<document ID-DOI="http://dx.doi.org/10.3897/zookeys.1128.82176" ID-Pensoft-Pub="1313-2970-1128-63" ID-Pensoft-UUID="6A7F3348684E5426AF476BB0CC7CC490" ID-ZooBank="74F7E96B1198487C9EAF56C63E958610" ModsDocID="1313-2970-1128-63" checkinTime="1668024550964" checkinUser="pensoft" docAuthor="Goutte, Sandra, Reyes-Velasco, Jacobo, Kassie, Abeje &amp; Boissinot, Stephane" docDate="2022" docId="399B4B5BCC0356439F520B7C70952E91" docLanguage="en" docName="ZooKeys 1128: 63-97" docOrigin="ZooKeys 1128" docPubDate="2022-11-08" docSource="http://dx.doi.org/10.3897/zookeys.1128.82176" docTitle="Leptopelis rugosus" docType="treatment" docVersion="2" id="6A7F3348684E5426AF476BB0CC7CC490" lastPageNumber="63" masterDocId="6A7F3348684E5426AF476BB0CC7CC490" masterDocTitle="Genetic and morphometric analyses of historical type specimens clarify the taxonomy of the Ethiopian Leptopelis gramineus species complex (Anura, Arthroleptidae)" masterLastPageNumber="97" masterPageNumber="63" pageNumber="63" updateTime="1668024852452" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>Genetic and morphometric analyses of historical type specimens clarify the taxonomy of the Ethiopian Leptopelis gramineus species complex (Anura, Arthroleptidae)</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Goutte, Sandra</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-8939-3773</mods:nameIdentifier>
<mods:affiliation>New York University Abu Dhabi, Saadiyat Island, Abu Dhabi, United Arab Emirates</mods:affiliation>
<mods:nameIdentifier type="email">sg5533@nyu.edu</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Reyes-Velasco, Jacobo</mods:namePart>
<mods:affiliation>New York University Abu Dhabi, Saadiyat Island, Abu Dhabi, United Arab Emirates &amp; Entorno Biotico A. C., Colima, Colima, Mexico</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Kassie, Abeje</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-9451-7910</mods:nameIdentifier>
<mods:affiliation>Ethiopian Biodiversity Institute, Addis Ababa, Ethiopia &amp; Addis Ababa University, Addis Ababa, Ethiopia</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Boissinot, Stephane</mods:namePart>
<mods:nameIdentifier type="ORCID">https://orcid.org/0000-0002-8760-1284</mods:nameIdentifier>
<mods:affiliation>New York University Abu Dhabi, Saadiyat Island, Abu Dhabi, United Arab Emirates</mods:affiliation>
<mods:nameIdentifier type="email">stephane.boissinot@nyu.edu</mods:nameIdentifier>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>ZooKeys</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2022</mods:date>
<mods:detail type="pubDate">
<mods:number>2022-11-08</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>1128</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>63</mods:start>
<mods:end>97</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location>
<mods:url>http://dx.doi.org/10.3897/zookeys.1128.82176</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.1128.82176</mods:identifier>
<mods:identifier type="Pensoft-Pub">1313-2970-1128-63</mods:identifier>
<mods:identifier type="ZooBank">74F7E96B1198487C9EAF56C63E958610</mods:identifier>
<mods:identifier type="Pensoft-UUID">6A7F3348684E5426AF476BB0CC7CC490</mods:identifier>
</mods:mods>
<treatment LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91" httpUri="http://treatment.plazi.org/id/399B4B5BCC0356439F520B7C70952E91" lastPageNumber="63" pageId="0" pageNumber="63">
<subSubSection pageId="0" pageNumber="63" type="nomenclature">
<paragraph pageId="0" pageNumber="63">
<taxonomicName LSID="399B4B5B-CC03-5643-9F52-0B7C70952E91" authority="(Ahl, 1924)" baseAuthorityName="Ahl" baseAuthorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis rugosus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosus">Leptopelis rugosus (Ahl, 1924)</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="reference_group">
<paragraph pageId="0" pageNumber="63">
<taxonomicName authorityName="Ahl" authorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Pseudocassina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudocassina rugosa" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosa">Pseudocassina rugosa</taxonomicName>
Ahl, 1924.
</paragraph>
<paragraph pageId="0" pageNumber="63">
<taxonomicName authorityName="Ahl" authorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Pseudocassina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudocassina ocellata" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="ocellata">Pseudocassina ocellata</taxonomicName>
Ahl, 1924.
</paragraph>
<paragraph pageId="0" pageNumber="63">
<taxonomicName authorityName="Tiutenko &amp; Zinenko" authorityYear="2021" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis montanus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="montanus">Leptopelis montanus</taxonomicName>
Tiutenko &amp; Zinenko, 2021.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="type material">
<paragraph pageId="0" pageNumber="63">Type material.</paragraph>
<paragraph pageId="0" pageNumber="63">
<emphasis bold="true" pageId="0" pageNumber="63">
<emphasis italics="true" pageId="0" pageNumber="63">Holotype</emphasis>
.
</emphasis>
Adult female (ZMB-26915) collected during Oscar 
<normalizedToken originalValue="Neumann’s">Neumann's</normalizedToken>
and Carlo von 
<normalizedToken originalValue="Erlanger’s">Erlanger's</normalizedToken>
expedition in 1900 in Meta, Kolla (approx. 
<geoCoordinate degrees="9.17" direction="north" orientation="latitude" precision="555" value="9.17">9.17°N</geoCoordinate>
, 
<geoCoordinate degrees="38.25" direction="east" orientation="longitude" precision="555" value="38.25">38.25°E</geoCoordinate>
, 2650 m a.s.l.).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="materials_examined">
<paragraph pageId="0" pageNumber="63">Material examined.</paragraph>
<paragraph pageId="0" pageNumber="63">
In addition to the 
<typeStatus>holotype</typeStatus>
, we examined 
<materialsCitation collectingDate="1900-07" collectorName="O. Neumann's, C. von Erlanger's" location="Hochebene Didda" specimenCount="one" specimenCount-female="1">
<specimenCount type="female">one female</specimenCount>
collected at the end of 
<collectingDate value="1900-07">July 1900</collectingDate>
during 
<collectorName>
O. 
