treatments-xml/data/40/75/99/407599A19958667C10123AF0DFD13FF7.xml

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<mods:title id="4EA9CED99A9F8A7323DCDE05DDDA4D94">Life in the spray zone - overlooked diversity in West African torrent-frogs (Anura, Odontobatrachidae, Odontobatrachus)</mods:title>
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<mods:namePart id="9E8DFCF86C2C92655D8DA956B63997B3">Brede, Christian</mods:namePart>
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<mods:date id="60B3967803FC68F4939B338106CD8ECA">2015</mods:date>
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<mods:number id="74D8470EE81EB22EAEAA588E711A4A12">91</mods:number>
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<treatment id="407599A19958667C10123AF0DFD13FF7" ID-GBIF-Taxon="127890132" LSID="urn:lsid:zoobank.org:act:542C46CE-2B91-41AC-8314-5F84469AED04" httpUri="http://treatment.plazi.org/id/407599A19958667C10123AF0DFD13FF7" lastPageId="23" lastPageNumber="138" pageId="19" pageNumber="134">
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<paragraph id="2909221F3261F0A1532F6DA9BB6C4963" pageId="19" pageNumber="134">Taxon classification Animalia Anura Odontobatrachidae</paragraph>
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<paragraph id="7FD0DBE952B8148473C2D4FC2B29BE88" pageId="19" pageNumber="134">
<taxonomicName id="C6A48D6295ED8578FB9A4CBF95751D7E" ID-CoL="48P5D" LSID="http://zoobank.org/542C46CE-2B91-41AC-8314-5F84469AED04" authority="Barej, Schmitz, Penner, Doumbia, Sandberger-Loua, Emmrich, Adeba &amp; Roedel" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">
Odontobatrachus arndti Barej, Schmitz, Penner, Doumbia, Sandberger-Loua, Emmrich, Adeba &amp; 
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<paragraph id="801D343C4E6CED0EAF0F1276107A3C5B" pageId="19" pageNumber="134">
<taxonomicName id="5E04C592B3D3A2EA09ED1C099571D91F" LSID="http://zoobank.org/542C46CE-2B91-41AC-8314-5F84469AED04" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
OTU4 sensu 
<bibRefCitation id="1A1C16DFE85A63B60E2313C9612A5D63" author="Barej, MF" journalOrPublisher="BMC Evolutionary Biology" pageId="25" pageNumber="140" title="Multiple genetic lineages challenge the monospecific status of the West African endemic frog-family Odontobatrachidae." url="10.1186/s12862-015-0346-9" volume="15" year="2015">Barej et al. (2015)</bibRefCitation>
</paragraph>
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<subSubSection id="93D42E49A1261F5AAE0F17FF66115CEC" pageId="19" pageNumber="134" type="holotype">
<paragraph id="53FBB61FF41CB674F621E0C477EB5301" pageId="19" pageNumber="134">Holotype.</paragraph>
<paragraph id="B29EF35845BB86CACC100F98E171CCFC" pageId="19" pageNumber="134">ZMB 78355 (male), Republic of Guinea, Nimba Mts., River Mandey (Latitude: 7.64786; Longitude: -8.42397), 694 m a.s.l., 18 June 2009, coll. L. Sandberger-Loua and J. Doumbia.</paragraph>
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<subSubSection id="59F86FF4DC3D0AD99E4199824CE24863" pageId="19" pageNumber="134" type="paratypes">
<paragraph id="943AB217BBDEECEDE4D3643666812366" pageId="19" pageNumber="134">Paratypes.</paragraph>
<paragraph id="342B431027416D3F8F21A3F2EA3351C8" pageId="19" pageNumber="134">
Guinea: MHNG 2731.49 (male), ZMB 78356 (female), 
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Region (7.65; -8.42), 670 m a.s.l., 18 June 2009; MHNG 2731.50, ZMB 78357 (2 females), 
<normalizedToken id="69FA81152B35A54B26940763C1D78E50" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.63; -8.41), 1121 m a.s.l., 4 November 2011; ZFMK 95467 (female), ZFMK 95468 (male), 
<normalizedToken id="47B745AE4435E6168BC1E896813F0A24" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.65; -8.42), 674 m a.s.l., 2 January 2011; ZMB 78354 (female), same data as holotype.
