treatments-xml/data/5F/21/46/5F2146EA658E52AD95C9522DBD826471.xml
2024-06-21 12:37:57 +02:00

849 lines
77 KiB
XML
Raw Blame History

This file contains ambiguous Unicode characters

This file contains Unicode characters that might be confused with other characters. If you think that this is intentional, you can safely ignore this warning. Use the Escape button to reveal them.

<document ID-DOI="http://dx.doi.org/10.3897/zse.97.73630" ID-GBIF-Dataset="3a1cc506-9f3c-4bc2-b12c-02675b0a0279" ID-GBIF-Taxon="190875615" ID-Pensoft-Pub="1860-0743-2-483" ID-Pensoft-UUID="A35E13FBED76532F9DAFD0A2AC6487B4" ID-ZooBank="4DC62CDBA6FB472D9E6DFDBF4E101785" LSID="urn:lsid:zoobank.org:act:3A22A655-D3B9-4C69-BF6F-C8F1E9595D84" ModsDocID="1860-0743-97-2-483" checkinTime="1638504930080" checkinUser="pensoft" docAuthor="Hutter, Carl R., Andriampenomanana, Zo F., Andrianasolo, Ginah Tsiorisoa, Cobb, Kerry A., Razafindraibe, Jary H., Abraham, Robin K. &amp; Lambert, Shea M." docDate="2021" docId="5F2146EA658E52AD95C9522DBD826471" docLanguage="en" docName="ZoosystEvol 97(2): 483-495" docOrigin="Zoosystematics and Evolution 97 (2)" docPubDate="2021-12-02" docSource="http://dx.doi.org/10.3897/zse.97.73630" docTitle="Gephyromantis marokoroko Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert 2021, sp. nov." docType="treatment" docUuid="3A22A655-D3B9-4C69-BF6F-C8F1E9595D84" docUuidSource="ZooBank" docVersion="4" id="A35E13FBED76532F9DAFD0A2AC6487B4" lastPageNumber="483" masterDocId="A35E13FBED76532F9DAFD0A2AC6487B4" masterDocTitle="A fantastic new species of secretive forest frog discovered from forest fragments near Andasibe, Madagascar" masterLastPageNumber="495" masterPageNumber="483" pageId="0" pageNumber="483" updateTime="1638550538227" updateUser="ExternalLinkService">
<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
<mods:titleInfo>
<mods:title>A fantastic new species of secretive forest frog discovered from forest fragments near Andasibe, Madagascar</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Hutter, Carl R.</mods:namePart>
<mods:affiliation>Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66045 - 7561, USA &amp; Museum of Natural Sciences and Department of Biological Sciences, Louisiana State University, Baton Rouge, LA 70803, USA</mods:affiliation>
<mods:nameIdentifier type="email">carl.hutter@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Andriampenomanana, Zo F.</mods:namePart>
<mods:affiliation>Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66045 - 7561, USA &amp; Departement de Biologie Animale, Universit d'Antananarivo, BP 906, Antananarivo 101, Madagascar</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Andrianasolo, Ginah Tsiorisoa</mods:namePart>
<mods:affiliation>Departement de Biologie Animale, Universit d'Antananarivo, BP 906, Antananarivo 101, Madagascar</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Cobb, Kerry A.</mods:namePart>
<mods:affiliation>Department of Biological Sciences &amp; Museum of Natural History, Auburn University, Auburn, Alabama 36849, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Razafindraibe, Jary H.</mods:namePart>
<mods:affiliation>Departement de Biologie Animale, Universit d'Antananarivo, BP 906, Antananarivo 101, Madagascar</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Abraham, Robin K.</mods:namePart>
<mods:affiliation>Biodiversity Institute and Department of Ecology and Evolutionary Biology, University of Kansas, Lawrence, KS 66045 - 7561, USA</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Lambert, Shea M.</mods:namePart>
<mods:affiliation>Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, AZ 85721, USA</mods:affiliation>
</mods:name>
<mods:typeOfResource>text</mods:typeOfResource>
<mods:relatedItem type="host">
<mods:titleInfo>
<mods:title>Zoosystematics and Evolution</mods:title>
</mods:titleInfo>
<mods:part>
<mods:date>2021</mods:date>
<mods:detail type="pubDate">
<mods:number>2021-12-02</mods:number>
</mods:detail>
<mods:detail type="volume">
<mods:number>97</mods:number>
</mods:detail>
<mods:detail type="issue">
<mods:number>2</mods:number>
</mods:detail>
<mods:extent unit="page">
<mods:start>483</mods:start>
<mods:end>495</mods:end>
</mods:extent>
</mods:part>
</mods:relatedItem>
<mods:location>
<mods:url>http://dx.doi.org/10.3897/zse.97.73630</mods:url>
</mods:location>
<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">http://dx.doi.org/10.3897/zse.97.73630</mods:identifier>
<mods:identifier type="Pensoft-Pub">1860-0743-2-483</mods:identifier>
<mods:identifier type="ZooBank">4DC62CDBA6FB472D9E6DFDBF4E101785</mods:identifier>
<mods:identifier type="Pensoft-UUID">A35E13FBED76532F9DAFD0A2AC6487B4</mods:identifier>
</mods:mods>
<treatment ID-GBIF-Taxon="190875615" LSID="urn:lsid:zoobank.org:act:3A22A655-D3B9-4C69-BF6F-C8F1E9595D84" httpUri="http://treatment.plazi.org/id/5F2146EA658E52AD95C9522DBD826471" lastPageNumber="483" pageId="0" pageNumber="483">
<subSubSection pageId="0" pageNumber="483" type="nomenclature">
<paragraph pageId="0" pageNumber="483">
<taxonomicName LSID="http://zoobank.org/3A22A655-D3B9-4C69-BF6F-C8F1E9595D84" authority="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert, 2021" authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko" status="sp. nov.">Gephyromantis marokoroko</taxonomicName>
<taxonomicNameLabel pageId="0" pageNumber="483">sp. nov.</taxonomicNameLabel>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="holotype">
<paragraph pageId="0" pageNumber="483">Holotype.</paragraph>
<paragraph pageId="0" pageNumber="483">
KU 343230 (field number CRH 1110), an adult male collected by Carl R. Hutter, Shea M. Lambert and Zo F. Andriampenomanana collected on 5 January 2016, at Vohidrazana Forest (
<geoCoordinate degrees="18.976" direction="south" orientation="latitude" precision="55" value="-18.976">18.976°S</geoCoordinate>
,
<geoCoordinate degrees="48.499" direction="east" orientation="longitude" precision="55" value="48.499">48.499°E</geoCoordinate>
; ca. 1150 m a.s.l.) in mid-altitude rainforest near Andasibe in Northern Central East Madagascar (Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. The distribution of Gephyromantis marokoroko sp. nov. in east-central Madagascar, view from above (A.) and from a profile view (B.). The black star marker indicates the type locality at Vohidrazana Forest where the black circle &quot; locality &quot; markers indicate other confirmed localities for the new species. Gephyromantis marokoroko sp. nov. is also found at high elevations and, thus, is likely distributed at other high elevation sites not surveyed. Elevational and satellite imagery data acquired from the USGS Earth Explorer (http: // earthexplorer. usgs. gov)." figureDoi="10.3897/zse.97.73630.figure1" httpUri="https://binary.pensoft.net/fig/617720" pageId="0" pageNumber="483">1</figureCitation>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="paratypes">
<paragraph pageId="0" pageNumber="483">Paratypes.</paragraph>
<paragraph pageId="0" pageNumber="483">
Adult male KU 343229 (CRH 1108), adult female UADBA-CRH 472 and adult male UADBA-CRH 1061 collected on 5 January 2016, with same collection data as holotype. Adult female KU 343218 (CRH 1397) collected on 18 January 2016, at Tavolobe (
<geoCoordinate degrees="19.005" direction="south" orientation="latitude" precision="55" value="-19.005">19.005°S</geoCoordinate>
,
<geoCoordinate degrees="48.461" direction="east" orientation="longitude" precision="55" value="48.461">48.461°E</geoCoordinate>
; ca. 1100 m a.s.l.) by Carl R. Hutter, Shea M. Lambert, Ginah Tsiorisoa Andrianasolo and Kerry A. Cobb. Adult male UADBA-CRH 1626, Adult female UADBA-CRH 1819 collected on 6 January 2017 at Vohidrazana Forest, adult female KU 347328 (CRH 1923) collected on 14 January 2017 at Andasibe-Mantadia National Park (Belakato:
<geoCoordinate degrees="18.821" direction="south" orientation="latitude" precision="55" value="-18.821">18.821°S</geoCoordinate>
,
<geoCoordinate degrees="48.439" direction="east" orientation="longitude" precision="55" value="48.439">48.439°E</geoCoordinate>
; ca. 1050 m a.s.l.) and adult female KU 347329 (CRH 2019) collected on 21 January 2017 at Vohimana (
<geoCoordinate degrees="18.926" direction="south" orientation="latitude" precision="55" value="-18.926">18.926°S</geoCoordinate>
,
<geoCoordinate degrees="48.489" direction="east" orientation="longitude" precision="55" value="48.489">48.489°E</geoCoordinate>
; ca. 1050 m a.s.l.), collected by Carl R. Hutter, Jary Harinarivo and Robin K. Abraham.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="available names">
<paragraph pageId="0" pageNumber="483">Available names.</paragraph>
<paragraph pageId="0" pageNumber="483">
There are no junior synonyms available that could be assigned to the new species from the subgenus
<taxonomicName class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="(Laurentomantis)" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="subGenus" subGenus="Laurentomantis">Laurentomantis</taxonomicName>
.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="etymology">
<paragraph pageId="0" pageNumber="483">Etymology.</paragraph>
<paragraph pageId="0" pageNumber="483">
The specific epithet
<emphasis italics="true" pageId="0" pageNumber="483">marokoroko</emphasis>
is a Malagasy word meaning
<normalizedToken originalValue="“rugose”">&quot;rugose&quot;</normalizedToken>
or
<normalizedToken originalValue="“rugged”">&quot;rugged&quot;</normalizedToken>
. The name was chosen to describe the rugose skin texture of this species. The name is to be treated as an invariable noun in apposition.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="diagnosis">
<paragraph pageId="0" pageNumber="483">Diagnosis.</paragraph>
<paragraph pageId="0" pageNumber="483">
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
(Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Ex-situ dorsal-lateral, dorsal and ventral photographs of A. Male Gephyromantis marokoroko sp. nov. (holotype, KU 343230) and B. Female (paratype, KU 343218) in life." figureDoi="10.3897/zse.97.73630.figure2" httpUri="https://binary.pensoft.net/fig/617721" pageId="0" pageNumber="483">2</figureCitation>
) is a member of the family
<taxonomicName authorityName="Laurent" authorityYear="1946" class="Amphibia" family="Mantellidae" kingdom="Animalia" lsidName="" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="family">Mantellidae</taxonomicName>
, subfamily
<taxonomicName authorityName="Laurent" authorityYear="1946" lsidName="" pageId="0" pageNumber="483" rank="subFamily" subFamily="Mantellinae">Mantellinae</taxonomicName>
, as diagnosed by
<bibRefCitation DOI="https://doi.org/10.1016/j.ode.2005.12.001" author="Glaw, F" journalOrPublisher="Organisms Diversity and Evolution" pageId="0" pageNumber="483" pagination="236 - 253" refId="B12" refString="Glaw, F, Vences, M, 2006. Phylogeny and genus-level classification of mantellid frogs. Organisms Diversity and Evolution 6: 236 - 253, DOI: https://doi.org/10.1016/j.ode.2005.12.001" title="Phylogeny and genus-level classification of mantellid frogs." url="https://doi.org/10.1016/j.ode.2005.12.001" volume="6" year="2006">Glaw and Vences (2006)</bibRefCitation>
. The new species can be diagnosed to the genus
<taxonomicName authorityName="Methuen" authorityYear="1920" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis</emphasis>
</taxonomicName>
morphologically through its granular dorsum, moderately enlarged fingertips, absence of foot webbing, bifid tongue and small femoral glands present only in males as a small number of large granules (type 2;
<bibRefCitation DOI="https://doi.org/10.1080/00222930050020140" author="Glaw, F" journalOrPublisher="Journal of Natural History" pageId="0" pageNumber="483" pagination="1135 - 1154" refId="B11" refString="Glaw, F, Vences, M, Gossmann, V, 2000. A new species of Mantidactylus (subgenus Guibemantis) from Madagascar, with a comparative survey of internal femoral gland structure in the genus (Amphibia: Ranidae: Mantellinae). Journal of Natural History 34: 1135 - 1154, DOI: https://doi.org/10.1080/00222930050020140" title="A new species of Mantidactylus (subgenus Guibemantis) from Madagascar, with a comparative survey of internal femoral gland structure in the genus (Amphibia: Ranidae: Mantellinae)." url="https://doi.org/10.1080/00222930050020140" volume="34" year="2000">Glaw et al. 2000</bibRefCitation>
). Within
<taxonomicName authorityName="Methuen" authorityYear="1920" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis</emphasis>
</taxonomicName>
, the new species can be diagnosed to the subgenus
<taxonomicName class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="(Laurentomantis)" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="subGenus" subGenus="Laurentomantis">Laurentomantis</taxonomicName>
through its irregular and rough granular dorsum, single subgular vocal sac in males, completely connected lateral metatarsalia, inner and outer metatarsal tubercle present and tympanum is the same size in male and female.
</paragraph>
<paragraph pageId="0" pageNumber="483">
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
is characterised by bright red eyes, prominent ridge elements on dorsum, life colouration with a dark brown ground colour with mottled red and grey, hind-limbs dark brown containing red crossbands, absence of red colouration on the sides of thighs and ventre, white spots on grey-coloured ventre and males with bulbous type 2 femoral glands with eight granules in two rows of four on each thigh. Furthermore, the new species is characterised by an advertisement call with a moderately long call duration (1095-1431 ms), 22-28 notes/call, 2-4 strong amplitude-modulated pulses per note and a dominant frequency of 2250-2812 Hz. Finally,
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
has a large genetic distance of 6% or greater amongst related species in the 16S rRNA marker and has strongly supported reciprocal monophyly to all other species in
<taxonomicName authorityName="Dubois" authorityYear="1980" class="Amphibia" family="Mantellidae" genus="Laurentomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Laurentomantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Laurentomantis</emphasis>
</taxonomicName>
(Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Results of phylogenetic analysis of the mitochondrial 16 S rRNA barcode 3 ' marker for Maximum Likelihood (ML) and Bayesian Inference (BI). Topology is a consensus tree from IQ-Tree. The support values are shown as Bootstrap on top and Posterior Probability on the bottom only for nodes that were not perfectly supported. Note that Gephyromantis marokoroko sp. nov. placement in the clade is weakly supported in both analyses." figureDoi="10.3897/zse.97.73630.figure3" httpUri="https://binary.pensoft.net/fig/617722" pageId="0" pageNumber="483">3</figureCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="483">
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
can be distinguished from other members of
<taxonomicName authorityName="Dubois" authorityYear="1980" class="Amphibia" family="Mantellidae" genus="Laurentomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Laurentomantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Laurentomantis</emphasis>
</taxonomicName>
morphologically (Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Comparison of distinguishing characters used to differentiate species within Laurentomantis. Table adapted from Vences et al. (2002) combined with new data. Genetic distances are uncorrected and taken from the 16 S rRNA mitochondrial marker." httpUri="http://table.plazi.org/id/C4E1602CA50A38DA33029116486A58AE" pageId="0" pageNumber="483" tableUuid="C4E1602CA50A38DA33029116486A58AE">1</tableCitation>
; Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Ex-situ dorsal-lateral, dorsal and ventral photographs of A. Male Gephyromantis marokoroko sp. nov. (holotype, KU 343230); B. Gephyromantis striatus (Marojejy, ZCMV 15140; photographs by Mark D. Scherz); and C. Gephyromantis ventrimaculatus (Ranomanfana, KU 340917)." figureDoi="10.3897/zse.97.73630.figure5" httpUri="https://binary.pensoft.net/fig/617724" pageId="0" pageNumber="483">5</figureCitation>
). The rugose and granular dorsal texture with prominent ridge elements and red mottled colouration and the larger number of eight prominent femoral gland granules per femur readily characterise this species from other
<taxonomicName authorityName="Dubois" authorityYear="1980" class="Amphibia" family="Mantellidae" genus="Laurentomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Laurentomantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Laurentomantis</emphasis>
</taxonomicName>
(Figs
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Results of phylogenetic analysis of the mitochondrial 16 S rRNA barcode 3 ' marker for Maximum Likelihood (ML) and Bayesian Inference (BI). Topology is a consensus tree from IQ-Tree. The support values are shown as Bootstrap on top and Posterior Probability on the bottom only for nodes that were not perfectly supported. Note that Gephyromantis marokoroko sp. nov. placement in the clade is weakly supported in both analyses." figureDoi="10.3897/zse.97.73630.figure3" httpUri="https://binary.pensoft.net/fig/617722" pageId="0" pageNumber="483">3</figureCitation>
and
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Ex-situ dorsal-lateral, dorsal and ventral photographs of A. Male Gephyromantis marokoroko sp. nov. (holotype, KU 343230); B. Gephyromantis striatus (Marojejy, ZCMV 15140; photographs by Mark D. Scherz); and C. Gephyromantis ventrimaculatus (Ranomanfana, KU 340917)." figureDoi="10.3897/zse.97.73630.figure5" httpUri="https://binary.pensoft.net/fig/617724" pageId="0" pageNumber="483">5</figureCitation>
). The new species is easily distinguished from
<taxonomicName genus="G." lsidName="G. horridus" pageId="0" pageNumber="483" rank="species" species="horridus">
<emphasis italics="true" pageId="0" pageNumber="483">G. horridus</emphasis>
</taxonomicName>
(
<bibRefCitation author="Boettger, O" journalOrPublisher="Zoologischer Anzeiger" pageId="0" pageNumber="483" pagination="279 - 283" refId="B3" refString="Boettger, O, 1880. Diagnoses reptilium et batrachiorum novorum a Carolo Ebanau in insula Nossi-Be Madagascariensis lectorum. Zoologischer Anzeiger 3: 279 - 283" title="Diagnoses reptilium et batrachiorum novorum a Carolo Ebanau in insula Nossi-Be Madagascariensis lectorum." volume="3" year="1880">Boettger 1880</bibRefCitation>
),
<taxonomicName genus="G." lsidName="G. malagasius" pageId="0" pageNumber="483" rank="species" species="malagasius">
<emphasis italics="true" pageId="0" pageNumber="483">G. malagasius</emphasis>
</taxonomicName>
(
<bibRefCitation author="Methuen, PA" journalOrPublisher="Annals of the Transvaal Museum" pageId="0" pageNumber="483" pagination="49 - 64" refId="B29" refString="Methuen, PA, Hewitt, J, 1913. On a collection of Batrachia from Madagascar made during the year 1911. Annals of the Transvaal Museum 4: 49 - 64" title="On a collection of Batrachia from Madagascar made during the year 1911." volume="4" year="1913">Methuen and Hewitt 1913</bibRefCitation>
) and
<taxonomicName genus="G." lsidName="G. ranjomavo" pageId="0" pageNumber="483" rank="species" species="ranjomavo">
<emphasis italics="true" pageId="0" pageNumber="483">G. ranjomavo</emphasis>
</taxonomicName>
(
<bibRefCitation author="Glaw, F" journalOrPublisher="Spixiana" pageId="0" pageNumber="483" pagination="121 - 127" refId="B15" refString="Glaw, F, Vences, M, 2011. Description of a new frog species of Gephyromantis (subgenus Laurentomantis) with tibial glands from Madagascar. Spixiana 34: 121 - 127" title="Description of a new frog species of Gephyromantis (subgenus Laurentomantis) with tibial glands from Madagascar." volume="34" year="2011">Glaw and Vences 2011</bibRefCitation>
) by lacking tibial glands, its larger number of femoral gland granules and its rugose and granular dorsal texture with prominent ridge elements. Furthermore, the new species is easily distinguished from
<taxonomicName genus="G." lsidName="G. ventrimaculatus" pageId="0" pageNumber="483" rank="species" species="ventrimaculatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. ventrimaculatus</emphasis>
</taxonomicName>
, where
<taxonomicName genus="G." lsidName="G. marokoroko" pageId="0" pageNumber="483" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">G. marokoroko</emphasis>
</taxonomicName>
has eight distinct femoral gland granules on each thigh (eight irregularly-shaped femoral gland granules in
<taxonomicName genus="G." lsidName="G. ventrimaculatus" pageId="0" pageNumber="483" rank="species" species="ventrimaculatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. ventrimaculatus</emphasis>
</taxonomicName>
), by the dark grey and red dorsal colouration (light brown in
<taxonomicName genus="G." lsidName="G. ventrimaculatus" pageId="0" pageNumber="483" rank="species" species="ventrimaculatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. ventrimaculatus</emphasis>
</taxonomicName>
) and by lacking blue marbling on the ventral surfaces (Fig.
<figureCitation captionStart="Figure 5" captionStartId="F5" captionText="Figure 5. Ex-situ dorsal-lateral, dorsal and ventral photographs of A. Male Gephyromantis marokoroko sp. nov. (holotype, KU 343230); B. Gephyromantis striatus (Marojejy, ZCMV 15140; photographs by Mark D. Scherz); and C. Gephyromantis ventrimaculatus (Ranomanfana, KU 340917)." figureDoi="10.3897/zse.97.73630.figure5" httpUri="https://binary.pensoft.net/fig/617724" pageId="0" pageNumber="483">5</figureCitation>
). The most similar species morphologically is
<taxonomicName genus="G." lsidName="G. striatus" pageId="0" pageNumber="483" rank="species" species="striatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. striatus</emphasis>
</taxonomicName>
(
<bibRefCitation DOI="https://doi.org/10.1163/18759866-07004001" author="Vences, M" journalOrPublisher="Contributions to Zoology" pageId="0" pageNumber="483" pagination="191 - 212" refId="B46" refString="Vences, M, Glaw, F, Andreone, F, Jesu, R, Schimmenti, G, 2002. Systematic revision of the enigmatic Malagasy broad-headed frogs (Laurentomantis Dubois, 1980), and their phylogenetic position within the endemic mantellid radiation of Madagascar. Contributions to Zoology 70: 191 - 212, DOI: https://doi.org/10.1163/18759866-07004001" title="Systematic revision of the enigmatic Malagasy broad-headed frogs (Laurentomantis Dubois, 1980), and their phylogenetic position within the endemic mantellid radiation of Madagascar." url="https://doi.org/10.1163/18759866-07004001" volume="70" year="2002">Vences et al. 2002</bibRefCitation>
), but the new species differs from
<taxonomicName genus="G." lsidName="G. striatus" pageId="0" pageNumber="483" rank="species" species="striatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. striatus</emphasis>
</taxonomicName>
through its larger number of femoral gland granules (8 vs. 3-6), the vertebral stripe is absent or indistinct and short (always distinct in
<taxonomicName genus="G." lsidName="G. striatus" pageId="0" pageNumber="483" rank="species" species="striatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. striatus</emphasis>
</taxonomicName>
), bright red eye (orange-brown in
<taxonomicName genus="G." lsidName="G. striatus" pageId="0" pageNumber="483" rank="species" species="striatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. striatus</emphasis>
</taxonomicName>
) and its prominent and strong ridge elements, as well as the dark grey and red colouration on the dorsum (weak ridge elements and brown and orange colouration on the dorsum in
<taxonomicName genus="G." lsidName="G. striatus" pageId="0" pageNumber="483" rank="species" species="striatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. striatus</emphasis>
</taxonomicName>
).
</paragraph>
<caption doi="10.3897/zse.97.73630.figure5" httpUri="https://binary.pensoft.net/fig/617724" pageId="0" pageNumber="483" start="Figure 5" startId="F5">
<paragraph pageId="0" pageNumber="483">
<emphasis bold="true" pageId="0" pageNumber="483">Figure 5.</emphasis>
Ex-situ dorsal-lateral, dorsal and ventral photographs of
<emphasis bold="true" pageId="0" pageNumber="483">A.</emphasis>
Male
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
sp. nov. (holotype, KU 343230);
<emphasis bold="true" pageId="0" pageNumber="483">B.</emphasis>
<taxonomicName baseAuthorityName="Vences, Glaw, Andreone, Jesu &amp; Schimmenti" baseAuthorityYear="2002" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis striatus" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="striatus">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis striatus</emphasis>
</taxonomicName>
(Marojejy, ZCMV 15140; photographs by Mark D. Scherz); and
<emphasis bold="true" pageId="0" pageNumber="483">C.</emphasis>
<taxonomicName baseAuthorityName="Angel" baseAuthorityYear="1935" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis ventrimaculatus" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="ventrimaculatus">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis ventrimaculatus</emphasis>
</taxonomicName>
(Ranomanfana, KU 340917).
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="483">
Bioacoustically, the advertisement call of
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
is similar to other species in
<taxonomicName authorityName="Dubois" authorityYear="1980" class="Amphibia" family="Mantellidae" genus="Laurentomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Laurentomantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Laurentomantis</emphasis>
</taxonomicName>
and can be distinguished from all other species in this subgenus through the following combination of continuous call characters: (1) moderately long call duration (1095-1431 ms); (2) 2-4 strongly amplitude-modulated pulses per note; and (3) a note repetition rate of 14-20 notes/s.
<taxonomicName baseAuthorityName="Vences, Glaw, Andreone, Jesu &amp; Schimmenti" baseAuthorityYear="2002" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis striatus" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="striatus">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis striatus</emphasis>
</taxonomicName>
,
<taxonomicName genus="G." lsidName="G. malagasius" pageId="0" pageNumber="483" rank="species" species="malagasius">
<emphasis italics="true" pageId="0" pageNumber="483">G. malagasius</emphasis>
</taxonomicName>
and
<taxonomicName genus="G." lsidName="G. horridus" pageId="0" pageNumber="483" rank="species" species="horridus">
<emphasis italics="true" pageId="0" pageNumber="483">G. horridus</emphasis>
</taxonomicName>
have overlapping call durations with the new species and overlapping note repetition rates, except for
<taxonomicName genus="G." lsidName="G. striatus" pageId="0" pageNumber="483" rank="species" species="striatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. striatus</emphasis>
</taxonomicName>
, which has the fastest note repetition rate (Table
<tableCitation captionStart="Table 1" captionStartId="T1" captionText="Table 1. Comparison of distinguishing characters used to differentiate species within Laurentomantis. Table adapted from Vences et al. (2002) combined with new data. Genetic distances are uncorrected and taken from the 16 S rRNA mitochondrial marker." httpUri="http://table.plazi.org/id/C4E1602CA50A38DA33029116486A58AE" pageId="0" pageNumber="483" tableUuid="C4E1602CA50A38DA33029116486A58AE">1</tableCitation>
). Despite these similarities, the clearly pulsed notes alone distinguish the new species from all other
<taxonomicName authorityName="Dubois" authorityYear="1980" class="Amphibia" family="Mantellidae" genus="Laurentomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Laurentomantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Laurentomantis</emphasis>
</taxonomicName>
, except
<taxonomicName genus="G." lsidName="G. ventrimaculatus" pageId="0" pageNumber="483" rank="species" species="ventrimaculatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. ventrimaculatus</emphasis>
</taxonomicName>
(
<bibRefCitation author="Angel, F" journalOrPublisher="Bulletin de la Societe Zoologique de France" pageId="0" pageNumber="483" pagination="202 - 207" refId="B2" refString="Angel, F, 1935. Batraciens nouveaux de Madagascar recoltes par M. R. Catala. Bulletin de la Societe Zoologique de France 60: 202 - 207" title="Batraciens nouveaux de Madagascar recoltes par M. R. Catala." volume="60" year="1935">Angel 1935</bibRefCitation>
), which has ca. 5-6 pulses/note; however,
<taxonomicName genus="G." lsidName="G. ventrimaculatus" pageId="0" pageNumber="483" rank="species" species="ventrimaculatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. ventrimaculatus</emphasis>
</taxonomicName>
differs by having the shortest call duration non-overlapping with other
<taxonomicName authorityName="Dubois" authorityYear="1980" class="Amphibia" family="Mantellidae" genus="Laurentomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Laurentomantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Laurentomantis</emphasis>
</taxonomicName>
species at 407-455 ms and a slightly faster note repetition rate of 21-24 notes/s. Temperature is not likely to be an important factor in the characteristic differences described here, as structural characters, such as clearly defined pulses, would not be affected by temperature (
<bibRefCitation DOI="https://doi.org/10.1007/BF00344901" author="Schneider, H" journalOrPublisher="Oecologia" pageId="0" pageNumber="483" pagination="99 - 110" refId="B44" refString="Schneider, H, 1974. Structure of the mating calls and relationships of the European tree frogs (Hylidae, Anura). Oecologia 14: 99 - 110, DOI: https://doi.org/10.1007/BF00344901" title="Structure of the mating calls and relationships of the European tree frogs (Hylidae, Anura)." url="https://doi.org/10.1007/BF00344901" volume="14" year="1974">Schneider 1974</bibRefCitation>
).
</paragraph>
<paragraph pageId="0" pageNumber="483">
Motivation might affect number of notes emitted and, thus, call duration; however, the recording of
<taxonomicName genus="G." lsidName="G. ventrimaculatus" pageId="0" pageNumber="483" rank="species" species="ventrimaculatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. ventrimaculatus</emphasis>
</taxonomicName>
is of a highly motivated male (i.e. many calls emitted in a short time) while the call of the new species was recorded from males which did not appear to be very motivated, emitting only 1-2 calls within an hour. Finally, comparisons could not be made to
<taxonomicName genus="G." lsidName="G. ranjomavo" pageId="0" pageNumber="483" rank="species" species="ranjomavo">
<emphasis italics="true" pageId="0" pageNumber="483">G. ranjomavo</emphasis>
</taxonomicName>
as calls were not available; however, the new species is clearly morphologically distinct (see above).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="description">
<paragraph pageId="0" pageNumber="483">Description of the holotype.</paragraph>
<paragraph pageId="0" pageNumber="483">Fixed in 10% buffered formalin solution, preserved in 70% ethanol, in good state of preservation, except for skin loss near the anterior dorsum, with left thigh muscle removed for tissue sample. Adult male, SVL 26.0 mm. Body very slender; head longer than wide HL 33.4% of SVL; slightly wider than body, HW 33.7% of SVL; snout of moderate length, ESD 16.2% of SVL; snout rounded in dorsal and lateral view; nostrils directed laterally, slightly protuberant, nearer to snout tip than eye; ED larger than END; canthus rostralis indistinct, concave; loreal region slightly concave; single subgular vocal sac; gular glands absent. Tympanic annulus distinct and round, small, TD 64.5% of ED; supratympanic fold indistinct and irregular, tympanic membrane lighter than ground colouration. Vomerine teeth not visible on the buccal roof, present under mucosal skin; choanae small, rounded. Tongue longer than wide; ovoid in shape, posteriorly bifid. Dermal fold along lower jaw absent. Arms slender, subarticular tubercles single; outer and inner metacarpal tubercles present, indistinct. Fingers without webbing; nuptial pads absent; relative finger length 2 &lt;1 &lt;4 &lt;3; second finger distinctly shorter than fourth finger, only slightly shorter than finger one; finger discs distinctly enlarged, larger on third and fourth finger. Hind limbs slender; lateral metatarsalia connected; inner metatarsal tubercle distinct, outer metatarsal tubercle small, but recognisable; TIBL 55.2% of SVL; FOL 45.2% of SVL. Tibial glands absent. Toes without webbing; relative toe length 1 &lt;2 &lt;5 &lt;3 &lt;4; toe three distinctly longer and broader than toe five; toe discs distinctly enlarged. Femoral glands large, well delimited, having eight distinct clusters on each femur of almost the same size, in two rows of four. Skin coarsely granular and heavily rugose on dorsal surfaces; large and sharply elevated tubercles and ridges irregularly distributed across dorsal surfaces, with less distinct ridges on the lower back; some ridges are symmetrical, larger tubercles and short ridges present on head and anterior dorsal region. Ventral skin granular on stomach, throat and limbs.</paragraph>
<paragraph pageId="0" pageNumber="483">After four years in preservative, dorsal ground colouration is a uniform dull brown including forelimbs and hind-limbs. The red colouration has faded to become light brown. Lighter coloured spots on ventral surfaces are still present.</paragraph>
<paragraph pageId="0" pageNumber="483">
In life (Fig.
<figureCitation captionStart="Figure 2" captionStartId="F2" captionText="Figure 2. Ex-situ dorsal-lateral, dorsal and ventral photographs of A. Male Gephyromantis marokoroko sp. nov. (holotype, KU 343230) and B. Female (paratype, KU 343218) in life." figureDoi="10.3897/zse.97.73630.figure2" httpUri="https://binary.pensoft.net/fig/617721" pageId="0" pageNumber="483">2</figureCitation>
), dorsal colouration is a dark grey ground colour with thick, bright red mottling distributed on the dorsum. Many of the raised ridges are dark grey with bright red edges. Lighter red stripe present short distance up the dorsum. Lateral head the same as dorsum, tympanum a lighter brown. Flanks are also dark grey, but have less bright red colouration, typically only found on ridges. Forelimbs have same colouration as dorsum, except bright red colouration is more spotted, with a few lighter red spots. Hind-limbs have same colouration as forelimbs, except with red crossbands present on the dorsal surface. A whitish annulus is present before the terminal disc on fingers and toes, fingers and toes light brown. Ventral surfaces brown, with no red present. White and light-yellow spots are present and scattered moderately along the ventre. Ventral sides of arms and hind-limbs brownish-grey, with light red spotting. Femoral glands lighter brown than surrounding limb surfaces. Single subgular vocal sac is light grey, with some light-yellow spotting down the centre. Jaw has scattered light-red spots along the lip. The pupil is black with a bright red iris, with black reticulations around the outer margin of the iris.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="variation">
<paragraph pageId="0" pageNumber="483">Variation.</paragraph>
<paragraph pageId="0" pageNumber="483">All paratypes resemble the holotype in morphology and colouration. In life, dorsal colouration varies slightly in the amount and intensity of red present. Spotting on the ventral surfaces varies in the colouration of the spots being white, light-yellow, light-orange or light-red. The vertebral stripe varies from being absent in some individuals to indistinct in others. Females lack femoral glands and have a granular texture on the femur.</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="morphometry of type series">
<paragraph pageId="0" pageNumber="483">Morphometry of type series.</paragraph>
<paragraph pageId="0" pageNumber="483">
Measurements of the holotype and paratypes are shown in Table
<tableCitation captionStart="Table 2" captionStartId="T2" captionText="Table 2. Morphometric measurements (in mm) of the holotype and paratypes of Gephyromantis marokoroko sp. nov. Femoral Gland Clusters (FGC) shown as &quot; left, right &quot; count." httpUri="http://table.plazi.org/id/78CC3AED4A67386F71E9365BFBCC8386" pageId="0" pageNumber="483" tableUuid="78CC3AED4A67386F71E9365BFBCC8386">2</tableCitation>
.
</paragraph>
<caption ID-Table-UUID="78CC3AED4A67386F71E9365BFBCC8386" httpUri="http://table.plazi.org/id/78CC3AED4A67386F71E9365BFBCC8386" pageId="0" pageNumber="483" start="Table 2" startId="T2">
<paragraph pageId="0" pageNumber="483">
<emphasis bold="true" pageId="0" pageNumber="483">Table 2.</emphasis>
Morphometric measurements (in mm) of the holotype and paratypes of
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
sp. nov. Femoral Gland Clusters (FGC) shown as &quot;left, right&quot; count.
</paragraph>
</caption>
<paragraph pageId="0" pageNumber="483">
<table pageId="0" pageNumber="483">
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="2">
<emphasis bold="true" pageId="0" pageNumber="483">Type status</emphasis>
</td>
<td colspan="6" pageId="0" pageNumber="483" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="483">Specimen</emphasis>
</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="483">Holotype</emphasis>
</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="483">Paratype</emphasis>
</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="483">Paratype</emphasis>
</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="483">Paratype</emphasis>
</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="483">Paratype</emphasis>
</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">
<emphasis bold="true" pageId="0" pageNumber="483">Paratype</emphasis>
</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">Field Number</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">CRH 1110</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">CRH 1108</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">CRH 1397</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">CRH 1061</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">CRH 1923</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">CRH 2019</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">Museum Number</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">KU 343230</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">KU 343229</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">KU 343218</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">UADBA CRH1061</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">KU 347328</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">KU 347329</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">Sex</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">M</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">F</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">F</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">SVL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">26.0</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">25.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">27.0</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">24.0</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">24.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">23.9</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">HW</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">9.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.6</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">HL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.7</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">10.0</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.7</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.6</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">ED</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.5</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.9</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">IOD</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.5</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.7</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.7</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.5</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.5</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">ESD</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">4.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">4.0</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">4.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">4.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">4.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">4.3</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">END</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.0</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.5</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.9</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">NSD</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">1.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">1.5</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">1.7</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">1.5</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">1.9</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">1.4</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">NND</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.5</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.2</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">TD</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.5</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">1.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.5</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.0</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.2</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">FIL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">5.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">14.9</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">4.9</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">5.0</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">FIIL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">5.7</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.6</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">FEML</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">13.9</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">14.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">14.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">13.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">14.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">14.1</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">TIBL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">14.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">14.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">14.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">13.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">14.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">13.8</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">FOL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">11.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">11.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">12.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">11.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">12.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">11.3</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">TARL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">7.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.0</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">HAL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">7.9</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">7.6</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">LAL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">7.7</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">7.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">7.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8.2</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">UAL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.0</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.0</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">FORL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">22.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">21.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">22.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">21.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">22.6</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">21.7</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">FGL</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">6.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">5.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">10.1</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">5.7</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">-</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">-</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">FGW</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">2.4</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.3</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">3.2</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">-</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">-</td>
</tr>
<tr pageId="0" pageNumber="483">
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">FGC</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8; 8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8; 8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8; 8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">8; 8</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">-</td>
<td colspan="1" pageId="0" pageNumber="483" rowspan="1">-</td>
</tr>
</table>
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="bioacoustics">
<paragraph pageId="0" pageNumber="483">Bioacoustics.</paragraph>
<paragraph pageId="0" pageNumber="483">We recorded three calls from two males at Vohidrazana Forest after collection at ca. 02:00 hr on 6 January 2016. Males call infrequently with extremely quiet calls from the upper surfaces of leaves up to 50 cm above the ground. The recorded male was captured and placed in a separate plastic collecting bag. Males would not call when we were within recording distance, so we placed the microphone 100 cm away from the bag near where it was captured and moved several metres away. Calls were recorded during light rain at a temperature of 20.4 °C.</paragraph>
<paragraph pageId="0" pageNumber="483">
The advertisement call of this species sounds like a heavily pulsed trill or
<normalizedToken originalValue="groan">'groan'</normalizedToken>
to the human observer, emitted irregularly. We define each groan as a call (Fig.
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Oscillograms and spectrograms of the call of Gephyromantis marokoroko sp. nov. (Holotype: KU 343230). A. The entire call spectrogram and B. Entire call oscillogram; C. Power spectra / frequency spectrogram of a single note; D. A close-up spectrogram of four notes and E. Corresponding oscillogram; and F. an individual note taken from the middle of the call." figureDoi="10.3897/zse.97.73630.figure6" httpUri="https://binary.pensoft.net/fig/617725" pageId="0" pageNumber="483">6A-C</figureCitation>
) with a duration of 1095.1-1431.9 (1221.5
<normalizedToken originalValue="±">+/-</normalizedToken>
183.5;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 3) ms. Each call consisted of a series of 22-28 (24
<normalizedToken originalValue="±">+/-</normalizedToken>
3.46;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 72) short notes with a duration of 12-29 (19.9
<normalizedToken originalValue="±">+/-</normalizedToken>
4.3;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 72) ms and an inter-note duration of 15.5-43.7 (32.3
<normalizedToken originalValue="±">+/-</normalizedToken>
5.6;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 71) ms. Note rate within each call was 14.4-20.1 (17.9
<normalizedToken originalValue="±">+/-</normalizedToken>
3.1;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 3) note/s. Each note was strongly pulsed, with 2-4 (3.1
<normalizedToken originalValue="±">+/-</normalizedToken>
0.783;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 72) pulses per note and a pulse rate of 111.1-235.3 (156.3
<normalizedToken originalValue="±">+/-</normalizedToken>
25.8) pulses/s (Fig.
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Oscillograms and spectrograms of the call of Gephyromantis marokoroko sp. nov. (Holotype: KU 343230). A. The entire call spectrogram and B. Entire call oscillogram; C. Power spectra / frequency spectrogram of a single note; D. A close-up spectrogram of four notes and E. Corresponding oscillogram; and F. an individual note taken from the middle of the call." figureDoi="10.3897/zse.97.73630.figure6" httpUri="https://binary.pensoft.net/fig/617725" pageId="0" pageNumber="483">6D-F</figureCitation>
). The call was strongly amplitude-modulated, beginning at a lower amplitude and increasing to the middle of the call, where the amplitude then decreased until the end of the call. The dominant frequency measured at peak amplitude of the call was 2390-2672 (2483
<normalizedToken originalValue="±">+/-</normalizedToken>
162;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 3) Hz, while the dominant frequency at the peak amplitude of the note was 2250-2813 (2458
<normalizedToken originalValue="±">+/-</normalizedToken>
149;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 72) Hz. For notes, the 90% bandwidth was from 1453-2297 (1942
<normalizedToken originalValue="±">+/-</normalizedToken>
177;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 72) Hz to 3000-4125 (3749
<normalizedToken originalValue="±">+/-</normalizedToken>
290;
<emphasis italics="true" pageId="0" pageNumber="483">n</emphasis>
= 72) Hz. No harmonic frequencies were visible on the spectrogram (Fig.
