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<document ID-DOI="http://dx.doi.org/10.3897/zookeys.868.26766" ID-GBIF-Dataset="43d18445-d3e8-4e30-bdcd-7f82bde7616a" ID-PMC="PMC6687670" ID-Pensoft-Pub="1313-2970-868-1" ID-Pensoft-UUID="679C0640C93B545BAD44B1013B0B12E5" ID-PubMed="31406482" ID-ZooBank="ED7FD98F964D402FAB705FC9B4CA4851" ModsDocID="1313-2970-868-1" checkinTime="1564724838606" checkinUser="pensoft" docAuthor="Paez, Nadia B. &amp; Ron, Santiago R." docDate="2019" docId="DE830EC800215B529F39DED3730E3CCE" docLanguage="en" docName="ZooKeys 868: 1-112" docOrigin="ZooKeys 868" docSource="http://dx.doi.org/10.3897/zookeys.868.26766" docTitle="(Huicundomantis) Paez &amp; Ron, 2019, subgenus nov." docType="treatment" docVersion="4" lastPageNumber="1" masterDocId="FFB7FFF73E69FFFBFFD8FF95FFBEB241" masterDocTitle="Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new species" masterLastPageNumber="112" masterPageNumber="1" pageNumber="1" updateTime="1668167659737" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Systematics of Huicundomantis, a new subgenus of Pristimantis (Anura, Strabomantidae) with extraordinary cryptic diversity and eleven new species</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Paez, Nadia B.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ron, Santiago R.</mods:namePart>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2019</mods:date>
<mods:detail type="volume">
<mods:number>868</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>1</mods:start>
<mods:end>112</mods:end>
</mods:extent>
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<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.868.26766</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.868.26766</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-868-1</mods:identifier>
<mods:identifier type="ZooBank">ED7FD98F964D402FAB705FC9B4CA4851</mods:identifier>
<mods:identifier type="Pensoft-UUID">679C0640C93B545BAD44B1013B0B12E5</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="159293047" LSID="urn:lsid:plazi:treatment:DE830EC800215B529F39DED3730E3CCE" httpUri="http://treatment.plazi.org/id/DE830EC800215B529F39DED3730E3CCE" lastPageNumber="1" pageId="0" pageNumber="1">
<subSubSection pageId="0" pageNumber="1" type="nomenclature">
<paragraph pageId="0" pageNumber="1">
<taxonomicName LSID="de830ec8-0021-5b52-9f39-ded3730e3cce" family="Strabomantidae" higherTaxonomySource="treatment-meta" kingdom="Animalia" lsidName="(Huicundomantis)" order="Anura" pageId="0" pageNumber="1" rank="subGenus" subGenus="Huicundomantis">Huicundomantis</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="1">subgenus nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="type species">
<paragraph pageId="0" pageNumber="1">Type species.</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis phoxocephalus" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis phoxocephalus</emphasis>
</taxonomicName>
(Lynch, 1979).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="definition">
<paragraph pageId="0" pageNumber="1">Definition.</paragraph>
<paragraph pageId="0" pageNumber="1">
This clade is strongly supported by genetic evidence (
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Phylogenetic relationships of Pristimantis, sugenus Huicundomantis. Maximum likelihood tree for genes 16 S, ND 1 and RAG 1. Bootstrap values (%) followed by Bayesian posterior probabilities are shown under the corresponding branches. Asterisks indicate support values of 100 (bootstrap) or 1 (posterior probabilities); missing values indicate values below 50 (bootstrap) or 0.5 (posterior probability). The collection number, identification, province and locality of the samples are shown next to each terminal; all samples are from Ecuador. Outgroup is not shown. Abbreviations: CCS = confirmed candidate species, NP = national park, PF = protected forest, UCS = unconfirmed candidate species." figureDoi="10.3897/zookeys.868.26766.figure3" httpUri="https://binary.pensoft.net/fig/322395" pageId="0" pageNumber="1">Fig. 3</figureCitation>
