSection>
<subSubSection id="C492C5E78B3BB54C9DB963F8FF1656C7" pageId="6" pageNumber="493" type="discussion">
<paragraph id="8C37966C8B3BB54C9DB963F8FF1656C7" blockId="6.[151,1437,329,714]" pageId="6" pageNumber="493">
<emphasis id="BEFC4A7E8B3BB54C9DB963F8FE5A569F" bold="true" box="[199,425,401,426]" pageId="6" pageNumber="493">Generic Placement</emphasis>
. Assigned to the genus 
<taxonomicName id="4B88EDEF8B3BB54C9FCD63FBFCDB569C" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[691,808,402,425]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B3BB54C9FCD63FBFCDB569C" box="[691,808,402,425]" italics="true" pageId="6" pageNumber="493">Nototriton</emphasis>
</taxonomicName>
due to having fewer than 14 costal grooves and reduced 
<emphasis id="BEFC4A7E8B3BB54C9DE963DFFF1356F8" box="[151,224,438,461]" italics="true" pageId="6" pageNumber="493">manus</emphasis>
and 
<emphasis id="BEFC4A7E8B3BB54C9C6763DFFECD56F8" box="[281,318,438,461]" italics="true" pageId="6" pageNumber="493">pes</emphasis>
that are longer than wide, and to the subgenus 
<taxonomicName id="4B88EDEF8B3BB54C9E2263DFFC2256F8" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[860,977,438,461]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B3BB54C9E2263DFFC2256F8" box="[860,977,438,461]" italics="true" pageId="6" pageNumber="493">Nototriton</emphasis>
</taxonomicName>
based the molecular evidence presented herein.
</paragraph>
</subSubSection>
<subSubSection id="C492C5E78B3BB54C9DB96394FAB752C8" pageId="6" pageNumber="493" type="diagnosis">
<paragraph id="8C37966C8B3BB54C9DB96394FE5655FF" blockId="6.[151,1437,329,714]" pageId="6" pageNumber="493">
<emphasis id="BEFC4A7E8B3BB54C9DB96394FEB25523" bold="true" box="[199,321,509,534]" pageId="6" pageNumber="493">Diagnosis.</emphasis>
The combination of the following characteristics can be used to distinguish 
<taxonomicName id="4B88EDEF8B3BB54C99EA6397FA6F5520" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[1172,1436,510,533]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B3BB54C99EA6397FA6F5520" box="[1172,1436,510,533]" italics="true" pageId="6" pageNumber="493">Nototriton costaricense</emphasis>
</taxonomicName>
from other described species in the subgenus 
<taxonomicName id="4B88EDEF8B3BB54C9FE2604BFCE3550C" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[668,784,546,569]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B3BB54C9FE2604BFCE3550C" box="[668,784,546,569]" italics="true" pageId="6" pageNumber="493">Nototriton</emphasis>
</taxonomicName>
: (1) having at least the distal phalanges on fingers II, III, and IV in addition to the distal phalanges on toes II, III, IV, and V free of webbing; (2) large nostril openings, greater than 
<quantity id="4B703B898B3BB54C9C5A6003FE8955B4" box="[292,378,618,642]" metricMagnitude="-4" metricUnit="m" metricValue="3.0" pageId="6" pageNumber="493" unit="mm" value="0.3">0.3 mm</quantity>
in diameter (
<figureCitation id="14B38AE98B3BB54C9F6E6000FDA755B7" box="[528,596,617,642]" captionStart="FIGURE 5" captionStartId="9.[151,250,1002,1024]" captionTargetBox="[151,1435,447,977]" captionTargetId="figure@9.[151,1435,447,977]" captionTargetPageId="9" captionText="FIGURE 5. Views of the nostrils of Nototriton costaricense sp. nov. Photograph taken by BK." httpUri="https://zenodo.org/record/1136412/files/figure.png" pageId="6" pageNumber="493">Fig. 5</figureCitation>
); (3) robust habitus, TW = 14.6 % of SL; (4) long hind limbs, HLL = 21.9 % of SL; (4) differentiated from other members of the genus 
<taxonomicName id="4B88EDEF8B3BB54C9E3960E7FC4F5590" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[839,956,654,677]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B3BB54C9E3960E7FC4F5590" box="[839,956,654,677]" italics="true" pageId="6" pageNumber="493">Nototriton</emphasis>
</taxonomicName>
, subgenus 
<taxonomicName id="4B88EDEF8B3BB54C994460E7FB5C5590" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[1082,1199,654,677]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B3BB54C994460E7FB5C5590" box="[1082,1199,654,677]" italics="true" pageId="6" pageNumber="493">Nototriton</emphasis>
</taxonomicName>
by its 16S, COI, and cyt 
<emphasis id="BEFC4A7E8B3BB54C9DBE60D8FF3D55FF" box="[192,206,689,714]" italics="true" pageId="6" pageNumber="493">b</emphasis>
mtDNA distances.
</paragraph>
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<emphasis id="BEFC4A7E8B3BB54C9DE96752FEE15265" bold="true" box="[151,274,1339,1361]" pageId="6" pageNumber="493">FIGURE 3.</emphasis>
<taxonomicName id="4B88EDEF8B3BB54C9C676755FDF45264" authority="Arias &amp; Kubicki, 2018" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[281,519,1340,1361]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="costaricense" status="sp. nov.">
<emphasis id="BEFC4A7E8B3BB54C9C676755FDF45264" box="[281,519,1340,1361]" italics="true" pageId="6" pageNumber="493">Nototriton costaricense</emphasis>
</taxonomicName>
<emphasis id="BEFC4A7E8B3BB54C9F736755FDAD5264" bold="true" box="[525,606,1340,1361]" pageId="6" pageNumber="493">
<taxonomicNameLabel id="A5CFF7058B3BB54C9F736755FDAD5264" box="[525,606,1340,1361]" pageId="6" pageNumber="493" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
Photograph taken by BK.
</paragraph>
</caption>
<paragraph id="8C37966C8B3BB54C9DB96711FAB752C8" blockId="6.[151,1437,1400,2037]" pageId="6" pageNumber="493">
<emphasis id="BEFC4A7E8B3BB54C9DB96711FE9F52A4" bold="true" box="[199,364,1400,1425]" pageId="6" pageNumber="493">Comparisons.</emphasis>
Since 
<taxonomicName id="4B88EDEF8B3BB54C9CC06713FD3B52A4" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[446,712,1402,1425]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B3BB54C9CC06713FD3B52A4" box="[446,712,1402,1425]" italics="true" pageId="6" pageNumber="493">Nototriton costaricense</emphasis>
</taxonomicName>
is only known to occur in 
<collectingCountry id="F49FD6FC8B3BB54C99786710FB7052A4" box="[1030,1155,1400,1425]" name="Costa Rica" pageId="6" pageNumber="493">Costa Rica</collectingCountry>
and molecular evidence strongly supports it forming part of the 
<taxonomicName id="4B88EDEF8B3BB54C9F1F67F4FCC05280" box="[609,819,1436,1461]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B3BB54C9F1F67F4FCC05280" box="[609,819,1436,1461]" italics="true" pageId="6" pageNumber="493">Nototriton picadoi</emphasis>
</taxonomicName>
species group, phenotypic comparisons are presented here only with respect to members of that clade (i.e., 
<emphasis id="BEFC4A7E8B3BB54C9E6B67ABFF0B52C8" italics="true" pageId="6" pageNumber="493">
<taxonomicName id="4B88EDEF8B3BB54C9E6B67ABFC4C52EC" baseAuthorityName="Taylor" baseAuthorityYear="1948" box="[789,959,1472,1497]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="abscondens">N. abscondens</taxonomicName>
, 
<taxonomicName id="4B88EDEF8B3BB54C9EB067ABFBB552EC" authorityName="Good &amp; Wake" authorityYear="1993" box="[974,1094,1474,1497]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="gamezi">N. gamezi</taxonomicName>
, 
<taxonomicName id="4B88EDEF8B3BB54C992B67ABFB0E52EC" authorityName="Garcia-Paris &amp; Wake" authorityYear="2000" box="[1109,1277,1474,1497]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="guanacaste">N. guanacaste</taxonomicName>
, 
<taxonomicName id="4B88EDEF8B3BB54C987267ABFA8052EC" authorityName="Good &amp; Wake" authorityYear="1993" box="[1292,1395,1474,1497]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="major">N. major</taxonomicName>
, 
<taxonomicName id="4B88EDEF8B3BB54C98FC67ABFF0752C8" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="matama">N. matama</taxonomicName>
,
</emphasis>
and 
<taxonomicName id="4B88EDEF8B3BB54C9C50678FFE5052C8" box="[302,419,1508,1533]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B3BB54C9C50678FFE5052C8" box="[302,419,1508,1533]" italics="true" pageId="6" pageNumber="493">N. picadoi</emphasis>
</taxonomicName>
), which are all endemic to 
<collectingCountry id="F49FD6FC8B3BB54C9FAA678CFCBD52C8" box="[724,846,1508,1533]" name="Costa Rica" pageId="6" pageNumber="493">Costa Rica</collectingCountry>
(
<bibRefCitation id="E819EB9D8B3BB54C9E23678DFB8652C8" author="Boza-Oviedo" box="[861,1141,1508,1533]" pageId="6" pageNumber="493" refString="Boza-Oviedo, E., Rovito, S. M., Chaves, G., Garcia-Rodriguez, A., Artavia, L. G., Bolanos, F. &amp; Wake, D. B. (2012) Salamanders from the eastern Cordillera de Talamanca, Costa Rica, with descriptions of five new species (Plethodontidae: Bolitoglossa, Nototriton, and Oedipina) and natural history notes from recent expeditions. Zootaxa, 3309, 36 - 61." type="journal article" year="2012">
Boza-Oviedo 
<emphasis id="BEFC4A7E8B3BB54C9E85678FFBDF52C8" box="[1019,1068,1508,1533]" italics="true" pageId="6" pageNumber="493">et al</emphasis>
. 2012
</bibRefCitation>
; 
<bibRefCitation id="E819EB9D8B3BB54C99FF678DFACB52C8" author="Townsend" box="[1153,1336,1508,1533]" pageId="6" pageNumber="493" refString="Townsend, J. H. (2016) Taxonomic revision of the moss salamander Nototriton barbouri (Schmidt) (Caudata: Plethodontidae), with description of two new species from the Cordillera Nombre de Dios, Honduras. Zootaxa, 4196 (4), 511 - 528. https: // doi. org / 10.11646 / zootaxa. 4196.4.3" type="journal article" year="2016">Townsend 2016</bibRefCitation>
).
</paragraph>
</subSubSection>
<subSubSection id="C492C5E78B3BB5439DB96461FE975298" lastPageId="9" lastPageNumber="496" pageId="6" pageNumber="493" type="description">
<paragraph id="8C37966C8B3BB54D9DB96461FF0A56C1" blockId="6.[151,1437,1400,2037]" lastBlockId="7.[151,1436,151,500]" lastPageId="7" lastPageNumber="494" pageId="6" pageNumber="493">
Contrasting characteristics for 
<taxonomicName id="4B88EDEF8B3BB54C9F556463FCC45114" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[555,823,1546,1569]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B3BB54C9F556463FCC45114" box="[555,823,1546,1569]" italics="true" pageId="6" pageNumber="493">Nototriton costaricense</emphasis>
</taxonomicName>
are presented in parentheses. 
