638 lines
61 KiB
XML
638 lines
61 KiB
XML
<document ID-DOI="http://dx.doi.org/10.3897/zookeys.824.31570" ID-GBIF-Dataset="f81fc57b-58da-4ab1-bc1b-65edd3bb55b0" ID-PMC="PMC6381079" ID-Pensoft-Pub="1313-2970-824-53" ID-PubMed="30799972" ID-ZBK="BDDAF7D8F5E5406DA485E4D062271833" ModsDocAuthor="" ModsDocDate="2019" ModsDocID="1313-2970-824-53" ModsDocOrigin="ZooKeys 824" ModsDocTitle="A new species of puddle frog from an unexplored mountain in southwestern Ethiopia (Anura, Phrynobatrachidae, Phrynobatrachus)" checkinTime="1551186949441" checkinUser="pensoft" docAuthor="Goutte, Sandra, Reyes-Velasco, Jacobo & Boissinot, Stephane" docDate="2019" docId="187C1C031426DEDEE45151AD9005587E" docLanguage="en" docName="ZooKeys 824: 53-70" docOrigin="ZooKeys 824" docSource="http://dx.doi.org/10.3897/zookeys.824.31570" docTitle="Phrynobatrachus bibita Goutte, Reyes-Velasco & Boissinot, sp. n." docType="treatment" docUuid="3B046650-2B48-4740-9ED9-A29A4EC10A72" docUuidSource="ZooBank" docVersion="5" lastPageNumber="53" masterDocId="FFF269106F5A547C6C6FFFAD5C53CC62" masterDocTitle="A new species of puddle frog from an unexplored mountain in southwestern Ethiopia (Anura, Phrynobatrachidae, Phrynobatrachus)" masterLastPageNumber="70" masterPageNumber="53" pageNumber="53" updateTime="1668166927814" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>A new species of puddle frog from an unexplored mountain in southwestern Ethiopia (Anura, Phrynobatrachidae, Phrynobatrachus)</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Goutte, Sandra</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Reyes-Velasco, Jacobo</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Boissinot, Stephane</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>ZooKeys</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2019</mods:date>
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<mods:detail type="volume">
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<mods:number>824</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>53</mods:start>
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<mods:end>70</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:location>
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<mods:url>http://dx.doi.org/10.3897/zookeys.824.31570</mods:url>
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</mods:location>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">http://dx.doi.org/10.3897/zookeys.824.31570</mods:identifier>
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<mods:identifier type="Pensoft-Pub">1313-2970-824-53</mods:identifier>
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<mods:identifier type="ZBK">BDDAF7D8F5E5406DA485E4D062271833</mods:identifier>
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<mods:identifier type="ZooBank">BDDAF7D8F5E5406DA485E4D062271833</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="154127856" LSID="urn:lsid:zoobank.org:act:3B046650-2B48-4740-9ED9-A29A4EC10A72" httpUri="http://treatment.plazi.org/id/187C1C031426DEDEE45151AD9005587E" lastPageNumber="53" pageId="0" pageNumber="53">
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<subSubSection pageId="0" pageNumber="53" type="nomenclature">
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<paragraph pageId="0" pageNumber="53">
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<taxonomicName LSID="http://zoobank.org/3B046650-2B48-4740-9ED9-A29A4EC10A72" authority="Goutte, Reyes-Velasco & Boissinot" class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita Goutte, Reyes-Velasco & Boissinot</taxonomicName>
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<taxonomicNameLabel pageId="0" pageNumber="53">sp. n.</taxonomicNameLabel>
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="53" type="materials_examined">
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<paragraph pageId="0" pageNumber="53">Material.</paragraph>
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<paragraph pageId="0" pageNumber="53">
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Holotype. A male (SB440; Figure 2), collected on 17 June 2018, at night, in Bibita Mountain, southwestern Ethiopia (
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<geoCoordinate direction="north" orientation="latitude" precision="5" value="6.8229">6.8229N</geoCoordinate>
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,
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<geoCoordinate direction="east" orientation="longitude" precision="5" value="35.0938">35.0938E</geoCoordinate>
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, datum = WGS84; 1972 m a.s.l.). The specimen was collected while in amplexus (female paratopotype SB439), on vegetation, in overgrown pond in primary forest. Paratopotypes. Nine specimens: eight females collected in the same pond as the holotype on 16 June 2018 (SB418-SB420, SB424-SB427) and on 17 June 2018 (SB439); another male (SB421) collected at night in the water (probably calling), at same locality as the holotype on 16 June 2018.
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</paragraph>
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<caption pageId="0" pageNumber="53">
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<paragraph pageId="0" pageNumber="53">
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Figure 2.
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<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita</taxonomicName>
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sp. n. A Live pictures of
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<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
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sp. n. Male holotype (left; SB440) and female paratopotype (right; SB424) B Ventral and dorsal views of the same individuals, with male on the left and female on the right. Scale bar: 10 mm.
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</paragraph>
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</caption>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="53" type="diagnosis">
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<paragraph pageId="0" pageNumber="53">Diagnosis.</paragraph>
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<paragraph pageId="0" pageNumber="53">
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Small species (SVL = 16.8
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<normalizedToken originalValue="±">+/-</normalizedToken>
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0.1 mm for males, 20.3
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<normalizedToken originalValue="±">+/-</normalizedToken>
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0.9 mm for females) attributed to the genus
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<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
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by the presence of tarsal and outer metatarsal tubercles (Suppl. material 2: Figure S1A). Body slender, with long legs (tibia length/SVL = 0.6 in both sexes) rather long snout for the genus and very elongated fingers (hand length/SVL = 0.3 in both sexes) and toes (foot length/SVL = 0.6 in both sexes) in comparison to its congeners. Webbing absent between fingers and minimal between toes. Tympanum not visible. Canthus rostralis marked and concave from nostril to eye. Snout pointed. Nostrils not visible from above. Eyelid spine absent. Throat of adult males white with light grey freckles on the anterior third, without any spinulae. Femoral glands hardly distinguishable but present in adult males. Two ridges in the scapular region and two short, oblique ridges behind the eyes. These four ridges may be all disjointed, the two scapular ridges may be jointed to form a chevron shape, or the ridges may be jointed laterally in an hourglass shape.