<figureCitation captionStart="Figure 6" captionStartId="F6" captionText="Figure 6. Oscillograms and spectrograms of the call of Gephyromantis marokoroko sp. nov. (Holotype: KU 343230). A. The entire call spectrogram and B. Entire call oscillogram; C. Power spectra / frequency spectrogram of a single note; D. A close-up spectrogram of four notes and E. Corresponding oscillogram; and F. an individual note taken from the middle of the call." figureDoi="10.3897/zse.97.73630.figure6" httpUri="https://binary.pensoft.net/fig/617725" pageId="0" pageNumber="483">6</figureCitation>
).
</paragraph>
<caption doi="10.3897/zse.97.73630.figure6" httpUri="https://binary.pensoft.net/fig/617725" pageId="0" pageNumber="483" start="Figure 6" startId="F6">
<paragraph pageId="0" pageNumber="483">
<emphasis bold="true" pageId="0" pageNumber="483">Figure 6.</emphasis>
Oscillograms and spectrograms of the call of
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
sp. nov. (Holotype: KU 343230).
<emphasis bold="true" pageId="0" pageNumber="483">A.</emphasis>
The entire call spectrogram and
<emphasis bold="true" pageId="0" pageNumber="483">B.</emphasis>
Entire call oscillogram;
<emphasis bold="true" pageId="0" pageNumber="483">C.</emphasis>
Power spectra/frequency spectrogram of a single note;
<emphasis bold="true" pageId="0" pageNumber="483">D.</emphasis>
A close-up spectrogram of four notes and
<emphasis bold="true" pageId="0" pageNumber="483">E.</emphasis>
Corresponding oscillogram; and
<emphasis bold="true" pageId="0" pageNumber="483">F.</emphasis>
an individual note taken from the middle of the call.
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="phylogenetics">
<paragraph pageId="0" pageNumber="483">Phylogenetics.</paragraph>
<paragraph pageId="0" pageNumber="483">
The phylogenetic results support the morphological diagnosis by placing
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
within the
<taxonomicName authorityName="Dubois" authorityYear="1980" class="Amphibia" family="Mantellidae" genus="Laurentomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Laurentomantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">Laurentomantis</taxonomicName>
subgenus with strong support. At the species level,
<taxonomicName genus="G." lsidName="G. marokoroko" pageId="0" pageNumber="483" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">G. marokoroko</emphasis>
</taxonomicName>
is monophyletic with strong support in ML and BI analyses (BS = 100, PP = 1.00; Fig.
<figureCitation captionStart="Figure 3" captionStartId="F3" captionText="Figure 3. Results of phylogenetic analysis of the mitochondrial 16 S rRNA barcode 3 ' marker for Maximum Likelihood (ML) and Bayesian Inference (BI). Topology is a consensus tree from IQ-Tree. The support values are shown as Bootstrap on top and Posterior Probability on the bottom only for nodes that were not perfectly supported. Note that Gephyromantis marokoroko sp. nov. placement in the clade is weakly supported in both analyses." figureDoi="10.3897/zse.97.73630.figure3" httpUri="https://binary.pensoft.net/fig/617722" pageId="0" pageNumber="483">3</figureCitation>
). Uncorrected p-distances, using the 16S fragment, indicate that
<taxonomicName genus="G." lsidName="G. ventrimaculatus" pageId="0" pageNumber="483" rank="species" species="ventrimaculatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. ventrimaculatus</emphasis>
</taxonomicName>
has the lowest distance to the new species, at ~ 6-9%. The combined nine marker multi-locus dataset places the new species sister to
<taxonomicName genus="G." lsidName="G. striatus" pageId="0" pageNumber="483" rank="species" species="striatus">
<emphasis italics="true" pageId="0" pageNumber="483">G. striatus</emphasis>
</taxonomicName>
with strong support (BS = 98; PP = 1.00) in both BI and ML analyses (Fig.
<figureCitation captionStart="Figure 4" captionStartId="F4" captionText="Figure 4. Results of phylogenetic analyses of the concatenated alignment of five mitochondrial and four nuclear markers for Maximum Likelihood (ML) and Bayesian Inference (BI). Topology is a consensus tree from IQ-Tree. On the right, the dots represent markers that were present in blue and absent in orange for each sample. The notes marked with a circle are those that did not receive perfect support (Bootstrap = 100; Posterior Probability = 1.00) from ML and BI, with the support values as BS on top and PP on the bottom. Note that Gephyromantis marokoroko sp. nov. has strong support in both analyses for a sister relationship to G. striatus." figureDoi="10.3897/zse.97.73630.figure4" httpUri="https://binary.pensoft.net/fig/617723" pageId="0" pageNumber="483">4</figureCitation>
). Overall, these results provide strong evidence that the species is a separately evolving lineage and strong evidence for the new species phylogenetic placement.
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="distribution">
<paragraph pageId="0" pageNumber="483">Distribution.</paragraph>
<paragraph pageId="0" pageNumber="483">
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
is known from several sites in the forests in the vicinity of Andasibe, but has only been found at high elevation sites (~ 1000-1200 m a.s.l.; Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. The distribution of Gephyromantis marokoroko sp. nov. in east-central Madagascar, view from above (A.) and from a profile view (B.). The black star marker indicates the type locality at Vohidrazana Forest where the black circle &quot; locality &quot; markers indicate other confirmed localities for the new species. Gephyromantis marokoroko sp. nov. is also found at high elevations and, thus, is likely distributed at other high elevation sites not surveyed. Elevational and satellite imagery data acquired from the USGS Earth Explorer (http: // earthexplorer. usgs. gov)." figureDoi="10.3897/zse.97.73630.figure1" httpUri="https://binary.pensoft.net/fig/617720" pageId="0" pageNumber="483">1</figureCitation>
). The new species is known from the following localities: Vohidrazana Forest (
<geoCoordinate degrees="18.976" direction="south" orientation="latitude" precision="55" value="-18.976">18.976°S</geoCoordinate>
,
<geoCoordinate degrees="48.499" direction="east" orientation="longitude" precision="55" value="48.499">48.499°E</geoCoordinate>
), Tavalobe (
<geoCoordinate degrees="19.005" direction="south" orientation="latitude" precision="55" value="-19.005">19.005°S</geoCoordinate>
,
<geoCoordinate degrees="48.461" direction="east" orientation="longitude" precision="55" value="48.461">48.461°E</geoCoordinate>
), Vohimana (
<geoCoordinate degrees="18.926" direction="south" orientation="latitude" precision="55" value="-18.926">18.926°S</geoCoordinate>
,
<geoCoordinate degrees="48.489" direction="east" orientation="longitude" precision="55" value="48.489">48.489°E</geoCoordinate>
) and Andasibe-Mantadia National Park (Belakato:
<geoCoordinate degrees="18.821" direction="south" orientation="latitude" precision="55" value="-18.821">18.821°S</geoCoordinate>
,
<geoCoordinate degrees="48.439" direction="east" orientation="longitude" precision="55" value="48.439">48.439°E</geoCoordinate>
).