). Morphological synapomorphies are unknown. Members of this clade are characterized by: (i) dorsolateral folds absent (except for
<taxonomicName lsidName="P. atratus" pageId="0" pageNumber="1" rank="species" species="atratus">
<emphasis italics="true" pageId="0" pageNumber="1">P. atratus</emphasis>
</taxonomicName>
); (ii) cranial crests absent (except for
<taxonomicName lsidName="P. atratus" pageId="0" pageNumber="1" rank="species" species="atratus">
<emphasis italics="true" pageId="0" pageNumber="1">P. atratus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. percultus" pageId="0" pageNumber="1" rank="species" species="percultus">
<emphasis italics="true" pageId="0" pageNumber="1">P. percultus</emphasis>
</taxonomicName>
, and
<taxonomicName lsidName="P. spinosus" pageId="0" pageNumber="1" rank="species" species="spinosus">
<emphasis italics="true" pageId="0" pageNumber="1">P. spinosus</emphasis>
</taxonomicName>
); (iii) tympanic membrane and tympanic annulus prominent (both absent in
<taxonomicName lsidName="P. philipi" pageId="0" pageNumber="1" rank="species" species="philipi">
<emphasis italics="true" pageId="0" pageNumber="1">P. philipi</emphasis>
</taxonomicName>
); (iv) dentigerous processes of vomer present; (v) small to prominent tubercles on heel; (vi) fingers with lateral fringes; (vii) basal webbing between toes (except for
<taxonomicName lsidName="P. balionotus" pageId="0" pageNumber="1" rank="species" species="balionotus">
<emphasis italics="true" pageId="0" pageNumber="1">P. balionotus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. hampatusami" pageId="0" pageNumber="1" rank="species" species="hampatusami">
<emphasis italics="true" pageId="0" pageNumber="1">P. hampatusami</emphasis>
</taxonomicName>
, and
<taxonomicName lsidName="P. philipi" pageId="0" pageNumber="1" rank="species" species="philipi">
<emphasis italics="true" pageId="0" pageNumber="1">P. philipi</emphasis>
</taxonomicName>
which lack webbing); (viii) Toe V longer or much longer than Toe III (
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Palmar and plantar surfaces of Pristimantis atillo sp. nov., Pristimantis chomskyi sp. nov., Pristimantis gloria sp. nov., and Pristimantis jimenezi sp. nov. Photographs of left hand and foot of the holotypes A Pristimantis atillo sp. nov.: QCAZ 42500, male B Pristimantis chomskyi sp. nov.: QCAZ 47515, male C Pristimantis gloria sp. nov.: QCAZ 57201, female D Pristimantis jimenezi sp. nov.: QCAZ 45170, male. All specimens are shown at the same scale." figureDoi="10.3897/zookeys.868.26766.figure7" httpUri="https://binary.pensoft.net/fig/322399" pageId="0" pageNumber="1">Figs 7</figureCitation>
-
<figureCitation captionStart="Figure 9" captionStartId="F9" captionText="Figure 9. Palmar and plantar surfaces of Pristimantis teslai sp. nov., Pristimantis torresi sp. nov., Pristimantis totoroi sp. nov., and Pristimantis verrucolatus sp. nov. Photographs of hand and foot of the holotypes of the following species: A Pristimantis teslai sp. nov.: QCAZ 46213, male B Pristimantis torresi sp. nov.: QCAZ 47342, female C Pristimantis totoroi sp. nov.: QCAZ 25105, male D Pristimantis verrucolatus sp. nov.: QCAZ 46982, male. All specimens are shown at the same scale." figureDoi="10.3897/zookeys.868.26766.figure9" httpUri="https://binary.pensoft.net/fig/322401" pageId="0" pageNumber="1">9</figureCitation>
); (ix) in life, groins and concealed surfaces of thighs with distinctive coloration patterns including flash colors and light or bright colored flecks or spots on a darker background; colors, shapes and sizes of these ornaments are variable among species; (x) SVL females 24.9-46.8 mm; SVL males 16.1-34.5 mm. Species of this clade may bear a fleshy keel or a papilla at the tip of the snout, and lateral, middorsal, or postocular folds.