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</paragraph>
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</subSubSection>
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<subSubSection pageId="0" pageNumber="53" type="comparisons">
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<paragraph pageId="0" pageNumber="53">
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<taxonomicName genus="Comparisons" lsidName="Comparisons" pageId="0" pageNumber="53" rank="genus">Comparisons</taxonomicName>
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.
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</paragraph>
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<paragraph pageId="0" pageNumber="53">
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This species can easily be distinguished from other Ethiopian
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<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
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: the body is slenderer, the hind limbs, fingers and toes are longer than all four other described Ethiopian
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<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
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,
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<taxonomicName lsidName="P. bullans" pageId="0" pageNumber="53" rank="species" species="bullans">P. bullans</taxonomicName>
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,
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<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
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,
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<taxonomicName lsidName="P. minutus" pageId="0" pageNumber="53" rank="species" species="minutus">P. minutus</taxonomicName>
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, and
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<taxonomicName lsidName="P. inexpectatus" pageId="0" pageNumber="53" rank="species" species="inexpectatus">P. inexpectatus</taxonomicName>
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. Additionally, the tips of fingers and toes are more enlarged than in any of these species, particularly in females. It is similar in size as
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<taxonomicName lsidName="P. minutus" pageId="0" pageNumber="53" rank="species" species="minutus">P. minutus</taxonomicName>
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, slightly larger than
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<taxonomicName lsidName="P. inexpectatus" pageId="0" pageNumber="53" rank="species" species="inexpectatus">P. inexpectatus</taxonomicName>
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and much smaller than
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<taxonomicName lsidName="P. bullans" pageId="0" pageNumber="53" rank="species" species="bullans">P. bullans</taxonomicName>
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and
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<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
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. It can be further distinguished from
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<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
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by a more marked canthus rostralis and a completely hidden tympanum. Adult males of
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<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
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sp. n. can be distinguished from
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<taxonomicName lsidName="P. bullans" pageId="0" pageNumber="53" rank="species" species="bullans">P. bullans</taxonomicName>
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,
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<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
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,
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<taxonomicName lsidName="P. minutus" pageId="0" pageNumber="53" rank="species" species="minutus">P. minutus</taxonomicName>
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, and
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<taxonomicName lsidName="P. inexpectatus" pageId="0" pageNumber="53" rank="species" species="inexpectatus">P. inexpectatus</taxonomicName>
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by the white coloration of their throat (
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<bibRefCitation author="Largen, MJ" journalOrPublisher="Tropical Zoology" pageId="0" pageNumber="53" pagination="287 - 306" title="The status of the genus Phrynobatrachus Guenther 1862 in Ethiopia and Eritrea, including description of a new species (Amphibia, Anura, Ranidae)." url="https://doi.org/10.1080/03946975.2001.10531158" volume="14" year="2001">Largen 2001</bibRefCitation>
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,
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<bibRefCitation author="Crutsinger, G" journalOrPublisher="African Zoology" pageId="0" pageNumber="53" pagination="19 - 23" title="A new species of Phrynobatrachus (Anura: Ranidae) from Tanzania." volume="39" year="2004">Crutsinger et al. 2004</bibRefCitation>
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). The new species is also distinct from the three junior synonyms of
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<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
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described from Ethiopia:
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<taxonomicName class="Amphibia" family="Arthroleptidae" genus="Arthroleptis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Arthroleptis bottegi" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bottegi">Arthroleptis bottegi</taxonomicName>
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Boulenger, 1895, described from Auta, Bale province, possesses a distinct tympanum, a rounded snout, and is much larger than
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<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
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(SVL of female holotype = 31 mm vs. 20.3
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<normalizedToken originalValue="±">+/-</normalizedToken>
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0.9 mm in female
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<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
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).
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<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus sciangallarum" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="sciangallarum">Phrynobatrachus sciangallarum</taxonomicName>
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Scortecci, 1943, described from a presumably adult male from Murle, Gemu-Gofa province, differs from
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<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
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by the presence of a considerably darkened throat and the lack of femoral glands. Additionally, it has more extensive foot webbing than
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<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
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(
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<bibRefCitation author="Largen, MJ" journalOrPublisher="Tropical Zoology" pageId="0" pageNumber="53" pagination="287 - 306" title="The status of the genus Phrynobatrachus Guenther 1862 in Ethiopia and Eritrea, including description of a new species (Amphibia, Anura, Ranidae)." url="https://doi.org/10.1080/03946975.2001.10531158" volume="14" year="2001">Largen 2001</bibRefCitation>
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).
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<taxonomicName genus="Arthroleptis-Phrynobatrachus" lsidName="Arthroleptis-Phrynobatrachus zavattarii" pageId="0" pageNumber="53" rank="species" species="zavattarii">Arthroleptis-Phrynobatrachus zavattarii</taxonomicName>
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Scortecci, 1943, described from Caschei, Gemu-Gofa province, only differs from
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<taxonomicName lsidName="P. sciangallarum" pageId="0" pageNumber="53" rank="species" species="sciangallarum">P. sciangallarum</taxonomicName>
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by the extent of the foot webbing, and all measurements of the type specimens fall within the range of
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<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
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(
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<bibRefCitation author="Largen, MJ" journalOrPublisher="Tropical Zoology" pageId="0" pageNumber="53" pagination="287 - 306" title="The status of the genus Phrynobatrachus Guenther 1862 in Ethiopia and Eritrea, including description of a new species (Amphibia, Anura, Ranidae)." url="https://doi.org/10.1080/03946975.2001.10531158" volume="14" year="2001">Largen 2001</bibRefCitation>
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). Blackburn (2014) provided color photographs of the holotypes of these last two junior synonyms of
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<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
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. Based on those photographs, it appears that
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<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
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has a slenderer head and more elongated fingers than these individuals. Finally, the type localities for these three junior synonyms of
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<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
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are in the southern part of the country, an area that is dry and at low elevation, in contrast with the habitat occupied by
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<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
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sp. n.