<taxonomicName id="4B88EDEF8B3BB54C99E46463FEC45170" authority="(Taylor, 1948)" authorityName="Taylor" authorityYear="1948" baseAuthorityName="Taylor" baseAuthorityYear="1948" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="abscondens">
<emphasis id="BEFC4A7E8B3BB54C99E46463FA6F5114" box="[1178,1436,1544,1569]" italics="true" pageId="6" pageNumber="493">Nototriton abscondens</emphasis>
(
<bibRefCitation id="E819EB9D8B3BB54C9DE06445FEDC5170" author="Taylor" box="[158,303,1580,1605]" pageId="6" pageNumber="493" refString="Taylor, E. H. (1948) New Costa Rican salamanders. Proceedings of Biological Society of Washington, 61, 177 - 180." type="journal article" year="1948">Taylor, 1948</bibRefCitation>
)
</taxonomicName>
can be distinguished from 
<taxonomicName id="4B88EDEF8B3BB54C9F166447FCE65171" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[616,789,1581,1605]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B3BB54C9F166447FCE65171" box="[616,789,1581,1605]" italics="true" pageId="6" pageNumber="493">N. costaricense</emphasis>
</taxonomicName>
by having relatively smaller nostrils openings, 0.3–0.8 % of SL (RNW = 1.3 % of SL); relatively narrower head, 12–14 % of SL (broader head, HeW = 15.1 % of SL); slender trunk, TW 8–10 % of SL (thick trunk, TW 14.6 % of SL). 
<taxonomicName id="4B88EDEF8B3BB54C9EDB641CFA9E51B8" authority="Good &amp; Wake, 1993" authorityName="Good &amp; Wake" authorityYear="1993" box="[933,1389,1652,1677]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="gamezi">
<emphasis id="BEFC4A7E8B3BB54C9EDB641CFB8051B9" box="[933,1139,1653,1676]" italics="true" pageId="6" pageNumber="493">Nototriton gamezi</emphasis>
<bibRefCitation id="E819EB9D8B3BB54C9903641DFA9E51B8" author="Good" box="[1149,1389,1652,1677]" pageId="6" pageNumber="493" refString="Good, D. A. &amp; Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159. https: // doi. org / 10.2307 / 1466956" type="journal article" year="1993">
Good &amp; 
<collectingCountry id="F49FD6FC8B3BB54C999B641DFAD451B8" box="[1253,1319,1652,1677]" name="Wake Island" pageId="6" pageNumber="493">Wake</collectingCountry>
, 1993
</bibRefCitation>
</taxonomicName>
has relatively shorter hind limbs, 16–17 % of SL (longer hind limb, HLL = 21.9 % of SL); narrower head, 13–14 % of SL (broader head, HeW = 15.1 % of SL); slender trunk, TW 8–10 % of SL (thick trunk, TW 14.6 % of SL). 
<taxonomicName id="4B88EDEF8B3BB54C9DE9648BFD3851CC" authority="Garcia-Paris &amp; Wake, 2000" authorityName="Garcia-Paris &amp; Wake" authorityYear="2000" box="[151,715,1760,1785]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="guanacaste">
<emphasis id="BEFC4A7E8B3BB54C9DE9648BFE6151CC" box="[151,402,1762,1785]" italics="true" pageId="6" pageNumber="493">Nototriton guanacaste</emphasis>
<bibRefCitation id="E819EB9D8B3BB54C9CE66488FD3851CC" author="Garcia-Paris" box="[408,715,1760,1785]" pageId="6" pageNumber="493" refString="Garcia-Paris, M. &amp; Wake, D. B. (2000) Molecular phylogenetic analysis of relationships of the tropical salamander genera Oedipina and Nototriton, with descriptions of a new genus and three new species. Copeia, 2000, 42 - 70. https: // doi. org / 10.1643 / 0045 - 8511 (2000) 2000 [0042: MPAORO] 2.0. CO; 2" type="journal article" year="2000">
García-París &amp; 
<collectingCountry id="F49FD6FC8B3BB54C9F386489FD7B51CC" box="[582,648,1760,1785]" name="Wake Island" pageId="6" pageNumber="493">Wake</collectingCountry>
, 2000
</bibRefCitation>
</taxonomicName>
has relatively smaller nostril openings, 0.4–0.95 % of SL (RNW = 1.3 % of SL); relatively shorter hind limb, 17–20 % of SL (longer hind limb, HLL = 21.9 % of SL). 
<taxonomicName id="4B88EDEF8B3BB54C9859656FFE3C5074" authority="Good &amp; Wake, 1993" authorityName="Good &amp; Wake" authorityYear="1993" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="major">
<emphasis id="BEFC4A7E8B3BB54C9859656FFF295074" italics="true" pageId="6" pageNumber="493">Nototriton major</emphasis>
<bibRefCitation id="E819EB9D8B3BB54C9D9D6541FE3C5074" author="Good" box="[227,463,1832,1857]" pageId="6" pageNumber="493" refString="Good, D. A. &amp; Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159. https: // doi. org / 10.2307 / 1466956" type="journal article" year="1993">
Good &amp; 
<collectingCountry id="F49FD6FC8B3BB54C9C366541FE795074" box="[328,394,1832,1857]" name="Wake Island" pageId="6" pageNumber="493">Wake</collectingCountry>
, 1993
</bibRefCitation>
</taxonomicName>
has relatively smaller nostrils openings, 0.3 % of SL (RNW = 1.3 % of SL); shorter snout-gular length, 17.7 % of SL (longer snout-gular length, LGFS = 21 % of SL); relatively narrower head, 11.3 % of SL (broader head, HeW = 15.1 % of SL); slender trunk, TW 9–10 % of SL (trunk robust, TW 14.6 % of SL); relatively shorter hind limb, 19.3 % of SL (longer hind limb, HLL = 21.9 % of SL); relatively shorter fore limb, 14.8 % of SL (longer front limb, FLL = 20.1 % of SL). 
<taxonomicName id="4B88EDEF8B3BB54C9E7965D3FB0C50E4" authority="Boza-Oviedo et al., 2012" authorityName="Boza-Oviedo et al." authorityYear="2012" box="[775,1279,1976,2001]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="6" pageNumber="493" phylum="Chordata" rank="species" species="matama">
<emphasis id="BEFC4A7E8B3BB54C9E7965D3FC2E50E4" box="[775,989,1978,2001]" italics="true" pageId="6" pageNumber="493">Nototriton matama</emphasis>
<bibRefCitation id="E819EB9D8B3BB54C9E9A65D1FB0C50E4" author="Boza-Oviedo" box="[996,1279,1976,2001]" pageId="6" pageNumber="493" refString="Boza-Oviedo, E., Rovito, S. M., Chaves, G., Garcia-Rodriguez, A., Artavia, L. G., Bolanos, F. &amp; Wake, D. B. (2012) Salamanders from the eastern Cordillera de Talamanca, Costa Rica, with descriptions of five new species (Plethodontidae: Bolitoglossa, Nototriton, and Oedipina) and natural history notes from recent expeditions. Zootaxa, 3309, 36 - 61." type="journal article" year="2012">
Boza-Oviedo 
<emphasis id="BEFC4A7E8B3BB54C99FE65D3FB4250E4" box="[1152,1201,1976,2001]" italics="true" pageId="6" pageNumber="493">et al</emphasis>
., 2012
</bibRefCitation>
</taxonomicName>
has relatively larger nostrils openings, 2 % of SL (RNW = 1.3 % of SL); pointed outer toe tips (rounded outer toe tips); relatively narrower head, 14 % of SL (broader head, HeW = 15.1 % of SL). 
<taxonomicName id="4B88EDEF8B3AB54D9ECC62F0FAA75785" authority="(Stejneger, 1911)" authorityName="Stejneger" authorityYear="1911" baseAuthorityName="Stejneger" baseAuthorityYear="1911" box="[946,1364,151,176]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="7" pageNumber="494" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B3AB54D9ECC62F0FB765785" box="[946,1157,151,176]" italics="true" pageId="7" pageNumber="494">Nototriton picadoi</emphasis>
(
<bibRefCitation id="E819EB9D8B3AB54D99E662FEFABF5785" author="Stejneger" box="[1176,1356,151,176]" pageId="7" pageNumber="494" refString="Stejneger, L. (1911) Description of three new batrachians from Costa Rica and Panama. Proceedings of the United States National Museum, 41, 285 - 288. https: // doi. org / 10.5479 / si. 00963801.1857.285" type="journal article" year="1911">Stejneger, 1911</bibRefCitation>
)
</taxonomicName>
has a relatively narrower head, 12–14 % of SL (broader head, HeW = 15.1 % of SL); slender trunk, TW 10–11 % of SL (trunk robust, TW 14.6 % of SL); relatively shorter hind limb, 18–19 % of SL (longer hind limb, HLL = 21.9 % of SL); relatively shorter fore limb, 16.9–18.2 % of SL (longer front limb, FLL = 20.1 % of SL). Although 
<taxonomicName id="4B88EDEF8B3AB54D9859636CFE1A5675" authority="Good &amp; Wake, 1993" authorityName="Good &amp; Wake" authorityYear="1993" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="7" pageNumber="494" phylum="Chordata" rank="species" species="tapanti">
<emphasis id="BEFC4A7E8B3AB54D9859636CFF155675" italics="true" pageId="7" pageNumber="494">Nototriton tapanti</emphasis>
<bibRefCitation id="E819EB9D8B3AB54D9D8C634EFE1A5675" author="Good" box="[242,489,295,320]" pageId="7" pageNumber="494" refString="Good, D. A. &amp; Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159. https: // doi. org / 10.2307 / 1466956" type="journal article" year="1993">
Good &amp; 
<collectingCountry id="F49FD6FC8B3AB54D9C21634EFE525675" box="[351,417,295,320]" name="Wake Island" pageId="7" pageNumber="494">Wake</collectingCountry>
, 1993
</bibRefCitation>
</taxonomicName>
is not a member of the 
<taxonomicName id="4B88EDEF8B3AB54D9E696340FC625675" baseAuthorityName="Stejneger" baseAuthorityYear="1911" box="[791,913,295,320]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="7" pageNumber="494" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B3AB54D9E696340FC625675" box="[791,913,295,320]" italics="true" pageId="7" pageNumber="494">N. picadoi</emphasis>
</taxonomicName>
species group, but we decided to include a phenotypic comparison here due to the fact that no DNA sequences of 
<taxonomicName id="4B88EDEF8B3AB54D9EAA6324FBB45651" authorityName="Good &amp; Wake" authorityYear="1993" box="[980,1095,333,356]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="7" pageNumber="494" phylum="Chordata" rank="species" species="tapanti">
<emphasis id="BEFC4A7E8B3AB54D9EAA6324FBB45651" box="[980,1095,333,356]" italics="true" pageId="7" pageNumber="494">N. tapanti</emphasis>
</taxonomicName>
were available for molecular comparison with 
<taxonomicName id="4B88EDEF8B3AB54D9C1B6318FDEB56BD" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[357,536,369,392]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="7" pageNumber="494" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B3AB54D9C1B6318FDEB56BD" box="[357,536,369,392]" italics="true" pageId="7" pageNumber="494">N. costaricense</emphasis>
</taxonomicName>
. 
<taxonomicName id="4B88EDEF8B3AB54D9F526318FD0F56BD" authorityName="Good &amp; Wake" authorityYear="1993" box="[556,764,369,392]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="7" pageNumber="494" phylum="Chordata" rank="species" species="tapanti">
<emphasis id="BEFC4A7E8B3AB54D9F526318FD0F56BD" box="[556,764,369,392]" italics="true" pageId="7" pageNumber="494">Nototriton tapanti</emphasis>
</taxonomicName>
can easily be distinguished from 
<taxonomicName id="4B88EDEF8B3AB54D99E76318FABF56BD" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[1177,1356,369,392]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="7" pageNumber="494" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B3AB54D99E76318FABF56BD" box="[1177,1356,369,392]" italics="true" pageId="7" pageNumber="494">N. costaricense</emphasis>
</taxonomicName>
by its syndactylous hands and feet (having at least the distal phalanges on fingers II, III, and IV in and toes II, III, IV, and V free of webbing); tips of free digits pointed and without subterminal pads (rounded tips and terminal pads present).