</paragraph>
<caption doi="10.3897/zookeys.868.26766.figure7" httpUri="https://binary.pensoft.net/fig/322399" pageId="0" pageNumber="1" start="Figure 7" startId="F7">
<paragraph pageId="0" pageNumber="1">
Figure 7. Palmar and plantar surfaces of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis atillo" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="atillo">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis atillo</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis chomskyi" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="chomskyi">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis chomskyi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis gloria" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="gloria">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis gloria</emphasis>
</taxonomicName>
sp. nov., and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis jimenezi" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="jimenezi">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis jimenezi</emphasis>
</taxonomicName>
sp. nov. Photographs of left hand and foot of the holotypes
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis atillo" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="atillo">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis atillo</emphasis>
</taxonomicName>
sp. nov.: QCAZ 42500, male
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis chomskyi" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="chomskyi">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis chomskyi</emphasis>
</taxonomicName>
sp. nov.: QCAZ 47515, male
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis gloria" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="gloria">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis gloria</emphasis>
</taxonomicName>
sp. nov.: QCAZ 57201, female
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis jimenezi" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="jimenezi">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis jimenezi</emphasis>
</taxonomicName>
sp. nov.: QCAZ 45170, male. All specimens are shown at the same scale.
</paragraph>
</caption>
<caption doi="10.3897/zookeys.868.26766.figure8" httpUri="https://binary.pensoft.net/fig/322400" pageId="0" pageNumber="1" start="Figure 8" startId="F8">
<paragraph pageId="0" pageNumber="1">
Figure 8. Palmar and plantar surfaces of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis lutzae" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="lutzae">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis lutzae</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis multicolor" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="multicolor">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis multicolor</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis nangaritza" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="nangaritza">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis nangaritza</emphasis>
</taxonomicName>
sp. nov., and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis phoxocephalus" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis phoxocephalus</emphasis>
</taxonomicName>
. Photographs of hand and foot of the holotypes (except for
<taxonomicName lsidName="P. phoxocephalus" pageId="0" pageNumber="1" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">P. phoxocephalus</emphasis>
</taxonomicName>
) of the following species:
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis lutzae" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="lutzae">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis lutzae</emphasis>
</taxonomicName>
sp. nov.: QCAZ 37546, female
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis multicolor" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="multicolor">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis multicolor</emphasis>
</taxonomicName>
sp. nov.: QCAZ 47213, male
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis nangaritza" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="nangaritza">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis nangaritza</emphasis>
</taxonomicName>
sp. nov.: QCAZ 41710, female
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis phoxocephalus" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis phoxocephalus</emphasis>
</taxonomicName>
: QCAZ 58463, female from the type locality. All specimens are shown at the same scale.
</paragraph>
</caption>
<caption doi="10.3897/zookeys.868.26766.figure9" httpUri="https://binary.pensoft.net/fig/322401" pageId="0" pageNumber="1" start="Figure 9" startId="F9">
<paragraph pageId="0" pageNumber="1">
Figure 9. Palmar and plantar surfaces of
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis teslai" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="teslai">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis teslai</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis torresi" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="torresi">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis torresi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis totoroi" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="totoroi">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis totoroi</emphasis>
</taxonomicName>
sp. nov., and
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis verrucolatus" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="verrucolatus">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis verrucolatus</emphasis>
</taxonomicName>
sp. nov. Photographs of hand and foot of the holotypes of the following species:
<emphasis bold="true" pageId="0" pageNumber="1">A</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis teslai" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="teslai">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis teslai</emphasis>
</taxonomicName>
sp. nov.: QCAZ 46213, male
<emphasis bold="true" pageId="0" pageNumber="1">B</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis torresi" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="torresi">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis torresi</emphasis>
</taxonomicName>
sp. nov.: QCAZ 47342, female
<emphasis bold="true" pageId="0" pageNumber="1">C</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis totoroi" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="totoroi">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis totoroi</emphasis>
</taxonomicName>
sp. nov.: QCAZ 25105, male
<emphasis bold="true" pageId="0" pageNumber="1">D</emphasis>