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</paragraph>
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<paragraph pageId="0" pageNumber="53">
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Among Eastern African
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<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
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,
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<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
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sp. n. can be distinguished from
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<taxonomicName lsidName="P. acridoides" pageId="0" pageNumber="53" rank="species" species="acridoides">P. acridoides</taxonomicName>
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(Cope, 1867),
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<taxonomicName lsidName="P. auritus" pageId="0" pageNumber="53" rank="species" species="auritus">P. auritus</taxonomicName>
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Boulenger, 1900,
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<taxonomicName lsidName="P. dendrobates" pageId="0" pageNumber="53" rank="species" species="dendrobates">P. dendrobates</taxonomicName>
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(Boulenger, 1919),
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<taxonomicName lsidName="P. graueri" pageId="0" pageNumber="53" rank="species" species="graueri">P. graueri</taxonomicName>
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(Nieden, 1911),
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<taxonomicName lsidName="P. keniensis" pageId="0" pageNumber="53" rank="species" species="keniensis">P. keniensis</taxonomicName>
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Barbour & Loveridge, 1928,
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<taxonomicName lsidName="P. krefftii" pageId="0" pageNumber="53" rank="species" species="krefftii">P. krefftii</taxonomicName>
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Boulenger, 1909,
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<taxonomicName lsidName="P. perpalmatus" pageId="0" pageNumber="53" rank="species" species="perpalmatus">P. perpalmatus</taxonomicName>
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Boulenger, 1898,
|
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<taxonomicName lsidName="P. petropedetoides" pageId="0" pageNumber="53" rank="species" species="petropedetoides">P. petropedetoides</taxonomicName>
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Ahl, 1924, and
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<taxonomicName lsidName="P. versicolor" pageId="0" pageNumber="53" rank="species" species="versicolor">P. versicolor</taxonomicName>
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Ahl, 1924, by a completely hidden tympanum (
|
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<bibRefCitation author="Cope, ED" journalOrPublisher="Journal of the Academy of Natural Sciences of Philadelphia" pageId="0" pageNumber="53" pagination="189 - 206" title="On the families of raniform Anura." volume="6" year="1867">Cope 1867</bibRefCitation>
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,
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<bibRefCitation author="Boulenger, GA" journalOrPublisher=", British Museum (Natural History), London" pageId="0" pageNumber="53" title="Catalogue of the BatrachiaSalientia. Ecaudata in the collection of the British Museum. 2 d ed." year="1882">Boulenger 1882</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Boulenger, GA" journalOrPublisher="Annals and Magazine of Natural History" pageId="0" pageNumber="53" pagination="496 - 497" title="Descriptions of three new frogs discovered by Dr. P. Krefft in Usambara, German East Africa." url="https://doi.org/10.1080/00222930908692705" volume="4" year="1909">1909</bibRefCitation>
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||
,
|
||
<bibRefCitation author="Razzetti, E" journalOrPublisher="Pubblinova Edizioni Negri and Instituto OIKOS, city" pageId="0" pageNumber="53" title="Field guide to the amphibians and reptiles of Arusha national park (Tanzania)." year="2002">Razzetti and Msuya 2002</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Pickersgill, M" journalOrPublisher="Serpent's Tale NHBD / Edition Chimaira, Frankfurt am Main" pageId="0" pageNumber="53" title="Frog search results of expeditions to southern and eastern Africa." year="2007">Pickersgill 2007</bibRefCitation>
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). This species can further be distinguished from
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<taxonomicName lsidName="P. auritus" pageId="0" pageNumber="53" rank="species" species="auritus">P. auritus</taxonomicName>
|
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Boulenger, 1900,
|
||
<taxonomicName lsidName="P. dendrobates" pageId="0" pageNumber="53" rank="species" species="dendrobates">P. dendrobates</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. graueri" pageId="0" pageNumber="53" rank="species" species="graueri">P. graueri</taxonomicName>
|
||
,
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<taxonomicName lsidName="P. kinagopensis" pageId="0" pageNumber="53" rank="species" species="kinagopensis">P. kinagopensis</taxonomicName>
|
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Angel, 1924,
|
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<taxonomicName lsidName="P. krefftii" pageId="0" pageNumber="53" rank="species" species="krefftii">P. krefftii</taxonomicName>
|
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,
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<taxonomicName lsidName="P. mababiensis" pageId="0" pageNumber="53" rank="species" species="mababiensis">P. mababiensis</taxonomicName>
|
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FritzSimons, 1932,
|
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<taxonomicName lsidName="P. parvulus" pageId="0" pageNumber="53" rank="species" species="parvulus">P. parvulus</taxonomicName>
|
||
(Boulenger, 1905),
|
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<taxonomicName lsidName="P. rouxi" pageId="0" pageNumber="53" rank="species" species="rouxi">P. rouxi</taxonomicName>
|
||
(Nieden,1912),
|
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<taxonomicName lsidName="P. scheffleri" pageId="0" pageNumber="53" rank="species" species="scheffleri">P. scheffleri</taxonomicName>
|
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(Nieden, 1911), and
|
||
<taxonomicName lsidName="P. ukigensis" pageId="0" pageNumber="53" rank="species" species="ukigensis">P. ukigensis</taxonomicName>
|
||
Grandison & Howell, 1983, by the white and light grey coloration of adult
|
||
<normalizedToken originalValue="males’">males'</normalizedToken>
|
||
throat (
|
||
<bibRefCitation author="Boulenger, GA" journalOrPublisher="Annals and Magazine of Natural History" pageId="0" pageNumber="53" pagination="496 - 497" title="Descriptions of three new frogs discovered by Dr. P. Krefft in Usambara, German East Africa." url="https://doi.org/10.1080/00222930908692705" volume="4" year="1909">Boulenger 1909</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Angel, F" journalOrPublisher="Bulletin du Museum national d'Histoire naturelle" pageId="0" pageNumber="53" pagination="130 - 132" title="Description de deux Batraciens nouveaux, d'Afrique Orientale anglaise, appartenant au genre Phrynobatrachus (Mission Alluaud et Jeannel, 1911 - 1912)." volume="30" year="1924">Angel 1924</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="FritzSimons, V" journalOrPublisher="Annals of the Transvaal Museum" pageId="0" pageNumber="53" pagination="35 - 40" title="Preliminary description of new forms of South African Reptilia and Amphibia, from the Vernay-Lang Kalahari Expedition, 1930." volume="15" year="1932">FritzSimons 1932</bibRefCitation>