</paragraph>
<caption id="D8F7C6E48B3AB54D9DE96459FE505151" httpUri="https://zenodo.org/record/1136410/files/figure.png" pageId="7" pageNumber="494" targetBox="[150,1437,555,1559]" targetPageId="7">
<paragraph id="8C37966C8B3AB54D9DE96459FE505151" blockId="7.[151,1435,1584,1636]" pageId="7" pageNumber="494">
<emphasis id="BEFC4A7E8B3AB54D9DE96459FEE75170" bold="true" box="[151,276,1584,1606]" pageId="7" pageNumber="494">FIGURE 4.</emphasis>
Images demonstrating the dorsal, lateral, and ventral aspects of the holotype of 
<taxonomicName id="4B88EDEF8B3AB54D992C6458FAB25173" authority="Arias &amp; Kubicki, 2018" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[1106,1345,1585,1606]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="7" pageNumber="494" phylum="Chordata" rank="species" species="costaricense" status="sp. nov.">
<emphasis id="BEFC4A7E8B3AB54D992C6458FAB25173" box="[1106,1345,1585,1606]" italics="true" pageId="7" pageNumber="494">Nototriton costaricense</emphasis>
</taxonomicName>
<emphasis id="BEFC4A7E8B3AB54D98346459FA685170" bold="true" box="[1354,1435,1584,1605]" pageId="7" pageNumber="494">
<taxonomicNameLabel id="A5CFF7058B3AB54D98346459FA685170" box="[1354,1435,1584,1605]" pageId="7" pageNumber="494" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
Photographs taken by BK.
</paragraph>
</caption>
<paragraph id="8C37966C8B3AB5429DB964E5FC215656" blockId="7.[151,1437,1676,2022]" lastBlockId="8.[151,1437,151,2038]" lastPageId="8" lastPageNumber="495" pageId="7" pageNumber="494">
<emphasis id="BEFC4A7E8B3AB54D9DB964E5FE185190" bold="true" box="[199,491,1676,1701]" pageId="7" pageNumber="494">
Description of 
<typeStatus id="533328CE8B3AB54D9C0364E5FE155190" box="[381,486,1676,1701]" pageId="7" pageNumber="494" type="holotype">holotype</typeStatus>
.
</emphasis>
Subadult having a SL of 
<quantity id="4B703B898B3AB54D9E6364E5FC7B5191" box="[797,904,1676,1700]" metricMagnitude="-2" metricUnit="m" metricValue="2.19" pageId="7" pageNumber="494" unit="mm" value="21.9">21.9 mm</quantity>
. Head slightly wider than neck and shoulders (HeW 
<quantity id="4B703B898B3AB54D9D9A64D9FECD51F2" box="[228,318,1712,1736]" metricMagnitude="-3" metricUnit="m" metricValue="3.3" pageId="7" pageNumber="494" unit="mm" value="3.3">3.3 mm</quantity>
, NeW 3.0 mm, ShW 
<quantity id="4B703B898B3AB54D9F4664D9FD6751FD" box="[568,660,1712,1736]" metricMagnitude="-3" metricUnit="m" metricValue="2.9" pageId="7" pageNumber="494" unit="mm" value="2.9">2.9 mm</quantity>
), with greatest width of head just posterior to the articulation of the jaws; snout raised anterodorsally, spadate to rounded in dorsal outline, and rounded in profile; snout relatively short (SnL 
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, 3.2 % of SL), with nearly terminal non-protruding large nostrils (LNH 
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, RNW 
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) directed anterolaterally; internarial area convex in dorsal outline. Eyes relatively large (EW = 142 % of SnL), weakly protruding beyond dorsal and ventral outline of head, directed anterolaterally, with a distinct suborbital groove. Top of head flat and smooth, tapering slightly toward anterior terminus, lacking contrasting interorbital or other dermal structures. Canthus rostralis weakly rounded; intercanthal area flat to slightly convex; and loreal region slightly concave. No obvious cirri (nasolabial protuberances), but nasolabial grooves weakly discernable on tip of snout; nasolabial grooves start at lateral margins of nares and extend ventrally, with a slight outward orientation, and terminate prior to reaching upper lip margin. Gular fold well-defined, starting on dorsolateral portion of neck, and wrapping around lateral section of head and crossing venter as a smooth anterior-oriented curve. Upper lip markedly protruding beyond edge of lower lip in ventral view; and no mental gland visible under skin of anterior intermandibular region. A very weakly discernible groove starts at posterior dorsolateral margin of orbit and extends dorsolaterally to anterior margin of gular fold. Pair of weak, but discernably raised parotoid glands present on dorsolateral margin of head between orbits and occiput.
</paragraph>
<paragraph id="8C37966C8B35B5429DB96306FF185591" blockId="8.[151,1437,151,2038]" pageId="8" pageNumber="495">
Arms relatively long and slender (FLL = 
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, 20.1% of SL), without noticeable hypertrophied forearm compared to upper arm. Hands small and slender (HaL = 
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, 29.3% of VGS; HaW = 1.0 mm, 30.3 % of HeW). Fingers II, III, and IV relatively long (LF2 
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, LF3 
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) and robust with rounded tips; fingers II and IV nearly equal in width, but Finger III noticeably wider (WF3 
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); terminal pads poorly discernable, appearing to be present on ventral tips of fingers I, II, III, and IV; Finger I barely protruding beyond palmar tissue; webbing absent between fingers II and III and fingers III and IV. Palmar surface appearing to be smooth overall, but with an evident dermal crease extending transversally near proximal edge. Dorsal surfaces of hand with well defined interdigital grooves extending across metacarpal region. Relative lengths of fingers on right hand I &lt;IV &lt;II &lt;III.
</paragraph>
<paragraph id="8C37966C8B35B5429DB960D8FF2254D3" blockId="8.[151,1437,151,2038]" pageId="8" pageNumber="495">
Legs relatively long and slender (HLL 
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, 21.9 % of SL), with little perceivable difference between thickness of upper and lower leg. Feet small and slender (FoL 
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, 39.0 % of VGS; FoW 
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, 34.1 % of VGS). Toes II, III, IV, and V relatively long and slender with rounded tips nearly equal in width (WT3 
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) and length, toes III and IV slightly longer than others (LT2 
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, LT3 
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); terminal pads poorly discernable, appearing to be present on ventral tips of toes I, II, III, IV, and V; Toe I barely protruding beyond plantar tissue; webbing absent between toes II, III, IV, and V. Plantar surface appearing to be smooth overall, but with an evident dermal crease extending transversally near proximal edge. Dorsal surfaces of foot with well defined interdigital grooves extending across metatarsal region. Relative lengths of toes on right foot I &lt;V&lt;IV &lt;II &lt;III.
</paragraph>
<paragraph id="8C37966C8B35B5429DB96198FE64521E" blockId="8.[151,1437,151,2038]" pageId="8" pageNumber="495">
Body subcylindrical (wider than high) in cross section, and relatively robust (TW = 
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). Between axilla and groin, 11 costal grooves visible, 13 if counting axillary and inguinal grooves; costal grooves most visible on ventral and lateral portions of body. Adpressed limbs separated by 5 costal folds; 12 costal folds between axilla and groin. Slight middorsal depression extends longitudinally along length of body, starting at base of head and becoming indiscernible on anterior portion of tail. Tail long, cylindrical in cross section, lacking discernable caudal grooves or constriction at base, and tapering evenly to a pointed terminus. Tail long, cylindrical in cross section, lacking an evident constriction at base, and evenly tapering to pointed terminus; at least 15 caudal grooves discernible on anterior 3/4ths of tail, no grooves discernible on posterior 1/4th of tail. Skin on surfaces of head, body, limbs, and tail smooth.
</paragraph>
<paragraph id="8C37966C8B35B5429DB9675CFC165111" blockId="8.[151,1437,151,2038]" pageId="8" pageNumber="495">
<emphasis id="BEFC4A7E8B35B5429DB9675CFE6A527B" bold="true" box="[199,409,1333,1358]" pageId="8" pageNumber="495">Coloration in life.</emphasis>
The ground color of the dorsal surfaces of the head and trunk consisted of a mixture of pale earthy tones (Light Flesh Color [250], Light Pratt’s Rufous [71], and Flame Scarlet [73]). Secondary coloration on the dorsal surfaces of the head and trunk consisted of irregular dark markings (Marron [39] and Dark Grayish Brown [284]), grayish white dashes, and numerous very fine circular Marron (39) chromatophores. On the posterior half of the parotoid glands there was a pale flesh colored patch. On the center of the head, between the orbits and occiput, there was a concentration of dark (Dark Grayish Brown [284]) irregular markings. On the left parotoid gland there was a dark wedge shaped mark, appearing to be a scar.
</paragraph>
<paragraph id="8C37966C8B35B5429DB96459FB6651A5" blockId="8.[151,1437,151,2038]" pageId="8" pageNumber="495">The upper surfaces of the arms and legs have a Dark Grayish Brown (284) ground color. Secondary colors on the upper surfaces of the arms and legs are in the form of irregular spots either pearly white, Light Buff (2), Light Flesh Color (250), and Flame Scarlet (73). The hands and feet were a pale translucent gray.</paragraph>
<paragraph id="8C37966C8B35B5429DB964F2FCF351CE" blockId="8.[151,1437,151,2038]" pageId="8" pageNumber="495">The dorsal surface of the tails was a mottled mixture of Flame Scarlet (73), Light Flesh Color (250), and Light Buff (2). The dominant color of the dorsal surface of the tail was Flame Scarlet. Additionally, there were numerous white spots and dashes on the dorsal surface of the tail.</paragraph>
<paragraph id="8C37966C8B35B5429DB9656FFDC250C3" blockId="8.[151,1437,151,2038]" pageId="8" pageNumber="495">The colorations on the lateral surfaces of the body and tail were divided by a concentration of white markings or dashes that especially on the trunk formed what could be weakly interpreted as a longitudinal band. The inferior half of the lateral body and tail was Dark Grayish Brown (284) with scattered white markings. The superior half of the body and tail consisted of a mixture of earthy tones typical of the overall adjoining dorsal surfaces. The lateral surfaces of the head, between the orbits and the gular fold, were dominated by Flame Scarlet chromatophores (73); the mandibular region was dark gray with some scattered white markings or spots. The iris was a coppery-red with scattered dark spots and reticulation.</paragraph>
<paragraph id="8C37966C8B34B5439DB962FEFE8156BD" blockId="9.[151,1437,151,392]" pageId="9" pageNumber="496">The ventral skin of the head, body, and limbs was pale brown to gray with a very fine dark gray to black reticulation. Numerous white spots and irregular markings were also scattered throughout the ventral surfaces of the head and body. On the ventrolateral surface of the head, between the nuchal groove and gular fold, there was a patch of flame Scarlet skin marginally wrapping onto the ventral surface. Overall, the tone of the gular region was noticeably paler than the rest of the ventral surfaces. The palmar and plantar surfaces were pale grayish-white. The ventral surface of the tail is similar in general appearance to that of the body, but the ground color is noticeably a darker tone of gray.</paragraph>
<caption id="D8F7C6E48B34B5439DE96183FBAA5335" box="[151,1113,1002,1024]" httpUri="https://zenodo.org/record/1136412/files/figure.png" pageId="9" pageNumber="496" targetBox="[151,1435,447,977]" targetPageId="9">
<paragraph id="8C37966C8B34B5439DE96183FBAA5335" blockId="9.[151,1113,1002,1024]" box="[151,1113,1002,1024]" pageId="9" pageNumber="496">
<emphasis id="BEFC4A7E8B34B5439DE96183FEE154CA" bold="true" box="[151,274,1002,1024]" pageId="9" pageNumber="496">FIGURE 5.</emphasis>
Views of the nostrils of 
<taxonomicName id="4B88EDEF8B34B5439F756182FD0A5335" authority="Arias &amp; Kubicki, 2018" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[523,761,1003,1024]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="9" pageNumber="496" phylum="Chordata" rank="species" species="costaricense" status="sp. nov.">
<emphasis id="BEFC4A7E8B34B5439F756182FD0A5335" box="[523,761,1003,1024]" italics="true" pageId="9" pageNumber="496">Nototriton costaricense</emphasis>
</taxonomicName>
<emphasis id="BEFC4A7E8B34B5439E7E6183FCA354CA" bold="true" box="[768,848,1002,1023]" pageId="9" pageNumber="496">
<taxonomicNameLabel id="A5CFF7058B34B5439E7E6183FCA354CA" box="[768,848,1002,1023]" pageId="9" pageNumber="496" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
Photograph taken by BK.