<normalizedToken originalValue="Neumann’s">Neumann's</normalizedToken>
</collectorName>
and 
<collectorName>
C. von 
<normalizedToken originalValue="Erlanger’s">Erlanger's</normalizedToken>
</collectorName>
expedition in &quot; 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:52E56D3294722470F497069663A59F15" name="Hochebene Didda">Hochebene Didda</location>
&quot; (ZMB-26913), which is one of the
</materialsCitation>
<materialsCitation collectingDate="2022-01-01" collectingDateMax="2022-12-31" collectingDateMin="2022-01-01" collectorName="Tiutenko, Zinenko, As" location="We" specimenCount="two" typeStatus="holotype">
<specimenCount type="generic" typeStatus="syntypes">two syntypes</specimenCount>
of 
<taxonomicName authorityName="Tiutenko &amp; Zinenko" authorityYear="2021" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis montanus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="montanus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis montanus</emphasis>
</taxonomicName>
<collectorName>Tiutenko</collectorName>
&amp; 
<collectorName>Zinenko</collectorName>
, 2021 (= 
<taxonomicName authorityName="Ahl" authorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Pseudocassina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudocassina ocellata" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="ocellata">
<emphasis italics="true" pageId="0" pageNumber="63">Pseudocassina ocellata</emphasis>
</taxonomicName>
), synonymised here with 
<taxonomicName lsidName="L. rugosus" pageId="0" pageNumber="63" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">L. rugosus</emphasis>
</taxonomicName>
. 
<collectorName>As</collectorName>
this specimen was mentioned as &quot;the 
<typeStatus>holotype</typeStatus>
&quot; by 
<bibRefCitation DOI="https://doi.org/10.1080/03946975.2001.10531159" author="Largen, MJ" journalOrPublisher="Tropical Zoology" pageId="0" pageNumber="63" pagination="307 - 402" refId="B24" refString="Largen, MJ, 2001. Catalogue of the amphibians of Ethiopia, including a key for their identification. Tropical Zoology 14 (2): 307 - 402, DOI: https://doi.org/10.1080/03946975.2001.10531159" title="Catalogue of the amphibians of Ethiopia, including a key for their identification." url="https://doi.org/10.1080/03946975.2001.10531159" volume="14" year="2001">Largen (2001)</bibRefCitation>
and has a collection date and a more precise locality than the other 
<typeStatus>syntype</typeStatus>
(ZMB-26914), which only states 
<normalizedToken originalValue="“Somaliand”">&quot;Somaliand&quot;</normalizedToken>
, considered erroneous by (
<bibRefCitation DOI="https://doi.org/10.1080/03749444.1977.10736845" author="Largen, MJ" journalOrPublisher="Supplemento" pageId="0" pageNumber="63" pagination="85 - 136" refId="B23" refString="Largen, MJ, 1977. The Status of the Genus Leptopelis (amphibia Anura Hyperoliidae) in Ethiopia, Including Descriptions of Two New Species. Monitore Zoologico Italiano. Supplemento 9: 85 - 136, DOI: https://doi.org/10.1080/03749444.1977.10736845" title="The Status of the Genus Leptopelis (amphibia Anura Hyperoliidae) in Ethiopia, Including Descriptions of Two New Species. Monitore Zoologico Italiano." url="https://doi.org/10.1080/03749444.1977.10736845" volume="9" year="1977">Largen 1977</bibRefCitation>
), we formally designate ZMB-26913 as the 
<typeStatus>lectotype</typeStatus>
of 
<taxonomicName authorityName="Ahl" authorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Pseudocassina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudocassina ocellata" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="ocellata">
<emphasis italics="true" pageId="0" pageNumber="63">Pseudocassina ocellata</emphasis>
</taxonomicName>
here (see remarks below regarding the locality and taxonomic status). 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:20D5D11D43A21B853F83D1AAA439C819" name="We">We</location>
also examined
</materialsCitation>
<materialsCitation collectingDate="2016-07-12" collectorName="J. Reyes-Velasco, S. Boissinot" elevation="3265" latitude="10.3316" location="Mehal Meda" longLatPrecision="7" longitude="39.7812" specimenCount="one" specimenCount-female="1" typeStatus="holotype">
<specimenCount type="female">one female</specimenCount>
(16-130), collected on 
<collectingDate value="2016-07-12">12 July 2016</collectingDate>
by 
<collectorName>J. Reyes-Velasco</collectorName>
and 
<collectorName>S. Boissinot</collectorName>
east of 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:4A67F9032F9093F3A3A1B508D0E66E15" latitude="10.3316" longLatPrecision="7" longitude="39.7812" name="Mehal Meda">Mehal Meda</location>
(
<geoCoordinate degrees="10.3316" direction="north" orientation="latitude" precision="5" value="10.3316">10.3316°N</geoCoordinate>
, 
<geoCoordinate degrees="39.7812" direction="east" orientation="longitude" precision="5" value="39.7812">39.7812°E</geoCoordinate>
, 
<elevation metricMagnitude="3" metricUnit="m" metricValue="3.2649999999999997" unit="m" value="3265.0">
<quantity metricMagnitude="3" metricUnit="m" metricValue="3.2649999999999997" unit="m" value="3265.0">3265 m</quantity>
a.s.l.
</elevation>
)
</materialsCitation>
, 
<materialsCitation collectingDate="2016-07-12" collectorName="J. Reyes-Velasco, S. Boissinot" elevation="3017" latitude="9.9894" location="Debre Sina" longLatPrecision="7" longitude="39.7452" specimenCount="one" specimenCount-male="1" typeStatus="holotype">
<specimenCount type="male">one male</specimenCount>
(16-103), collected on 
<collectingDate value="2016-07-12">12 July 2016</collectingDate>
by 
<collectorName>J. Reyes-Velasco</collectorName>
and 
<collectorName>S. Boissinot</collectorName>
north of 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:057BFCD2DAAAAA9892968C56F75A69E2" latitude="9.9894" longLatPrecision="7" longitude="39.7452" name="Debre Sina">Debre Sina</location>
(
<geoCoordinate degrees="9.9894" direction="north" orientation="latitude" precision="5" value="9.9894">9.9894°N</geoCoordinate>
, 
<geoCoordinate degrees="39.7452" direction="east" orientation="longitude" precision="5" value="39.7452">39.7452°E</geoCoordinate>
, 
<elevation metricMagnitude="3" metricUnit="m" metricValue="3.017" unit="m" value="3017.0">
<quantity metricMagnitude="3" metricUnit="m" metricValue="3.017" unit="m" value="3017.0">3017 m</quantity>
a.s.l.
