treatments-xml/data/03/F2/87/03F287F8FF93FF98CA17C225F9E4BD16.xml

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<mods:title>Miniaturization in Direct-Developing Frogs from Mexico with the Description of Six New Species</mods:title>
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<mods:namePart>Jameson, Tom J. M.</mods:namePart>
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<mods:namePart>Streicher, Jeffrey W.</mods:namePart>
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<mods:namePart>Manuelli, Luigi</mods:namePart>
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<mods:namePart>Head, Jason J.</mods:namePart>
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<mods:namePart>Smith, Eric N.</mods:namePart>
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<mods:title>Herpetological Monographs</mods:title>
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<mods:date>2022</mods:date>
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<heading box="[266,689,1119,1143]" centered="true" fontSize="10" level="4" pageId="27" pageNumber="27" reason="4">
<taxonomicName authority="(Brocchi 1877)" baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[266,689,1119,1143]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[266,513,1119,1142]" italics="true" pageId="27" pageNumber="27">Craugastormexicanus</emphasis>
(
<bibRefCitation author="Brocchi, P." box="[529,680,1119,1143]" pageId="27" pageNumber="27" pagination="175 - 179" refId="ref35609" refString="Brocchi, P. 1877. Sur quelques batraciens raniformes et bufoniformes de l'Amerique Centrale. Bulletin de La Societe Philomathique de Paris 1: 175 - 179. [In French.]" type="journal article" year="1877">Brocchi 1877</bibRefCitation>
)
</taxonomicName>
</heading>
</paragraph>
</subSubSection>
<subSubSection pageId="27" pageNumber="27" type="materials_examined">
<paragraph blockId="27.[142,813,1164,1804]" pageId="27" pageNumber="27">
<materialsCitation collectionCode="MNHNP" country="Mexico" location="Smith and Taylor" pageId="27" pageNumber="27" specimenCode="MNHNP 6218" specimenCount="1" stateProvince="Oaxaca">
<taxonomicName authority="Brocchi 1877: 184" authorityName="Brocchi" authorityPageNumber="184" authorityYear="1877" box="[142,592,1164,1188]" class="Amphibia" family="Leptodactylidae" genus="Leiuperus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<originalDetermination box="[142,547,1164,1188]" pageId="27" pageNumber="27">
<emphasis box="[142,379,1164,1187]" italics="true" pageId="27" pageNumber="27">Leiuperus mexicanus</emphasis>
<bibRefCitation author="Brocchi, P." box="[391,592,1164,1188]" pageId="27" pageNumber="27" pagination="175 - 179" refId="ref35609" refString="Brocchi, P. 1877. Sur quelques batraciens raniformes et bufoniformes de l'Amerique Centrale. Bulletin de La Societe Philomathique de Paris 1: 175 - 179. [In French.]" type="journal article" year="1877">Brocchi 1877:184</bibRefCitation>
</originalDetermination>
</taxonomicName>
. Holotype unsexed (
<specimenCode box="[198,368,1193,1217]" pageId="27" pageNumber="27">MNHNP 6218</specimenCode>
) from ‘‘ 
<collectingCountry box="[463,543,1193,1217]" name="Mexico" pageId="27" pageNumber="27">Mexico</collectingCountry>
’’ (
<emphasis box="[577,597,1192,1217]" italics="true" pageId="27" pageNumber="27">¼</emphasis>
Cerro San Felipe, 
<collectingRegion box="[190,271,1223,1247]" country="Mexico" name="Oaxaca" pageId="27" pageNumber="27">Oaxaca</collectingRegion>
, 
<collectingCountry box="[285,365,1222,1246]" name="Mexico" pageId="27" pageNumber="27">Mexico</collectingCountry>
[
<bibRefCitation author="Smith and Taylor" box="[380,645,1222,1246]" firstAuthor="Smith" journalOrPublisher="The University of Kansas Science Bulletin" pageId="27" pageNumber="27" pagination="313 - 339" part="33" refId="ref39066" refString="Smith, H. M., and E. H. Taylor. 1950. Type localities of Mexican reptiles and amphibians. The University of Kansas Science Bulletin 33: 313 - 339." title="Type localities of Mexican reptiles and amphibians" type="journal article" year="1950">
<location LSID="urn:lsid:plazi:treatment:03F287F8FF93FF98CA17C225F9E4BD16:8E846035FF93FF9ECA61C2BCF953BD9A" box="[380,578,1222,1246]" country="Mexico" name="Smith and Taylor" pageId="27" pageNumber="27" stateProvince="Oaxaca">Smith and Taylor</location>
1950
</bibRefCitation>
]). [Examined].
</materialsCitation>
</paragraph>
<paragraph blockId="27.[142,813,1164,1804]" pageId="27" pageNumber="27">
<treatmentCitationGroup pageId="27" pageNumber="27">
<taxonomicName authority="(Brocchi)" baseAuthorityName="Brocchi" box="[142,482,1252,1276]" class="Amphibia" family="Leptodactylidae" genus="Paludicola" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicana">
<emphasis box="[142,367,1252,1276]" italics="true" pageId="27" pageNumber="27">Paludicola mexicana</emphasis>
(Brocchi)
</taxonomicName>
: 
<bibRefCitation author="Boulenger, G. A." box="[493,718,1252,1276]" pageId="27" pageNumber="27" refId="ref35578" refString="Boulenger, G. A. 1882. Catalogue of the Batrachia, Salientia and Ecaudata in the Collection of the British Museum. Natural History Museum Publications, UK." type="book" year="1882">Boulenger 1882:237</bibRefCitation>
; 
<treatmentCitation author="Neiden, F." page="513" pageId="27" pageNumber="27" year="1923">
<bibRefCitation author="Neiden" firstAuthor="Neiden" journalOrPublisher="Das Tierreich" page="513" pageId="27" pageNumber="27" pagination="32 - 584" part="46" refId="ref38124" refString="Neiden, F. 1923. Anura. I. Subordo Aglossa und Phaneroglossa, Sectio 1. Arcifera. Das Tierreich 46: 32 - 584. [In German.]" title="Anura. I. Subordo Aglossa und Phaneroglossa, Sectio 1. Arcifera" type="journal article" year="1923">Neiden 1923:513</bibRefCitation>
</treatmentCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph blockId="27.[142,813,1164,1804]" box="[142,701,1310,1335]" pageId="27" pageNumber="27">
<treatmentCitationGroup box="[142,701,1310,1335]" pageId="27" pageNumber="27">
<taxonomicName authority="(Brocchi)" baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[142,500,1311,1335]" class="Amphibia" family="Leptodactylidae" genus="Pleurodema" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicana">
<emphasis box="[142,384,1311,1335]" italics="true" pageId="27" pageNumber="27">Pleurodema mexicana</emphasis>
(Brocchi)
</taxonomicName>
: 
<treatmentCitation author="Parker, H. W." box="[512,698,1310,1335]" page="475" pageId="27" pageNumber="27" year="1927">
<bibRefCitation author="Parker" box="[512,698,1310,1335]" firstAuthor="Parker" journalOrPublisher="Journal of Natural History" page="475" pageId="27" pageNumber="27" pagination="450 - 478" part="20" refId="ref38228" refString="Parker, H. W. 1927. A revision of the frogs of the genera Pseudopaludicola, Physalaemus, and Pleurodema. Journal of Natural History 20: 450 - 478." title="A revision of the frogs of the genera Pseudopaludicola, Physalaemus, and Pleurodema" type="journal article" year="1927">Parker 1927:475</bibRefCitation>
</treatmentCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph blockId="27.[142,813,1164,1804]" pageId="27" pageNumber="27">
<treatmentCitationGroup pageId="27" pageNumber="27">
<treatmentCitation author="Taylor, E. H." box="[142,630,1340,1364]" page="505" pageId="27" pageNumber="27" year="1940">
<taxonomicName authority="Taylor 1940: 505" authorityName="Taylor" authorityPageNumber="505" authorityYear="1940" box="[142,630,1340,1364]" class="Amphibia" family="Craugastoridae" genus="Microbatrachylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="oaxacae">
<emphasis box="[142,433,1340,1364]" italics="true" pageId="27" pageNumber="27">Microbatrachylus oaxacae</emphasis>
<bibRefCitation author="Taylor" box="[445,630,1340,1364]" firstAuthor="Taylor" journalOrPublisher="University of Kansas Science Bulletin" page="505" pageId="27" pageNumber="27" pagination="489 - 571" part="26" refId="ref39294" refString="Taylor, E. H. 1940. Herpetological miscellany No. I. University of Kansas Science Bulletin 26: 489 - 571." title="Herpetological miscellany No. I" type="journal article" year="1940">Taylor 1940:505</bibRefCitation>
</taxonomicName>
</treatmentCitation>
. 