|
||
). The new species differs from
|
||
<taxonomicName lsidName="P. bullans" pageId="0" pageNumber="53" rank="species" species="bullans">P. bullans</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. kakamikro" pageId="0" pageNumber="53" rank="species" species="kakamikro">P. kakamikro</taxonomicName>
|
||
Schick, Zimkus, Channing,
|
||
<normalizedToken originalValue="Köhler">Koehler</normalizedToken>
|
||
, &
|
||
<normalizedToken originalValue="Lötters">Loetters</normalizedToken>
|
||
, 2010, and
|
||
<taxonomicName lsidName="P. perpalmatus" pageId="0" pageNumber="53" rank="species" species="perpalmatus">P. perpalmatus</taxonomicName>
|
||
by the presence of femoral glands in adult males (
|
||
<bibRefCitation author="Crutsinger, G" journalOrPublisher="African Zoology" pageId="0" pageNumber="53" pagination="19 - 23" title="A new species of Phrynobatrachus (Anura: Ranidae) from Tanzania." volume="39" year="2004">Crutsinger et al. 2004</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Pickersgill, M" journalOrPublisher="Serpent's Tale NHBD / Edition Chimaira, Frankfurt am Main" pageId="0" pageNumber="53" title="Frog search results of expeditions to southern and eastern Africa." year="2007">Pickersgill 2007</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Schick, S" journalOrPublisher="Salamandra" pageId="0" pageNumber="53" pagination="24 - 36" title="Systematics of " Little Brown Frogs " from East Africa: recognition of Phrynobatrachusscheffleri and description of a new species from the Kakamega Forest, Kenya (Amphibia: Phrynobatrachidae)." volume="46" year="2010">Schick et al. 2010</bibRefCitation>
|
||
). The new species is also larger than
|
||
<taxonomicName lsidName="P. pallidus" pageId="0" pageNumber="53" rank="species" species="pallidus">P. pallidus</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. parvulus" pageId="0" pageNumber="53" rank="species" species="parvulus">P. parvulus</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. scheffleri" pageId="0" pageNumber="53" rank="species" species="scheffleri">P. scheffleri</taxonomicName>
|
||
, and
|
||
<taxonomicName lsidName="P. ungujae" pageId="0" pageNumber="53" rank="species" species="ungujae">P. ungujae</taxonomicName>
|
||
Pickersgill, 2007 (
|
||
<bibRefCitation author="Boulenger, GA" journalOrPublisher="Annals and Magazine of Natural History; Zoology, Botany, and Geology" pageId="0" pageNumber="53" pagination="105 - 115" title="A list of the batrachians and reptiles collected by Dr. W. J. Ansorge in Angola with descriptions of new species." url="https://doi.org/10.1080/03745480509443656" volume="16" year="1905">Boulenger 1905</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Pickersgill, M" journalOrPublisher="Serpent's Tale NHBD / Edition Chimaira, Frankfurt am Main" pageId="0" pageNumber="53" title="Frog search results of expeditions to southern and eastern Africa." year="2007">Pickersgill 2007</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Schick, S" journalOrPublisher="Salamandra" pageId="0" pageNumber="53" pagination="24 - 36" title="Systematics of " Little Brown Frogs " from East Africa: recognition of Phrynobatrachusscheffleri and description of a new species from the Kakamega Forest, Kenya (Amphibia: Phrynobatrachidae)." volume="46" year="2010">Schick et al. 2010</bibRefCitation>
|
||
), and possesses longer legs than
|
||
<taxonomicName lsidName="P. kakamikro" pageId="0" pageNumber="53" rank="species" species="kakamikro">P. kakamikro</taxonomicName>
|
||
and
|
||
<taxonomicName lsidName="P. pallidus" pageId="0" pageNumber="53" rank="species" species="pallidus">P. pallidus</taxonomicName>
|
||
Pickersgill, 2007, as well as longer feet than
|
||
<taxonomicName lsidName="P. pallidus" pageId="0" pageNumber="53" rank="species" species="pallidus">P. pallidus</taxonomicName>
|
||
and
|
||
<taxonomicName lsidName="P. ungujae" pageId="0" pageNumber="53" rank="species" species="ungujae">P. ungujae</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Pickersgill, M" journalOrPublisher="Serpent's Tale NHBD / Edition Chimaira, Frankfurt am Main" pageId="0" pageNumber="53" title="Frog search results of expeditions to southern and eastern Africa." year="2007">Pickersgill 2007</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Schick, S" journalOrPublisher="Salamandra" pageId="0" pageNumber="53" pagination="24 - 36" title="Systematics of " Little Brown Frogs " from East Africa: recognition of Phrynobatrachusscheffleri and description of a new species from the Kakamega Forest, Kenya (Amphibia: Phrynobatrachidae)." volume="46" year="2010">Schick et al. 2010</bibRefCitation>
|
||
). Finally,
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
sp. n. can be distinguished from
|
||
<taxonomicName lsidName="P. acridoides" pageId="0" pageNumber="53" rank="species" species="acridoides">P. acridoides</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. bullans" pageId="0" pageNumber="53" rank="species" species="bullans">P. bullans</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. mababiensis" pageId="0" pageNumber="53" rank="species" species="mababiensis">P. mababiensis</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. pallidus" pageId="0" pageNumber="53" rank="species" species="pallidus">P. pallidus</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. ungujae" pageId="0" pageNumber="53" rank="species" species="ungujae">P. ungujae</taxonomicName>
|
||
, and
|
||
<taxonomicName lsidName="P. versicolor" pageId="0" pageNumber="53" rank="species" species="versicolor">P. versicolor</taxonomicName>
|
||
Ahl, 1924, by the absence of dark bars on the lower jaw (
|
||
<bibRefCitation author="Cope, ED" journalOrPublisher="Journal of the Academy of Natural Sciences of Philadelphia" pageId="0" pageNumber="53" pagination="189 - 206" title="On the families of raniform Anura." volume="6" year="1867">Cope 1867</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Boulenger, GA" journalOrPublisher=", British Museum (Natural History), London" pageId="0" pageNumber="53" title="Catalogue of the BatrachiaSalientia. Ecaudata in the collection of the British Museum. 2 d ed." year="1882">Boulenger 1882</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="FritzSimons, V" journalOrPublisher="Annals of the Transvaal Museum" pageId="0" pageNumber="53" pagination="35 - 40" title="Preliminary description of new forms of South African Reptilia and Amphibia, from the Vernay-Lang Kalahari Expedition, 1930." volume="15" year="1932">Fritzsimons 1932</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Crutsinger, G" journalOrPublisher="African Zoology" pageId="0" pageNumber="53" pagination="19 - 23" title="A new species of Phrynobatrachus (Anura: Ranidae) from Tanzania." volume="39" year="2004">Crutsinger et al. 2004</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Pickersgill, M" journalOrPublisher="Serpent's Tale NHBD / Edition Chimaira, Frankfurt am Main" pageId="0" pageNumber="53" title="Frog search results of expeditions to southern and eastern Africa." year="2007">Pickersgill 2007</bibRefCitation>
|
||
).