</elevation>
)
</materialsCitation>
, 
<materialsCitation collectingDate="2016-07-12" collectorName="J. Reyes-Velasco, S. Boissinot" elevation="3216" latitude="10.33" location="Mehal Meda" longLatPrecision="779" longitude="39.8" specimenCount="three" specimenCount-male="1" typeStatus="holotype">
<specimenCount type="male">three males</specimenCount>
(16-109, 16-116, 16-129), collected on 
<collectingDate value="2016-07-12">12 July 2016</collectingDate>
by 
<collectorName>J. Reyes-Velasco</collectorName>
and 
<collectorName>S. Boissinot</collectorName>
east of 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:632CFE2B60691D83A579DAD2276CD854" latitude="10.33" longLatPrecision="779" longitude="39.8" name="Mehal Meda">Mehal Meda</location>
(10.16- 
<geoCoordinate degrees="10.33" direction="north" orientation="latitude" precision="555" value="10.33">10.33°N</geoCoordinate>
, 39.76- 
<geoCoordinate degrees="39.80" direction="east" orientation="longitude" precision="555" value="39.8">39.80°E</geoCoordinate>
, 
<elevation metricMagnitude="3" metricUnit="m" metricValue="3.216" metricValueMax="3.2649999999999997" metricValueMin="3.167" unit="m" value="3216.0" valueMax="3265.0" valueMin="3167.0">
<quantity metricMagnitude="3" metricUnit="m" metricValue="3.216" metricValueMax="3.2649999999999997" metricValueMin="3.167" unit="m" value="3216.0" valueMax="3265.0" valueMin="3167.0">3167-3265 m</quantity>
a.s.l.
</elevation>
)
</materialsCitation>
, 
<materialsCitation collectingDate="2016-07-13" collectorName="J. Reyes-Velasco, S. Boissinot" elevation="2833" latitude="9.6979" location="Debre Birhan" longLatPrecision="7" longitude="39.5628" specimenCount="five" specimenCount-male="1" typeStatus="holotype">
<specimenCount type="male">five males</specimenCount>
(16-150, 16-151, 16-152, 16-153, 16-154), collected on 
<collectingDate value="2016-07-13">13 July 2016</collectingDate>
by 
<collectorName>J. Reyes-Velasco</collectorName>
and 
<collectorName>S. Boissinot</collectorName>
east of 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:DC8F2C356CAC92BA4A81A874DC0974BE" latitude="9.6979" longLatPrecision="7" longitude="39.5628" name="Debre Birhan">Debre Birhan</location>
(
<geoCoordinate degrees="9.6979" direction="north" orientation="latitude" precision="5" value="9.6979">9.6979°N</geoCoordinate>
, 
<geoCoordinate degrees="39.5628" direction="east" orientation="longitude" precision="5" value="39.5628">39.5628°E</geoCoordinate>
, 
<elevation metricMagnitude="3" metricUnit="m" metricValue="2.833" unit="m" value="2833.0">
<quantity metricMagnitude="3" metricUnit="m" metricValue="2.833" unit="m" value="2833.0">2833 m</quantity>
a.s.l.
</elevation>
)
</materialsCitation>
, 
<materialsCitation collectingDate="2016-07-14" collectorName="J. Reyes-Velasco, S. Boissinot" elevation="2692" latitude="9.75" location="Fiche" longLatPrecision="779" longitude="38.74" specimenCount="three" specimenCount-male="1" typeStatus="holotype">
<specimenCount type="male">three males</specimenCount>
(16-164, 16-168, 16-172), collected on 
<collectingDate value="2016-07-14">14 July 2016</collectingDate>
by 
<collectorName>J. Reyes-Velasco</collectorName>
and 
<collectorName>S. Boissinot</collectorName>
south of 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:88F5322D0651A0129DDA50AA2616AB40" latitude="9.75" longLatPrecision="779" longitude="38.74" name="Fiche">Fiche</location>
(9.73- 
<geoCoordinate degrees="9.75" direction="north" orientation="latitude" precision="555" value="9.75">9.75°N</geoCoordinate>
, 
<geoCoordinate degrees="38.74" direction="east" orientation="longitude" precision="555" value="38.74">38.74°E</geoCoordinate>
, 
<elevation metricMagnitude="3" metricUnit="m" metricValue="2.6915" metricValueMax="2.726" metricValueMin="2.657" unit="m" value="2691.5" valueMax="2726.0" valueMin="2657.0">
<quantity metricMagnitude="3" metricUnit="m" metricValue="2.6915" metricValueMax="2.726" metricValueMin="2.657" unit="m" value="2691.5" valueMax="2726.0" valueMin="2657.0">2657-2726 m</quantity>
a.s.l.
</elevation>
)
</materialsCitation>
, 
<materialsCitation collectingDate="2018-07-01" collectingDateMax="2018-07-02" collectingDateMin="2018-07-01" collectorName="S. Goutte, Y. Bourgeois" elevation="2339" latitude="9.6979" location="Debre Birhan" longLatPrecision="7" longitude="39.5628" specimenCount="five" specimenCount-male="1" typeStatus="holotype">
<specimenCount type="male">five males</specimenCount>
(SB530, SB531, SB532, SB533, SB540), collected on 
<collectingDate value="2018-07-01" valueMax="2018-07-02" valueMin="2018-07-01">1 and 2 July 2018</collectingDate>
by 
<collectorName>S. Goutte</collectorName>
and 
<collectorName>Y. Bourgeois</collectorName>
east of 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:3B03C1B5B2584FE48F968FE4FDC45D9B" latitude="9.6979" longLatPrecision="7" longitude="39.5628" name="Debre Birhan">Debre Birhan</location>
(
<geoCoordinate degrees="9.6979" direction="north" orientation="latitude" precision="5" value="9.6979">9.6979°N</geoCoordinate>
, 
<geoCoordinate degrees="39.5628" direction="east" orientation="longitude" precision="5" value="39.5628">39.5628°E</geoCoordinate>
, 
<elevation metricMagnitude="3" metricUnit="m" metricValue="2.339" unit="m" value="2339.0">
<quantity metricMagnitude="3" metricUnit="m" metricValue="2.339" unit="m" value="2339.0">2339 m</quantity>
a.s.l.
</elevation>
)
</materialsCitation>
, 
<materialsCitation collectingDate="2018-07-02" collectorName="S. Goutte, Y. Bourgeois" elevation="3408" latitude="9.682" location="Debre Birhan" longLatPrecision="7" longitude="39.739" municipality="Ankober" specimenCount="four" specimenCount-male="1" typeStatus="holotype">
<specimenCount type="male">four males</specimenCount>
(SB541, SB542, SB544, SB545), collected on 
<collectingDate value="2018-07-02">2 July 2018</collectingDate>
by 
<collectorName>S. Goutte</collectorName>
and 
<collectorName>Y. Bourgeois</collectorName>
between 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:CBD8A8069A1288932A5743F9B845804E" latitude="9.682" longLatPrecision="7" longitude="39.739" municipality="Ankober" name="Debre Birhan">Debre Birhan</location>
and 
<collectingMunicipality>Ankober</collectingMunicipality>
(
<geoCoordinate degrees="9.6820" direction="north" orientation="latitude" precision="5" value="9.682">9.6820°N</geoCoordinate>
, 
<geoCoordinate degrees="39.7390" direction="east" orientation="longitude" precision="5" value="39.739">39.7390°E</geoCoordinate>
, 
<elevation metricMagnitude="3" metricUnit="m" metricValue="3.408" unit="m" value="3408.0">
<quantity metricMagnitude="3" metricUnit="m" metricValue="3.408" unit="m" value="3408.0">3408 m</quantity>
a.s.l.
</elevation>
)
</materialsCitation>
, 
<specimenCount type="female">two females</specimenCount>
(SB555, SB558) and 
<materialsCitation collectingDate="2018-07-04" collectorName="S. Goutte, Y. Bourgeois" elevation="3383" latitude="10.33" location="Mehal Meda" longLatPrecision="779" longitude="39.8" specimenCount="one" specimenCount-male="1" typeStatus="holotype">
<specimenCount type="male">one male</specimenCount>
(SB556), collected on 
<collectingDate value="2018-07-04">4 July 2018</collectingDate>
by 
<collectorName>S. Goutte</collectorName>
and 
<collectorName>Y. Bourgeois</collectorName>
east of 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:60122AAFB7DA20F5C5003914D21C3400" latitude="10.33" longLatPrecision="779" longitude="39.8" name="Mehal Meda">Mehal Meda</location>
(10.31- 
<geoCoordinate degrees="10.33" direction="north" orientation="latitude" precision="555" value="10.33">10.33°N</geoCoordinate>
, 39.78- 
<geoCoordinate degrees="39.80" direction="east" orientation="longitude" precision="555" value="39.8">39.80°E</geoCoordinate>
, 
<elevation metricMagnitude="3" metricUnit="m" metricValue="3.383" metricValueMax="3.429" metricValueMin="3.3369999999999997" unit="m" value="3383.0" valueMax="3429.0" valueMin="3337.0">
<quantity metricMagnitude="3" metricUnit="m" metricValue="3.383" metricValueMax="3.429" metricValueMin="3.3369999999999997" unit="m" value="3383.0" valueMax="3429.0" valueMin="3337.0">3337-3429 m</quantity>
a.s.l.
</elevation>
) and
</materialsCitation>
<materialsCitation collectingDate="2018-07-11" collectorName="S. Goutte, Y. Bourgeois" elevation="2365" latitude="9.7307" location="Fiche" longLatPrecision="7" longitude="38.7439" specimenCount="three" specimenCount-male="1" typeStatus="holotype">
<specimenCount type="male">three males</specimenCount>
(SB608, SB609, SB610), collected on 
<collectingDate value="2018-07-11">11 July 2018</collectingDate>
by 
<collectorName>S. Goutte</collectorName>
and 
<collectorName>Y. Bourgeois</collectorName>
south of 
<location LSID="urn:lsid:plazi:treatment:399B4B5BCC0356439F520B7C70952E91:CA21C6EE5434C702C2FFAC59ABFDBC47" latitude="9.7307" longLatPrecision="7" longitude="38.7439" name="Fiche">Fiche</location>
(
<geoCoordinate degrees="9.7307" direction="north" orientation="latitude" precision="5" value="9.7307">9.7307°N</geoCoordinate>
, 
<geoCoordinate degrees="38.7439" direction="east" orientation="longitude" precision="5" value="38.7439">38.7439°E</geoCoordinate>
, 
<elevation metricMagnitude="3" metricUnit="m" metricValue="2.3649999999999998" unit="m" value="2365.0">
<quantity metricMagnitude="3" metricUnit="m" metricValue="2.3649999999999998" unit="m" value="2365.0">2365 m</quantity>
a.s.l.
</elevation>
)
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="diagnosis">
<paragraph pageId="0" pageNumber="63">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="63">
A large (male (n = 22) SVL 38.7 
<normalizedToken originalValue="±">+/-</normalizedToken>
2.7 mm, female (n = 2) SVL 52.4 
<normalizedToken originalValue="±">+/-</normalizedToken>
0.2 mm) species of the 
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis gramineus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="gramineus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis gramineus</emphasis>
</taxonomicName>
species complex (Figs 
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Dorsal and ventral views of the female holotype (ZMB- 26915) of Leptopelis rugosus." figureDoi="10.3897/zookeys.1128.82176.figure5" httpUri="https://binary.pensoft.net/fig/766798" pageId="0" pageNumber="63">5</figureCitation>
, 
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Leptopelis rugosus A live male (SB 610; left) and female (SB 558; right) B dorsal and ventral views of male (SB 610) after euthanasia and before fixation C dorsal and ventral views of female (SB 558) after euthanasia and before fixation." figureDoi="10.3897/zookeys.1128.82176.figure6" httpUri="https://binary.pensoft.net/fig/766799" pageId="0" pageNumber="63">6</figureCitation>
). Robust, semi-fossorial form. It differs from other members of the 
<taxonomicName baseAuthorityName="Boulenger" baseAuthorityYear="1898" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis gramineus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="gramineus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis gramineus</emphasis>
</taxonomicName>
species complex by the following combination of characters: (1) large tympanum (male TD/ED 0.57 
<normalizedToken originalValue="±">+/-</normalizedToken>
0.11, female TD/ED 0.54), (2) long snout (male SL/HL 0.25 
<normalizedToken originalValue="±">+/-</normalizedToken>
0.02, female SL/HL 0.27 
<normalizedToken originalValue="±">+/-</normalizedToken>
0.01), (3) well-developed metatarsal tubercle (male MTL/FL 0.17 
<normalizedToken originalValue="±">+/-</normalizedToken>
0.02, female MTL/FL 0.18 
<normalizedToken originalValue="±">+/-</normalizedToken>
0.02), (4) ventrum lacking any brown spots, (5) yellow colouration on the side of the ventrum and the inner thighs almost always present.
</paragraph>
<caption doi="10.3897/zookeys.1128.82176.figure5" httpUri="https://binary.pensoft.net/fig/766798" pageId="0" pageNumber="63" start="Figure 5" startId="F5">
<paragraph pageId="0" pageNumber="63">
<emphasis bold="true" pageId="0" pageNumber="63">Figure 5.</emphasis>
Dorsal and ventral views of the female holotype (ZMB-26915) of 
<taxonomicName baseAuthorityName="Ahl" baseAuthorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis rugosus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis rugosus</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<caption doi="10.3897/zookeys.1128.82176.figure6" httpUri="https://binary.pensoft.net/fig/766799" pageId="0" pageNumber="63" start="Figure 6" startId="F6">
<paragraph pageId="0" pageNumber="63">
<emphasis bold="true" pageId="0" pageNumber="63">Figure 6.</emphasis>
<taxonomicName baseAuthorityName="Ahl" baseAuthorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis rugosus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis rugosus</emphasis>
</taxonomicName>
<emphasis bold="true" pageId="0" pageNumber="63">A</emphasis>
live male (SB610; left) and female (SB558; right) 
<emphasis bold="true" pageId="0" pageNumber="63">B</emphasis>
dorsal and ventral views of male (SB610) after euthanasia and before fixation 
<emphasis bold="true" pageId="0" pageNumber="63">C</emphasis>
dorsal and ventral views of female (SB558) after euthanasia and before fixation.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="comparisons">
<paragraph pageId="0" pageNumber="63">Comparison.</paragraph>
<paragraph pageId="0" pageNumber="63">
Larger body size, longer head and snout and greater snout-nostril distance and larger tympanum and metatarsal tubercle than 
<taxonomicName lsidName="L. gramineus" pageId="0" pageNumber="63" rank="species" species="gramineus">
<emphasis italics="true" pageId="0" pageNumber="63">L. gramineus</emphasis>
</taxonomicName>
, 
<taxonomicName lsidName="L. diffidens" pageId="0" pageNumber="63" rank="species" species="diffidens">
<emphasis italics="true" pageId="0" pageNumber="63">L. diffidens</emphasis>
</taxonomicName>
and 
<taxonomicName genus="L." lsidName="L." pageId="0" pageNumber="63" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="63">L.</emphasis>
</taxonomicName>
sp. Kibre Mengist (Table 
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Summary of the linear morphometric measurement for the Leptopelis gramineus species complex." httpUri="http://table.plazi.org/id/BF4B1B424886D2055504A1AEA5E8D7AC" pageId="0" pageNumber="63" tableUuid="BF4B1B424886D2055504A1AEA5E8D7AC">1</tableCitation>
, Suppl. material 5: table S9). Longer hind-limbs than the Bale/Assela clade, but shorter than 
<taxonomicName lsidName="L. susanae" pageId="0" pageNumber="63" rank="species" species="susanae">
<emphasis italics="true" pageId="0" pageNumber="63">L. susanae</emphasis>
</taxonomicName>
(Table 
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Summary of the linear morphometric measurement for the Leptopelis gramineus species complex." httpUri="http://table.plazi.org/id/BF4B1B424886D2055504A1AEA5E8D7AC" pageId="0" pageNumber="63" tableUuid="BF4B1B424886D2055504A1AEA5E8D7AC">1</tableCitation>
, Suppl. material 5: table S9). Finger and toe discs less developed and head narrower and shorter than 
<taxonomicName lsidName="L. susanae" pageId="0" pageNumber="63" rank="species" species="susanae">
<emphasis italics="true" pageId="0" pageNumber="63">L. susanae</emphasis>
</taxonomicName>
(Table 
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Summary of the linear morphometric measurement for the Leptopelis gramineus species complex." httpUri="http://table.plazi.org/id/BF4B1B424886D2055504A1AEA5E8D7AC" pageId="0" pageNumber="63" tableUuid="BF4B1B424886D2055504A1AEA5E8D7AC">1</tableCitation>
, Suppl. material 5: table S9). 
<taxonomicName baseAuthorityName="Ahl" baseAuthorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis rugosus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis rugosus</emphasis>
</taxonomicName>
is distinguished from the Bale/Assela clade by the lack of dark pigmentation on the ventrum, throat and ventral side of the limbs (Figs 
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Leptopelis rugosus A live male (SB 610; left) and female (SB 558; right) B dorsal and ventral views of male (SB 610) after euthanasia and before fixation C dorsal and ventral views of female (SB 558) after euthanasia and before fixation." figureDoi="10.3897/zookeys.1128.82176.figure6" httpUri="https://binary.pensoft.net/fig/766799" pageId="0" pageNumber="63">6</figureCitation>
, 
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. Leptopelis shebellensis sp. nov. A male holotype (SB 482; left) and live female paratype (SB 41; right) B male holotype (SB 482; left) and dorsal and ventral views of female paratype (SB 39; right) after fixation." figureDoi="10.3897/zookeys.1128.82176.figure7" httpUri="https://binary.pensoft.net/fig/766800" pageId="0" pageNumber="63">7</figureCitation>
).
</paragraph>
<caption doi="10.3897/zookeys.1128.82176.figure7" httpUri="https://binary.pensoft.net/fig/766800" pageId="0" pageNumber="63" start="Figure 7" startId="F7">
<paragraph pageId="0" pageNumber="63">
<emphasis bold="true" pageId="0" pageNumber="63">Figure 7.</emphasis>
<taxonomicName authorityName="Goutte, Reyes-Velasco, Kassie &amp; Boissinot" authorityYear="2022" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis shebellensis" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="shebellensis">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis shebellensis</emphasis>
</taxonomicName>
sp. nov. 
<emphasis bold="true" pageId="0" pageNumber="63">A</emphasis>
male holotype (SB482; left) and live female paratype (SB41; right) 
<emphasis bold="true" pageId="0" pageNumber="63">B</emphasis>
male holotype (SB482; left) and dorsal and ventral views of female paratype (SB39; right) after fixation.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="description">
<paragraph pageId="0" pageNumber="63">Description of the holotype.</paragraph>
<paragraph pageId="0" pageNumber="63">
Relatively large adult female (SVL 44.7 mm) in good condition of preservation (Fig. 
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Dorsal and ventral views of the female holotype (ZMB- 26915) of Leptopelis rugosus." figureDoi="10.3897/zookeys.1128.82176.figure5" httpUri="https://binary.pensoft.net/fig/766798" pageId="0" pageNumber="63">5</figureCitation>
). Body robust and round. Head a third of body size in length, wider than long (HW/HL 1.21). Canthus rostralis obtuse and snout rounded and wide (IND/IOD 0.96). Nostril half-way between the tip of the snout and the eye (NS/SL 0.53). Tympanum partially hidden by flank skin rugosities and barely visible. Hind-limbs relatively long (TL/SVL 0.38 and THL/SVL 0.43). Finger and toe discs barely expanded, but distinct, ovoid. Finger formula: I &lt;II &lt;IV &lt;III. Hand free of webbing. Foot longer than tibia (FL/TL 1.3). Inner metatarsal tubercle well-developed, oval in shape, 0.20 
<normalizedToken originalValue="×">x</normalizedToken>
foot length. Outer metatarsal tubercle absent. Toe formula: I &lt;II &lt;V &lt;III &lt;IV. Toe webbing formula (toe internal/external sides, number of phalanges webbed): Ie(1), IIi/e(1-1), IIIi/e(1-2), IVi/e(2-2), Vi(2). Skin of the dorsum, flanks and ventrum highly rugose.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="colouration of the holotype in preservative">
<paragraph pageId="0" pageNumber="63">Colouration of the holotype in preservative.</paragraph>
<paragraph pageId="0" pageNumber="63">
Dorsal ground colour and canthal region dark olive brown with no visible pattern, except for a large light brown blotch covering about a third of the dorsum (Fig. 
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Dorsal and ventral views of the female holotype (ZMB- 26915) of Leptopelis rugosus." figureDoi="10.3897/zookeys.1128.82176.figure5" httpUri="https://binary.pensoft.net/fig/766798" pageId="0" pageNumber="63">5</figureCitation>
). This discolouration probably appeared during the specimen preservation and after its original description, where Ahl described the dorsum as &quot;solid dark olive-brown&quot;. The thin light yellowish line noted by 
<bibRefCitation DOI="https://doi.org/10.1002/mmnz.4830110102" author="Ahl, E" journalOrPublisher="Mitteilungen aus dem Zoologischen Museum in Berlin" pageId="0" pageNumber="63" pagination="1 - 12" refId="B1" refString="Ahl, E, 1924. Ueber eine Froschsammlung aus Nordost-Afrika und Arabien. Mitteilungen aus dem Zoologischen Museum in Berlin 11: 1 - 12, DOI: https://doi.org/10.1002/mmnz.4830110102" title="Ueber eine Froschsammlung aus Nordost-Afrika und Arabien." url="https://doi.org/10.1002/mmnz.4830110102" volume="11" year="1924">Ahl (1924)</bibRefCitation>
to extend to the upper arm is mostly faded away behind the tympanic region. Upper lip and flanks light yellowish-brown. Throat, ventrum, ventral side of the thighs and tibias light yellowish-brown. Front and hind-limbs olive brown without any marking.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="variation">
<paragraph pageId="0" pageNumber="63">Variations.</paragraph>
<paragraph pageId="0" pageNumber="63">
<taxonomicName baseAuthorityName="Ahl" baseAuthorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis rugosus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis rugosus</emphasis>
</taxonomicName>
presents less colour polymorphism than the smaller members of the 
<taxonomicName lsidName="L. gramineus" pageId="0" pageNumber="63" rank="species" species="gramineus">
<emphasis italics="true" pageId="0" pageNumber="63">L. gramineus</emphasis>
</taxonomicName>
species complex. Dorsum is green to dark green and can be completely uniform or have a few to many irregular brown blotches (Fig. 
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Leptopelis rugosus A live male (SB 610; left) and female (SB 558; right) B dorsal and ventral views of male (SB 610) after euthanasia and before fixation C dorsal and ventral views of female (SB 558) after euthanasia and before fixation." figureDoi="10.3897/zookeys.1128.82176.figure6" httpUri="https://binary.pensoft.net/fig/766799" pageId="0" pageNumber="63">6</figureCitation>
). In some individuals, these blotches form an irregular dorsal stripe extending from the top of the head to the lower back. A dark brown to black bar covers the canthal region and extends behind the eye, over the tympanum and sometimes behind the arm junction. This canthal stripe is overlined by a thin, more or less visible yellowish line. Flanks are the same colour as the dorsal ground colour and, in all individuals examined, except for the female SB558, have more or less well-defined brown ocelli. Limbs are the same colouration as the dorsum and rarely have irregular brown markings. Upper lip may be cream or a lighter shade of green than the dorsum without any markings. Iris sand colour to brown. Tympanum may be uniformly green or partially covered by a brown blotch joining the brown bar behind the eye. Throat and chest uniformly white to pale yellow. Ventrum generally white or cream with light to deep yellow zones on the sides extending to the ventral side of the thighs. In some individuals, the ventrum may be completely yellow. In gravid females, yellow eggs are visible through the thick ventral skin. In most individuals, the palms of the hands, ventral and inner sides of the limbs show very little to no dark pigmentation. Some individuals have a few irregular brown or black blotches on the inner tibia, forearm, hand and foot. Dorsal skin may be smooth, slightly or very rugose.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="habitat">
<paragraph pageId="0" pageNumber="63">Habitat, distribution and natural history.</paragraph>
<paragraph pageId="0" pageNumber="63">
<taxonomicName baseAuthorityName="Ahl" baseAuthorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis rugosus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis rugosus</emphasis>
</taxonomicName>
is found in grassy meadows of the Ethiopian Highlands north of the GRV at mid- to high elevations (2,339-3,337 m a.s.l.). This species occurs notably near Debre Birhan, Debre Sina, Fiche, Addis Ababa, Holeta and Ambo (Fig. 
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. Distribution ranges of six species of the Leptopelis gramineus species complex. Small, more arboreal forms are represented by green (L. diffidens), yellow (L. gramineus) and purple (L. xeniae sp. nov.) triangles. Larger, semi-fossorial forms are represented by red (L. rugosus) and white (L. shebellensis sp. nov.) circles. The large arboreal Leptopelis susanae is represented by a light blue triangle. Stars indicate type localities given in the original descriptions of L. gramineus (yellow; dashed line between Badditu and Dime), Pseudocassina rugosa (red) and Pseudocassina ocellata (grey), both synonymised here with L. rugosus (see the discussion in the main text)." figureDoi="10.3897/zookeys.1128.82176.figure1" httpUri="https://binary.pensoft.net/fig/766794" pageId="0" pageNumber="63">1</figureCitation>
, Suppl. material 5: table S1). The northernmost population was found near Mehal Meda (
<geoCoordinate degrees="10.3171" direction="north" orientation="latitude" precision="5" value="10.3171">10.3171°N</geoCoordinate>
, 
<geoCoordinate degrees="39.8024" direction="east" orientation="longitude" precision="5" value="39.8024">39.8024°E</geoCoordinate>
), while the southernmost individuals were found between Ambo and Wonchi (
<geoCoordinate degrees="8.9007" direction="north" orientation="latitude" precision="5" value="8.9007">8.9007°N</geoCoordinate>
, 
<geoCoordinate degrees="37.8928" direction="east" orientation="longitude" precision="5" value="37.8928">37.8928°E</geoCoordinate>
). One of the two types in 
<normalizedToken originalValue="Ahl’s">Ahl's</normalizedToken>
original description of 
<taxonomicName authorityName="Ahl" authorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Pseudocassina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudocassina ocellata" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="ocellata">
<emphasis italics="true" pageId="0" pageNumber="63">Pseudocassina ocellata</emphasis>
</taxonomicName>
was collected on the Arussi Plateau, which, if the locality is correct, is the only known specimen east of the GRV (see remark on the type locality below).
</paragraph>
<paragraph pageId="0" pageNumber="63">Males are heard calling at night and sometimes during the day for extended periods of time. Advertisement calls are emitted from the ground, either on the grass or from a cavity in the ground or under a rock, generally near a stream or a flooded area. Several males were found calling from the banks of a completely dried-out stream, although it is unknown to us whether the reproduction period extends to the dry season or whether males keep calling only during shorter dry periods.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="advertisement call">
<paragraph pageId="0" pageNumber="63">Advertisement call.</paragraph>
<paragraph pageId="0" pageNumber="63">
The call of 
<taxonomicName baseAuthorityName="Ahl" baseAuthorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis rugosus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis rugosus</emphasis>
</taxonomicName>
is a short rattle composed of a single note of 65 
<normalizedToken originalValue="±">+/-</normalizedToken>
21 ms in duration, containing 4 
<normalizedToken originalValue="±">+/-</normalizedToken>
1 pulses (Fig. 
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Advertisement calls of six Leptopelis species inhabiting the Ethiopian Highlands A L. shebellensis sp. nov. (SB 61) B L. rugosus (SB 609) C L. gramineus (SB 212) D L. xeniae sp. nov. (SB 169) E L. diffidens (SB 134) F L. susanae (SB 223)." figureDoi="10.3897/zookeys.1128.82176.figure4" httpUri="https://binary.pensoft.net/fig/766797" pageId="0" pageNumber="63">4B</figureCitation>
). In most individuals, the two first pulses are emitted at very short intervals, while the subsequent pulses are more spaced (average inter-pulse interval 20 
<normalizedToken originalValue="±">+/-</normalizedToken>
5 ms). Other individuals produce notes with regularly-spaced pulses. Amplitude is highest at the beginning of the note and decreases gradually. Within a call bout, calls are spaced by 13 
<normalizedToken originalValue="±">+/-</normalizedToken>
7 seconds, often with an acceleration of the call rate from a call every 10 seconds to one call per second. Call dominant frequency is 1,769 
<normalizedToken originalValue="±">+/-</normalizedToken>
60 Hz, with a bandwidth of 742 
<normalizedToken originalValue="±">+/-</normalizedToken>
82 Hz.
</paragraph>
<paragraph pageId="0" pageNumber="63">
The call of 
<taxonomicName baseAuthorityName="Ahl" baseAuthorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis rugosus" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">Leptopelis rugosus</emphasis>
</taxonomicName>
is distinguishable from the calls of 
<taxonomicName lsidName="L. gramineus" pageId="0" pageNumber="63" rank="species" species="gramineus">
<emphasis italics="true" pageId="0" pageNumber="63">L. gramineus</emphasis>
</taxonomicName>
, 
<taxonomicName lsidName="L. diffidens" pageId="0" pageNumber="63" rank="species" species="diffidens">
<emphasis italics="true" pageId="0" pageNumber="63">L. diffidens</emphasis>
</taxonomicName>
and the Kibre Mengist and Bale/Assela clades by its lower number of pulses per note and narrower frequency band width. It is further distinguished from the call of 
<taxonomicName lsidName="L. gramineus" pageId="0" pageNumber="63" rank="species" species="gramineus">
<emphasis italics="true" pageId="0" pageNumber="63">L. gramineus</emphasis>
</taxonomicName>
, 
<taxonomicName lsidName="L. diffidens" pageId="0" pageNumber="63" rank="species" species="diffidens">
<emphasis italics="true" pageId="0" pageNumber="63">L. diffidens</emphasis>
</taxonomicName>
and 
<taxonomicName genus="L." lsidName="L." pageId="0" pageNumber="63" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="63">L.</emphasis>
</taxonomicName>
sp. Kibre Mengist by its shorter note duration and from 
<taxonomicName genus="L." lsidName="L." pageId="0" pageNumber="63" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="63">L.</emphasis>
</taxonomicName>
sp. Kibre Mengist by its lower peak frequency and higher pulse rate. Finally, it is distinguished from the call of 
<taxonomicName lsidName="L. susanae" pageId="0" pageNumber="63" rank="species" species="susanae">
<emphasis italics="true" pageId="0" pageNumber="63">L. susanae</emphasis>
</taxonomicName>
by its longer duration and lower pulse rate (Table 
<tableCitation captionStart="Table 2" captionStartId="T2" captionText="Table 2. Summary of call acoustic characters for the Leptopelis gramineus species complex." httpUri="http://table.plazi.org/id/612A31E07847DDFCD1055D8D4BAD5511" pageId="0" pageNumber="63" tableUuid="612A31E07847DDFCD1055D8D4BAD5511">2</tableCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="63" type="remarks">
<paragraph pageId="0" pageNumber="63">Remarks.</paragraph>
<paragraph pageId="0" pageNumber="63">Diagnostic characters used in the original description.</paragraph>
<paragraph pageId="0" pageNumber="63">
Ahl described 
<taxonomicName authorityName="Ahl" authorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Pseudocassina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudocassina rugosa" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="rugosa">
<emphasis italics="true" pageId="0" pageNumber="63">Pseudocassina rugosa</emphasis>
</taxonomicName>
, based on a single female and provided three main diagnostic characters when compared with 
<taxonomicName authorityName="Ahl" authorityYear="1924" class="Amphibia" family="Arthroleptidae" genus="Pseudocassina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Pseudocassina ocellata" order="Anura" pageId="0" pageNumber="63" phylum="Chordata" rank="species" species="ocellata">
<emphasis italics="true" pageId="0" pageNumber="63">Pseudocassina ocellata</emphasis>
</taxonomicName>
, which he described in the same article, based on one female and one male: (1) the texture of the skin (rugose for 
<taxonomicName lsidName="P. rugosa" pageId="0" pageNumber="63" rank="species" species="rugosa">
<emphasis italics="true" pageId="0" pageNumber="63">P. rugosa</emphasis>
</taxonomicName>
and smooth for 
<taxonomicName lsidName="P. ocellata" pageId="0" pageNumber="63" rank="species" species="ocellata">
<emphasis italics="true" pageId="0" pageNumber="63">P. ocellata</emphasis>
</taxonomicName>
), (2) the length of the tibia (
<taxonomicName lsidName="P. rugosa" pageId="0" pageNumber="63" rank="species" species="rugosa">
<emphasis italics="true" pageId="0" pageNumber="63">P. rugosa</emphasis>
</taxonomicName>
TL/SVL 1/3.5 and 
<taxonomicName lsidName="P. ocellata" pageId="0" pageNumber="63" rank="species" species="ocellata">
<emphasis italics="true" pageId="0" pageNumber="63">P. ocellata</emphasis>
</taxonomicName>
TL/SVL 1/3), (3) the visibility of the tympanum (hidden for 
<taxonomicName lsidName="P. rugosa" pageId="0" pageNumber="63" rank="species" species="rugosa">
<emphasis italics="true" pageId="0" pageNumber="63">P. rugosa</emphasis>
</taxonomicName>
and visible for 
<taxonomicName lsidName="P. ocellata" pageId="0" pageNumber="63" rank="species" species="ocellata">
<emphasis italics="true" pageId="0" pageNumber="63">P. ocellata</emphasis>
</taxonomicName>
). Ahl also named 
<taxonomicName lsidName="P. ocellata" pageId="0" pageNumber="63" rank="species" species="ocellata">
<emphasis italics="true" pageId="0" pageNumber="63">P. ocellata</emphasis>
</taxonomicName>
after the presence of ocelli on the flanks of the individuals he examined, which reflects on its specificity, even though he did not use this trait as a diagnostic character.
</paragraph>
<paragraph pageId="0" pageNumber="63">
We found individuals of 
<taxonomicName lsidName="L. rugosus" pageId="0" pageNumber="63" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">L. rugosus</emphasis>
</taxonomicName>
and the Bale/Assela clade with either a completely smooth, slightly rugose or coarsely rugose dorsum. The rugosity of the skin thus seems to be variable amongst individuals and, perhaps, age or season. Additionally, we have noticed that the rugosity of the skin may disappear after euthanasia and/or fixation of the specimen. Finally, even though almost all individuals of 
<taxonomicName lsidName="L. diffidens" pageId="0" pageNumber="63" rank="species" species="diffidens">
<emphasis italics="true" pageId="0" pageNumber="63">L. diffidens</emphasis>
</taxonomicName>
, 
<taxonomicName lsidName="L. gramineus" pageId="0" pageNumber="63" rank="species" species="gramineus">
<emphasis italics="true" pageId="0" pageNumber="63">L. gramineus</emphasis>
</taxonomicName>
and the 
<taxonomicName genus="L." lsidName="L." pageId="0" pageNumber="63" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="63">L.</emphasis>
</taxonomicName>
sp. Kibre Mengist examined had smooth skin, we found two females 
<taxonomicName lsidName="L. diffidens" pageId="0" pageNumber="63" rank="species" species="diffidens">
<emphasis italics="true" pageId="0" pageNumber="63">L. diffidens</emphasis>
</taxonomicName>
and one female 
<taxonomicName genus="L." lsidName="L." pageId="0" pageNumber="63" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="63">L.</emphasis>
</taxonomicName>
sp. Kibre Mengist with slightly rugose skin. The visibility of the tympanum seems to be variable across the individuals as well, perhaps linked to the size of the individual and the rugosity of the skin. While some female 
<taxonomicName lsidName="L. rugosus" pageId="0" pageNumber="63" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">L. rugosus</emphasis>
</taxonomicName>
have a partially hidden tympanum, all males examined had a visible tympanum. The difference in tibia length between the specimens described by Ahl reflects individual variations as shown in our dataset. Finally, ocelli on the flanks and/or dorsum are present in certain individuals of 
<taxonomicName lsidName="L. rugosus" pageId="0" pageNumber="63" rank="species" species="rugosus">
<emphasis italics="true" pageId="0" pageNumber="63">L. rugosus</emphasis>
</taxonomicName>
and the Bale/Assela clade and is not a diagnostic character.
</paragraph>
</subSubSection>
</treatment>
</document>