</paragraph>
</caption>
<paragraph id="8C37966C8B34B5439DB96645FDDF53B8" blockId="9.[151,1437,1068,2029]" pageId="9" pageNumber="496">
<emphasis id="BEFC4A7E8B34B5439DB96645FE205370" bold="true" box="[199,467,1068,1093]" pageId="9" pageNumber="496">Coloration in ethanol.</emphasis>
After nearly two years in ethanol (70%), the overall coloration of the 
<typeStatus id="533328CE8B34B54398756645FA9E5370" box="[1291,1389,1068,1093]" pageId="9" pageNumber="496" type="holotype">holotype</typeStatus>
has darkened throughout and contains a mixture of four principal colors; Chestnut (30), Flame Scarlet (73), Raw Umber (280), and a pale fleshy gray
</paragraph>
<paragraph id="8C37966C8B34B5439DB966F1FE975298" blockId="9.[151,1437,1068,2029]" pageId="9" pageNumber="496">
<emphasis id="BEFC4A7E8B34B5439DB966F1FB365384" bold="true" box="[199,1221,1176,1201]" pageId="9" pageNumber="496">
Measurements (in mm), tooth counts, limb interval, and percentages of the 
<typeStatus id="533328CE8B34B543992966F1FB335384" box="[1111,1216,1176,1201]" pageId="9" pageNumber="496" type="holotype">holotype</typeStatus>
.
</emphasis>
SL 21.9; Tal 25.0; TL 46.9; ShW 2.9; HeW 3.3; NeW 3.0; EW 1.1; SnL 0.7; JSL 2.5; LGFS 4.6; LNH 0.34; RNW 0.31; IND 0.7; NLP 0.4; ICD 1.6; HLL 4.8; FLL 4.4; TW 3.2; VGS 4.1; FSL 6.4; UHL 2.7; AGL 12.0; VL 1.4; HaW 1.0; HaL 1.2; LF2 0.47; LF3 0.47; WF3 0.22; FoW 1.4; FoL 1.6; LT2 0.71; LT3 0.54; WT3 0.26. Limb interval 5. Measurements in related percentages: VGS/SL 18.7 %; IND/HeW 21.2 %; AGL/SL 54.8 %; HeW/SL 15.1 %; Hew/AGL 27.5 %; SnL/ HeW 21.2 %; LNH/HeW 10.3 %; LNH/SL 1.6 %; RNW/HeW, 9.4 %; RNW/SL 1.4 %; HLL/SL 21.9 %; FLL/SL 20.1 %; HaL/VGS 29.3 %; FoL/VGS 39.0 %; Haw/HeW 30.3 %; FoW/HeW 42.4 %; LT3/HeW 16.4 %.
</paragraph>
</subSubSection>
<subSubSection id="C492C5E78B34B5439DB967D1FEBF5108" pageId="9" pageNumber="496" type="etymology">
<paragraph id="8C37966C8B34B5439DB967D1FEBF5108" blockId="9.[151,1437,1068,2029]" pageId="9" pageNumber="496">
<emphasis id="BEFC4A7E8B34B5439DB967D1FEBD52E4" bold="true" box="[199,334,1464,1489]" pageId="9" pageNumber="496">Etymology.</emphasis>
The specific epithet refers to the Spanish word meaning Costa Rican, “ 
<taxonomicName id="4B88EDEF8B34B54399FA67D1FAE652E4" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[1156,1301,1464,1489]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="9" pageNumber="496" phylum="Chordata" rank="species" species="costaricense">costaricense</taxonomicName>
”. The name represents the fact that the 
<typeStatus id="533328CE8B34B5439CB067B5FDC352C0" box="[462,560,1500,1525]" pageId="9" pageNumber="496" type="holotype">holotype</typeStatus>
and species was discovered in 
<collectingCountry id="F49FD6FC8B34B5439EE467B4FBE952C0" box="[922,1050,1500,1525]" name="Costa Rica" pageId="9" pageNumber="496">Costa Rica</collectingCountry>
. Given the close proximity of the 
<typeStatus id="533328CE8B34B5439DE96468FF34512C" box="[151,199,1537,1561]" pageId="9" pageNumber="496">type</typeStatus>
locality to the Costa Rica-Panama border, we speculate that one day it may indeed be discovered within 
<collectingCountry id="F49FD6FC8B34B5439DE9644CFF035109" box="[151,240,1573,1596]" name="Panama" pageId="9" pageNumber="496">
<collectingRegion id="4E4C588E8B34B5439DE9644CFF035109" box="[151,240,1573,1596]" country="Panama" name="Panama" pageId="9" pageNumber="496">Panama</collectingRegion>
</collectingCountry>
as well.
</paragraph>
</subSubSection>
<subSubSection id="C492C5E78B34B5439DB96421FDD550D8" pageId="9" pageNumber="496" type="biology_ecology">
<paragraph id="8C37966C8B34B5439DB96421FDD550D8" blockId="9.[151,1437,1068,2029]" pageId="9" pageNumber="496">
<emphasis id="BEFC4A7E8B34B5439DB96421FD495154" bold="true" box="[199,698,1608,1633]" pageId="9" pageNumber="496">Habitat and natural history observations.</emphasis>
The 
<typeStatus id="533328CE8B34B5439F886421FCAB5154" box="[758,856,1608,1633]" pageId="9" pageNumber="496" type="holotype">holotype</typeStatus>
of 
<taxonomicName id="4B88EDEF8B34B5439E016423FB7B5154" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[895,1160,1610,1633]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="9" pageNumber="496" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B34B5439E016423FB7B5154" box="[895,1160,1610,1633]" italics="true" pageId="9" pageNumber="496">Nototriton costaricense</emphasis>
</taxonomicName>
was discovered inside a moss mat that was growing approximately one meter above the ground on the trunk of a tree. The 
<typeStatus id="533328CE8B34B543998B6404FAD651B0" box="[1269,1317,1645,1669]" pageId="9" pageNumber="496">type</typeStatus>
locality is ca. 
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SSW of a peak known as Cerro Pat; the general site is a relatively flat ridge, ranging from 
<quantity id="4B703B898B34B543987C64F9FA6F519D" box="[1282,1436,1680,1705]" metricMagnitude="3" metricUnit="m" metricValue="1.45" metricValueMax="1.5" metricValueMin="1.4" pageId="9" pageNumber="496" unit="m" value="1450.0" valueMax="1500.0" valueMin="1400.0">1400–1500 m</quantity>
asl, which is covered by mature forest. Despite the elevation of the site typically being dominated by cloud forest with an abundance of bryophytes and epiphytes, it was nearly devoid of such floral conditions; bryophytes and epiphytes were uncommon there. The 
<typeStatus id="533328CE8B34B5439F3C6494FD815020" box="[578,626,1789,1813]" pageId="9" pageNumber="496">type</typeStatus>
locality is located on a ridgeline that extends out towards the north from the Kamuk massif. Just south of the 
<typeStatus id="533328CE8B34B5439F376548FD8A500C" box="[585,633,1825,1849]" pageId="9" pageNumber="496">type</typeStatus>
locality, the topography of the ridgeline leading towards the Kamuk massif drastically rises until reaching an unnamed peak with an elevation of approximately 
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asl. The unnamed peak and its slopes is covered in mature forest, and in contrast to that of the 
<typeStatus id="533328CE8B34B54399F96500FB4450B4" box="[1159,1207,1897,1921]" pageId="9" pageNumber="496">type</typeStatus>
locality, there is an abundance of bryophytes and other epiphytes covering the trees. Despite an intense search by EA through moss, bromeliads, and leaf litter, in what appeared to be more optimal habitat along the slopes of the unnamed peak, no additional salamanders were found.
</paragraph>
</subSubSection>
<subSubSection id="C492C5E78B37B5409DB962FEFD6856C0" pageId="10" pageNumber="497" type="discussion">
<paragraph id="8C37966C8B37B5409DB962FEFD6856C0" blockId="10.[151,1437,151,789]" pageId="10" pageNumber="497">
Inasmuch as 
<taxonomicName id="4B88EDEF8B37B5409C2462F0FD915785" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[346,610,153,176]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B5409C2462F0FD915785" box="[346,610,153,176]" italics="true" pageId="10" pageNumber="497">Nototriton costaricense</emphasis>
</taxonomicName>
is only known from a single specimen, any assumptions about its natural history are highly speculative. With that said, however, like other members of the 
<taxonomicName id="4B88EDEF8B37B540995862D4FB0657E0" baseAuthorityName="Stejneger" baseAuthorityYear="1911" box="[1062,1269,188,213]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B37B540995862D4FB0657E0" box="[1062,1269,188,213]" italics="true" pageId="10" pageNumber="497">Nototriton picadoi</emphasis>
</taxonomicName>
species group, it is probable that this species is prone to living within moss mats, both in terrestrial and arboreal situations, within cloud forest. All known Costa Rican species of 
<taxonomicName id="4B88EDEF8B37B5409FD2636CFCD25629" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[684,801,261,284]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B37B5409FD2636CFCD25629" box="[684,801,261,284]" italics="true" pageId="10" pageNumber="497">Nototriton</emphasis>
</taxonomicName>
are known to principally inhabit moss, unlike congeners of the subgenus 
<emphasis id="BEFC4A7E8B37B5409C306340FE305675" box="[334,451,297,320]" italics="true" pageId="10" pageNumber="497">Bryotriton</emphasis>
found in Nuclear Central America that principally known to inhabit microhabitats such as leaf litter, rotting logs, and bromeliads (
<bibRefCitation id="E819EB9D8B37B5409FFB6325FC705651" author="Townsend" box="[645,899,332,357]" pageId="10" pageNumber="497" refString="Townsend, J. H., Butler, J. M., Wilson, L. D. &amp; Austin, J. D. (2010) A distinctive new species of moss salamander (Caudata: Plethodontidae: Nototriton) from an imperiled Honduran endemism hotspot. Zootaxa, 2434, 1 - 16." type="journal article" year="2010">
Townsend 
<emphasis id="BEFC4A7E8B37B5409E7D6324FCCD5650" box="[771,830,332,357]" italics="true" pageId="10" pageNumber="497">et al.</emphasis>
2010
</bibRefCitation>
, 
<bibRefCitation id="E819EB9D8B37B5409EEF6325FC385651" author="Townsend" box="[913,971,332,356]" pageId="10" pageNumber="497" refString="Townsend, J. H., Medina-Flores, M., Murillo, J. L. &amp; Austin, J. D. (2011) Cryptic diversity in Chortis Highland moss salamanders (Caudata: Plethodontidae: Nototriton) revealed using mtDNA barcodes and phylogenetics, with a new species from eastern Honduras. Systematics and Biodiversity, 9, 275 - 287. https: // doi. org / 10.1080 / 14772000.2011.611188" type="journal article" year="2011">2011</bibRefCitation>
, 
<bibRefCitation id="E819EB9D8B37B5409EA76325FBE75651" author="Townsend" box="[985,1044,332,356]" pageId="10" pageNumber="497" refString="Townsend, J. H., Medina-Flores, M., Reyes-Calderon, O. &amp; Austin, J. D. (2013) A new Nototriton (Caudata: Plethodontidae) from Parque Nacional Montana de Botaderos in northeastern Honduras. Zootaxa, 3666 (3), 358 - 368. https: // doi. org / 10.11646 / zootaxa. 3666.3.6" type="journal article" year="2013">2013</bibRefCitation>
; 
<bibRefCitation id="E819EB9D8B37B540995D6325FB2F5650" author="Townsend" box="[1059,1244,332,357]" pageId="10" pageNumber="497" refString="Townsend, J. H. (2016) Taxonomic revision of the moss salamander Nototriton barbouri (Schmidt) (Caudata: Plethodontidae), with description of two new species from the Cordillera Nombre de Dios, Honduras. Zootaxa, 4196 (4), 511 - 528. https: // doi. org / 10.11646 / zootaxa. 4196.4.3" type="journal article" year="2016">Townsend 2016</bibRefCitation>
). Within another patch of moss on the same tree where the 
<typeStatus id="533328CE8B37B5409FFD6306FD1656BD" box="[643,741,367,392]" pageId="10" pageNumber="497" type="holotype">holotype</typeStatus>
of 
<taxonomicName id="4B88EDEF8B37B5409E6E6318FC4C56BD" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[784,959,369,392]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B5409E6E6318FC4C56BD" box="[784,959,369,392]" italics="true" pageId="10" pageNumber="497">N. costaricense</emphasis>
</taxonomicName>
was found, an individual of 
<taxonomicName id="4B88EDEF8B37B540986E6306FF085698" class="Amphibia" family="Plethodontidae" genus="Bolitoglossa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="minutula">
<emphasis id="BEFC4A7E8B37B540986E6306FF085698" italics="true" pageId="10" pageNumber="497">Bolitoglossa minutula</emphasis>
</taxonomicName>
was also discovered (
<bibRefCitation id="E819EB9D8B37B5409F6B63FDFD525699" author="Arias" box="[533,673,404,429]" pageId="10" pageNumber="497" refString="Arias, E. (2017) New distribution records for two uncommon amphibian species on the Caribbean slopes of the Cordillera de Talamanca, Costa Rica. Mesoamerican Herpetology, 4, 446 - 449." type="journal article" year="2017">Arias 2017</bibRefCitation>
). Other amphibian species found in sympatry with 
<taxonomicName id="4B88EDEF8B37B540985963FCFED056E5" authorityName="Arias &amp; Kubicki" authorityYear="2018" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B540985963FCFED056E5" italics="true" pageId="10" pageNumber="497">Nototriton costaricense</emphasis>
</taxonomicName>
are 
<emphasis id="BEFC4A7E8B37B5409C1263D1FD4756FA" box="[364,692,439,464]" italics="true" pageId="10" pageNumber="497">
<taxonomicName id="4B88EDEF8B37B5409C1263D1FD7156E5" baseAuthorityName="Cope" baseAuthorityYear="1875" box="[364,642,439,464]" class="Amphibia" family="Craugastoridae" genus="Craugastor" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="podiciferus">Craugastor podiciferus</taxonomicName>
, C.
</emphasis>
aff. 
<emphasis id="BEFC4A7E8B37B5409E7E63DEFE1656C0" italics="true" pageId="10" pageNumber="497">underwoodi, Diasporus hylaeformis, Incilius epioticus, Pristimantis caryophyllaceus,</emphasis>
and 
<emphasis id="BEFC4A7E8B37B5409F6263B4FD6856C0" box="[540,667,476,501]" italics="true" pageId="10" pageNumber="497">P. pardalis.</emphasis>
</paragraph>
</subSubSection>
<subSubSection id="C492C5E78B37B5409DB96396FE815420" pageId="10" pageNumber="497" type="distribution">
<paragraph id="8C37966C8B37B5409DB96396FE815420" blockId="10.[151,1437,151,789]" pageId="10" pageNumber="497">
<emphasis id="BEFC4A7E8B37B5409DB96396FE93552D" bold="true" box="[199,352,511,536]" pageId="10" pageNumber="497">Distribution.</emphasis>
<taxonomicName id="4B88EDEF8B37B5409C166068FD81552D" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[360,626,513,536]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B5409C166068FD81552D" box="[360,626,513,536]" italics="true" pageId="10" pageNumber="497">Nototriton costaricense</emphasis>
</taxonomicName>
is known from a single site within the Lower Montane Rain Forest life zone (
<bibRefCitation id="E819EB9D8B37B5409DA4604DFE7E5508" author="Holdridge" box="[218,397,548,573]" pageId="10" pageNumber="497" refString="Holdridge, L. R. (1967) Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica, 206 pp." type="book" year="1967">Holdridge 1967</bibRefCitation>
) along the mid-elevation slopes of the extreme southeastern Cordillera de Talamanca, within Parque Internacional La Amistad, in the vicinity of Cerro Pat, ca. 
<quantity id="4B703B898B37B5409ED9602EFBF7556A" box="[935,1028,583,608]" metricMagnitude="3" metricUnit="m" metricValue="1.5" pageId="10" pageNumber="497" unit="m" value="1500.0">1500 m</quantity>
(
<figureCitation id="14B38AE98B37B5409967602EFB935555" box="[1049,1120,583,608]" captionStart="FIGURE 1" captionStartId="1.[151,250,1670,1692]" captionTargetBox="[150,1436,735,1642]" captionTargetId="figure@1.[150,1436,734,1644]" captionTargetPageId="1" captionText="FIGURE 1. Map showing the type locality for Nototriton costaricense sp. nov. (red star) in the Cordillera de Talamanca.The localities for the other known members of the N. picadoi species group are shown for comparison.The green area indicates the suitable area for Nototriton." httpUri="https://zenodo.org/record/1136404/files/figure.png" pageId="10" pageNumber="497">Fig. 1</figureCitation>
). Further explorations and encounters with this species are needed before we will be able to further understand its potential range. We speculate that it is very possible that 
<taxonomicName id="4B88EDEF8B37B5409F3160F8FD0C559D" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[591,767,656,680]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B5409F3160F8FD91559D" box="[591,610,657,680]" italics="true" pageId="10" pageNumber="497">N</emphasis>
. 
<emphasis id="BEFC4A7E8B37B5409F0D60F8FD0C559D" box="[627,767,657,680]" italics="true" pageId="10" pageNumber="497">costaricense</emphasis>
</taxonomicName>
extends into 
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<collectingRegion id="4E4C588E8B37B5409EDF60F9FC095592" box="[929,1018,656,679]" country="Panama" name="Panama" pageId="10" pageNumber="497">Panama</collectingRegion>
</collectingCountry>
due to the proximity and relatively homogenous habitat lying between the 
<typeStatus id="533328CE8B37B5409F1360DCFD6E55F8" box="[621,669,693,717]" pageId="10" pageNumber="497">type</typeStatus>
locality and the border (~ 11 kilometers straight line distance). 
<taxonomicName id="4B88EDEF8B37B5409DE960B0FE5055C5" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[151,419,729,752]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B5409DE960B0FE5055C5" box="[151,419,729,752]" italics="true" pageId="10" pageNumber="497">Nototriton costaricense</emphasis>
</taxonomicName>
, if found to inhabit 
<collectingCountry id="F49FD6FC8B37B5409FE460B1FD0555DA" box="[666,758,728,751]" name="Panama" pageId="10" pageNumber="497">
<collectingRegion id="4E4C588E8B37B5409FE460B1FD0555DA" box="[666,758,728,751]" country="Panama" name="Panama" pageId="10" pageNumber="497">Panama</collectingRegion>
</collectingCountry>
, would represent the first species of this genus to extend beyond 
<collectingCountry id="F49FD6FC8B37B5409D8E6095FE9D5420" box="[240,366,764,789]" name="Costa Rica" pageId="10" pageNumber="497">Costa Rica</collectingCountry>
.
</paragraph>
</subSubSection>
</subSection>
<subSection id="E5078F878B37B5419DE96101FB5852F5" lastPageId="11" lastPageNumber="498" pageId="10" pageNumber="497" type="discussion">
<subSubSection id="C492C5E78B37B5419DE96101FB5852F5" lastPageId="11" lastPageNumber="498" pageId="10" pageNumber="497" type="discussion">
<paragraph id="8C37966C8B37B5409DE96101FEE854B7" blockId="10.[151,283,872,898]" box="[151,283,872,898]" pageId="10" pageNumber="497">
<heading id="D77F21008B37B5409DE96101FEE854B7" bold="true" box="[151,283,872,898]" fontSize="11" level="1" pageId="10" pageNumber="497" reason="1">
<emphasis id="BEFC4A7E8B37B5409DE96101FEE854B7" bold="true" box="[151,283,872,898]" pageId="10" pageNumber="497">Discussion</emphasis>
</heading>
</paragraph>
<paragraph id="8C37966C8B37B5409DE961C7FEC85266" blockId="10.[151,1437,942,2011]" pageId="10" pageNumber="497">
With the discovery and description of 
<taxonomicName id="4B88EDEF8B37B5409F3661C6FCA354F3" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[584,848,943,966]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B5409F3661C6FCA354F3" box="[584,848,943,966]" italics="true" pageId="10" pageNumber="497">Nototriton costaricense</emphasis>
</taxonomicName>
, the number of species known for this genus within 
<collectingCountry id="F49FD6FC8B37B5409DE961BAFEE354DE" box="[151,272,978,1003]" name="Costa Rica" pageId="10" pageNumber="497">Costa Rica</collectingCountry>
is now elevated to nine, and the total diversity of salamanders for the tiny country 52. In the last decade 11 species (23.5% of total) of salamanders were described from 
<collectingCountry id="F49FD6FC8B37B5409E0C619EFC1E533A" box="[882,1005,1014,1039]" name="Costa Rica" pageId="10" pageNumber="497">Costa Rica</collectingCountry>
(
<bibRefCitation id="E819EB9D8B37B5409E82619FFB30533A" author="Wake" box="[1020,1219,1014,1039]" pageId="10" pageNumber="497" refString="Wake, D. B., Savage, J. M. &amp; Hanken, J. (2007) Montane salamanders from the Costa Rica - Panama border region, with descriptions of two new species of Bolitoglossa. Copeia, 2007, 556 - 565." type="journal article" year="2007">
<collectingCountry id="F49FD6FC8B37B5409E82619FFBCE533A" box="[1020,1085,1014,1039]" name="Wake Island" pageId="10" pageNumber="497">Wake</collectingCountry>
<emphasis id="BEFC4A7E8B37B540993B619EFB8C533A" box="[1093,1151,1014,1039]" italics="true" pageId="10" pageNumber="497">et al.</emphasis>
2007
</bibRefCitation>
; 
<bibRefCitation id="E819EB9D8B37B54099B1619EFF205306" author="Garcia-Paris" pageId="10" pageNumber="497" refString="Garcia-Paris, M., Parra-Olea, G. &amp; Wake, D. B. (2008) Description of a new species of the Bolitoglossa subpalmata group (Caudata: Plethodontidae) from Costa Rica. Herpetological Journal, 18, 23 - 31." type="journal article" year="2008">
García-París 
<emphasis id="BEFC4A7E8B37B540981D619EFA6F533A" box="[1379,1436,1014,1039]" italics="true" pageId="10" pageNumber="497">et al.</emphasis>
2008
</bibRefCitation>
; 
<bibRefCitation id="E819EB9D8B37B5409DA06673FE115306" author="Bolanos" box="[222,482,1050,1075]" pageId="10" pageNumber="497" refString="Bolanos, F. &amp; Wake, D. B. (2009) Two new species of montane web-footed salamanders (Plethodontidae: Bolitoglossa) from the Costa Rica-Panama border region. Zootaxa, 1981, 57 - 68." type="journal article" year="2009">
Bolaños &amp; 
<collectingCountry id="F49FD6FC8B37B5409C216673FE6D5306" box="[351,414,1050,1075]" name="Wake Island" pageId="10" pageNumber="497">Wake</collectingCountry>
2009
</bibRefCitation>
; 
<bibRefCitation id="E819EB9D8B37B5409C906673FCF55306" author="Boza-Oviedo" box="[494,774,1050,1075]" pageId="10" pageNumber="497" refString="Boza-Oviedo, E., Rovito, S. M., Chaves, G., Garcia-Rodriguez, A., Artavia, L. G., Bolanos, F. &amp; Wake, D. B. (2012) Salamanders from the eastern Cordillera de Talamanca, Costa Rica, with descriptions of five new species (Plethodontidae: Bolitoglossa, Nototriton, and Oedipina) and natural history notes from recent expeditions. Zootaxa, 3309, 36 - 61." type="journal article" year="2012">
Boza-Oviedo 
<emphasis id="BEFC4A7E8B37B5409FF56675FD375306" box="[651,708,1050,1075]" italics="true" pageId="10" pageNumber="497">et al.</emphasis>
2012
</bibRefCitation>
; Kubicki 2016; 
<bibRefCitation id="E819EB9D8B37B5409EC26673FB4F5306" author="Kubicki" box="[956,1212,1050,1075]" pageId="10" pageNumber="497" refString="Kubicki, B. &amp; Arias, E. (2016) A beautiful new yellow salamander, genus Bolitoglossa (Caudata: Plethodontidae), from the northeastern slopes of the Cordillera de Talamanca, Costa Rica. Zootaxa, 4184 (2), 329 - 346. https: // doi. org / 10.11646 / zootaxa. 4184.2.5" type="journal article" year="2016">Kubicki &amp; Arias 2016</bibRefCitation>
), indicating that the inventory of salamanders from the country is incomplete. The bolitoglossines recently described from 
<collectingCountry id="F49FD6FC8B37B540985F6656FA6F5362" box="[1313,1436,1086,1111]" name="Costa Rica" pageId="10" pageNumber="497">Costa Rica</collectingCountry>
are only known from a few localities and relatively few specimens, and half of these species are known from the Caribbean slopes of 
<collectingCountry id="F49FD6FC8B37B5409CF666EEFDF953AA" box="[392,522,1158,1183]" name="Costa Rica" pageId="10" pageNumber="497">Costa Rica</collectingCountry>
. We would like to emphasize, again, that special attention needs to placed on improving the knowledge of the diversity of not only salamanders, but amphibians in general along the midelevation Caribbean slopes of the Cordillera de Talamanca of 
<collectingCountry id="F49FD6FC8B37B5409E0F66A6FC0053D2" box="[881,1011,1230,1255]" name="Costa Rica" pageId="10" pageNumber="497">Costa Rica</collectingCountry>
. The mid-elevation slopes along the Caribbean versant of Talamanca, due to the difficult terrain and over all limited access, harbor some of the poorest known and least explored forests within 
<collectingCountry id="F49FD6FC8B37B5409F0B677EFD05521A" box="[629,758,1302,1327]" name="Costa Rica" pageId="10" pageNumber="497">Costa Rica</collectingCountry>
; further inventories should be viewed as a priority in this overall region.