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis verrucolatus" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="verrucolatus">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis verrucolatus</emphasis>
</taxonomicName>
sp. nov.: QCAZ 46982, male. All specimens are shown at the same scale.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="content">
<paragraph pageId="0" pageNumber="1">Content.</paragraph>
<paragraph pageId="0" pageNumber="1">
This clade comprises 23 described species (11 of them described below):
<taxonomicName lsidName="P. atillo" pageId="0" pageNumber="1" rank="species" species="atillo">
<emphasis italics="true" pageId="0" pageNumber="1">P. atillo</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. atratus" pageId="0" pageNumber="1" rank="species" species="atratus">
<emphasis italics="true" pageId="0" pageNumber="1">P. atratus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. balionotus" pageId="0" pageNumber="1" rank="species" species="balionotus">
<emphasis italics="true" pageId="0" pageNumber="1">P. balionotus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. chomskyi" pageId="0" pageNumber="1" rank="species" species="chomskyi">
<emphasis italics="true" pageId="0" pageNumber="1">P. chomskyi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. cryptomelas" pageId="0" pageNumber="1" rank="species" species="cryptomelas">
<emphasis italics="true" pageId="0" pageNumber="1">P. cryptomelas</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. gagliardoi" pageId="0" pageNumber="1" rank="species" species="gagliardoi">
<emphasis italics="true" pageId="0" pageNumber="1">P. gagliardoi</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. gloria" pageId="0" pageNumber="1" rank="species" species="gloria">
<emphasis italics="true" pageId="0" pageNumber="1">P. gloria</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. hampatusami" pageId="0" pageNumber="1" rank="species" species="hampatusami">
<emphasis italics="true" pageId="0" pageNumber="1">P. hampatusami</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. jimenezi" pageId="0" pageNumber="1" rank="species" species="jimenezi">
<emphasis italics="true" pageId="0" pageNumber="1">P. jimenezi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. lutzae" pageId="0" pageNumber="1" rank="species" species="lutzae">
<emphasis italics="true" pageId="0" pageNumber="1">P. lutzae</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. multicolor" pageId="0" pageNumber="1" rank="species" species="multicolor">
<emphasis italics="true" pageId="0" pageNumber="1">P. multicolor</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. muscosus" pageId="0" pageNumber="1" rank="species" species="muscosus">
<emphasis italics="true" pageId="0" pageNumber="1">P. muscosus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. nangaritza" pageId="0" pageNumber="1" rank="species" species="nangaritza">
<emphasis italics="true" pageId="0" pageNumber="1">P. nangaritza</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. percultus" pageId="0" pageNumber="1" rank="species" species="percultus">
<emphasis italics="true" pageId="0" pageNumber="1">P. percultus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. philipi" pageId="0" pageNumber="1" rank="species" species="philipi">
<emphasis italics="true" pageId="0" pageNumber="1">P. philipi</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. phoxocephalus" pageId="0" pageNumber="1" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">P. phoxocephalus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. spinosus" pageId="0" pageNumber="1" rank="species" species="spinosus">
<emphasis italics="true" pageId="0" pageNumber="1">P. spinosus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. teslai" pageId="0" pageNumber="1" rank="species" species="teslai">
<emphasis italics="true" pageId="0" pageNumber="1">P. teslai</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. tinguichaca" pageId="0" pageNumber="1" rank="species" species="tinguichaca">
<emphasis italics="true" pageId="0" pageNumber="1">P. tinguichaca</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. torresi" pageId="0" pageNumber="1" rank="species" species="torresi">
<emphasis italics="true" pageId="0" pageNumber="1">P. torresi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. totoroi" pageId="0" pageNumber="1" rank="species" species="totoroi">
<emphasis italics="true" pageId="0" pageNumber="1">P. totoroi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. versicolor" pageId="0" pageNumber="1" rank="species" species="versicolor">
<emphasis italics="true" pageId="0" pageNumber="1">P. versicolor</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. verrucolatus" pageId="0" pageNumber="1" rank="species" species="verrucolatus">
<emphasis italics="true" pageId="0" pageNumber="1">P. verrucolatus</emphasis>
</taxonomicName>
sp. nov. This clade encompasses the
<taxonomicName lsidName="P. phoxocephalus" pageId="0" pageNumber="1" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">P. phoxocephalus</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="P. cryptomelas" pageId="0" pageNumber="1" rank="species" species="cryptomelas">
<emphasis italics="true" pageId="0" pageNumber="1">P. cryptomelas</emphasis>
</taxonomicName>
species groups.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="distribution">
<paragraph pageId="0" pageNumber="1">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="1">
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Huicundomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Huicundomantis" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Huicundomantis</emphasis>
</taxonomicName>
occurs in Eastern and Western Andean slopes and Inter-Andean valleys of southern and central Ecuador, and Eastern Andean slopes of northern Peru. They inhabit the following Natural Regions: Deciduous Costa Forest, Western Foothill Forest, Western Montane Forest, Paramo, Inter-Andean Shrub, Eastern Montane Forest, and Eastern Foothill Forest, between elevations of 230 and 4200 m a.s.l.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="1" type="etymology">
<paragraph pageId="0" pageNumber="1">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="1">
We name this clade
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Huicundomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Huicundomantis" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="1">Huicundomantis</emphasis>
</taxonomicName>
because these frogs are frequently found inside bromeliad plants. Huicundo is a word in Quechua, an indigenous South American language, locally used to referring to bromeliads.