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="53" type="description">
|
||
<paragraph pageId="0" pageNumber="53">Description of the holotype.</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">
|
||
Adult male (Figure 2; Table 1), body slender, head longer than wide (HW/HL = 0.96), the snout is pointed and the canthus rostralis is marked and concave from nostril to eye. Tympanum not visible. Pupil horizontal. Eyelid spine absent. Maxillary teeth present, vomerine teeth absent. Tongue elongate, free for two thirds of its length, tip divided in two short lobes. Legs and feet long for the genus (TL/SVL = 0.6; FL/SVL = 0.6). Tarsal tubercle and outer metatarsal tubercle present, but inner metatarsal tubercle not distinguishable. Subarticular tubercles absent on the hands and small and barely distinguishable on the feet. Palmar tubercle absent. Fingers and toes elongated and their tips slightly swollen but not forming discs. First fingers shorter and swollen but nuptial pads absent. Fingers free of webbing. No webbing between toe I and II, and minimal webbing between the other toes. Finger formula: I<II<IV<III. Toe formula: I<II<V<III<IV. Skin of dorsum and dorsal side of limbs covered by small pointy asperities. Femoral glands hardly visible but present (Suppl. material 2: Figure S1B). Chevron-shaped gland in scapular region extending from above the shoulder girdle to approx. mid-body. Two rows of small warts form faint ridges between the eye and the scapular region but do not reach the chevron-shaped gland. The skin of the throat is not thin and loose as in many other
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
|
||
species and gular folds and spinulae are absent.
|
||
</paragraph>
|
||
<caption pageId="0" pageNumber="53">
|
||
<paragraph pageId="0" pageNumber="53">
|
||
Table 1. Morphometric measurements of
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita</taxonomicName>
|
||
sp. n.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="0" pageNumber="53">
|
||
<table pageId="0" pageNumber="53">
|
||
<tr pageId="0" pageNumber="53">
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">Specimen</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 440</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 421</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 418</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 419</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 420</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 424</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 425</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 426</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 427</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">SB 439</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">Males (n = 2)</th>
|
||
<th colspan="1" pageId="0" pageNumber="53" rowspan="1">Females (n = 8)</th>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">SVL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">HW</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">HL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">SL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">NS</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">IND</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">EN</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">IOD</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">ED</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">UEW</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">FLL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">HAL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">FinDW</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">THL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">TL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">FL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">Toe4DW</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">MTL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">TLSVL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">HWHL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">ENIND</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">HWSVL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">FLTL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">FLSVL</td>
|
||
</tr>
|
||
<tr pageId="0" pageNumber="53">
|
||
<td colspan="1" pageId="0" pageNumber="53" rowspan="1">HLSVL</td>
|
||
</tr>
|
||
</table>
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="53" type="coloration of holotype in life">
|
||
<paragraph pageId="0" pageNumber="53">Coloration of holotype in life.</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">The body is golden (Figure 2A); a dark chevron is present in the scapular region underlining two short back ridges. A faint dark triangle is present between the eyes and a faint dark bar is present between the eye and the shoulder. The upper eyelids are darker than the rest of the head. Loreal region faintly darker than the rest of the head. Upper and lower jaws lack any blotching. The flanks and the sides of the head present numerous small white spots, extending to the upper lip below the eye. Iris dark gold with a golden ring around the iris interrupted by a black dot at the bottom of the iris. The thighs and legs are faintly and irregularly banded. The back of the thighs is yellowish with irregular coverage of melanophores. A dark triangle is present in the vent region. Ventral region is white molted with light grey. The throat is white, with the anterior third light grey with white freckles. Ventral side of the thighs and legs shows irregular melanophores and white spots.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="53" type="coloration of holotype in preservative">
|
||
<paragraph pageId="0" pageNumber="53">Coloration of holotype in preservative.</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">
|
||
After euthanasia, the
|
||
<normalizedToken originalValue="individual’s">individual's</normalizedToken>
|
||
coloration darkened notably (Figure 2B). In preservative, the dorsum presents a grey coloration.
|
||
</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="53" type="variation">
|
||
<paragraph pageId="0" pageNumber="53">Variation.</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">Morphometric variations of the type series are summarized in Table 1. Inner metatarsal tubercles are not visible in all individuals. Subarticular tubercles are more pronounced in some individuals. Webbing of the feet presents small variation, ranging from minimal to no webbing for toes I and II, and webbed to the first phalange for toes III to V. The dorsal and leg skin is mostly smooth with small warts more pronounced in some individuals. The two ridges in the scapular region and two oblique ridges behind the eyes were visible in all examined specimens. In four specimens, they formed a chevron in the scapular region, in three specimens the ridges jointed laterally, forming an hourglass, and in two specimens all four ridges were disjointed.</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">
|
||
Males and females differ in size and in proportions (Figure 2). Finger and toe tips are enlarged in females and fingertips are slightly enlarged in males.
|
||
<normalizedToken originalValue="Males’">Males'</normalizedToken>
|
||
first finger is short and swollen but no distinct nuptial pad is visible. Adult males present femoral glands, which are lacking in females. Bicolor eggs are visible through the skin of gravid females.