</paragraph>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="natural history">
<paragraph pageId="0" pageNumber="483">Natural history.</paragraph>
<paragraph pageId="0" pageNumber="483">
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
is apparently locally rare and, thus far, only found within undisturbed, primary forests at highland elevations (ca. 1000-1200 m). Individuals of the species were perched on the surfaces of vegetation less than 50 cm in height (Fig.
<figureCitation captionStart="Figure 7" captionStartId="F7" captionText="Figure 7. In-situ photograph of Gephyromantis marokoroko sp. nov. (UADBA-CRH 1629)." figureDoi="10.3897/zse.97.73630.figure7" httpUri="https://binary.pensoft.net/fig/617726" pageId="0" pageNumber="483">7</figureCitation>
). The species was infrequently encountered, always after moderate to heavy rain, with multiple individuals occasionally grouped in small clusters (~ 20 m2). The
<normalizedToken originalValue="species">species'</normalizedToken>
call is very quiet and irregular and is barely audible to a human observer, even within three metres of a calling individual. Individuals of the new species were often found syntopically with another
<taxonomicName authorityName="Dubois" authorityYear="1980" class="Amphibia" family="Mantellidae" genus="Laurentomantis" higherTaxonomySource="GBIF" kingdom="Animalia" lsidName="Laurentomantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Laurentomantis</emphasis>
</taxonomicName>
,
<taxonomicName genus="G." lsidName="G." pageId="0" pageNumber="483" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">G.</emphasis>
</taxonomicName>
sp. Ca13, which is a candidate species identified in
<bibRefCitation DOI="https://doi.org/10.1073/pnas.0810821106" author="Vieites, DR" journalOrPublisher="Proceedings of the National Academy of Sciences" pageId="0" pageNumber="483" pagination="8267 - 8272" refId="B50" refString="Vieites, DR, Wollenberg, KC, Andreone, F, Koehler, J, Glaw, F, Vences, M, 2009. Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences 106: 8267 - 8272, DOI: https://doi.org/10.1073/pnas.0810821106" title="Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory." url="https://doi.org/10.1073/pnas.0810821106" volume="106" year="2009">Vieites et al. (2009)</bibRefCitation>
. Other syntopic
<taxonomicName authorityName="Methuen" authorityYear="1920" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis</emphasis>
</taxonomicName>
include
<taxonomicName genus="G." lsidName="G. eiselti" pageId="0" pageNumber="483" rank="species" species="eiselti">
<emphasis italics="true" pageId="0" pageNumber="483">G. eiselti</emphasis>
</taxonomicName>
,
<taxonomicName genus="G." lsidName="G. salegy" pageId="0" pageNumber="483" rank="species" species="salegy">
<emphasis italics="true" pageId="0" pageNumber="483">G. salegy</emphasis>
</taxonomicName>
,
<taxonomicName genus="G." lsidName="G. plicifer" pageId="0" pageNumber="483" rank="species" species="plicifer">G. sp. aff. plicifer</taxonomicName>
(not yet assessed for a candidate species number) and
<taxonomicName genus="G." lsidName="G. cornutus" pageId="0" pageNumber="483" rank="species" species="cornutus">
<emphasis italics="true" pageId="0" pageNumber="483">G. cornutus</emphasis>
</taxonomicName>
.
</paragraph>
<caption doi="10.3897/zse.97.73630.figure7" httpUri="https://binary.pensoft.net/fig/617726" pageId="0" pageNumber="483" start="Figure 7" startId="F7">
<paragraph pageId="0" pageNumber="483">
<emphasis bold="true" pageId="0" pageNumber="483">Figure 7.</emphasis>
In-situ photograph of
<taxonomicName authorityName="Hutter &amp; Andriampenomanana &amp; Andrianasolo &amp; Cobb &amp; Razafindraibe &amp; Abraham &amp; Lambert" authorityYear="2021" class="Amphibia" family="Mantellidae" genus="Gephyromantis" higherTaxonomySource="CoL" kingdom="Animalia" lsidName="Gephyromantis marokoroko" order="Anura" pageId="0" pageNumber="483" phylum="Chordata" rank="species" species="marokoroko">
<emphasis italics="true" pageId="0" pageNumber="483">Gephyromantis marokoroko</emphasis>
</taxonomicName>
sp. nov. (UADBA-CRH1629).
</paragraph>
</caption>
</subSubSection>
<subSubSection pageId="0" pageNumber="483" type="conservation status">
<paragraph pageId="0" pageNumber="483">Conservation status.</paragraph>
<paragraph pageId="0" pageNumber="483">
The new species is known from Andasibe-Mantadia National Park and several other managed areas (e.g. Vohimana, the community managed Vohidrazana Forest and Tavalobe). However, as currently understood, the distribution of this species is severely fragmented and restricted to only four known high-elevation localities (~ 1000-1200 m), which are very small patches with no connectivity (Fig.
<figureCitation captionStart="Figure 1" captionStartId="F1" captionText="Figure 1. The distribution of Gephyromantis marokoroko sp. nov. in east-central Madagascar, view from above (A.) and from a profile view (B.). The black star marker indicates the type locality at Vohidrazana Forest where the black circle &quot; locality &quot; markers indicate other confirmed localities for the new species. Gephyromantis marokoroko sp. nov. is also found at high elevations and, thus, is likely distributed at other high elevation sites not surveyed. Elevational and satellite imagery data acquired from the USGS Earth Explorer (http: // earthexplorer. usgs. gov)." figureDoi="10.3897/zse.97.73630.figure1" httpUri="https://binary.pensoft.net/fig/617720" pageId="0" pageNumber="483">1</figureCitation>
). Many other high elevation sites in the region have been surveyed by the authors over three field seasons. Furthermore, Vohidrazana Forest and Tavalobe face ongoing threats that result in the reduction of quality and extent of habitat. For example, slash-and-burn agriculture and forest products are frequently extracted directly from this
<normalizedToken originalValue="species">species'</normalizedToken>
habitats that are outside protected areas. Given this information, we categorise this species as
<normalizedToken originalValue="“Endangered”">&quot;Endangered&quot;</normalizedToken>
[B1ab(iii-iv)] following IUCN Criteria (
<bibRefCitation author="IUCN" journalOrPublisher="Molecular Ecology Resources" pageId="0" pageNumber="483" refId="B22" refString="IUCN, 2001. IUCN Red List Categories and Criteria: Version 3.1. IUCN, Switzerland and Cambridge." title="IUCN Red List Categories and Criteria: Version 3.1. IUCN, Switzerland and Cambridge." year="2001">IUCN 2001</bibRefCitation>
).
</paragraph>
</subSubSection>
</treatment>
</document>