</paragraph>
<paragraph id="8C37966C8B37B5409DB96737FE23511F" blockId="10.[151,1437,942,2011]" pageId="10" pageNumber="497">
With our discovery of 
<taxonomicName id="4B88EDEF8B37B5409CBC6736FD395243" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[450,714,1375,1398]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B5409CBC6736FD395243" box="[450,714,1375,1398]" italics="true" pageId="10" pageNumber="497">Nototriton costaricense</emphasis>
</taxonomicName>
, the distribution of genus is now known to extend ~ 
<quantity id="4B703B898B37B54098716737FAA55242" box="[1295,1366,1374,1399]" metricMagnitude="4" metricUnit="m" metricValue="5.0" pageId="10" pageNumber="497" unit="km" value="50.0">50 km</quantity>
to the south-southeast from the previously known southern limit of the genus, which was the 
<typeStatus id="533328CE8B37B540991067EAFB6D52AE" box="[1134,1182,1411,1435]" pageId="10" pageNumber="497">type</typeStatus>
locality of 
<taxonomicName id="4B88EDEF8B37B540985E67EDFA6F52AE" box="[1312,1436,1412,1435]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="matama">
<emphasis id="BEFC4A7E8B37B540985E67EDFA6F52AE" box="[1312,1436,1412,1435]" italics="true" pageId="10" pageNumber="497">N. matama</emphasis>
</taxonomicName>
on the southeast terminus of the Matama ridge, 
<collectingRegion id="4E4C588E8B37B5409FB867CFFC72528A" box="[710,897,1446,1471]" country="Costa Rica" name="Limon" pageId="10" pageNumber="497">Limón Province</collectingRegion>
, 
<collectingCountry id="F49FD6FC8B37B5409EEE67CEFBE2528A" box="[912,1041,1446,1471]" name="Costa Rica" pageId="10" pageNumber="497">Costa Rica</collectingCountry>
. Between the 
<typeStatus id="533328CE8B37B54099C967CEFB14528A" box="[1207,1255,1447,1471]" pageId="10" pageNumber="497">type</typeStatus>
localities of 
<taxonomicName id="4B88EDEF8B37B54098FC67CEFF0252D6" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="matama">
<emphasis id="BEFC4A7E8B37B54098FC67CEFF0252D6" italics="true" pageId="10" pageNumber="497">N. matama</emphasis>
</taxonomicName>
and 
<taxonomicName id="4B88EDEF8B37B5409C5067A5FE2D52D6" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[302,478,1484,1507]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B5409C5067A5FE2D52D6" box="[302,478,1484,1507]" italics="true" pageId="10" pageNumber="497">N. costaricense</emphasis>
</taxonomicName>
, there is an abundance of suitable habitat for the genus, and this area needs to be surveyed further in attempt to learn more about the actual distributional boundaries between these two enigmatic and recently described taxa.
</paragraph>
<paragraph id="8C37966C8B37B5419DB9645FFA8C5675" blockId="10.[151,1437,942,2011]" lastBlockId="11.[151,1437,151,1472]" lastPageId="11" lastPageNumber="498" pageId="10" pageNumber="497">
The genetic distances between 
<taxonomicName id="4B88EDEF8B37B5409F5B645EFCDE517B" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[549,813,1591,1614]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B37B5409F5B645EFCDE517B" box="[549,813,1591,1614]" italics="true" pageId="10" pageNumber="497">Nototriton costaricense</emphasis>
</taxonomicName>
and the other members of the 
<taxonomicName id="4B88EDEF8B37B54099FB645EFB09517A" baseAuthorityName="Stejneger" baseAuthorityYear="1911" box="[1157,1274,1590,1615]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B37B54099FB645EFB09517A" box="[1157,1274,1590,1615]" italics="true" pageId="10" pageNumber="497">N. picadoi</emphasis>
</taxonomicName>
species group range from 1.8–2.4 % for 16S; this distance is relatively low in comparison to those suggested by 
<bibRefCitation id="E819EB9D8B37B54099806432FF1251A2" pageId="10" pageNumber="497" refString="Fouquet, A., Gilles, A., Vences, M., Marty, C., Blanc, M. &amp; Gemmell, N. J. (2007) Underestimation of species richness in Neotropical frogs revealed by mtDNA analyses. PLoS one, 2, e 1109. https: // doi. org / 10.1371 / journal. pone. 0001109" type="journal article">
Fouquet 
<emphasis id="BEFC4A7E8B37B540981C6435FA6F5146" box="[1378,1436,1626,1651]" italics="true" pageId="10" pageNumber="497">et al.</emphasis>
(2007)
</bibRefCitation>
for Neotropical frogs, in which the authors recommend a minimum of 3% 16S divergence be recognized as threshold to define candidate species. We recently commented that this threshold possibly underestimates the true diversity of Neotropical salamanders (
<bibRefCitation id="E819EB9D8B37B5409F3F64AFFCB351EA" author="Kubicki" box="[577,832,1734,1759]" pageId="10" pageNumber="497" refString="Kubicki, B. &amp; Arias, E. (2016) A beautiful new yellow salamander, genus Bolitoglossa (Caudata: Plethodontidae), from the northeastern slopes of the Cordillera de Talamanca, Costa Rica. Zootaxa, 4184 (2), 329 - 346. https: // doi. org / 10.11646 / zootaxa. 4184.2.5" type="journal article" year="2016">Kubicki &amp; Arias 2016</bibRefCitation>
), and that each taxon should be carefully evaluated on an individual basis considering all the phenotypic, geographic, and molecular evidence at hand. 
<bibRefCitation id="E819EB9D8B37B54099A16483FF125012" author="Kubicki" pageId="10" pageNumber="497" refString="Kubicki, B. &amp; Arias, E. (2016) A beautiful new yellow salamander, genus Bolitoglossa (Caudata: Plethodontidae), from the northeastern slopes of the Cordillera de Talamanca, Costa Rica. Zootaxa, 4184 (2), 329 - 346. https: // doi. org / 10.11646 / zootaxa. 4184.2.5" type="journal article" year="2016">Kubicki &amp; Arias (2016)</bibRefCitation>
described a well-supported morphologically distinctive new species of 
<taxonomicName id="4B88EDEF8B37B540993D6567FB3C5012" box="[1091,1231,1806,1831]" class="Amphibia" family="Plethodontidae" genus="Bolitoglossa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B37B540993D6567FB3C5012" box="[1091,1231,1806,1831]" italics="true" pageId="10" pageNumber="497">Bolitoglossa</emphasis>
</taxonomicName>
that had genetic distances as low as 2.0 % (16S) among its closest related taxa. 
<taxonomicName id="4B88EDEF8B37B5409E20655BFA72507E" authority="Boza-Oviedo et al., 2012" authorityName="Boza-Oviedo et al." authorityYear="2012" box="[862,1409,1842,1867]" class="Amphibia" family="Plethodontidae" genus="Bolitoglossa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="splendida">
<emphasis id="BEFC4A7E8B37B5409E20655BFBAD507E" box="[862,1118,1842,1867]" italics="true" pageId="10" pageNumber="497">Bolitoglossa splendida</emphasis>
<bibRefCitation id="E819EB9D8B37B5409918655BFA72507E" author="Boza-Oviedo" box="[1126,1409,1842,1867]" pageId="10" pageNumber="497" refString="Boza-Oviedo, E., Rovito, S. M., Chaves, G., Garcia-Rodriguez, A., Artavia, L. G., Bolanos, F. &amp; Wake, D. B. (2012) Salamanders from the eastern Cordillera de Talamanca, Costa Rica, with descriptions of five new species (Plethodontidae: Bolitoglossa, Nototriton, and Oedipina) and natural history notes from recent expeditions. Zootaxa, 3309, 36 - 61." type="journal article" year="2012">
Boza-Oviedo 
<emphasis id="BEFC4A7E8B37B540987D655DFAB1507E" box="[1283,1346,1842,1867]" italics="true" pageId="10" pageNumber="497">et al.,</emphasis>
2012
</bibRefCitation>
</taxonomicName>
is another example of salamander that is well recognized for its phenotypic uniqueness, but presents once again a relatively low (2.0 % 16S) genetic divergence from of its closest related congeners (
<bibRefCitation id="E819EB9D8B37B54099136513FA6250A6" author="Boza-Oviedo" box="[1133,1425,1914,1939]" pageId="10" pageNumber="497" refString="Boza-Oviedo, E., Rovito, S. M., Chaves, G., Garcia-Rodriguez, A., Artavia, L. G., Bolanos, F. &amp; Wake, D. B. (2012) Salamanders from the eastern Cordillera de Talamanca, Costa Rica, with descriptions of five new species (Plethodontidae: Bolitoglossa, Nototriton, and Oedipina) and natural history notes from recent expeditions. Zootaxa, 3309, 36 - 61." type="journal article" year="2012">
Boza-Oviedo 
<emphasis id="BEFC4A7E8B37B54098706515FAB950A6" box="[1294,1354,1914,1939]" italics="true" pageId="10" pageNumber="497">et al.</emphasis>
2012
</bibRefCitation>
). Within the genus 
<taxonomicName id="4B88EDEF8B37B5409C1D65F6FE2B5083" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[355,472,1951,1974]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B37B5409C1D65F6FE2B5083" box="[355,472,1951,1974]" italics="true" pageId="10" pageNumber="497">Nototriton</emphasis>
</taxonomicName>
it has been previously published and justified that low genetic distances are known exist among well-recognized species, i.e. 1.9 % in 16S between 
<taxonomicName id="4B88EDEF8B37B5409EFB65ADFBFE50EE" baseAuthorityName="Schmidt" baseAuthorityYear="1936" box="[901,1037,1986,2011]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="barbouri">
<emphasis id="BEFC4A7E8B37B5409EFB65ADFBFE50EE" box="[901,1037,1986,2011]" italics="true" pageId="10" pageNumber="497">N. barbouri</emphasis>
</taxonomicName>
and 
<taxonomicName id="4B88EDEF8B37B541993465ADFF2B5785" authority="(Townsend 2016)" authorityName="Townsend" authorityYear="2016" baseAuthorityName="Townsend" baseAuthorityYear="2016" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" lastPageId="11" lastPageNumber="498" order="Caudata" pageId="10" pageNumber="497" phylum="Chordata" rank="species" species="limnospectator">
<emphasis id="BEFC4A7E8B37B540993465ADFAE650EE" box="[1098,1301,1986,2011]" italics="true" pageId="10" pageNumber="497">N. limnospectator</emphasis>
(
<bibRefCitation id="E819EB9D8B37B541985965ABFF225785" author="Townsend" lastPageId="11" lastPageNumber="498" pageId="10" pageNumber="497" refString="Townsend, J. H. (2016) Taxonomic revision of the moss salamander Nototriton barbouri (Schmidt) (Caudata: Plethodontidae), with description of two new species from the Cordillera Nombre de Dios, Honduras. Zootaxa, 4196 (4), 511 - 528. https: // doi. org / 10.11646 / zootaxa. 4196.4.3" type="journal article" year="2016">Townsend 2016</bibRefCitation>
)
</taxonomicName>
. 
<bibRefCitation id="E819EB9D8B36B5419D9962FEFE3B5785" box="[231,456,151,176]" pageId="11" pageNumber="498" refString="Rovito, S. M., Parra-Olea, G., Hanken, J., Bonett, R. M. &amp; Wake, D. B. (2013) Adaptive radiation in miniature: the minute salamanders of the Mexican highlands (Amphibia: Plethodontidae: Thorius). Biological Journal of the Linnean Society, 109, 622 - 643. https: // doi. org / 10.1111 / bij. 12083" type="journal article">
Rovito 
<emphasis id="BEFC4A7E8B36B5419C4262F0FE9D5785" box="[316,366,151,176]" italics="true" pageId="11" pageNumber="498">et al</emphasis>
. (2013)
</bibRefCitation>
used morphological and molecular (two mitochondrial and one nuclear genes) data to define the species within another genus of miniaturized salamanders in 
<collectingCountry id="F49FD6FC8B36B5419E9C62D5FBC957E0" box="[994,1082,188,213]" name="Mexico" pageId="11" pageNumber="498">
<collectingRegion id="4E4C588E8B36B5419E9C62D5FBC957E0" box="[994,1082,188,213]" country="Mexico" name="Mexico" pageId="11" pageNumber="498">Mexico</collectingRegion>
</collectingCountry>
, 
<taxonomicName id="4B88EDEF8B36B541993762D5FB5357E0" authorityName="Cope" authorityYear="1869" box="[1097,1184,188,213]" class="Amphibia" family="Plethodontidae" genus="Thorius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B36B541993762D5FB5357E0" box="[1097,1184,188,213]" italics="true" pageId="11" pageNumber="498">Thorius</emphasis>
</taxonomicName>
. 
<bibRefCitation id="E819EB9D8B36B54199CD62D5FA6F57E1" box="[1203,1436,188,213]" pageId="11" pageNumber="498" refString="Rovito, S. M., Parra-Olea, G., Hanken, J., Bonett, R. M. &amp; Wake, D. B. (2013) Adaptive radiation in miniature: the minute salamanders of the Mexican highlands (Amphibia: Plethodontidae: Thorius). Biological Journal of the Linnean Society, 109, 622 - 643. https: // doi. org / 10.1111 / bij. 12083" type="journal article">
Rovito 
<emphasis id="BEFC4A7E8B36B541987462D4FAB557E0" box="[1290,1350,188,213]" italics="true" pageId="11" pageNumber="498">et al.</emphasis>
(2013)
</bibRefCitation>
showed that an integrative approach (i.e. morphological and molecular data) could provide robust support for complex groups, despite the fact that the 16S divergences among several species of 
<taxonomicName id="4B88EDEF8B36B541992F636DFB5B5628" authorityName="Cope" authorityYear="1869" box="[1105,1192,260,285]" class="Amphibia" family="Plethodontidae" genus="Thorius" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B36B541992F636DFB5B5628" box="[1105,1192,260,285]" italics="true" pageId="11" pageNumber="498">Thorius</emphasis>
</taxonomicName>
were equal to or less than 2%. More studies evaluating the validity of species weakly supported with mitochondrial DNA are needed.
</paragraph>
<paragraph id="8C37966C8B36B5419DB96325FE115491" blockId="11.[151,1437,151,1472]" pageId="11" pageNumber="498">
According to 
<bibRefCitation id="E819EB9D8B36B5419C126325FDAB5650" box="[364,600,332,357]" pageId="11" pageNumber="498" refString="Rovito, S. M., Parra-Olea, G., Recuero, E. &amp; Wake, D. B. (2015) Diversification and biogeographical history of Neotropical plethodontid salamanders. Zoological Journal of the Linnean Society, 175, 167 - 188. https: // doi. org / 10.1111 / zoj. 12271" type="journal article">
Rovito 
<emphasis id="BEFC4A7E8B36B5419CBA6324FDF15650" box="[452,514,332,357]" italics="true" pageId="11" pageNumber="498">et al.</emphasis>
(2015)
</bibRefCitation>
the genus 
<taxonomicName id="4B88EDEF8B36B5419F9F6324FCA55651" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[737,854,333,356]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B36B5419F9F6324FCA55651" box="[737,854,333,356]" italics="true" pageId="11" pageNumber="498">Nototriton</emphasis>
</taxonomicName>
arose 
<emphasis id="BEFC4A7E8B36B5419ED46324FC385651" box="[938,971,333,356]" italics="true" pageId="11" pageNumber="498">ca.</emphasis>
10 Ma and due the Nearctic origin of Bolitoglossini (Shen 
<emphasis id="BEFC4A7E8B36B5419CF96318FE3156BD" box="[391,450,367,392]" italics="true" pageId="11" pageNumber="498">et al.</emphasis>
2016) it is highly probable that the genus arose in Nuclear Central America and only recently dispersed to Isthmian Central America –affected possibly by their low dispersal potential. According to this scenario, we speculate that the species of 
<taxonomicName id="4B88EDEF8B36B5419FA463D0FCBC56E5" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[730,847,441,464]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B36B5419FA463D0FCBC56E5" box="[730,847,441,464]" italics="true" pageId="11" pageNumber="498">Nototriton</emphasis>
</taxonomicName>
inhabiting 
<collectingCountry id="F49FD6FC8B36B5419EA063D1FB9356E5" box="[990,1120,439,464]" name="Costa Rica" pageId="11" pageNumber="498">Costa Rica</collectingCountry>
are the result of a rapid diversification during the Pliocene-Pleistocene, which would explain the relatively low genetic distance among members of the subgenus 
<emphasis id="BEFC4A7E8B36B5419CC56068FDC4552D" box="[443,567,513,536]" italics="true" pageId="11" pageNumber="498">
<taxonomicName id="4B88EDEF8B36B5419CC56068FDC0552D" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[443,563,513,536]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="genus">Nototriton</taxonomicName>
.
</emphasis>
<bibRefCitation id="E819EB9D8B36B5419F436396FCD7552D" box="[573,804,511,536]" pageId="11" pageNumber="498" refString="Nicolas, V., Schaeffer, B., Missoup, A. D., Kennis, J., Colyn, M., Denys, C., Tatard, C., Cruaud, C. &amp; Laredo, C. (2012) Assessment of three mitochondrial genes (16 S, Cytb, CO 1) for identifying species in the Praomyini tribe (Rodentia: Muridae). PLoS one, 7, e 36586. https: // doi. org / 10.1371 / journal. pone. 0036586" type="journal article">
Nicolas 
<emphasis id="BEFC4A7E8B36B5419FE46068FD21552D" box="[666,722,511,536]" italics="true" pageId="11" pageNumber="498">et al.</emphasis>
(2012)
</bibRefCitation>
used three mitochondrial genes (16S, COI, and cyt b) to identify species within of a group of Old World rodents that arose during the Pleistocene; they founded very low genetic distances between the described species (as low as 0.49 % in 16S and 2.94 % in COI). Although, there is not a minimal divergence value suggested to differentiate species for cyt 
<emphasis id="BEFC4A7E8B36B5419EB46005FC2B55B0" box="[970,984,620,645]" italics="true" pageId="11" pageNumber="498">b</emphasis>
the divergence is known to be as low as 5 % between species that are well supported morphologically, i.e. 
<taxonomicName id="4B88EDEF8B36B5419EFA60F8FBC7559D" box="[900,1076,655,680]" class="Amphibia" family="Plethodontidae" genus="Bolitoglossa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="aureogularis">
<emphasis id="BEFC4A7E8B36B5419EFA60F8FBC7559D" box="[900,1076,655,680]" italics="true" pageId="11" pageNumber="498">B. aureogularis</emphasis>
</taxonomicName>
and 
<taxonomicName id="4B88EDEF8B36B541991560F8FEEE55F9" authority="(Boza-Oviedo et al. 2012)" authorityName="Boza-Oviedo et al." authorityYear="2012" class="Amphibia" family="Plethodontidae" genus="Bolitoglossa" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="robinsoni">
<emphasis id="BEFC4A7E8B36B541991560F8FB06559D" box="[1131,1269,655,680]" italics="true" pageId="11" pageNumber="498">B. robinsoni</emphasis>
(
<bibRefCitation id="E819EB9D8B36B541987B60E6FEE555F9" author="Boza-Oviedo" pageId="11" pageNumber="498" refString="Boza-Oviedo, E., Rovito, S. M., Chaves, G., Garcia-Rodriguez, A., Artavia, L. G., Bolanos, F. &amp; Wake, D. B. (2012) Salamanders from the eastern Cordillera de Talamanca, Costa Rica, with descriptions of five new species (Plethodontidae: Bolitoglossa, Nototriton, and Oedipina) and natural history notes from recent expeditions. Zootaxa, 3309, 36 - 61." type="journal article" year="2012">
Boza-Oviedo 
<emphasis id="BEFC4A7E8B36B5419DE960DCFF2255F8" box="[151,209,692,717]" italics="true" pageId="11" pageNumber="498">et al.</emphasis>
2012
</bibRefCitation>
)
</taxonomicName>
. 
<bibRefCitation id="E819EB9D8B36B5419C5760DDFDD755F9" author="Johns" box="[297,548,692,717]" pageId="11" pageNumber="498" refString="Johns, G. C. &amp; Avise, J. C. (1998) A comparative summary of genetic distances in the vertebrates from the mitochondrial cytochrome b gene. Molecular Biology and Evolution, 15, 1481 - 1490. https: // doi. org / 10.1093 / oxfordjournals. molbev. a 025875" type="journal article" year="1998">Johns &amp; Avice (1998)</bibRefCitation>
reviewed the cyt 
<emphasis id="BEFC4A7E8B36B5419F8F60DDFD0C55F8" box="[753,767,692,717]" italics="true" pageId="11" pageNumber="498">b</emphasis>
distances among vertebrates and they found distances less than 5 % between different species of amphibians and reptiles of the same genus. It is also important mention that the use of a threshold for species delimitation has been criticized if it is not accompanied by other evidence such as morphological, behavioral or ecological characteristics (
<bibRefCitation id="E819EB9D8B36B5419E6C6149FBE1540D" author="Meyer" box="[786,1042,799,824]" pageId="11" pageNumber="498" refString="Meyer, C. P. &amp; Paulay, G. (2005) DNA barcoding: error rates based on comprehensive sampling. PLoS biology, 3, e 422. https: // doi. org / 10.1371 / journal. pbio. 0030422" type="journal article" year="2005">Meyer &amp; Paulay 2005</bibRefCitation>
; 
<bibRefCitation id="E819EB9D8B36B54199606176FB13540D" author="Clare" box="[1054,1248,799,824]" pageId="11" pageNumber="498" refString="Clare, E. L., Lim, B. K., Fenton, M. B. &amp; Hebert, P. D. N. (2011) Neotropical bats: estimating species diversity with DNA barcodes. PloS one, 6, e 22648. https: // doi. org / 10.1371 / journal. pone. 0022648" type="journal article" year="2011">
Clare 
<emphasis id="BEFC4A7E8B36B541991D6148FB6F540D" box="[1123,1180,799,824]" italics="true" pageId="11" pageNumber="498">et al.</emphasis>
2011
</bibRefCitation>
). Padial &amp; De la Riva (2011) pointed out that thresholds may be useful to identify groups of interest for further study, but minimum levels of divergence for certain traits cannot be defined as a prerequisite for species recognition under the evolutionary species concept.
</paragraph>
<paragraph id="8C37966C8B36B5419DB961D9FDB95395" blockId="11.[151,1437,151,1472]" pageId="11" pageNumber="498">
Due to the relatively few sequences available for members of the subgenus 
<taxonomicName id="4B88EDEF8B36B541996661D8FB7E54FD" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[1048,1165,945,968]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B36B541996661D8FB7E54FD" box="[1048,1165,945,968]" italics="true" pageId="11" pageNumber="498">Nototriton</emphasis>
</taxonomicName>
, details surrounding the phylogenetic relationships within this taxon are for the most part poorly known. The two species groups within the subgenus 
<taxonomicName id="4B88EDEF8B36B5419C796190FE8F5325" baseAuthorityName="Dubois &amp; Raffaelli" baseAuthorityYear="2012" box="[263,380,1017,1040]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="genus">
<emphasis id="BEFC4A7E8B36B5419C796190FE8F5325" box="[263,380,1017,1040]" italics="true" pageId="11" pageNumber="498">Nototriton</emphasis>
</taxonomicName>
are well supported (i.e. 
<taxonomicName id="4B88EDEF8B36B5419FF06190FCF75325" baseAuthorityName="Stejneger" baseAuthorityYear="1911" box="[654,772,1015,1040]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B36B5419FF06190FCF75325" box="[654,772,1015,1040]" italics="true" pageId="11" pageNumber="498">N. picadoi</emphasis>
</taxonomicName>
s.g. and 
<taxonomicName id="4B88EDEF8B36B5419E146190FC145325" baseAuthorityName="Taylor" baseAuthorityYear="1949" box="[874,999,1015,1040]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="richardi">
<emphasis id="BEFC4A7E8B36B5419E146190FC145325" box="[874,999,1015,1040]" italics="true" pageId="11" pageNumber="498">N. richardi</emphasis>
</taxonomicName>
s.g.), but the relationships within each group are poorly understood, especially so with the 
<taxonomicName id="4B88EDEF8B36B5419F986674FC905300" baseAuthorityName="Taylor" baseAuthorityYear="1949" box="[742,867,1052,1077]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="richardi">
<emphasis id="BEFC4A7E8B36B5419F986674FC905300" box="[742,867,1052,1077]" italics="true" pageId="11" pageNumber="498">N. richardi</emphasis>
</taxonomicName>
species group (i.e. 
<taxonomicName id="4B88EDEF8B36B541993C6674FB4C5300" baseAuthorityName="Taylor" baseAuthorityYear="1949" box="[1090,1215,1052,1077]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="richardi">
<emphasis id="BEFC4A7E8B36B541993C6674FB4C5300" box="[1090,1215,1052,1077]" italics="true" pageId="11" pageNumber="498">N. richardi</emphasis>
</taxonomicName>
and 
<taxonomicName id="4B88EDEF8B36B54199866674FA9A5301" authorityName="Good &amp; Wake" authorityYear="1993" box="[1272,1385,1053,1076]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="tapanti">
<emphasis id="BEFC4A7E8B36B54199866674FA9A5301" box="[1272,1385,1053,1076]" italics="true" pageId="11" pageNumber="498">N. tapanti</emphasis>
</taxonomicName>
) for which sequences are only available for 
<taxonomicName id="4B88EDEF8B36B5419F166628FD1B536D" baseAuthorityName="Taylor" baseAuthorityYear="1949" box="[616,744,1087,1112]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="richardi">
<emphasis id="BEFC4A7E8B36B5419F166628FD1B536D" box="[616,744,1087,1112]" italics="true" pageId="11" pageNumber="498">N. richardi</emphasis>
</taxonomicName>
. 
<taxonomicName id="4B88EDEF8B36B5419F846628FC3B536D" authorityName="Good &amp; Wake" authorityYear="1993" box="[762,968,1089,1112]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="tapanti">
<emphasis id="BEFC4A7E8B36B5419F846628FC3B536D" box="[762,968,1089,1112]" italics="true" pageId="11" pageNumber="498">Nototriton tapanti</emphasis>
</taxonomicName>
is currently only known from the adult female 
<typeStatus id="533328CE8B36B5419D93660DFEBC5348" box="[237,335,1124,1149]" pageId="11" pageNumber="498" type="holotype">holotype</typeStatus>
which was collected in 1986 and unfortunately no tissues from this specimen are available for genetic analysis (
<bibRefCitation id="E819EB9D8B36B5419C2666EEFDCE5395" author="Good" box="[344,573,1159,1184]" pageId="11" pageNumber="498" refString="Good, D. A. &amp; Wake, D. B. (1993) Systematic studies of the Costa Rican moss salamanders, genus Nototriton, with descriptions of three new species. Herpetological Monographs, 7, 131 - 159. https: // doi. org / 10.2307 / 1466956" type="journal article" year="1993">
Good &amp; 
<collectingCountry id="F49FD6FC8B36B5419CC566EEFE095395" box="[443,506,1159,1184]" name="Wake Island" pageId="11" pageNumber="498">Wake</collectingCountry>
1993
</bibRefCitation>
).
</paragraph>
<paragraph id="8C37966C8B36B5419DB966C4FB5852F5" blockId="11.[151,1437,151,1472]" pageId="11" pageNumber="498">
<taxonomicName id="4B88EDEF8B36B5419DB966C4FE2153F1" authorityName="Arias &amp; Kubicki" authorityYear="2018" box="[199,466,1197,1220]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="costaricense">
<emphasis id="BEFC4A7E8B36B5419DB966C4FE2153F1" box="[199,466,1197,1220]" italics="true" pageId="11" pageNumber="498">Nototriton costaricense</emphasis>
</taxonomicName>
was found to be the sister clade to all species of the 
<taxonomicName id="4B88EDEF8B36B541993766C4FB3253F0" baseAuthorityName="Stejneger" baseAuthorityYear="1911" box="[1097,1217,1196,1221]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B36B541993766C4FB3253F0" box="[1097,1217,1196,1221]" italics="true" pageId="11" pageNumber="498">N. picadoi</emphasis>
</taxonomicName>
species group that inhabit 
<collectingCountry id="F49FD6FC8B36B5419D8F66B9FE8053DD" box="[241,371,1231,1256]" name="Costa Rica" pageId="11" pageNumber="498">Costa Rica</collectingCountry>
, however the support for that relationship is relatively weak. Additionally, the alpha-level distinction between 
<taxonomicName id="4B88EDEF8B36B5419C02669CFE005238" baseAuthorityName="Stejneger" baseAuthorityYear="1911" box="[380,499,1268,1293]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B36B5419C02669CFE005238" box="[380,499,1268,1293]" italics="true" pageId="11" pageNumber="498">N. picadoi</emphasis>
</taxonomicName>
and 
<taxonomicName id="4B88EDEF8B36B5419F53669CFD595239" box="[557,682,1269,1292]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="matama">
<emphasis id="BEFC4A7E8B36B5419F53669CFD595239" box="[557,682,1269,1292]" italics="true" pageId="11" pageNumber="498">N. matama</emphasis>
</taxonomicName>
is weakly supported with a 0.6% genetic distance for 16S (Boza- Oviedo 
<emphasis id="BEFC4A7E8B36B5419D8E6770FEDB5205" box="[240,296,1303,1328]" italics="true" pageId="11" pageNumber="498">et al.</emphasis>
2012) in addition to phenotypic support that is questionable due to possible artifact of preservation or desiccation that might have taken place during the manipulation of the specimens of the 
<typeStatus id="533328CE8B36B54199026754FB5F5260" box="[1148,1196,1341,1365]" pageId="11" pageNumber="498">type</typeStatus>
series of 
<taxonomicName id="4B88EDEF8B36B54198646754FA665261" box="[1306,1429,1341,1364]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="matama">
<emphasis id="BEFC4A7E8B36B54198646754FA665261" box="[1306,1429,1341,1364]" italics="true" pageId="11" pageNumber="498">N. matama</emphasis>
</taxonomicName>
. We feel that the relationship not only between 
<taxonomicName id="4B88EDEF8B36B5419FCA6708FCD8524D" baseAuthorityName="Stejneger" baseAuthorityYear="1911" box="[692,811,1375,1400]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B36B5419FCA6708FCD8524D" box="[692,811,1375,1400]" italics="true" pageId="11" pageNumber="498">N. picadoi</emphasis>
</taxonomicName>
and 
<taxonomicName id="4B88EDEF8B36B5419E186708FC10524D" box="[870,995,1377,1400]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="matama">
<emphasis id="BEFC4A7E8B36B5419E186708FC10524D" box="[870,995,1377,1400]" italics="true" pageId="11" pageNumber="498">N. matama</emphasis>
</taxonomicName>
requires a further detailed review, but those interspecific relationships among the members of the 
<taxonomicName id="4B88EDEF8B36B5419E4267ECFC4252A8" baseAuthorityName="Stejneger" baseAuthorityYear="1911" box="[828,945,1412,1437]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="picadoi">
<emphasis id="BEFC4A7E8B36B5419E4267ECFC4252A8" box="[828,945,1412,1437]" italics="true" pageId="11" pageNumber="498">N. picadoi</emphasis>
</taxonomicName>
and 
<taxonomicName id="4B88EDEF8B36B5419E9467ECFB9452A8" baseAuthorityName="Taylor" baseAuthorityYear="1949" box="[1002,1127,1412,1437]" class="Amphibia" family="Plethodontidae" genus="Nototriton" higherTaxonomySource="GBIF" kingdom="Animalia" order="Caudata" pageId="11" pageNumber="498" phylum="Chordata" rank="species" species="richardi">
<emphasis id="BEFC4A7E8B36B5419E9467ECFB9452A8" box="[1002,1127,1412,1437]" italics="true" pageId="11" pageNumber="498">N. richardi</emphasis>
</taxonomicName>
species groups as a whole, with an emphasis to include more localities, DNA data, ecological data, and phenotypic data.
</paragraph>
</subSubSection>
</subSection>
</treatment>
</document>