|
||
</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">Coloration of the body varies from golden to light brown, with few large light green blotches in some individuals. Most specimens present a dark chevron in the scapular region underlining two short back ridges that are either jointed or disjointed with two oblique ridges between the back of the eye and the shoulders. A more or less pronounced dark stripe is present between the nostril and the eye, and between the eye and the arm. A more or less distinctive dark bar is present between the eyes and some individuals present a lighter or green snout. The flanks and the sides of the head present numerous small white spots, extending to the upper lip in some individuals. The thighs and legs of some individuals are very faintly and irregularly banded. The back of the thighs is cream with small light grey spots. Vocal sac in adult males is white with the frontal third of the throat light grey with white freckles, while in females the throat and the ventral skin is light grey or yellowish molted with white. Some individuals have a thin light line on the backside of the thigh from the vent to the tarsal tubercle. A dark triangle is generally present in the back of the thigh, around the vent.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="53" type="etymology">
|
||
<paragraph pageId="0" pageNumber="53">Etymology.</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">The specific name refers to Bibita Mountain, the type and only known locality for the species. It is an invariable noun used in apposition.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="53" type="habitat">
|
||
<paragraph pageId="0" pageNumber="53">Habitat, distribution, and natural history.</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">
|
||
All individuals were collected in a single large overgrown forest pond (Figure 3A), at night. The surrounding forest consisted of large trees with overhanging epiphytes and dense undergrowth. All females and the amplected pair were found on vegetation ca. 30 cm above water (Figure 3B). A single male was found in the water, presumably while calling. All collected females were gravid, and bicolor eggs were visible through the skin. Females seemed to aggregate in specific areas of the pond, were numerous egg clutches were found on leaves overhanging the water (Figure 3B). Laying eggs on vegetation overhanging the water is unusual in
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
|
||
, most species laying their eggs directly in the water (
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Molecular Biology and Evolution" pageId="0" pageNumber="53" url="https://doi.org/10.1371/journal.pone.0035118" year="2012">Zimkus et al. 2012</bibRefCitation>
|
||
). We thus confirmed that these eggs belonged to
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita</taxonomicName>
|
||
sp. n. by sequencing their mitochondrial rRNA 16s. Various forms of terrestrial egg deposition have been described in the genus
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
|
||
(
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Molecular Biology and Evolution" pageId="0" pageNumber="53" url="https://doi.org/10.1371/journal.pone.0035118" year="2012">Zimkus et al. 2012</bibRefCitation>
|
||
): most similarly to
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. sandersoni" pageId="0" pageNumber="53" rank="species" species="sandersoni">P. sandersoni</taxonomicName>
|
||
(Parker, 1935) lays its eggs on vegetation up to 2 m above small puddles, small streams or water-saturated soil (
|
||
<bibRefCitation author="Amiet, J-L" journalOrPublisher="Amphibia-Reptilia" pageId="0" pageNumber="53" pagination="1 - 13" title="Ecologie, ethologie et developpement de Phrynodonsandersoni Parker, 1939 (Amphibia, Anura, Ranidae)." url="https://doi.org/10.1163/156853881X00249" volume="2" year="1981">Amiet 1981</bibRefCitation>
|
||
) and
|
||
<taxonomicName lsidName="P. krefftii" pageId="0" pageNumber="53" rank="species" species="krefftii">P. krefftii</taxonomicName>
|
||
lays its eggs above the water, on rocks or vegetation (
|
||
<bibRefCitation pageId="0" pageNumber="53">Harper and Vonesh 2010</bibRefCitation>
|
||
).
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus guineensis" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="guineensis">Phrynobatrachus guineensis</taxonomicName>
|
||
<normalizedToken originalValue="Guibé">Guibe</normalizedToken>
|
||
& Lamotte, 1961 lays its eggs on the bark of trees above water-filled tree holes (
|
||
<bibRefCitation pageId="0" pageNumber="53">
|
||
<normalizedToken originalValue="Rödel">Roedel</normalizedToken>
|
||
1998
|
||
</bibRefCitation>
|
||
) and
|
||
<taxonomicName lsidName="P. dendrobates" pageId="0" pageNumber="53" rank="species" species="dendrobates">P. dendrobates</taxonomicName>
|
||
lays its eggs in tree holes or above streams (
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Molecular Biology and Evolution" pageId="0" pageNumber="53" url="https://doi.org/10.1371/journal.pone.0035118" year="2012">Zimkus et al. 2012</bibRefCitation>
|
||
). Finally,
|
||
<taxonomicName lsidName="P. phyllophilus" pageId="0" pageNumber="53" rank="species" species="phyllophilus">P. phyllophilus</taxonomicName>
|
||
<normalizedToken originalValue="Rödel">Roedel</normalizedToken>
|
||
& Ernst, 2002,
|
||
<taxonomicName lsidName="P. tokba" pageId="0" pageNumber="53" rank="species" species="tokba">P. tokba</taxonomicName>
|
||
(Chabanaud, 1921), and
|
||
<taxonomicName lsidName="P. villiersi" pageId="0" pageNumber="53" rank="species" species="villiersi">P. villiersi</taxonomicName>
|
||
<normalizedToken originalValue="Guibé">Guibe</normalizedToken>
|
||
, 1969 lay their eggs on the leaf litter or the forest floor (
|
||
<bibRefCitation author="Roedel, MO" journalOrPublisher="Molecular Ecology" pageId="0" pageNumber="53" url="https://doi.org/10.1670/0022-1511(2002)036[0121:ANRMFT]2.0.CO;2" year="2002 a">
|
||
<normalizedToken originalValue="Rödel">Roedel</normalizedToken>
|
||
and Ernst 2002a
|
||
</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Roedel, MO" journalOrPublisher="Molecular Ecology" pageId="0" pageNumber="53" url="https://doi.org/10.1670/0022-1511(2002)036[0561:ANPFTU]2.0.CO;2" year="2002 b">2002b</bibRefCitation>
|
||
;
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Molecular Biology and Evolution" pageId="0" pageNumber="53" url="https://doi.org/10.1371/journal.pone.0035118" year="2012">Zimkus et al. 2012</bibRefCitation>
|
||
).