</paragraph>
<subSection pageId="0" pageNumber="1" type="pristimantis phoxocephalus species group new taxon">
<paragraph pageId="0" pageNumber="1">
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Pristimantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pristimantis phoxocephalus" order="Anura" pageId="0" pageNumber="1" phylum="Chordata" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">Pristimantis phoxocephalus</emphasis>
</taxonomicName>
species group new taxon
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Definition.</emphasis>
The
<taxonomicName lsidName="P. phoxocephalus" pageId="0" pageNumber="1" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">P. phoxocephalus</emphasis>
</taxonomicName>
species group is strongly supported in our phylogeny. Members of this group share the following morphological traits: (i) dorsolateral folds absent (except for
<taxonomicName lsidName="P. atratus" pageId="0" pageNumber="1" rank="species" species="atratus">
<emphasis italics="true" pageId="0" pageNumber="1">P. atratus</emphasis>
</taxonomicName>
); (ii) snout with a fleshy keel or papilla at the tip; (iii) cranial crests absent (except for
<taxonomicName lsidName="P. atratus" pageId="0" pageNumber="1" rank="species" species="atratus">
<emphasis italics="true" pageId="0" pageNumber="1">P. atratus</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="P. percultus" pageId="0" pageNumber="1" rank="species" species="percultus">
<emphasis italics="true" pageId="0" pageNumber="1">P. percultus</emphasis>
</taxonomicName>
); (iv) tympanic membrane and tympanic annulus prominent; (v) dentigerous processes of vomer present; (vi) males with vocal slits (except for
<taxonomicName lsidName="P. versicolor" pageId="0" pageNumber="1" rank="species" species="versicolor">
<emphasis italics="true" pageId="0" pageNumber="1">P. versicolor</emphasis>
</taxonomicName>
); (vii) fingers and toes with lateral fringes; (viii) basal webbing between toes (except for
<taxonomicName lsidName="P. balionotus" pageId="0" pageNumber="1" rank="species" species="balionotus">
<emphasis italics="true" pageId="0" pageNumber="1">P. balionotus</emphasis>
</taxonomicName>
and
<taxonomicName lsidName="P. hampatusami" pageId="0" pageNumber="1" rank="species" species="hampatusami">
<emphasis italics="true" pageId="0" pageNumber="1">P. hampatusami</emphasis>
</taxonomicName>
); (ix) Toe V longer or much longer than Toe III (
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Palmar and plantar surfaces of Pristimantis atillo sp. nov., Pristimantis chomskyi sp. nov., Pristimantis gloria sp. nov., and Pristimantis jimenezi sp. nov. Photographs of left hand and foot of the holotypes A Pristimantis atillo sp. nov.: QCAZ 42500, male B Pristimantis chomskyi sp. nov.: QCAZ 47515, male C Pristimantis gloria sp. nov.: QCAZ 57201, female D Pristimantis jimenezi sp. nov.: QCAZ 45170, male. All specimens are shown at the same scale." figureDoi="10.3897/zookeys.868.26766.figure7" httpUri="https://binary.pensoft.net/fig/322399" pageId="0" pageNumber="1">Fig. 7</figureCitation>
); (x) in life, groins and concealed surfaces of thighs with distinctive coloration, including flash colors and light or bright colored flecks or spots on a darker background; colors, shapes, and sizes of ornaments variable among species; (xi) SVL females 24.9-46.8 mm; SVL males 16.8-34.5 mm.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Content.</emphasis>
Currently, the
<taxonomicName lsidName="P. phoxocephalus" pageId="0" pageNumber="1" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">P. phoxocephalus</emphasis>
</taxonomicName>
group comprises 17 described species (10 of them are described below):
<taxonomicName lsidName="P. atillo" pageId="0" pageNumber="1" rank="species" species="atillo">
<emphasis italics="true" pageId="0" pageNumber="1">P. atillo</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. atratus" pageId="0" pageNumber="1" rank="species" species="atratus">
<emphasis italics="true" pageId="0" pageNumber="1">P. atratus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. balionotus" pageId="0" pageNumber="1" rank="species" species="balionotus">