</paragraph>
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<subSubSection id="FCF10B36B9E5E75B176C3A345EE32FD7" pageId="19" pageNumber="134" type="materials_examined">
<paragraph id="9A8DDAFC5AAFDF91701EB1FB7B134CE8" pageId="19" pageNumber="134">Additional material.</paragraph>
<paragraph id="6BAA53249CB317A073FB10873B1CE2EE" pageId="19" pageNumber="134">
<normalizedToken id="CB71B8CA46F8560838B04C1C93406F38" originalValue="Côte">Cote</normalizedToken>
<normalizedToken id="2C6A146E89403E001E168CE38E2EA902" originalValue="d’Ivoire">d'Ivoire</normalizedToken>
: ZMB 78326, ZMB 78329 (3 females), ZMB 78327-8 (2 males), Dix-Huit Montagnes Region (7.85; -7.39), app. 500 m a.s.l. Liberia: ZMB 78332 (male), Nimba County (7.56; -8.64), 647 m a.s.l.; ZMB 78333-35 (3 males), Nimba County (7.48; -8.58), 513 m a.s.l. Guinea: ZMB 78336 (female), ZMB 78337-39 (3 males), 
<normalizedToken id="51DA2205C6758924C46495DA8A7873F4" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.61; -8.27), 400 m a.s.l.; ZMB 78340-41 (2 females), 
<normalizedToken id="DDDB0D0E507AEADE7F8FA6B8F76C35BA" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.61; -8.26), 460 m a.s.l.; ZMB 78342 (juvenile), 
<normalizedToken id="12417C4E549EAF9455302BD7649D94B1" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.70; -8.40), 751 m a.s.l.; ZMB 78343, ZMB 78345 (2 females), ZMB 78344, ZMB 78346 (2 males), 
<normalizedToken id="D0499A8285B67128F60F7206B0EB27A8" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.70; -8.40), 760 m a.s.l.; ZMB 78347 (male), 
<normalizedToken id="343BF79DBC5628356C103F675035D6EA" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.71; -8.41), 518 m a.s.l.; ZMB 78348 (male), ZMB 78349 (female), 
<normalizedToken id="8DE76B865E3553205D829A4F98A7C780" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.70; -8.40), 764 m a.s.l.; ZMB 78350-1 (2 females), ZMB 78352 (male), 
<normalizedToken id="B70863539DB02BD706D8472648B1E9C0" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.68; -8.39), 1027 m a.s.l.; ZMB 78353 (juvenile), 
<normalizedToken id="08894217E26E25694BBFC655B6A4841F" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.65; -8.42), 670 m a.s.l.; ZMB 78358-59 (2 males), 
<normalizedToken id="6FB4BE26EC312A52809BDB2B5AEC5343" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.65; -8.34), 577 m a.s.l.; ZMB 78360 (female), ZMB 78361 (male), 
<normalizedToken id="643D63E380C65495CD6CE4D2A5F41860" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.65; -8.36), 815 m a.s.l.; ZMB 78362 (female), ZMB 78363 (male), 
<normalizedToken id="C039A714B61518974B0C923D52ACAECF" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.63; -8.35), 652 m a.s.l.; ZMB 78364, ZMB 78367 (2 females), ZMB 78365-6 (2 males), 
<normalizedToken id="BAF22BB7295D1BBA3BDFBA24A6B501F0" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.65; -8.37), 949 m a.s.l.; ZMB 78368 (female), ZMB 78369 (male), 
<normalizedToken id="87A42F5AB48D2FC400DC6000512B7C52" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.67; -8.37), 1317 m a.s.l.; ZMB 78370 (male), ZMB 78371 (female), 
<normalizedToken id="59193BF872C72C01AC002F05D919C918" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.67; -8.37), 1234 m a.s.l.; ZMB 78372 (female), ZMB 78373 (male), 
<normalizedToken id="CB328D5F25039D335122F42CCC9100A8" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.62; -8.42), 1154 m a.s.l.; ZMB 78374 (female), ZMB 78375 (male), 
<normalizedToken id="F346D20F47D6853A762DE03721639999" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.62; -8.45), 701 m a.s.l.; ZMB 78376 (female), ZMB 78377 (male), 
<normalizedToken id="987630AB6B5EA38E43BA9ABD787BB6E8" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.63; -8.44), 750 m a.s.l.; ZMB 78378 (female), ZMB 78379 (male), 
<normalizedToken id="54D3BADE9E54FF6E57176F3A34E8AC74" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.67; -8.35), 786 m a.s.l.; ZMB 78380 (female), ZMB 78381 (male), 
<normalizedToken id="A079B6D6FD3FACE156A51D22977BD2B6" originalValue="N’Zérékoré">N'Zerekore</normalizedToken>
Region (7.67; -8.40), 998 m a.s.l.
</paragraph>
</subSubSection>
<subSubSection id="65E4CDCD923FC20C86EAA2E380A00D75" pageId="19" pageNumber="134" type="diagnosis">
<paragraph id="A1D9D851322AC5E9A41F0775BAA2CD97" pageId="19" pageNumber="134">Diagnosis.</paragraph>
<paragraph id="3019EC95B04214DEC4D3724EE26ED555" pageId="19" pageNumber="134">
Medium to large sized frogs, robust body shape; head narrow, highest eye diameter/eye-naris-distance ratio in the family, low mean tympanum diameter orbita diameter ratio, webbing almost fully developed, leaving 0.25-0.5 of the distal phalange free at the inner side of toe II, leaving 0.75-1 of the distal phalange free at toe IV, belly pattern very variable, glandular lines on tibia contain mean conic glands forming frequently interrupted lines. Genetically 
<taxonomicName id="116F452CA31DF1B209E2A2E7B0B3B325" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
differs by a minimum of 2.89% in the mitochondrial 16S gene from its congeners.
</paragraph>
</subSubSection>
<subSubSection id="AF0B52B960998A862F65E67241D35506" pageId="19" pageNumber="134" type="differential diagnosis">
<paragraph id="93228AD9401A7E10B299B40C09198109" pageId="19" pageNumber="134">Differential diagnosis.</paragraph>
<paragraph id="9ACB135BA7F4463E7BDF472A3F619CEC" pageId="19" pageNumber="134">
<taxonomicName id="DB58E4D7D80FF1848AB3F318EC140BAD" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
can be distinguished from its congeners by a combination of characters (characters distinguishing 
<taxonomicName id="E61C2631E25AEEC408A73E95C2100EEE" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
vs. 
<taxonomicName id="A851EC459CBB22898593A81ED0FD5C14" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus ziama" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="ziama">Odontobatrachus ziama</taxonomicName>
, 
<taxonomicName id="20791FC74840AE732DF3C40DF3CA70B8" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus smithi" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="smithi">Odontobatrachus smithi</taxonomicName>
and 
<taxonomicName id="E039A1ED4AADC5D914D5DCEEF4432D41" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus fouta" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="fouta">Odontobatrachus fouta</taxonomicName>
see above; for all significant differences see Table 5): male 
<taxonomicName id="963171A14BEF183EAB0ECCFE553431FF" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
differ from 
<taxonomicName id="3F1E1394AB0B48F4023112FB609449F7" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus natator" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="natator">Odontobatrachus natator</taxonomicName>
by the following ratios (Table 1): larger TI/SUL, FL/SUL, O/EN and smaller HW/SUL, TD/O, IT/FL, TD/SUL in 
<taxonomicName id="C93F8034C597E6DF0646C56754AFB645" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
than in 
<taxonomicName id="34FD1A8ACCB69756194C472ACB5C6D2B" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus natator" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="natator">Odontobatrachus natator</taxonomicName>
; female 
<taxonomicName id="7DD772F822966736BB76E094B62C9FC3" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
differ from 
<taxonomicName id="D691D8DC7B73750A2B399FCC8AA91955" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus natator" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="natator">Odontobatrachus natator</taxonomicName>
by the following ratios (Table 2): smaller FM/TI, HW/SUL and O/EN larger in 
<taxonomicName id="03A8FE1B3D1F6761280F239879904B49" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
than in 
<taxonomicName id="DF0E2F89D80F624543073393CC1318CC" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus natator" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="natator">Odontobatrachus natator</taxonomicName>
. Webbing formulae are very similar in the two species (Table 7). Glandular lines on tibia contain small to mean conic glands forming hardly interrupted lines in 
<taxonomicName id="5023E53C14E46ED2681A48BFC220B50A" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
(Fig. 12b, c), while similar to 
<taxonomicName id="42F3ADBB918787F98B3E69848CA42717" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus fouta" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="fouta">Odontobatrachus fouta</taxonomicName>
small to large glands form more or less interrupted lines in 
<taxonomicName id="69BFB71CE8B8BF61A442FC073EE14676" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus natator" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="natator">Odontobatrachus natator</taxonomicName>
(Fig. 4 
<normalizedToken id="741BA7306545006F39E8EF8C1574F190" originalValue="a–e">a-e</normalizedToken>
).
</paragraph>
</subSubSection>
<subSubSection id="D3C478FB3F2F2D6C646C71F2D1B94347" pageId="19" pageNumber="134" type="genetics">
<paragraph id="AB86DFA1FBBD385EB44E901F6EB8EB5C" pageId="19" pageNumber="134">Genetics.</paragraph>
<paragraph id="CBDDED1E0FDE007EB75B7E333C690C76" pageId="19" pageNumber="134">
The species is genetically well differentiated from all congeners and known populations form a well-supported and monophyletic clade (
<bibRefCitation id="A444425931369F6650DAF09594040D15" author="Barej, MF" journalOrPublisher="BMC Evolutionary Biology" pageId="25" pageNumber="140" title="Multiple genetic lineages challenge the monospecific status of the West African endemic frog-family Odontobatrachidae." url="10.1186/s12862-015-0346-9" volume="15" year="2015">Barej et al. 2015</bibRefCitation>
). Uncorrected 16S p-distances between 
<taxonomicName id="CD277BEAE2C99FA11F39CB15DBE47A55" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
and other 
<taxonomicName id="0A859B891981EC99C71AB1ACAF42E1C6" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="genus">Odontobatrachus</taxonomicName>
species range from 2.89-5.55%, while maximum intrataxon differences of 
<taxonomicName id="2E0D4321FC666CCFDDDE4F2B4545DA52" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="19" pageNumber="134" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
add up to 0.58% (mean value 0.05%; N = 861; Appendix 1: Table A).
</paragraph>
</subSubSection>
<subSubSection id="6125BCE861E02CD038192ED244F68046" lastPageId="21" lastPageNumber="136" pageId="19" pageNumber="134" type="holotype description">
<paragraph id="469A348473BAA37B6284E16D00A77ED2" pageId="19" pageNumber="134">Holotype description.</paragraph>
<paragraph id="F07E65B518B77F86621E57FA30EA2709" lastPageId="21" lastPageNumber="136" pageId="19" pageNumber="134">
The male holotype has been assigned to this taxon in both DCA analyses (absolute values and ratios). The holotype is an adult male with a slightly robust body (Fig. 11): snout-urostyle length of 48.8 mm; head width 17.1 mm; head slightly longer than broad; snout in lateral view short, flattened and rounded at the snout tip; snout in dorsal view triangular, rounded; lower jaw with sharp tusk-like prolonga 
<pageBreakToken id="6E4CC9E62A66F93457FFFD9EA157CC5C" pageId="20" pageNumber="135" start="start">tions</pageBreakToken>
protruding the skin and single triangular knob at lower jaw symphysis, corresponding socket in between premaxillae; upper premaxillae and maxillae with numerous teeth, posteriorly curved; vomerine teeth present, arranged in two small odontophores, closer to each other than to choanae; tongue broadly heart shaped; horizontal eye diameter 7.6 mm; interorbital distance 5.2 mm; pupil horizontally elliptical; eye diameter distinctly larger than tympanum diameter; tympanum distinct (horizontal diameter 2.7 mm); nares closer to snout than to eye; snout shorter than eye diameter; canthus rostralis rounded; loreal region concave; paired lateral vocal sacs; forelimbs robust, forearms hypertrophied, fingers slender; prepollex absent; relative finger lengths III&gt;IV&gt;II&gt;I; velvety nuptial excrescences weakly developed on finger I; subarticular tubercles large, subconical; supernumerary tubercles absent; fingertips dilated, triangular, notched in the middle; femur length 24.6 mm; tibia length 26.3 mm; femoral glands large (length 
<normalizedToken id="FC1877D1E30DF48A30D49FCD15A0FE92" originalValue="×">x</normalizedToken>
width: left: 12.9 
<normalizedToken id="8D1A296762DC254E4EC998BACCB9E8F5" originalValue="×">x</normalizedToken>
8.0 mm, right: 13.1 
<normalizedToken id="8C3C854DA972CE5F994E942580588E18" originalValue="×">x</normalizedToken>
7.8 
<pageBreakToken id="39C09E3021140E9BF68DB177065FEBDD" pageId="21" pageNumber="136" start="start">mm</pageBreakToken>
); femoral glands positioned on the posterior part of the ventral side of femur; relation femoral gland length to femur length: 0.53; minuscule circular glands running along upper side of tibia; foot length (incl. longest toe) 34.9 mm; relative toe lengths IV&gt;III&gt;V&gt;II&gt;I; shortest toe 6.4 mm; inner metatarsal tubercle elliptical; toe tips broadened forming triangular dilated discs; inner metatarsal tubercle prominent (3.8 mm); number of subconical subarticular tubercles on toes I-V: 1, 1, 2, 3, 2; supernumerary tubercles absent; prominent skin fold on posterior side of feet; dorsal skin texture heterogeneous; dorsum and flanks covered with slender dorsal ridges of app. 2.5-3.5 mm (partially flattened); venter with fine granulation; flank texture rough and granular as dorsum; webbing fully developed (0-0/0-1/0-0.75/0.75-0); webbing between toes hardly concave. Damage of the male holotype: cut at pectoral region (liver tissue sampled) and skin cut on right lumbar region (in ventral view); glandular dorsal ridges partially not recognisable due to preservation.
</paragraph>
<caption id="230916306EA3608CF4074519B1932CE1" pageId="21" pageNumber="136">
<paragraph id="01EE386B888E4D357A79D10A359C13E5" pageId="21" pageNumber="136">
Figure 11. Male holotype of 
<taxonomicName id="2CD0CD29B20A418CB60D684C2ACD80C0" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="21" pageNumber="136" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
sp. n. (ZMB 78355) in ventral and dorsal view (top from left to right, scale bar: 10 mm); head in lateral view, details of pedal webbing, details of hand (bottom left to right, scale bar: 5 mm).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="0C500AE476E95B4465187F3CCFCB5BFC" pageId="21" pageNumber="136" type="colouration of holotype in alcohol">
<paragraph id="8E2C89BED041A82148EFDB020698AC6A" pageId="21" pageNumber="136">Colouration of holotype in alcohol</paragraph>
<paragraph id="D6A4A612A1FE246F619FC23A688FDEB4" pageId="21" pageNumber="136">(Fig. 11). Dorsum dark brownish, few paler marblings; hind limbs coloured as dorsum; throat pale with few darker marblings; venter pale, area around incision darker; pale colouration between axillaries and elbows, femoral glands pale, clearly silhouetted from femora, with blurred minuscule reticulation; femora and tibia pale as belly.</paragraph>
</subSubSection>
<subSubSection id="BED2062FFC93CC7978EF176112331548" lastPageId="22" lastPageNumber="137" pageId="21" pageNumber="136" type="variation">
<paragraph id="586B94C35D1816ADE60B7872ABC23092" pageId="21" pageNumber="136">Variation.</paragraph>
<paragraph id="B7E8C7E1DC94FF25B2F82E9116E02B4B" lastPageId="22" lastPageNumber="137" pageId="21" pageNumber="136">
Females are significantly larger than males (SUL: Z = -4.933, p &lt;0.001, Nmales = 26, Nfemales = 24), max SUL in females 64.0 mm and 53.6 mm in males, and consequently possess longer extremities (FM: Z = -3.894, p &lt;0.001; TI: Z = -4.458, p &lt;0.001; FL: Z = -4.264, p &lt;0.001), broader heads (HW: Z = -4.090, p &lt;0.001), longer snouts (EN: Z = -2.678, p &lt;0.01; ES: Z = -2.906, p &lt;0.01) and larger eyes (Z = -2.779, p &lt;0.01), larger TD (Z = -2.214, p &lt;0.05). However, ratios are predominantly similar between the two sexes, although males show higher values in FL/SUL (Z = -2.214, p &lt;0.05), FM/SUL (Z = -2.932, p &lt;0.01), FM/TI (Z = -3.010, p &lt;0.01) and HW/SUL (Z = -4.136, p &lt;0.001). Both sexes possess enlarged tusk-like prolongations in the lower jaw as well as the name-bearing 
<normalizedToken id="DD63B1773FF6A309868F37E599B493BB" originalValue="‘teeth’">'teeth'</normalizedToken>
on the upper jaw. Male secondary sexual characters are femoral glands, velvety nuptial excrescences on finger I and presence of vocal sacs. Webbing formulae showed little variance (Table 7). However, some specimen possess a more extensive webbing on toe IV (0.5/0.5). Dorsal ridges are elongated and slender (Fig. 12 
<normalizedToken id="283CBF8BD589E19FFDF4A3327E77E1CD" originalValue="a–c">a-c</normalizedToken>
). Number of distinct dorsal ridges (counted from spine to flank) ranges between three and six, usually four to five ridges per body site. Glandular ridges on tibia usually are built of small to mean conic glands and form hardly interrupted lines (Fig. 12b, c). However, both characters were not recognisable due to preservation artefacts in many specimens. Dorsal colouration (in life) varies from almost black, beige with reddish-brown spots ar 
<pageBreakToken id="CD8CEB80F8614AFA496A8DAD35FC6309" pageId="22" pageNumber="137" start="start">ranged</pageBreakToken>
in longitudinal lines (Fig. 12; 
<bibRefCitation id="CE4C588C84BA7C4F4807CD2C7E456613" author="Guibe, J" journalOrPublisher="Memoires de l'Institut fondamental d'Afrique noire" pageId="25" pageNumber="140" pagination="241 - 273" title="La reserve naturelle integrale du Mont Nimba. XII. Batraciens (sauf Arthroleptis, Phrynobatrachus et Hyperolius)." volume="53" year="1958">
<normalizedToken id="3D34E679DEDAE3229B660BD661C5EDD1" originalValue="Guibé">Guibe</normalizedToken>
and Lamotte 1958
</bibRefCitation>
; 
<bibRefCitation id="EDA0C16E9A6830306104A29C45501F79" author="Roedel, M-O" journalOrPublisher="Tropical Zoology" pageId="25" pageNumber="140" pagination="201 - 232" title="A conservation assessment of amphibians in the Foret Classee du Pic de Fon, Simandou Range, southeastern Republic of Guinea, with the description of a new Amnirana species (AmphibiaAnuraRanidae)." url="10.1080/03946975.2004.10531206" volume="17" year="2004">
<normalizedToken id="EF5D959994EF18A1A0076179B4F7E88B" originalValue="Rödel">Roedel</normalizedToken>
and Bangoura 2004
</bibRefCitation>
). Male femoral glands are yellow (
<bibRefCitation id="6937FD23909BDF28632A2765FB627A74" author="Roedel, M-O" journalOrPublisher="Salamandra" pageId="25" pageNumber="140" pagination="91 - 110" title="The amphibians of Mont Sangbe National Park, Ivory Coast." volume="39" year="2003">
<normalizedToken id="6428CA4B64016F69B1447FDA68E8E6E8" originalValue="Rödel">Roedel</normalizedToken>
2003
</bibRefCitation>
). Belly colouration (in alcohol) is very variable, ranging from completely whitish, dirty whitish, a distinct reticulation pattern, dark throat with marbling on belly, marbling on throat and belly blurring to paler colouration posteriorly, to completely dark throat and belly, showing no sex-dependant differentiation.
</paragraph>
<caption id="278DF9C953DF399FCE43D283C351D602" pageId="22" pageNumber="137">
<paragraph id="6A65BFC3AB3898FE93E18F41191AE1CD" pageId="22" pageNumber="137">
Figure 12. 
<taxonomicName id="562BAE4F3C623E98E59439F4780FBD54" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="22" pageNumber="137" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
sp. n. in life: a) and c) Nimba Mts, Guinea; b) Mt. 
<normalizedToken id="1A17511C1B8B54F1C74DB91D70F75F31" originalValue="Sangbé">Sangbe</normalizedToken>
, 
<normalizedToken id="273FDB747070D38CEBF6C113389D9380" originalValue="Côte">Cote</normalizedToken>
<normalizedToken id="70BC12BEA25080E2AAF2428BFCB8C274" originalValue="d’Ivoire">d'Ivoire</normalizedToken>
. Non vouchered specimens. Note parasitic mites (minuscule red dots) close to the cloaca in (c).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="8AC6841EB631E6D11785712D4F498872" lastPageId="23" lastPageNumber="138" pageId="22" pageNumber="137" type="acoustics">
<paragraph id="A361E9140712A0F57B7297424F9C8FD7" pageId="22" pageNumber="137">Acoustics.</paragraph>
<paragraph id="99251B6E11E1E9953B177A1775734102" lastPageId="23" lastPageNumber="138" pageId="22" pageNumber="137">
Three calls of 
<taxonomicName id="8B143F67BBCFAB379DD416EB6422359F" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="22" pageNumber="137" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
were recorded from specimens in terraria. Calls sound like a repeat of 
<normalizedToken id="DC00121F54F8372A547FA7A639693735" originalValue="“chucks”">&quot;chucks&quot;</normalizedToken>
, consisting of several tonal notes. Two harmonics were visible (Fig. 13), the second harmonic being the dominant frequency (2842.4-3359.2 Hz), the first being the fundamental frequency (1421.2-1679.6 Hz). The call duration ranged from 1.2 to 3.0 s. One call comprised 22 notes (Fig. 13a) and the other two comprised five notes each (Fig. 13b). Each note had a duration of 34.7 
<normalizedToken id="A8F4AFE8E4E7BD56D5520F27BB4962B3" originalValue="±">+/-</normalizedToken>
0.01 ms (N = 32). The notes were separated from each other by pauses of 238.0 
<normalizedToken id="5504457B8B27DA7B9C869AC3437F12EF" originalValue="±">+/-</normalizedToken>
0.01 ms (N = 12) with the two calls comprising five notes (Fig. 13b). Pause duration in the third call (22 notes) was decreasing from 160.0 to 67.0 ms from the beginning towards the end of the call (Fig. 13a). 
<bibRefCitation id="CCBB127D96D6A913FE947F1A60ACA46A" author="Roedel, M-O" journalOrPublisher="Salamandra" pageId="25" pageNumber="140" pagination="91 - 110" title="The amphibians of Mont Sangbe National Park, Ivory Coast." volume="39" year="2003">
<normalizedToken id="B403ADF29EAC1A8EC55496F3AF6446A0" originalValue="Rödel’s">Roedel's</normalizedToken>
(2003)
</bibRefCitation>
anecdotal report of a 
<pageBreakToken id="2F0B5EAF2FE219EEF3CB5F3423763EDA" pageId="23" pageNumber="138" start="start">torrent</pageBreakToken>
frog call from Mt. 
<normalizedToken id="E128C52AB8C20DC7F690EF7A9C8108A4" originalValue="Sangbé">Sangbe</normalizedToken>
(
<normalizedToken id="7A3957069103FAF8C509D228BEDEFFB3" originalValue="Côte">Cote</normalizedToken>
<normalizedToken id="C3FA913CDBE6F2E42D4F5349D80CFC80" originalValue="d’Ivoire">d'Ivoire</normalizedToken>
) comprising a series of click sounds with ever-shorter intervals corresponds to the second call type (22 notes).
</paragraph>
<caption id="E0B8E05D5BCD6520272061315F60465D" pageId="23" pageNumber="138">
<paragraph id="AE2AE5D8A8AFD4C25C18FB1AAD35DC70" pageId="23" pageNumber="138">
Figure 13. Spectrogram and oscillogram of two calls of 
<taxonomicName id="DA51793B2366C036B83FEC2C4BF064A0" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="23" pageNumber="138" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
sp. n. from Nimba Mts., Guinea with a dominant frequency of app. 2800-3400 Hz, a fundamental frequency of app. 1400-1700 Hz and 22 notes showing decreasing pause duration between notes (a) and 5 notes with constant pauses between notes (b).
</paragraph>
</caption>
</subSubSection>
<subSubSection id="D82BA4CA6265C1FDBD37BA310CDCD37F" pageId="23" pageNumber="138" type="distribution">
<paragraph id="712CFD9C872F1906FE58ED866EAC0E15" pageId="23" pageNumber="138">Distribution.</paragraph>
<paragraph id="C00D19EAE6501189248727A68212E86B" pageId="23" pageNumber="138">
<taxonomicName id="4FB8631920CA516D2DFEEAE2D3F60F23" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="23" pageNumber="138" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
is known to occur on the Nimba Mts. in Guinea and Liberia, the adjacent areas at Mt. Gangra (Liberia) and 
<normalizedToken id="118CCCA9367AAF6248D95621BBCB8816" originalValue="Déré">Dere</normalizedToken>
(Guinea), as well as the Mt. 