<typeStatus box="[645,748,1340,1364]" pageId="27" pageNumber="27">Holotype</typeStatus>
male (
<collectionCode box="[198,283,1369,1393]" collectionName="USA, Illinois, Chicago, Field Museum of Natural History (also used by Finnish Museum of Natural History)" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34795" lsid="urn:lsid:biocol.org:col:34795" name="Field Museum of Natural History" pageId="27" pageNumber="27" type="Museum">FMNH</collectionCode>
100001) from ‘‘Cerro San Flipe, near 
<collectingRegion box="[727,808,1369,1393]" country="Mexico" name="Oaxaca" pageId="27" pageNumber="27">Oaxaca</collectingRegion>
, 
<collectingRegion box="[190,271,1399,1423]" country="Mexico" name="Oaxaca" pageId="27" pageNumber="27">Oaxaca</collectingRegion>
, 
<collectingCountry box="[285,369,1399,1423]" name="Mexico" pageId="27" pageNumber="27">Mexico</collectingCountry>
.’’ [Examined].
</treatmentCitationGroup>
</paragraph>
<paragraph blockId="27.[142,813,1164,1804]" pageId="27" pageNumber="27">
<treatmentCitationGroup pageId="27" pageNumber="27">
<treatmentCitation author="Taylor, E. H." box="[142,689,1428,1452]" page="87" pageId="27" pageNumber="27" year="1941">
<taxonomicName authority="Taylor 1941: 87" authorityName="Taylor" authorityPageNumber="87" authorityYear="1941" box="[142,689,1428,1452]" class="Amphibia" family="Craugastoridae" genus="Microbatrachylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="lineatissimus">
<emphasis box="[142,495,1428,1452]" italics="true" pageId="27" pageNumber="27">Microbatrachylus lineatissimus</emphasis>
<bibRefCitation author="Taylor" box="[512,689,1428,1452]" firstAuthor="Taylor" journalOrPublisher="Proceedings of the Biological Society of Washington" page="87" pageId="27" pageNumber="27" pagination="87 - 94" part="54" refId="ref39319" refString="Taylor, E. H. 1941. Some Mexican frogs. Proceedings of the Biological Society of Washington 54: 87 - 94." title="Some Mexican frogs" type="journal article" year="1941">Taylor 1941:87</bibRefCitation>
</taxonomicName>
</treatmentCitation>
. 
<typeStatus box="[710,813,1428,1452]" pageId="27" pageNumber="27">Holotype</typeStatus>
male (
<collectionCode box="[264,349,1457,1481]" collectionName="USA, Illinois, Chicago, Field Museum of Natural History (also used by Finnish Museum of Natural History)" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34795" lsid="urn:lsid:biocol.org:col:34795" name="Field Museum of Natural History" pageId="27" pageNumber="27" type="Museum">FMNH</collectionCode>
1000036) from ‘‘Cerro San Flipe, near 
<collectingRegion box="[190,271,1487,1511]" country="Mexico" name="Oaxaca" pageId="27" pageNumber="27">Oaxaca</collectingRegion>
, 
<collectingRegion box="[285,367,1487,1511]" country="Mexico" name="Oaxaca" pageId="27" pageNumber="27">Oaxaca</collectingRegion>
, 
<collectingCountry box="[379,463,1487,1511]" name="Mexico" pageId="27" pageNumber="27">Mexico</collectingCountry>
.’’ [Examined].
</treatmentCitationGroup>
</paragraph>
<paragraph blockId="27.[142,813,1164,1804]" pageId="27" pageNumber="27">
<treatmentCitationGroup pageId="27" pageNumber="27">
<treatmentCitation author="Davis" box="[142,813,1516,1540]" page="146" pageId="27" pageNumber="27" year="1957">
<taxonomicName authority="Davis and Dixon 1957: 146" authorityName="Davis and Dixon" authorityPageNumber="146" authorityYear="1957" box="[142,813,1516,1540]" class="Amphibia" family="Craugastoridae" genus="Microbatrachylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="fuscatus">
<emphasis box="[142,454,1516,1540]" italics="true" pageId="27" pageNumber="27">Microbatrachylus fuscatus</emphasis>
<bibRefCitation author="Davis, W. B. &amp; J. R. Dixon" box="[472,813,1516,1540]" pageId="27" pageNumber="27" pagination="145 - 147" refId="ref35963" refString="Davis, W. B., and J. R. Dixon. 1957. Notes on Mexican amphibians, with description of a new Microbatrachylus. Herpetologica 13: 145 - 147." type="journal article" year="1957">Davis and Dixon 1957:146.</bibRefCitation>
</taxonomicName>
</treatmentCitation>
<typeStatus box="[190,293,1545,1569]" pageId="27" pageNumber="27">Holotype</typeStatus>
female (
<collectionCode box="[399,480,1545,1569]" collectionName="TCWC" country="USA" httpUri="http://grbio.org/cool/dj6p-xxc3" name="Texas Cooperative Wildlife Collection" pageId="27" pageNumber="27">TCWC</collectionCode>
12171) from ‘‘ 
<quantity box="[660,757,1545,1569]" metricMagnitude="4" metricUnit="m" metricValue="3.218688" pageId="27" pageNumber="27" unit="mi" value="20.0">20 miles</quantity>
east of Tulancingo, 
<collectingRegion box="[360,450,1574,1598]" country="Mexico" name="Hidalgo" pageId="27" pageNumber="27">Hidalgo</collectingRegion>
, 
<collectingCountry box="[464,548,1574,1598]" name="Mexico" pageId="27" pageNumber="27">Mexico</collectingCountry>
.’’ [Not examined; neartopotypic specimen from 
<collectingRegion box="[490,578,1604,1628]" country="Mexico" name="Hidalgo" pageId="27" pageNumber="27">Hidalgo</collectingRegion>
examined (
<collectionCode box="[717,773,1604,1628]" collectionName="UTA" country="USA" httpUri="http://biocol.org/urn:lsid:biocol.org:col:34873" lsid="urn:lsid:biocol.org:col:34873" name="University of Texas at Arlington" pageId="27" pageNumber="27">UTA</collectionCode>
<collectionCode box="[786,812,1604,1628]" pageId="27" pageNumber="27">A-66138</collectionCode>
)].
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection pageId="27" pageNumber="27" type="reference_group">
<paragraph blockId="27.[142,813,1164,1804]" box="[142,707,1662,1686]" pageId="27" pageNumber="27">
<treatmentCitationGroup box="[142,707,1662,1686]" pageId="27" pageNumber="27">
<taxonomicName authority="(Taylor)" baseAuthorityName="Taylor" baseAuthorityYear="1940" box="[142,537,1662,1686]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="oaxacae">
<emphasis box="[142,435,1662,1686]" italics="true" pageId="27" pageNumber="27">Eleutherodactylus oaxacae</emphasis>
(Taylor)
</taxonomicName>
: 
<treatmentCitation author="Lynch, J. D." box="[548,704,1662,1686]" page="3" pageId="27" pageNumber="27" year="1965">
<bibRefCitation author="Lynch" box="[548,704,1662,1686]" firstAuthor="Lynch" journalOrPublisher="Chicago Academy of Science, USA" page="3" pageId="27" pageNumber="27" refId="ref37877" refString="Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA." title="A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae)" type="book" year="1965">Lynch 1965:3</bibRefCitation>
</treatmentCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph blockId="27.[142,813,1164,1804]" box="[142,763,1692,1716]" pageId="27" pageNumber="27">
<treatmentCitationGroup box="[142,763,1692,1716]" pageId="27" pageNumber="27">
<taxonomicName authority="(Taylor)" baseAuthorityName="Taylor" baseAuthorityYear="1941" box="[142,593,1692,1716]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="lineatissimus">
<emphasis box="[142,491,1692,1716]" italics="true" pageId="27" pageNumber="27">Eleutherodactylus lineatissimus</emphasis>
(Taylor)
</taxonomicName>
: 
<treatmentCitation author="Lynch, J. D." box="[605,760,1692,1716]" page="3" pageId="27" pageNumber="27" year="1965">
<bibRefCitation author="Lynch" box="[605,760,1692,1716]" firstAuthor="Lynch" journalOrPublisher="Chicago Academy of Science, USA" page="3" pageId="27" pageNumber="27" refId="ref37877" refString="Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA." title="A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae)" type="book" year="1965">Lynch 1965:3</bibRefCitation>
</treatmentCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph blockId="27.[142,813,1164,1804]" box="[142,787,1721,1745]" pageId="27" pageNumber="27">
<treatmentCitationGroup box="[142,787,1721,1745]" pageId="27" pageNumber="27">
<taxonomicName authority="(Brocchi)" baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[142,581,1721,1745]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[142,464,1721,1745]" italics="true" pageId="27" pageNumber="27">Eleutherodactylus mexicanus</emphasis>
(Brocchi)
</taxonomicName>
: 
<treatmentCitation author="Gorham" box="[593,782,1721,1745]" page="86" pageId="27" pageNumber="27" year="1966">
<bibRefCitation author="Gorham" box="[593,782,1721,1745]" firstAuthor="Gorham" page="86" pageId="27" pageNumber="27" year="1966">Gorham 1966:86</bibRefCitation>
</treatmentCitation>
.