<emphasis italics="true" pageId="0" pageNumber="1">P. balionotus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. chomskyi" pageId="0" pageNumber="1" rank="species" species="chomskyi">
<emphasis italics="true" pageId="0" pageNumber="1">P. chomskyi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. gloria" pageId="0" pageNumber="1" rank="species" species="gloria">
<emphasis italics="true" pageId="0" pageNumber="1">P. gloria</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. hampatusami" pageId="0" pageNumber="1" rank="species" species="hampatusami">
<emphasis italics="true" pageId="0" pageNumber="1">P. hampatusami</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. jimenezi" pageId="0" pageNumber="1" rank="species" species="jimenezi">
<emphasis italics="true" pageId="0" pageNumber="1">P. jimenezi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. lutzae" pageId="0" pageNumber="1" rank="species" species="lutzae">
<emphasis italics="true" pageId="0" pageNumber="1">P. lutzae</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. multicolor" pageId="0" pageNumber="1" rank="species" species="multicolor">
<emphasis italics="true" pageId="0" pageNumber="1">P. multicolor</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. percultus" pageId="0" pageNumber="1" rank="species" species="percultus">
<emphasis italics="true" pageId="0" pageNumber="1">P. percultus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. phoxocephalus" pageId="0" pageNumber="1" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">P. phoxocephalus</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. teslai" pageId="0" pageNumber="1" rank="species" species="teslai">
<emphasis italics="true" pageId="0" pageNumber="1">P. teslai</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. tinguichaca" pageId="0" pageNumber="1" rank="species" species="tinguichaca">
<emphasis italics="true" pageId="0" pageNumber="1">P. tinguichaca</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. torresi" pageId="0" pageNumber="1" rank="species" species="torresi">
<emphasis italics="true" pageId="0" pageNumber="1">P. torresi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. totoroi" pageId="0" pageNumber="1" rank="species" species="totoroi">
<emphasis italics="true" pageId="0" pageNumber="1">P. totoroi</emphasis>
</taxonomicName>
sp. nov.,
<taxonomicName lsidName="P. versicolor" pageId="0" pageNumber="1" rank="species" species="versicolor">
<emphasis italics="true" pageId="0" pageNumber="1">P. versicolor</emphasis>
</taxonomicName>
,
<taxonomicName lsidName="P. verrucolatus" pageId="0" pageNumber="1" rank="species" species="verrucolatus">
<emphasis italics="true" pageId="0" pageNumber="1">P. verrucolatus</emphasis>
</taxonomicName>
sp. nov.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Distribution.</emphasis>
Eastern and Western Andean slopes and Inter-Andean valleys of central and southern Ecuador, in ten provinces: Azuay,
<normalizedToken originalValue="Bolívar">Bolivar</normalizedToken>
,
<normalizedToken originalValue="Cañar">Canar</normalizedToken>
, Chimborazo, Cotopaxi, El Oro, Loja, Morona Santiago, Tungurahua, and Zamora Chinchipe. They inhabit Deciduous Costa Forest, Western Foothill Forest, Western Montane Forest, Paramo, Inter-Andean Shrub, and Eastern Montane Forest Natural Regions, between 230 and 4100 m a.s.l.
</paragraph>
<paragraph pageId="0" pageNumber="1">
<emphasis bold="true" pageId="0" pageNumber="1">Remarks.</emphasis>
The
<taxonomicName lsidName="P. phoxocephalus" pageId="0" pageNumber="1" rank="species" species="phoxocephalus">
<emphasis italics="true" pageId="0" pageNumber="1">P. phoxocephalus</emphasis>
</taxonomicName>
species group is sister to the
<taxonomicName lsidName="P. cryptomelas" pageId="0" pageNumber="1" rank="species" species="cryptomelas">
<emphasis italics="true" pageId="0" pageNumber="1">P. cryptomelas</emphasis>
</taxonomicName>
species group.
</paragraph>
</subSection>
</subSubSection>
</treatment>
</document>