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita</taxonomicName>
|
||
sp. n. thus adds to the diversity of reproductive modes in the genus.
|
||
</paragraph>
|
||
<caption pageId="0" pageNumber="53">
|
||
<paragraph pageId="0" pageNumber="53">
|
||
Figure 3. Habitat of
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita</taxonomicName>
|
||
sp. n. A Type locality of
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
sp. n. Overgrown pond in primary forest B Two females
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
sp. n. in situ, next to a clutch of eggs, in vegetation at ca. 30 cm above the water. Multiple females and egg clutches were found in similar circumstances.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="0" pageNumber="53">
|
||
Five egg clutches were photographed and contained approximately 30 eggs each (range 22 to 33). All observed clutches were found between 20 - 40 cm above water on vegetation, and up to two clutches were found on a single leaf. When laid, the eggs are bicolor, heavily pigmented, and encased in a thin gelatinous layer; as the egg develops, the pigmentation is more evenly distributed at its surface, with dark brown freckle, and the gelatinous layer becomes much thicker. It is possible that female
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
sp. n. are guarding their egg clutches in a similar manner as
|
||
<taxonomicName lsidName="P. sandersoni" pageId="0" pageNumber="53" rank="species" species="sandersoni">P. sandersoni</taxonomicName>
|
||
females, which attend their eggs at night by standing over them or staying in the close proximity (
|
||
<bibRefCitation author="Amiet, J-L" journalOrPublisher="Amphibia-Reptilia" pageId="0" pageNumber="53" pagination="1 - 13" title="Ecologie, ethologie et developpement de Phrynodonsandersoni Parker, 1939 (Amphibia, Anura, Ranidae)." url="https://doi.org/10.1163/156853881X00249" volume="2" year="1981">Amiet 1981</bibRefCitation>
|
||
). Additional behavioral observations are necessary to determine whether females
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
sp. n. truly display such parental care. Other frog species were found in the same pond or nearby, including
|
||
<taxonomicName class="Amphibia" family="Hyperoliidae" genus="Afrixalus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Afrixalus enseticola" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="enseticola">Afrixalus enseticola</taxonomicName>
|
||
Largen, 1974,
|
||
<taxonomicName class="Amphibia" family="Arthroleptidae" genus="Leptopelis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Leptopelis vannutellii" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="vannutellii">Leptopelis vannutellii</taxonomicName>
|
||
(Boulenger, 1898),
|
||
<taxonomicName class="Amphibia" family="Hyperoliidae" genus="Paracassina" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Paracassina obscura" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="obscura">Paracassina obscura</taxonomicName>
|
||
(Boulenger, 1895), and
|
||
<taxonomicName class="Amphibia" family="Ptychadenidae" genus="Ptychadena" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Ptychadena erlangeri" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="erlangeri">Ptychadena cf. erlangeri</taxonomicName>
|
||
(Ahl, 1924).
|
||
</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">The forest in Bibita Mountain appears to be well preserved, as there are no settlements inside the forest, and a footpath is the only way to cross the forest. People from the surrounded villages harvest wild honey from the forest, but this is the only noticeable human activity there. Based on satellite imagery, no human disturbance is apparent at the higher elevations of the mountain; however, most areas below 1900 m have been transformed into agricultural land.</paragraph>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="53" type="call">
|
||
<paragraph pageId="0" pageNumber="53">Call.</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">
|
||
The advertisement call is composed of a series of pulsed notes with a slight upward frequency modulation within each note (Figure 4A). It is similar to that of
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus minutus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="minutus">Phrynobatrachus minutus</taxonomicName>
|
||
(Figure 4B) in spectral structure, note length, and dominant frequency. The vocalization of
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
sp. n. can be distinguished from that of
|
||
<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
|
||
(Figure 4C) by its higher dominant frequency, indicative of its smaller body size. Dominant frequency: 3318
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
94 Hz. Note duration = 630
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
80 ms. Interval inter-notes = 1085
|
||
<normalizedToken originalValue="±">+/-</normalizedToken>
|
||
630 ms. The mediocre quality of our recording prevents us from analyzing the advertisement call of
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
sp. n. in further details, and a more complete description of their vocalization will be needed in the future, in particular to distinguish it from other species of dwarf
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
<caption pageId="0" pageNumber="53">
|
||
<paragraph pageId="0" pageNumber="53">
|
||
Figure 4. Advertisement call of
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita</taxonomicName>
|
||
sp. n. Spectrograms (upper panels) and sonograms (lower panels; relative amplitude) of Ethiopian
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
|
||
advertisement calls. A
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita</taxonomicName>
|
||
sp. n. (specimen not collected) B
|
||
<taxonomicName lsidName="P. minutus" pageId="0" pageNumber="53" rank="species" species="minutus">P. minutus</taxonomicName>
|
||
(SB233) C
|
||
<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
|
||
(specimen not collected).
|
||
</paragraph>
|
||
</caption>
|
||
</subSubSection>
|
||
<subSubSection pageId="0" pageNumber="53" type="estimates of evolutionary relationships">
|
||
<paragraph pageId="0" pageNumber="53">Estimates of evolutionary relationships.</paragraph>
|
||
<paragraph pageId="0" pageNumber="53">
|
||
The goal of our phylogenetic analysis was to test the phylogenetic position of the new species, not to obtain a phylogeny of the genus, which has been previously done elsewhere (
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="53" pagination="883 - 900" title="Complex patterns of continental speciation: molecular phylogenetics and biogeography of sub-Saharan puddle frogs (Phrynobatrachus)." url="https://doi.org/10.1016/j.ympev.2009.12.012" volume="55" year="2010">Zimkus et al. 2010</bibRefCitation>
|
||
). Our phylogenetic estimates recovered three main lineages in the genus
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
|
||
(Figure 5); however, several of the nodes have low support. We briefly describe the resulting topology below.
|
||
</paragraph>
|
||
<caption pageId="0" pageNumber="53">
|
||
<paragraph pageId="0" pageNumber="53">
|
||
Figure 5. Phylogenetic placement of
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita</taxonomicName>
|
||
sp. n. Bayesian phylogenetic inference of the genus
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
|
||
based on the mitochondrial rRNA 16s. Nodes with a posterior support of 1 are marked with a black circle and nodes with high posterior support (>0.95) are marked with a white circle. Individuals of
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
|
||
species known to occur in Ethiopia are shown in boldface. Photos of Ethiopian representatives are displayed, from top to bottom:
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus minutus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="minutus">Phrynobatrachus minutus</taxonomicName>
|
||
(SB175; Kibre Mengist),
|
||
<taxonomicName lsidName="P. inexpectatus" pageId="0" pageNumber="53" rank="species" species="inexpectatus">P. inexpectatus</taxonomicName>
|
||
(SB143; Magnete, Harenna forest),
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
sp. n. (SB440; male holotype),
|
||
<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
|
||
(SB454; Mizan Teferi).