<normalizedToken id="EFA0972FB042B739DF97C263135D0B82" originalValue="Sangbé">Sangbe</normalizedToken>
in western 
<normalizedToken id="D4992710C4347FCDB6DF4FB3D07269B5" originalValue="Côte">Cote</normalizedToken>
<normalizedToken id="5691A0D0D1DF8587BEC2702A80C424B5" originalValue="d’Ivoire">d'Ivoire</normalizedToken>
(Fig. 1). This taxon represents the easternmost representative of the family. Localities at the southern end of the Nimba Mts. and along Mt. Gangra are in very close proximity to 
<taxonomicName id="EEA54FEA9BBFD06467D93C50C1D7743B" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus natator" order="Anura" pageId="23" pageNumber="138" phylum="Chordata" rank="species" species="natator">Odontobatrachus natator</taxonomicName>
. Both species inhabit similar altitudes at the foot of the Nimba Mts. However, at present no differing habitat requirements or ecological adaptations are known (
<bibRefCitation id="6BD87386ECEDED3CB40A69C413E841E9" author="Barej, MF" journalOrPublisher="BMC Evolutionary Biology" pageId="25" pageNumber="140" title="Multiple genetic lineages challenge the monospecific status of the West African endemic frog-family Odontobatrachidae." url="10.1186/s12862-015-0346-9" volume="15" year="2015">Barej et al. 2015</bibRefCitation>
), which could explain their spatial separation.
</paragraph>
</subSubSection>
<subSubSection id="AFA1511D98540E2DFB2F764FA469229A" pageId="23" pageNumber="138" type="etymology">
<paragraph id="68023D64A7493171E4663BA941E0EA17" pageId="23" pageNumber="138">Etymology.</paragraph>
<paragraph id="2D63B422714568C2C99623E68E62CCD8" pageId="23" pageNumber="138">The species epithet arndti was chosen in order to honour Prof. emerit. Dr. Rudolf G. Arndt, New Jersey USA, for his trust in young academics and his invaluable support of this study.</paragraph>
</subSubSection>
<subSubSection id="5F06082230E5C8EA101CCBA701B1C74C" pageId="23" pageNumber="138" type="common name">
<paragraph id="2E1AD6460F65CEA439B60C5345A7A9FE" pageId="23" pageNumber="138">Common name.</paragraph>
<paragraph id="DCB9A9223E3EF83E41CD434FC143AEF5" pageId="23" pageNumber="138">
We advise to use the term 
<normalizedToken id="283ED89A67AF94027EC922DDF34C670D" originalValue="‘‘Arndt’s">''Arndt's</normalizedToken>
<normalizedToken id="5DBB782BEF60B701630DB768E0248A81" originalValue="torrent-frog’’">torrent-frog''</normalizedToken>
in English and 
<normalizedToken id="BA3EB23D8D868D22CABCCB7C8406F505" originalValue="‘‘">''</normalizedToken>
grenouilles des torrents 
<normalizedToken id="07A2EC7C4546F55BF8FC0B77607564FB" originalValue="d’Arndt”">d'Arndt&quot;</normalizedToken>
in French.
</paragraph>
</subSubSection>
<subSubSection id="F1C8F61AEFE04CFE282C1F41435F715E" pageId="23" pageNumber="138" type="conservation status">
<paragraph id="62B4DD04409BCB50CA0E7C52FE00607A" pageId="23" pageNumber="138">Conservation status.</paragraph>
<paragraph id="F422534960A8F6B16A9B1876435506CC" pageId="23" pageNumber="138">
Both, the EOO of 2595 km2 and the AOO of 156 km2 classify 
<taxonomicName id="5273A396904FFE648BC728B8B554BE8F" class="Amphibia" family="Odontobatrachidae" genus="Odontobatrachus" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Odontobatrachus arndti" order="Anura" pageId="23" pageNumber="138" phylum="Chordata" rank="species" species="arndti">Odontobatrachus arndti</taxonomicName>
as &quot;Endangered (EN)&quot; (
<bibRefCitation id="0F075572406D4EA42D507E81AD5F8599" author="Barej, MF" journalOrPublisher="BMC Evolutionary Biology" pageId="25" pageNumber="140" title="Multiple genetic lineages challenge the monospecific status of the West African endemic frog-family Odontobatrachidae." url="10.1186/s12862-015-0346-9" volume="15" year="2015">Barej et al. 2015</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>