</treatmentCitationGroup>
</paragraph>
<paragraph blockId="27.[142,813,1164,1804]" pageId="27" pageNumber="27">
<treatmentCitationGroup pageId="27" pageNumber="27">
<taxonomicName authority="(Brocchi)" baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[142,536,1751,1775]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[142,406,1751,1774]" italics="true" pageId="27" pageNumber="27">Craugastor mexicanus</emphasis>
(Brocchi)
</taxonomicName>
: 
<treatmentCitation author="Crawford and Smith" page="536" pageId="27" pageNumber="27" year="2005">
<bibRefCitation author="Crawford, A. J. &amp; E. N. Smith" pageId="27" pageNumber="27" pagination="536 - 555" refId="ref35831" refString="Crawford, A. J., and E. N. Smith. 2005. Cenozoic biogeography and evolution in direct-developing frogs of Central America (Leptodactylidae: Eleutherodactylus) as inferred from a phylogenetic analysis of nuclear and mitochondrial genes. Molecular Phylogenetics and Evolution 35: 536 - 555." type="journal article" year="2005">Crawford and Smith 2005:536</bibRefCitation>
.
</treatmentCitation>
</treatmentCitationGroup>
</paragraph>
</subSubSection>
<subSubSection pageId="27" pageNumber="27" type="diagnosis">
<paragraph blockId="27.[142,813,1830,2001]" lastBlockId="27.[844,1514,188,2001]" pageId="27" pageNumber="27">
<emphasis bold="true" box="[169,300,1830,1854]" pageId="27" pageNumber="27">Diagnosis.</emphasis>
—Based on 
<specimenCount box="[463,631,1831,1855]" pageId="27" pageNumber="27" type="generic">26 specimens</specimenCount>
. Aspecies of 
<taxonomicName authorityName="Cope" authorityYear="1862" box="[142,266,1860,1883]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="genus">
<emphasis box="[142,266,1860,1883]" italics="true" pageId="27" pageNumber="27">Craugastor</emphasis>
</taxonomicName>
distinguished by the following combination of characters: (1) large adult size (maximum SVL 
<emphasis box="[670,690,1888,1913]" italics="true" pageId="27" pageNumber="27">¼</emphasis>
<quantity box="[698,800,1889,1913]" metricMagnitude="-2" metricUnit="m" metricValue="4.05" pageId="27" pageNumber="27" unit="mm" value="40.5">40.5 mm</quantity>
); (2) full ossification of the skeleton in adults; (3) presence of posterolateral projection of frontoparietal (
<figureCitation box="[647,754,1948,1972]" captionStart="FIG" captionStartId="26.[142,156,926,945]" captionTargetBox="[118,1493,188,904]" captionTargetId="figure-570@26.[118,1494,187,905]" captionTargetPageId="26" captionText="FIG. 26.—Skulls of ‘Microbatrachylus lineatissimus’ (A, paratype, FMNH 104548), Craugastor mexicanus (B, UTA A-56558) and C. pygmaeus (C, UTA A- 64272) in dorsal and lateral views. Posterolateral projection of the frontoparietal on C. mexicanus (B) is indicated by an arrow and grey shading. A color version of this figure is available online." figureDoi="http://doi.org/10.5281/zenodo.6518639" httpUri="https://zenodo.org/record/6518639/files/figure.png" pageId="27" pageNumber="27">Fig. 26B</figureCitation>
); (4) presence of vomerine odontophores (in larger individuals); (5) presence or absence of raised tubercles on eyelids, &lt;4 smooth to round and only slightly protruding tubercles; (6) supratympanic fold developed; (7) face flank, labium barred with or without distinctive canthal stripe; (8) one (or two fused) postrital tubercles; (9) gular region peppered with melanocytes; (10) dorsal surface extremely variable, ranging from dark or light stripes, dark hourglass, dark or pale dorsal stripe of different widths, being unicolored dark or pale brown, to bird-dropping coloration (black peppered grayish dorsum with a dirty white medial streak and bright-white heels), all color morphologies have variable presence of interocular band and suprascapular or rump blotching or stripping; in most snout colored as rest of body but sometimes pale; (11) variable middorsal ridge; (12) dorsal skin smooth or tubercular and may have hourglass and/or vertebral and/or paravertebral ridges; (13) body flank unicolored pale, slightly darker anteriorly, or spotted posteriorly and or anteriorly, supratympanic dark coloration sometimes reaching posterior axillary area; smooth to shagreened; (14) inguinal gland present and axillary gland sometimes present in adults; (15) when leg adpressed to body, heel reaches snout tip or beyond; (16) outer tarsal ridge with 0–5 rounded tubercles; smooth or with thick but only slightly raised fringe; (17) finger and toe tips round and expanded (rarely slightly spatulate or barely expanded and somewhat pointed); (18) inner metatarsal tubercle larger than outer metatarsal tubercle.
</paragraph>
</subSubSection>
<subSubSection pageId="27" pageNumber="27" type="discussion">
<paragraph blockId="27.[844,1514,188,2001]" pageId="27" pageNumber="27">
<emphasis bold="true" box="[871,1037,980,1004]" pageId="27" pageNumber="27">Comparisons.</emphasis>
— 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[1064,1311,980,1004]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[1064,1311,980,1004]" italics="true" pageId="27" pageNumber="27">Craugastor mexicanus</emphasis>
</taxonomicName>
can be differentiated from 
<emphasis box="[959,1466,1009,1033]" italics="true" pageId="27" pageNumber="27">
<taxonomicName authorityName="Jameson &amp; Streicher &amp; Manuelli &amp; Head &amp; Smith" authorityYear="2022" box="[959,1149,1009,1033]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="candelariensis">C. candelariensis</taxonomicName>
, 
<taxonomicName authorityName="Jameson &amp; Streicher &amp; Manuelli &amp; Head &amp; Smith" authorityYear="2022" box="[1161,1274,1009,1033]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="cueyatl">C. cueyatl</taxonomicName>
, 
<taxonomicName baseAuthorityName="Taylor" baseAuthorityYear="1936" box="[1285,1462,1009,1033]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="hobartsmithi">C. hobartsmithi</taxonomicName>
,
</emphasis>
and 
<taxonomicName authorityName="Jameson &amp; Streicher &amp; Manuelli &amp; Head &amp; Smith" authorityYear="2022" box="[844,1016,1039,1063]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="portilloensis">
<emphasis box="[844,1016,1039,1063]" italics="true" pageId="27" pageNumber="27">C. portilloensis</emphasis>
</taxonomicName>
by the equal sizes of the inner and outer metatarsal tubercles (unequal sizes in 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[1275,1424,1068,1092]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[1275,1424,1068,1092]" italics="true" pageId="27" pageNumber="27">C. mexicanus</emphasis>
</taxonomicName>
). It can be differentiated from 
<taxonomicName authorityName="Jameson &amp; Streicher &amp; Manuelli &amp; Head &amp; Smith" authorityYear="2022" box="[1100,1234,1097,1121]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="bitonium">
<emphasis box="[1100,1234,1097,1121]" italics="true" pageId="27" pageNumber="27">C. bitonium</emphasis>
</taxonomicName>
and 
<taxonomicName authorityName="Ahumada-Carrillo et al. 2013:" authorityYear="1338" box="[1291,1435,1098,1121]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="pygmaeus">
<emphasis box="[1291,1435,1098,1121]" italics="true" pageId="27" pageNumber="27">C. pygmaeus</emphasis>
</taxonomicName>
by the absence of a posterolateral projection of the frontoparietal (present in 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[977,1126,1156,1180]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="42" phylum="Chordata" rank="species" species="mexicanus">
C. 
<emphasis box="[1012,1126,1157,1180]" italics="true" pageId="27" pageNumber="27">mexicanus</emphasis>
</taxonomicName>
). It can be differentiated from 
<taxonomicName authorityName="Jameson &amp; Streicher &amp; Manuelli &amp; Head &amp; Smith" authorityYear="2022" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="polaclavus">
<emphasis italics="true" pageId="27" pageNumber="27">C. polaclavus</emphasis>
</taxonomicName>
and 
<taxonomicName authorityName="Jameson &amp; Streicher &amp; Manuelli &amp; Head &amp; Smith" authorityYear="2022" box="[1032,1158,1185,1209]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="rubinus">
<emphasis box="[1032,1158,1185,1209]" italics="true" pageId="27" pageNumber="27">C. rubinus</emphasis>
</taxonomicName>
by the absence of vomerine odontophores (present in 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[1157,1312,1215,1238]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[1157,1312,1215,1238]" italics="true" pageId="27" pageNumber="27">C. mexicanus</emphasis>
</taxonomicName>
). It can be differentiated from 
<taxonomicName baseAuthorityName="Taylor" baseAuthorityYear="1942" box="[1045,1191,1244,1268]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="montanus">
<emphasis box="[1045,1191,1244,1268]" italics="true" pageId="27" pageNumber="27">C. montanus</emphasis>
</taxonomicName>
by the condition of supratympanic folds in adults; developed in 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[1286,1435,1274,1297]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[1286,1435,1274,1297]" italics="true" pageId="27" pageNumber="27">C. mexicanus</emphasis>
</taxonomicName>
versus moderate to poorly developed in 
<taxonomicName baseAuthorityName="Taylor" baseAuthorityYear="1942" box="[1239,1385,1303,1326]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="montanus">
<emphasis box="[1239,1385,1303,1326]" italics="true" pageId="27" pageNumber="27">C. montanus</emphasis>
</taxonomicName>
. It can be differentiated from 
<taxonomicName baseAuthorityName="Gunther" baseAuthorityYear="1900" box="[1062,1234,1332,1356]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="omiltemanus">
<emphasis box="[1062,1234,1332,1356]" italics="true" pageId="27" pageNumber="27">C. omiltemanus</emphasis>
</taxonomicName>
by ventral skin texture in life; smooth to granular in 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[1136,1283,1362,1385]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[1136,1283,1362,1385]" italics="true" pageId="27" pageNumber="27">C. mexicanus</emphasis>
</taxonomicName>
versus areolate in 
<taxonomicName baseAuthorityName="Gunther" baseAuthorityYear="1900" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="omiltemanus">
<emphasis italics="true" pageId="27" pageNumber="27">C. omiltemanus</emphasis>
</taxonomicName>
. 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[1004,1256,1391,1414]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[1004,1256,1391,1414]" italics="true" pageId="27" pageNumber="27">Craugastor mexicanus</emphasis>
</taxonomicName>
is most similar to 
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="saltator">
<emphasis italics="true" pageId="27" pageNumber="27">C. saltator</emphasis>
</taxonomicName>
. We were unable to identify any reliable morphological characters to differentiate 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[1256,1413,1450,1473]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[1256,1413,1450,1473]" italics="true" pageId="27" pageNumber="27">C. mexicanus</emphasis>
</taxonomicName>
and 
<emphasis italics="true" pageId="27" pageNumber="27">
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="saltator">C. saltator</taxonomicName>
;
</emphasis>
however, they have nonoverlapping geographic distributions, with 
<taxonomicName box="[1068,1192,1508,1532]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="27" pageNumber="27" phylum="Chordata" rank="species" species="saltator">
<emphasis box="[1068,1192,1508,1532]" italics="true" pageId="27" pageNumber="27">C. saltator</emphasis>
</taxonomicName>
only occurring in western 
<collectingRegion box="[844,947,1537,1561]" country="Mexico" name="Guerrero" pageId="27" pageNumber="27">Guerrero</collectingRegion>
(
<figureCitation box="[965,1035,1537,1561]" captionStart="FIG" captionStartId="11.[166,180,657,676]" captionTargetBox="[142,817,180,644]" captionTargetId="figure-472@11.[143,812,187,636]" captionTargetPageId="11" captionText="FIG. 6.—Distribution of Craugastor mexicanus, C. montanus, C. pygmaeus, and C. saltator in Mexico and Guatemala. Dots inside symbols indicate individuals used in the molecular analyses in this study. Relevant type localities are indicated by text and arrows." figureDoi="http://doi.org/10.5281/zenodo.6518676" httpUri="https://zenodo.org/record/6518676/files/figure.png" pageId="27" pageNumber="27">Fig. 6</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="27" pageNumber="27" type="description">
<paragraph blockId="27.[844,1514,188,2001]" pageId="27" pageNumber="27">
<emphasis bold="true" box="[871,1022,1566,1590]" pageId="27" pageNumber="27">Description.</emphasis>
—In previous literature, described as largebodied, long-legged with row of small tubercles on outer edge of the tarsus (
<bibRefCitation author="Taylor" box="[1072,1211,1625,1649]" firstAuthor="Taylor" journalOrPublisher="Proceedings of the Biological Society of Washington" pageId="27" pageNumber="27" pagination="87 - 94" part="54" refId="ref39319" refString="Taylor, E. H. 1941. Some Mexican frogs. Proceedings of the Biological Society of Washington 54: 87 - 94." title="Some Mexican frogs" type="journal article" year="1941">Taylor 1941</bibRefCitation>
); variable palmar tubercle arrangements (palmar tubercle divided, single, or evaginated; 
<bibRefCitation author="Lynch" box="[886,1019,1684,1708]" firstAuthor="Lynch" journalOrPublisher="Chicago Academy of Science, USA" pageId="27" pageNumber="27" refId="ref37877" refString="Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA." title="A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae)" type="book" year="1965">Lynch 1965</bibRefCitation>
); inner metatarsal tubercle larger than outer metatarsal tubercle; lack of tarsal tubercles (
<bibRefCitation author="Lynch" box="[1342,1480,1713,1737]" firstAuthor="Lynch" journalOrPublisher="Revista de La Academia Colombiana de Ciencias Exactas, Fisicas y Naturales" pageId="27" pageNumber="27" pagination="129 - 156" part="24" refId="ref37957" refString="Lynch, J. D. 2000. The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia). Revista de La Academia Colombiana de Ciencias Exactas, Fisicas y Naturales 24: 129 - 156." title="The relationships of an ensemble of Guatemalan and Mexican frogs (Eleutherodactylus: Leptodactylidae: Amphibia)" type="journal article" year="2000">Lynch 2000</bibRefCitation>
).
</paragraph>
<paragraph blockId="27.[844,1514,188,2001]" pageId="27" pageNumber="27">
<typeStatus box="[876,979,1743,1767]" pageId="27" pageNumber="27">Holotype</typeStatus>
(MNHNP 6318) is large (~ 
<quantity box="[1292,1369,1743,1767]" metricMagnitude="-2" metricUnit="m" metricValue="4.0" pageId="27" pageNumber="27" unit="mm" value="40.0">40 mm</quantity>
SVL); owing to poor preservation, columella of 
<typeStatus box="[1300,1400,1772,1796]" pageId="27" pageNumber="27">holotype</typeStatus>
partially ruptured the tympana on both sides (
<figureCitation box="[1257,1340,1801,1825]" captionStart="FIG" captionStartId="3.[166,180,1153,1172]" captionTargetBox="[143,1516,188,1131]" captionTargetId="figure-192@3.[142,1518,187,1132]" captionTargetPageId="3" captionText="FIG. 1.—Male holotype of Eleutherodactylus hobartsmithi Taylor (A, FMNH 100114, SVL ¼ 14.4 mm); female holotype of Leiuperus mexicanus Brocchi (B, MNHNP 6318, SVL ~ 40 mm); male holotype of Microbatrachylus oaxacae Taylor (C, FMNH 100001, SVL ¼ 18.1 mm); male holotype of Microbatrachylus lineatissimus Taylor (D, FMNH 100036, SVL ¼ 20.0 mm); female holotype of Microbatrachylus montanus Taylor (E, USNM 115507, SVL ¼ 27.0 mm); female holotype of Eleutherodactylus pygmaeus Taylor (F, UIMNH 16125, SVL ¼ 17.0 mm); female holotype of Microbatrachylus albolabris Taylor (G, FMNH 100071, SVL ¼ 16.5 mm); male holotype of Microbatrachylus minimus Taylor (H, FMNH 100323, SVL ¼ 15.0 mm); female holotype of Microbatrachylus imitator Taylor (I, USNM 115508, SVL ¼ 14.2 mm); male lectotype of Syrrhaphus omiltemanus Günther (J, BMNH 1947.2.16.62, SVL ¼ 19.8 mm); female lectotype of Hylodes calcitrans Günther (K, BMNH 1947.2.16.47, SVL ¼ 34.9 mm); female holotype of Eleutherodactylus saltator Taylor (L, FMNH 100166, SVL ¼ 44.0 mm). Symbols correspond to current taxonomic assignments (square ¼ C. hobartsmithi, triangle ¼ C. mexicanus, crossedsquare ¼ C. montanus, pentagon ¼ C. omiltemanus, circle ¼ C. pygmaeus, rotated-triangle ¼ C. saltator). A color version of this figure is available online." figureDoi="http://doi.org/10.5281/zenodo.6518589" httpUri="https://zenodo.org/record/6518589/files/figure.png" pageId="27" pageNumber="27">Fig. 1B</figureCitation>
); finger lengths III&gt; IV&gt; II&gt; I; toe length IV&gt; V 
<emphasis box="[1282,1302,1829,1854]" italics="true" pageId="27" pageNumber="27">¼</emphasis>
III&gt; II&gt; I.
</paragraph>
</subSubSection>
<subSubSection lastPageId="28" lastPageNumber="28" pageId="27" pageNumber="27" type="distribution">
<paragraph blockId="27.[844,1514,188,2001]" lastBlockId="28.[118,790,1244,2001]" lastPageId="28" lastPageNumber="28" pageId="27" pageNumber="27">
<emphasis bold="true" box="[871,1028,1860,1884]" pageId="27" pageNumber="27">Distribution.</emphasis>
—This species is widespread throughout eastern 
<collectingCountry box="[933,1013,1889,1913]" name="Mexico" pageId="27" pageNumber="27">Mexico</collectingCountry>
in high-elevation habitats (
<quantity box="[1322,1476,1889,1913]" metricMagnitude="3" metricUnit="m" metricValue="2.127" metricValueMax="2.7" metricValueMin="1.554" pageId="27" pageNumber="27" unit="m" value="2127.0" valueMax="2700.0" valueMin="1554.0">1554–2700 m</quantity>
) of 
<collectingRegion box="[844,926,1919,1943]" country="Mexico" name="Oaxaca" pageId="27" pageNumber="27">Oaxaca</collectingRegion>
, 
<collectingRegion box="[947,1026,1919,1943]" country="Mexico" name="Puebla" pageId="27" pageNumber="27">Puebla</collectingRegion>
, 
<collectingRegion box="[1046,1136,1919,1943]" country="Mexico" name="Hidalgo" pageId="27" pageNumber="27">Hidalgo</collectingRegion>
, and 
<collectingRegion box="[1216,1315,1919,1942]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="27" pageNumber="27">Veracruz</collectingRegion>
(
<figureCitation box="[1340,1417,1919,1943]" captionStart="FIG" captionStartId="11.[166,180,657,676]" captionTargetBox="[142,817,180,644]" captionTargetId="figure-472@11.[143,812,187,636]" captionTargetPageId="11" captionText="FIG. 6.—Distribution of Craugastor mexicanus, C. montanus, C. pygmaeus, and C. saltator in Mexico and Guatemala. Dots inside symbols indicate individuals used in the molecular analyses in this study. Relevant type localities are indicated by text and arrows." figureDoi="http://doi.org/10.5281/zenodo.6518676" httpUri="https://zenodo.org/record/6518676/files/figure.png" pageId="27" pageNumber="27">Fig. 6</figureCitation>
). These habitats span the Sierra Madre Oriental and Sierra Madre del Sur. 
<bibRefCitation author="Canseco Marquez, L. &amp; M. G. Gutierrez Mayen" box="[953,1513,1977,2001]" pageId="27" pageNumber="27" refId="ref35670" refString="Canseco Marquez, L., and M. G. Gutierrez Mayen. 2010. Anfibios y Reptiles del Valle Tehuacan-Cuicatlan. Comision Nacional para el Conocimiento y Uso de la Biodiversidad (CONABIO), Mexico." type="book" year="2010">Canseco Márquez and Gutiérrez Mayén (2010)</bibRefCitation>
report that 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[243,390,1245,1268]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[243,390,1245,1268]" italics="true" pageId="28" pageNumber="28">C. mexicanus</emphasis>
</taxonomicName>
occurs in the forests adjacent to the Tehuacán-Cuicatlán Valley in 
<collectingRegion box="[453,529,1274,1298]" country="Mexico" name="Puebla" pageId="28" pageNumber="28">Puebla</collectingRegion>
and 
<collectingRegion box="[586,668,1274,1298]" country="Mexico" name="Oaxaca" pageId="28" pageNumber="28">Oaxaca</collectingRegion>
. It is likely this species occurs in 
<collectingRegion box="[375,481,1303,1327]" country="Mexico" name="Guerrero" pageId="28" pageNumber="28">Guerrero</collectingRegion>
; however, most specimens resembling 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[252,403,1333,1356]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[252,403,1333,1356]" italics="true" pageId="28" pageNumber="28">C. mexicanus</emphasis>
</taxonomicName>
we examined from 
<collectingRegion box="[638,741,1333,1357]" country="Mexico" name="Guerrero" pageId="28" pageNumber="28">Guerrero</collectingRegion>
are referrable to 
<taxonomicName box="[267,385,1362,1386]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="saltator">
<emphasis box="[267,385,1362,1386]" italics="true" pageId="28" pageNumber="28">C. saltator</emphasis>
</taxonomicName>
.
</paragraph>
</subSubSection>
<subSection box="[27,778,2060,2078]" pageId="27" pageNumber="27" type="multiple">
<paragraph blockId="27.[27,876,2060,2097]" box="[27,778,2060,2078]" pageId="27" pageNumber="27">
Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 
<date box="[670,778,2061,2078]" pageId="27" pageNumber="27" value="2022-05-04">04 May 2022</date>
</paragraph>
</subSection>
<caption ID-DOI="http://doi.org/10.5281/zenodo.6518641" ID-Zenodo-Dep="6518641" httpUri="https://zenodo.org/record/6518641/files/figure.png" pageId="28" pageNumber="28" startId="28.[142,156,990,1009]" targetBox="[119,1494,189,968]" targetPageId="28">
<paragraph blockId="28.[118,1491,989,1200]" pageId="28" pageNumber="28">
FIG. 27.—Examples of intraspecific polymorphism in 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[626,824,990,1009]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[626,824,990,1009]" italics="true" pageId="28" pageNumber="28">Craugastor mexicanus</emphasis>
</taxonomicName>
from Mexico; female (A, UTA A-56558, SVL 
<emphasis box="[1245,1261,989,1009]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
37.3 mm) from Putla de Guerrero municipality; Oaxaca, female (B, UTA A-64415, SVL 
<emphasis box="[669,685,1013,1033]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
28.3 mm) from Carretera Sola de Vega–Juquila, Oaxaca; subadult (C, UTA A-64271, SVL 
<emphasis box="[1473,1489,1013,1033]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
11.9 mm) from Carretera Sola de Vega–Juquila, Oaxaca; subadult and adult (D, UTA A-64258, SVL 
<emphasis box="[1002,1018,1037,1057]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
13.6 mm and MZFC-HE-35584, SVL 
<emphasis box="[1361,1377,1037,1057]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
unavailable) both from 8.1 mi south of Sola de Vega, Oaxaca; adult (E, UTA A-66107, SVL 
<emphasis box="[827,843,1061,1081]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
unavailable) from Carretera Sola de Vega–Juquila, Oaxaca; subadult (F, UTA A-64260, SVL 
<emphasis box="[301,317,1085,1105]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
16.6 mm) from Carretera Sola de Vega–Juquila, Oaxaca; female (G, UTA A-64413, SVL 
<emphasis box="[1124,1140,1085,1105]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
23.5) from Sierra Miahuatlán, Oaxaca; female (H, UTA A-56579, SVL 
<emphasis box="[399,415,1108,1128]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
21.3 mm) from Sierra Negra, Puebla; female (I, UTA A-56559, SVL 
<emphasis box="[1032,1048,1108,1128]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
35.4 mm) from Sierra Mazateca, Puerto Soledad, Oaxaca; subadult (J, UTA A-64257, SVL 
<emphasis box="[480,496,1132,1152]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
17.6 mm) from Sierra Mixe, Carretera Ayutla–Zacatepec, Oaxaca; subadult (K, UTA A-64259, SVL 
<emphasis box="[1382,1398,1132,1152]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
10.9 mm) from 6.1 mi south of San Miguel Suchixtepec, Oaxaca; subadult (L, UTA A-66138, SVL 
<emphasis box="[917,933,1156,1176]" italics="true" pageId="28" pageNumber="28">¼</emphasis>
unavailable) from 20 mi east of Tulancingo, Hidalgo. Acolor version of this figure is available online.
</paragraph>
</caption>
<subSubSection lastPageId="29" lastPageNumber="29" pageId="28" pageNumber="28" type="discussion">
<paragraph blockId="28.[118,790,1244,2001]" pageId="28" pageNumber="28">
<emphasis bold="true" box="[145,321,1391,1415]" pageId="28" pageNumber="28">Phylogenetics.</emphasis>
—The concatenated data set placed 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="mexicanus">
<emphasis italics="true" pageId="28" pageNumber="28">C. mexicanus</emphasis>
</taxonomicName>
as the sister taxon to a clade of 
<emphasis italics="true" pageId="28" pageNumber="28">
<taxonomicName baseAuthorityName="Gunther" baseAuthorityYear="1900" box="[591,765,1420,1444]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="omiltemanus">C. omiltemanus</taxonomicName>
þ 
<taxonomicName box="[118,236,1450,1474]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="saltator">C. saltator</taxonomicName>
</emphasis>
with high support in the BAYES analysis (0.98) but moderate support in the ML analysis (75; 
<figureCitation box="[648,716,1479,1503]" captionStart="FIG" captionStartId="8.[142,156,1857,1876]" captionTargetBox="[295,1312,189,1835]" captionTargetId="figure-1@8.[294,1312,187,1835]" captionTargetPageId="8" captionText="FIG. 3.—Concatenated Maximum Likelihood analyses of two mitochondrial (12S and 16S) and two nuclear (RAG1 and TYR) gene fragments sequenced from the Craugastor mexicanus series (total of 2230 base pairs). Node values correspond to bootstrap support from Maximum Likelihood analysis and posterior probabilities from a Bayesian analysis of the same data set, respectively. Support values are not reported for nodes that had,50/0.50. NS indicates no support in the Bayesian analysis for a depicted relationship. Locality abbreviations are as follows: COL ¼ Colima, HID ¼ Hidalgo, GRO ¼ Guerrero, JAL ¼ Jalisco, MEX ¼ Estado de México, OAX ¼ Oaxaca, PUE ¼ Puebla, and VER ¼ Veracruz." figureDoi="http://doi.org/10.5281/zenodo.6518593" httpUri="https://zenodo.org/record/6518593/files/figure.png" pageId="28" pageNumber="28">Fig. 3</figureCitation>
). This appears to be a relationship supported exclusively by the mtDNA data set because separate analysis of the nDNA markers did not confidently infer a sister taxon of 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="mexicanus">
<emphasis italics="true" pageId="28" pageNumber="28">C. mexicanus</emphasis>
</taxonomicName>
(
<figureCitation box="[260,347,1596,1620]" captionStart="FIG" captionStartId="9.[166,180,1708,1727]" captionTargetBox="[142,1514,192,1686]" captionTargetId="figure-77@9.[142,1518,189,1686]" captionTargetPageId="9" captionText="FIG. 4.—Maximum Likelihood analyses of mitochondrial DNA (mtDNA) markers sequenced from the Craugastor mexicanus series (total of 1047 base pairs). Black circles indicate nodes receiving bootstrap support values.90 from the Maximum Likelihood analysis and 0.90 posterior probabilities in corresponding Bayesian analyses. Where posterior probabilities were.0.90 but Maximum Likelihood bootstrap support was,90, a number indicates the bootstrap support. NS indicates no support in the Bayesian analysis for a depicted relationship. Locality abbreviations are as follows: COL ¼ Colima, HID ¼ Hidalgo, GRO ¼ Guerrero, JAL ¼ Jalisco, MEX ¼ Estado de México, OAX ¼ Oaxaca, PUE ¼ Puebla, and VER ¼ Veracruz." figureDoi="http://doi.org/10.5281/zenodo.6518597" httpUri="https://zenodo.org/record/6518597/files/figure.png" pageId="28" pageNumber="28">Figs. 4</figureCitation>
and 
<figureCitation box="[425,440,1596,1620]" captionStart="FIG" captionStartId="10.[142,156,1604,1623]" captionTargetBox="[118,1493,187,1579]" captionTargetId="figure-138@10.[118,1494,187,1582]" captionTargetPageId="10" captionText="FIG. 5.—Maximum Likelihood analysis of nuclear DNA (nDNA) markers sequenced from the Craugastor mexicanus series (total of 1183 base pairs). Black circles indicate nodes receiving bootstrap support values.90 from the Maximum Likelihood analysis and 0.90 posterior probabilities in corresponding Bayesian analyses. Where posterior probabilities were.0.90 but Maximum Likelihood bootstrap support was,90, a number indicates the bootstrap support. NS indicates no support in the Bayesian analysis for a depicted relationship. Locality abbreviations are as follows: COL ¼ Colima, GRO ¼ Guerrero, JAL ¼ Jalisco, MEX ¼ Estado de México, OAX ¼ Oaxaca, PUE ¼ Puebla, and VER ¼ Veracruz." figureDoi="http://doi.org/10.5281/zenodo.6518599" httpUri="https://zenodo.org/record/6518599/files/figure.png" pageId="28" pageNumber="28">5</figureCitation>
). Although BAYES analyses strongly supported the monophyly of 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[620,788,1626,1649]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[620,788,1626,1649]" italics="true" pageId="28" pageNumber="28">C. mexicanus</emphasis>
</taxonomicName>
(&gt; 0.90), the ML analyses recovered variable support ranging from 79 (concatenated analysis) to 54 (nDNA-only analysis). In terms of genetic distances (
<tableCitation box="[631,731,1713,1738]" captionStart="TABLE" captionStartId="7.[169,182,188,207]" captionTargetPageId="7" captionText="TABLE 4.—Genetic P-distances derived from the concatenated alignment 2230 base pairs of 12S and 16S (mtDNA) þ RAG1 and TYR (nDNA) for species contained within the Craugastor mexicanus series. Bolded distances on diagonal are within species P-distances (if more than a single individual was used to estimate mean genetic distance)." httpUri="http://table.plazi.org/id/DF246666FF8FFF82CBB4C6C6FAA2B9BB" pageId="28" pageNumber="28" tableUuid="DF246666FF8FFF82CBB4C6C6FAA2B9BB">Table 4</tableCitation>
), 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="mexicanus">
<emphasis italics="true" pageId="28" pageNumber="28">C. mexicanus</emphasis>
</taxonomicName>
was most similar to 
<taxonomicName baseAuthorityName="Gunther" baseAuthorityYear="1900" box="[508,691,1743,1767]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="omiltemanus">
<emphasis box="[508,691,1743,1767]" italics="true" pageId="28" pageNumber="28">C. omiltemanus</emphasis>
</taxonomicName>
(4.9%), followed by similarity to 
<taxonomicName box="[397,515,1772,1796]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="saltator">
<emphasis box="[397,515,1772,1796]" italics="true" pageId="28" pageNumber="28">C. saltator</emphasis>
</taxonomicName>
(5.1%).
</paragraph>
<paragraph blockId="28.[118,790,1244,2001]" lastBlockId="28.[820,1490,1244,2001]" pageId="28" pageNumber="28">
<emphasis bold="true" box="[145,424,1801,1825]" pageId="28" pageNumber="28">Intraspecific variation.</emphasis>
—We examined over 
<specimenCount pageId="28" pageNumber="28" type="generic">220 specimens</specimenCount>
of 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[223,375,1831,1855]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[223,375,1831,1855]" italics="true" pageId="28" pageNumber="28">C. mexicanus</emphasis>
</taxonomicName>
for this revision (Appendix I), and briefly provide some patterns of intraspecific variation that we observed. Many populations have substantial colorpattern polymorphism (
<figureCitation box="[383,465,1919,1943]" captionStart="FIG" captionStartId="28.[142,156,990,1009]" captionTargetBox="[119,1494,189,968]" captionTargetId="figure-477@28.[118,1495,188,968]" captionTargetPageId="28" captionText="FIG. 27.—Examples of intraspecific polymorphism in Craugastor mexicanus from Mexico; female (A, UTA A-56558, SVL ¼ 37.3 mm) from Putla de Guerrero municipality; Oaxaca, female (B, UTA A-64415, SVL ¼ 28.3 mm) from Carretera Sola de Vega–Juquila, Oaxaca; subadult (C, UTA A-64271, SVL ¼ 11.9 mm) from Carretera Sola de Vega–Juquila, Oaxaca; subadult and adult (D, UTA A-64258, SVL ¼ 13.6 mm and MZFC-HE-35584, SVL ¼ unavailable) both from 8.1 mi south of Sola de Vega, Oaxaca; adult (E, UTA A-66107, SVL ¼ unavailable) from Carretera Sola de Vega–Juquila, Oaxaca; subadult (F, UTA A-64260, SVL ¼ 16.6 mm) from Carretera Sola de Vega–Juquila, Oaxaca; female (G, UTA A-64413, SVL ¼ 23.5) from Sierra Miahuatlán, Oaxaca; female (H, UTA A-56579, SVL ¼ 21.3 mm) from Sierra Negra, Puebla; female (I, UTA A-56559, SVL ¼ 35.4 mm) from Sierra Mazateca, Puerto Soledad, Oaxaca; subadult (J, UTA A-64257, SVL ¼ 17.6 mm) from Sierra Mixe, Carretera Ayutla–Zacatepec, Oaxaca; subadult (K, UTA A-64259, SVL ¼ 10.9 mm) from 6.1 mi south of San Miguel Suchixtepec, Oaxaca; subadult (L, UTA A-66138, SVL ¼ unavailable) from 20 mi east of Tulancingo, Hidalgo. A color version of this figure is available online." figureDoi="http://doi.org/10.5281/zenodo.6518641" httpUri="https://zenodo.org/record/6518641/files/figure.png" pageId="28" pageNumber="28">Fig. 27</figureCitation>
), similar to what is observed in the 
<taxonomicName baseAuthorityName="Cope" baseAuthorityYear="1867" box="[189,319,1948,1972]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="28" pageNumber="28" phylum="Chordata" rank="species" species="rhodopis">
<emphasis box="[189,319,1948,1972]" italics="true" pageId="28" pageNumber="28">C. rhodopis</emphasis>
</taxonomicName>
series (
<bibRefCitation author="Lynch" box="[404,538,1948,1972]" firstAuthor="Lynch" journalOrPublisher="Systematic Biology" pageId="28" pageNumber="28" pagination="18 - 23" part="15" refId="ref37904" refString="Lynch, J. D. 1966. Multiple morphotypy and parallel polymorphism in some neotropical frogs. Systematic Biology 15: 18 - 23." title="Multiple morphotypy and parallel polymorphism in some neotropical frogs" type="journal article" year="1966">Lynch 1966</bibRefCitation>
; 
<bibRefCitation author="Streicher" box="[549,777,1948,1972]" etAl="et al." firstAuthor="Streicher" journalOrPublisher="Systematics and Biodiversity" pageId="28" pageNumber="28" pagination="1 - 22" part="12" refId="ref39200" refString="Streicher, J. W., U. O. Garcia-Vazquez, P. Ponce-Campos, O. Flores-Villela, J. A. Campbell, and E. N. Smith. 2014. Evolutionary relationships amongst polymorphic direct-developing frogs in the Craugastor rhodopis Species Group (Anura: Craugastoridae). Systematics and Biodiversity 12: 1 - 22." title="Evolutionary relationships amongst polymorphic direct-developing frogs in the Craugastor rhodopis Species Group (Anura: Craugastoridae)" type="journal article" year="2014">Streicher et al. 2014</bibRefCitation>
). In some northern populations (
<collectingRegion box="[489,578,1977,2001]" country="Mexico" name="Hidalgo" pageId="28" pageNumber="28">Hidalgo</collectingRegion>
and 
<collectingRegion box="[647,727,1977,2001]" country="Mexico" name="Puebla" pageId="28" pageNumber="28">Puebla</collectingRegion>
), the canthal mask is broken into a black spot posterior to the tympanum (often with a thin yet distinctive white margin). Specimens throughout 
<collectingRegion box="[1080,1159,1303,1327]" country="Mexico" name="Oaxaca" pageId="28" pageNumber="28">Oaxaca</collectingRegion>
varied in whether they had a single or divided palmar tubercle (as reported by 
<bibRefCitation author="Lynch" firstAuthor="Lynch" journalOrPublisher="Chicago Academy of Science, USA" pageId="28" pageNumber="28" refId="ref37877" refString="Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA." title="A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae)" type="book" year="1965">Lynch 1965</bibRefCitation>
, see his 
<figureCitation box="[966,1031,1362,1386]" captionStart="FIG" captionStartId="6.[142,156,1036,1055]" captionTargetBox="[118,1493,187,1014]" captionTargetId="figure-444@6.[118,1494,187,1014]" captionTargetPageId="6" captionText="FIG. 2.—Skull of Craugastor mexicanus (USNM 47905) in anterior, dorsal, lateral, and ventral views showing placements of landmarks. Landmarks are (1) anterior midline tip premaxilla, (2) maxilla–premaxilla suture, (3) anterior tip nasal, (4) anterior tip frontoparietal, (5) posterior lateral tip frontoparietal, (6) posterior dorsal tip squamosal, (7) anterior frontoparietal prootic suture, (8) lateral tip nasal, (9) squamosal maxillary suture, (10) posterior tip maxilla, (11) anterior tip parasphenoid (12) posterior tip parasphenoid, (13) lateral tip parasphenoid, and, (14) pterygoid maxillary suture. A color version of this figure is available online." figureDoi="http://doi.org/10.5281/zenodo.6518591" httpUri="https://zenodo.org/record/6518591/files/figure.png" pageId="28" pageNumber="28">Fig. 2</figureCitation>
); in 
<specimenCount box="[1079,1229,1362,1386]" pageId="28" pageNumber="28" type="generic">six specimens</specimenCount>
we examined there was asymmetry with a single palmar tubercle on one hand and divided on the other.
</paragraph>
<paragraph blockId="28.[820,1490,1244,2001]" pageId="28" pageNumber="28">Patterns of dorsal ridging and coloration often coincide, with observations of the following morphs: (1) straight raised ridges that are tubercular and darker in coloration than the rest of the dorsum; (2) straight raised dorsal ridges that are tubercular but of the same color as the rest of the dorsum; (3) straight lines of color that are dark but with no tubercular raised ridges; (4) ridges that are not straight but form an hourglass shape and are tubercular; (5) ridges that are not straight but form an hourglass shape and are not tubercular; (6) scattered tubercles on the dorsum, which are sometimes dark and sometimes same color as the background; (7) pale or dark unicolored dorsum; (8) a unicolored dorsum with a pale median stripe that can be ridged or smooth; (9) a wide dark band on the dorsum, which usually co-occurs with a pale upper labium; and, (10) white hills of coloration and a peppered grayish dorsum sometimes with a diffused lighter cream color, a pattern that may mimic bird droppings (appearing superficially as a smear of uric acid and digestive waste).</paragraph>
<subSection box="[27,778,2060,2078]" pageId="28" pageNumber="28" type="multiple">
<paragraph blockId="28.[27,876,2060,2097]" box="[27,778,2060,2078]" pageId="28" pageNumber="28">
Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 
<date box="[670,778,2061,2078]" pageId="28" pageNumber="28" value="2022-05-04">04 May 2022</date>
</paragraph>
</subSection>
<paragraph blockId="29.[142,813,188,1107]" pageId="29" pageNumber="29">
<emphasis bold="true" box="[169,285,188,212]" pageId="29" pageNumber="29">Remarks.</emphasis>
—The skull of 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[450,597,189,212]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[450,597,189,212]" italics="true" pageId="29" pageNumber="29">C. mexicanus</emphasis>
</taxonomicName>
is similar to that of 
<taxonomicName baseAuthorityName="Gunther" baseAuthorityYear="1900" box="[142,316,218,242]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="omiltemanus">
<emphasis box="[142,316,218,242]" italics="true" pageId="29" pageNumber="29">C. omiltemanus</emphasis>
</taxonomicName>
and 
<emphasis box="[373,495,218,242]" italics="true" pageId="29" pageNumber="29">
<taxonomicName box="[373,491,218,242]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="saltator">C. saltator</taxonomicName>
,
</emphasis>
with more anteriorly placed anterior suture of the frontoparietal and prootic than in other species. We noticed that the ‘‘ 
<taxonomicName baseAuthorityName="Taylor" baseAuthorityYear="1941" box="[478,656,277,301]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="lineatissimus">
<emphasis box="[478,656,277,301]" italics="true" pageId="29" pageNumber="29">M. lineatissimus</emphasis>
</taxonomicName>
’’ morphotype (raised and parallel ridges on the dorsum) occurs widely throughout the range of 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[442,595,338,361]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[442,595,338,361]" italics="true" pageId="29" pageNumber="29">C. mexicanus</emphasis>
</taxonomicName>
. Interestingly, the examined 
<typeStatus box="[257,352,367,391]" pageId="29" pageNumber="29">paratype</typeStatus>
of ‘‘ 
<taxonomicName baseAuthorityName="Taylor" baseAuthorityYear="1941" box="[406,586,367,391]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="lineatissimus">
<emphasis box="[406,586,367,391]" italics="true" pageId="29" pageNumber="29">M. lineatissimus</emphasis>
</taxonomicName>
’’ has a unique skull shape for 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[259,409,397,420]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[259,409,397,420]" italics="true" pageId="29" pageNumber="29">C. mexicanus</emphasis>
</taxonomicName>
. The shape is similar to that of 
<emphasis italics="true" pageId="29" pageNumber="29">
<taxonomicName baseAuthorityName="Taylor" baseAuthorityYear="1936" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="hobartsmithi">C. hobartsmithi</taxonomicName>
, 
<taxonomicName baseAuthorityName="Taylor" baseAuthorityYear="1942" box="[301,446,427,450]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="montanus">C. montanus</taxonomicName>
,
</emphasis>
and 
<emphasis box="[512,664,427,450]" italics="true" pageId="29" pageNumber="29">
<taxonomicName authorityName="Ahumada-Carrillo et al. 2013:" authorityYear="1338" box="[512,660,427,450]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="pygmaeus">C. pygmaeus</taxonomicName>
,
</emphasis>
with a more posteriorly placed anterior suture of the frontoparietal and prootic than in other species, coupled with a narrower back of the skull than in any other species (
<figureCitation box="[562,662,516,540]" captionStart="FIG" captionStartId="28.[142,156,990,1009]" captionTargetBox="[119,1494,189,968]" captionTargetId="figure-477@28.[118,1495,188,968]" captionTargetPageId="28" captionText="FIG. 27.—Examples of intraspecific polymorphism in Craugastor mexicanus from Mexico; female (A, UTA A-56558, SVL ¼ 37.3 mm) from Putla de Guerrero municipality; Oaxaca, female (B, UTA A-64415, SVL ¼ 28.3 mm) from Carretera Sola de Vega–Juquila, Oaxaca; subadult (C, UTA A-64271, SVL ¼ 11.9 mm) from Carretera Sola de Vega–Juquila, Oaxaca; subadult and adult (D, UTA A-64258, SVL ¼ 13.6 mm and MZFC-HE-35584, SVL ¼ unavailable) both from 8.1 mi south of Sola de Vega, Oaxaca; adult (E, UTA A-66107, SVL ¼ unavailable) from Carretera Sola de Vega–Juquila, Oaxaca; subadult (F, UTA A-64260, SVL ¼ 16.6 mm) from Carretera Sola de Vega–Juquila, Oaxaca; female (G, UTA A-64413, SVL ¼ 23.5) from Sierra Miahuatlán, Oaxaca; female (H, UTA A-56579, SVL ¼ 21.3 mm) from Sierra Negra, Puebla; female (I, UTA A-56559, SVL ¼ 35.4 mm) from Sierra Mazateca, Puerto Soledad, Oaxaca; subadult (J, UTA A-64257, SVL ¼ 17.6 mm) from Sierra Mixe, Carretera Ayutla–Zacatepec, Oaxaca; subadult (K, UTA A-64259, SVL ¼ 10.9 mm) from 6.1 mi south of San Miguel Suchixtepec, Oaxaca; subadult (L, UTA A-66138, SVL ¼ unavailable) from 20 mi east of Tulancingo, Hidalgo. A color version of this figure is available online." figureDoi="http://doi.org/10.5281/zenodo.6518641" httpUri="https://zenodo.org/record/6518641/files/figure.png" pageId="29" pageNumber="29">Fig. 27A</figureCitation>
). This unique condition likely explains the specimen as an outlier in several of our statistical analyses (
<figureCitation box="[434,512,575,600]" captionStart="FIG" captionStartId="12.[142,156,905,924]" captionTargetBox="[119,1494,187,883]" captionTargetId="figure-214@12.[118,1495,187,884]" captionTargetPageId="12" captionText="FIG. 9.—Body size and shape variation within the Craugastor mexicanus series as depicted by principal components analysis of 15 linear measurements. Principal component scores for the first two components (left) are depicted along with their residuals (right). Large-bodied species are those with an adult body size.20 mm SVL (C. mexicanus, C. montanus, C. omiltemanus, C. saltator), and small-bodied species are those with adult body sizes,20 mm (C. bitonium, C. candelariensis, C. cueyatl, C. hobartsmithi, C. polacalvus, C. portilloensis, C. pygmaeus, and C. rubinus)." figureDoi="http://doi.org/10.5281/zenodo.6518605" httpUri="https://zenodo.org/record/6518605/files/figure.png" pageId="29" pageNumber="29">Figs. 9</figureCitation>
, 
<figureCitation box="[525,553,576,600]" captionStart="FIG" captionStartId="12.[844,858,1933,1952]" captionTargetBox="[820,1488,1418,1911]" captionTargetId="figure-365@12.[820,1489,1417,1911]" captionTargetPageId="12" captionText="FIG. 11.—Ossification levels within the Craugastor mexicanus series compared with body size (estimated as log centroid size). Ossification scores are based on the six-stage ontogenetic sequence described in Table 3." figureDoi="http://doi.org/10.5281/zenodo.6518609" httpUri="https://zenodo.org/record/6518609/files/figure.png" pageId="29" pageNumber="29">11</figureCitation>
, and 12). However, we were unable to CT-scan other individuals with the ‘‘ 
<taxonomicName baseAuthorityName="Taylor" baseAuthorityYear="1941" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="lineatissimus">
<emphasis italics="true" pageId="29" pageNumber="29">M. lineatissimus</emphasis>
</taxonomicName>
’’ morphotype (or the 
<typeStatus box="[538,633,635,659]" pageId="29" pageNumber="29">holotype</typeStatus>
of 
<taxonomicName class="Amphibia" family="Craugastoridae" genus="Microbatrachylus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="lineatissmus">
<emphasis italics="true" pageId="29" pageNumber="29">M. lineatissmus</emphasis>
</taxonomicName>
), so future investigation is necessary to determine whether the examined 
<typeStatus box="[425,521,695,719]" pageId="29" pageNumber="29">paratype</typeStatus>
(FMNH 104548) is an aberrant individual of 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[406,558,725,748]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[406,558,725,748]" italics="true" pageId="29" pageNumber="29">C. mexicanus</emphasis>
</taxonomicName>
or represents a valid taxon.
</paragraph>
<paragraph blockId="29.[142,813,188,1107]" pageId="29" pageNumber="29">
<bibRefCitation author="Lynch" box="[174,331,784,808]" firstAuthor="Lynch" journalOrPublisher="Chicago Academy of Science, USA" pageId="29" pageNumber="29" refId="ref37877" refString="Lynch, J. D. 1965. A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae). Chicago Academy of Science, USA." title="A Review of the Eleutherodactylid Frog Genus Microbatrachylus (Leptodactylidae)" type="book" year="1965">Lynch (1965)</bibRefCitation>
reports that ‘‘ 
<taxonomicName baseAuthorityName="Taylor" baseAuthorityYear="1940" box="[514,638,785,808]" class="Amphibia" family="Craugastoridae" genus="Eleutherodactylus" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="oaxacae">
<emphasis box="[514,638,785,808]" italics="true" pageId="29" pageNumber="29">E. oaxacae</emphasis>
</taxonomicName>
’’ has parotoid glands. However, we saw no evidence of these glands in the 
<specimenCount box="[142,239,844,868]" pageId="29" pageNumber="29" type="generic" typeStatus="type">two type</specimenCount>
specimens that we examined (FMNH 100001 and FMNH 126638), nor are we aware of any species of 
<taxonomicName authorityName="Cope" authorityYear="1862" box="[142,266,904,927]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="genus">
<emphasis box="[142,266,904,927]" italics="true" pageId="29" pageNumber="29">Craugastor</emphasis>
</taxonomicName>
that possess parotoid glands (
<emphasis box="[636,656,902,927]" italics="true" pageId="29" pageNumber="29">¼</emphasis>
large poison glands on the nuchal region). 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[493,743,934,957]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[493,743,934,957]" italics="true" pageId="29" pageNumber="29">Craugastor mexicanus</emphasis>
</taxonomicName>
likely co-occurs with 
<taxonomicName baseAuthorityName="Gunther" baseAuthorityYear="1900" box="[312,486,963,987]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="omiltemanus">
<emphasis box="[312,486,963,987]" italics="true" pageId="29" pageNumber="29">C. omiltemanus</emphasis>
</taxonomicName>
at high-elevation localities of central 
<collectingRegion box="[232,310,993,1017]" country="Mexico" name="Oaxaca" pageId="29" pageNumber="29">Oaxaca</collectingRegion>
(
<figureCitation box="[331,412,993,1017]" captionStart="FIG" captionStartId="11.[166,180,657,676]" captionTargetBox="[142,817,180,644]" captionTargetId="figure-472@11.[143,812,187,636]" captionTargetPageId="11" captionText="FIG. 6.—Distribution of Craugastor mexicanus, C. montanus, C. pygmaeus, and C. saltator in Mexico and Guatemala. Dots inside symbols indicate individuals used in the molecular analyses in this study. Relevant type localities are indicated by text and arrows." figureDoi="http://doi.org/10.5281/zenodo.6518676" httpUri="https://zenodo.org/record/6518676/files/figure.png" pageId="29" pageNumber="29">Figs. 6</figureCitation>
and 
<figureCitation box="[479,494,993,1017]" captionStart="FIG" captionStartId="11.[868,882,673,692]" captionTargetBox="[829,1514,170,657]" captionTargetId="figure-580@11.[844,1513,186,651]" captionTargetPageId="11" captionText="FIG. 8.—Distribution of Craugastor hobartsmithi, C. cf. hobartsmithi, C. omiltemanus and six new species from Mexico. Dots inside symbols indicate individuals used in the molecular analyses in this study. Inset depicts region of Oaxaca where three of the new species exist in near sympatry. A question mark indicates uncertain georeferencing. Relevant type localities are indicated by text and arrows." figureDoi="http://doi.org/10.5281/zenodo.6518601" httpUri="https://zenodo.org/record/6518601/files/figure.png" pageId="29" pageNumber="29">8</figureCitation>
). At intermediate elevation localities in 
<collectingRegion box="[285,384,1023,1046]" country="Mexico" name="Veracruz de Ignacio de la Llave" pageId="29" pageNumber="29">Veracruz</collectingRegion>
and 
<collectingRegion box="[449,531,1023,1047]" country="Mexico" name="Oaxaca" pageId="29" pageNumber="29">Oaxaca</collectingRegion>
, it may overlap with 
<taxonomicName authorityName="Ahumada-Carrillo et al. 2013:" authorityYear="1338" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="pygmaeus">
<emphasis italics="true" pageId="29" pageNumber="29">C. pygmaeus</emphasis>
</taxonomicName>
(
<figureCitation box="[275,349,1052,1077]" captionStart="FIG" captionStartId="11.[166,180,1861,1880]" captionTargetBox="[143,812,1243,1840]" captionTargetId="figure-515@11.[143,812,1243,1840]" captionTargetPageId="11" captionText="FIG. 7.—Box plots depicting elevational distribution of Craugastor mexicanus and C. pygmaeus inferred from locality data associated with museum specimens. Results are shown for both specimens used in the molecular analysis (Molecules þ Morphology) and for specimens assigned to species using only morphology. Asterisks represent potential outliers, which are.1.5 times the interquartile range (¼ range inside the box)." figureDoi="http://doi.org/10.5281/zenodo.6518603" httpUri="https://zenodo.org/record/6518603/files/figure.png" pageId="29" pageNumber="29">Fig. 7</figureCitation>
). Male 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[447,601,1053,1076]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[447,601,1053,1076]" italics="true" pageId="29" pageNumber="29">C. mexicanus</emphasis>
</taxonomicName>
have significantly larger tympana than do female 
<taxonomicName baseAuthorityName="Brocchi" baseAuthorityYear="1877" box="[497,645,1083,1106]" class="Amphibia" family="Craugastoridae" genus="Craugastor" kingdom="Animalia" order="Anura" pageId="29" pageNumber="29" phylum="Chordata" rank="species" species="mexicanus">
<emphasis box="[497,645,1083,1106]" italics="true" pageId="29" pageNumber="29">C. mexicanus</emphasis>
</taxonomicName>
(
<figureCitation box="[662,744,1082,1106]" captionStart="FIG" captionStartId="13.[166,180,1957,1976]" captionTargetBox="[142,1518,1263,1934]" captionTargetId="figure-395@13.[142,1518,1263,1935]" captionTargetPageId="13" captionText="FIG. 13.—Evidence of sexual dimorphism of tympanum width in Craugastor mexicanus (left) and C. pygmaeus (right). Sex was determined by directly examining gonads for select specimens of C. mexicanus and is implied by scaling patterns in those specimens where sex was not directly determined. Downloaded From: https://bioone.org/journals/Herpetological-Monographs on 04 May 2022" figureDoi="http://doi.org/10.5281/zenodo.6518613" httpUri="https://zenodo.org/record/6518613/files/figure.png" pageId="29" pageNumber="29">Fig. 13</figureCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>