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph pageId="0" pageNumber="53">
|
||
We recovered a basal group of
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="genus">Phrynobatrachus</taxonomicName>
|
||
, which included the species
|
||
<taxonomicName lsidName="P. acutirostris" pageId="0" pageNumber="53" rank="species" species="acutirostris">P. acutirostris</taxonomicName>
|
||
Nieden, 1912,
|
||
<taxonomicName lsidName="P. krefftii" pageId="0" pageNumber="53" rank="species" species="krefftii">P. krefftii</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. dendrobates" pageId="0" pageNumber="53" rank="species" species="dendrobates">P. dendrobates</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. petropedetoides" pageId="0" pageNumber="53" rank="species" species="petropedetoides">P. petropedetoides</taxonomicName>
|
||
Ahl, 1924, and
|
||
<taxonomicName lsidName="P. sandersoni" pageId="0" pageNumber="53" rank="species" species="sandersoni">P. sandersoni</taxonomicName>
|
||
. This clade is consistent with the "Clade A" of
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="53" pagination="883 - 900" title="Complex patterns of continental speciation: molecular phylogenetics and biogeography of sub-Saharan puddle frogs (Phrynobatrachus)." url="https://doi.org/10.1016/j.ympev.2009.12.012" volume="55" year="2010">Zimkus et al. (2010)</bibRefCitation>
|
||
, however this clade received low support (posterior probability (pp) <0.5). All remaining species in the genus were grouped into two major clades, which received strong support (pp = 1). The first one of these two groups is similar to the "Clade B" of
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="53" pagination="883 - 900" title="Complex patterns of continental speciation: molecular phylogenetics and biogeography of sub-Saharan puddle frogs (Phrynobatrachus)." url="https://doi.org/10.1016/j.ympev.2009.12.012" volume="55" year="2010">Zimkus et al. (2010)</bibRefCitation>
|
||
, which includes the Ethiopian species
|
||
<taxonomicName lsidName="P. minutus" pageId="0" pageNumber="53" rank="species" species="minutus">P. minutus</taxonomicName>
|
||
,
|
||
<taxonomicName lsidName="P. inexpectatus" pageId="0" pageNumber="53" rank="species" species="inexpectatus">P. inexpectatus</taxonomicName>
|
||
, and
|
||
<taxonomicName genus="P." lsidName="P." pageId="0" pageNumber="53" rank="genus">P.</taxonomicName>
|
||
sp. n.
|
||
<normalizedToken originalValue="“Oromia”">"Oromia"</normalizedToken>
|
||
of
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Systematics and Biodiversity" pageId="0" pageNumber="53" pagination="39 - 47" title="Light at the end of the tunnel: insights into the molecular systematics of East African puddle frogs (Anura: Phrynobatrachidae)." url="https://doi.org/10.1080/14772000903543004" volume="8" year="2010">Zimkus and Schick (2010)</bibRefCitation>
|
||
, while the second group is consistent with the "Clade C" of
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="53" pagination="883 - 900" title="Complex patterns of continental speciation: molecular phylogenetics and biogeography of sub-Saharan puddle frogs (Phrynobatrachus)." url="https://doi.org/10.1016/j.ympev.2009.12.012" volume="55" year="2010">Zimkus et al. (2010)</bibRefCitation>
|
||
, and includes the Ethiopian species
|
||
<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
|
||
.
|
||
<taxonomicName class="Amphibia" family="Phrynobatrachidae" genus="Phrynobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Phrynobatrachus bibita" order="Anura" pageId="0" pageNumber="53" phylum="Chordata" rank="species" species="bibita">Phrynobatrachus bibita</taxonomicName>
|
||
sp. n. was not closely related to any of the other Ethiopian species, and instead basal to the last group ("Clade C" of
|
||
<bibRefCitation author="Zimkus, BM" journalOrPublisher="Molecular Phylogenetics and Evolution" pageId="0" pageNumber="53" pagination="883 - 900" title="Complex patterns of continental speciation: molecular phylogenetics and biogeography of sub-Saharan puddle frogs (Phrynobatrachus)." url="https://doi.org/10.1016/j.ympev.2009.12.012" volume="55" year="2010">Zimkus et al. (2010)</bibRefCitation>
|
||
), however with high support (pp = 1). Our analysis grouped the egg clutch collected (SB422) with the adults
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
sp. n. (holotype SB440 and paratopotypes SB419, SB420, SB425), ensuring that it belonged to the new species. We did not include samples of three junior synonyms of
|
||
<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
|
||
from Ethiopia (see
|
||
<taxonomicName genus="Comparisons" lsidName="Comparisons" pageId="0" pageNumber="53" rank="genus">Comparisons</taxonomicName>
|
||
), however, samples referable to these junior synonyms have previously been included on a phylogeny of members of the
|
||
<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
|
||
group (
|
||
<bibRefCitation pageId="0" pageNumber="53">Lara 2016</bibRefCitation>
|
||
), and they are well nested within
|
||
<taxonomicName lsidName="P. natalensis" pageId="0" pageNumber="53" rank="species" species="natalensis">P. natalensis</taxonomicName>
|
||
, we can thus conclude that they do not represent
|
||
<taxonomicName lsidName="P. bibita" pageId="0" pageNumber="53" rank="species" species="bibita">P. bibita</taxonomicName>
|
||
.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |