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<document ID-DOI="10.11646/megataxa.7.2.1" ID-ISSN="2703-3090" ID-Zenodo-Dep="7441023" ID-ZooBank="2FD8C310-6486-4592-92F6-5EB894EBD6AC" checkinTime="1671098794254" checkinUser="plazi" docAuthor="Scherz, Mark D., Crottini, Angelica, Hutter, Carl R., Hildenbrand, Andrea, Andreone, Franco, Fulgence, Thio Rosin, Köhler, Gunther, Ndriantsoa, Serge Herilala, Ohler, Annemarie, Preick, Michaela, Rakotoarison, Andolalao, Rancilhac, Loïs, Raselimanana, Achille P., Riemann, Jana C., Rödel, Mark-Oliver, Rosa, Gonçalo M., Streicher, Jeffrey W., Vieites, David R., Köhler, Jörn, Hofreiter, Michael, Glaw, Frank &amp; Vences, Miguel" docDate="2022" docId="5F25F715FF94FF874CB14C0F49827CAE" docLanguage="en" docName="megataxa.7.2.1.pdf" docOrigin="Megataxa 7 (2)" docSource="http://dx.doi.org/10.5281/zenodo.6687412" docStyle="DocumentStyle:50B6B96D96A55C21793B52C23E95B310.3:Megataxa.2020-.journal_article" docStyleId="50B6B96D96A55C21793B52C23E95B310" docStyleName="Megataxa.2020-.journal_article" docStyleVersion="3" docTitle="Mantidactylus ambohimitombi subsp. miloko Scherz &amp; Crottini &amp; Hutter &amp; Hildenbrand &amp; Andreone &amp; Fulgence &amp; Köhler &amp; Ndriantsoa &amp; Ohler &amp; Preick &amp; Rakotoarison &amp; Rancilhac &amp; Raselimanana &amp; Riemann &amp; Rödel &amp; Rosa &amp; Streicher &amp; Vieites &amp; Köhler &amp; Hofreiter &amp; Glaw &amp; Vences 2022, ssp. nov." docType="treatment" docVersion="1" lastPageNumber="186" masterDocId="A31C8F6DFFDCFFCE4C394B044D497F0B" masterDocTitle="An inordinate fondness for inconspicuous brown frogs: integration of phylogenomics, archival DNA analysis, morphology, and bioacoustics yields 24 new taxa in the subgenus Brygoomantis (genus Mantidactylus) from Madagascar" masterLastPageNumber="311" masterPageNumber="113" pageNumber="185" updateTime="1672856617530" updateUser="ExternalLinkService" zenodo-license-document="CC-BY-NC-4.0">
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<mods:titleInfo>
<mods:title>An inordinate fondness for inconspicuous brown frogs: integration of phylogenomics, archival DNA analysis, morphology, and bioacoustics yields 24 new taxa in the subgenus Brygoomantis (genus Mantidactylus) from Madagascar</mods:title>
</mods:titleInfo>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Scherz, Mark D.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Crottini, Angelica</mods:namePart>
<mods:affiliation>CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, InBIO Laboratório Associado, Campus de Vairão, Universidade do Porto, 4485 - 661 Vairão, Portugal</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Hutter, Carl R.</mods:namePart>
<mods:nameIdentifier type="ORCID">0000-0001-6381-6339</mods:nameIdentifier>
<mods:affiliation>Museum of Natural Sciences and Department of Biological Sciences, Louisiana State University, Baton Rouge, LA 70803, USA Ễcarl. hutter @ gmail. com; https: // orcid. org / 0000 - 0001 - 6381 - 6339</mods:affiliation>
<mods:nameIdentifier type="email">USAỄcarl.hutter@gmail.com</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Hildenbrand, Andrea</mods:namePart>
<mods:affiliation>Hauptstr. 13, 82234 Wessling, Germany Ễandrea @ gutachten-hildenbrand. de</mods:affiliation>
<mods:nameIdentifier type="email">GermanyỄandrea@gutachten-hildenbrand.de</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Andreone, Franco</mods:namePart>
<mods:affiliation>Museo Regionale di Scienze Naturali, Via G. Giolitti, 36, 10123 Torino, Italy</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Fulgence, Thio Rosin</mods:namePart>
<mods:nameIdentifier type="ORCID">0000-0001-7205-7282</mods:nameIdentifier>
<mods:affiliation>Natural and Environmental Sciences, Regional University Centre of the SAVA Region (CURSA), Antalaha, Madagascar Ễthiorosinf @ yahoo. fr; https: // orcid. org / 0000 - 0001 - 7205 - 7282</mods:affiliation>
<mods:nameIdentifier type="email">MadagascarỄthiorosinf@yahoo.fr</mods:nameIdentifier>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Köhler, Gunther</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ndriantsoa, Serge Herilala</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Ohler, Annemarie</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Preick, Michaela</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rakotoarison, Andolalao</mods:namePart>
<mods:affiliation>Mention Zoologie et Biodiversité Animale, Université dAntananarivo, BP 906, Antananarivo, 101 Madagascar</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rancilhac, Loïs</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Raselimanana, Achille P.</mods:namePart>
<mods:affiliation>Mention Zoologie et Biodiversité Animale, Université dAntananarivo, BP 906, Antananarivo, 101 Madagascar</mods:affiliation>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Riemann, Jana C.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rödel, Mark-Oliver</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rosa, Gonçalo M.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Streicher, Jeffrey W.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Vieites, David R.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Köhler, Jörn</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Hofreiter, Michael</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Glaw, Frank</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Vences, Miguel</mods:namePart>
</mods:name>
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<mods:titleInfo>
<mods:title>Megataxa</mods:title>
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<mods:date>2022</mods:date>
<mods:detail type="pubDate">
<mods:number>2022-12-15</mods:number>
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<mods:detail type="volume">
<mods:number>7</mods:number>
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<mods:number>2</mods:number>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.11646/megataxa.7.2.1</mods:identifier>
<mods:identifier type="ISSN">2703-3090</mods:identifier>
<mods:identifier type="Zenodo-Dep">7441023</mods:identifier>
<mods:identifier type="ZooBank">2FD8C310-6486-4592-92F6-5EB894EBD6AC</mods:identifier>
</mods:mods>
<treatment LSID="urn:lsid:plazi:treatment:5F25F715FF94FF874CB14C0F49827CAE" httpUri="http://treatment.plazi.org/id/5F25F715FF94FF874CB14C0F49827CAE" lastPageId="73" lastPageNumber="186" pageId="72" pageNumber="185">
<subSubSection box="[136,670,1803,1829]" pageId="72" pageNumber="185" type="nomenclature">
<paragraph blockId="72.[136,670,1803,1829]" box="[136,670,1803,1829]" pageId="72" pageNumber="185">
<taxonomicName authority="Scherz &amp; Crottini &amp; Hutter &amp; Hildenbrand &amp; Andreone &amp; Fulgence &amp; Köhler &amp; Ndriantsoa &amp; Ohler &amp; Preick &amp; Rakotoarison &amp; Rancilhac &amp; Raselimanana &amp; Riemann &amp; Rödel &amp; Rosa &amp; Streicher &amp; Vieites &amp; Köhler &amp; Hofreiter &amp; Glaw &amp; Vences, 2022" authorityName="Scherz &amp; Crottini &amp; Hutter &amp; Hildenbrand &amp; Andreone &amp; Fulgence &amp; Köhler &amp; Ndriantsoa &amp; Ohler &amp; Preick &amp; Rakotoarison &amp; Rancilhac &amp; Raselimanana &amp; Riemann &amp; Rödel &amp; Rosa &amp; Streicher &amp; Vieites &amp; Köhler &amp; Hofreiter &amp; Glaw &amp; Vences" authorityYear="2022" box="[136,562,1803,1829]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="subSpecies" species="ambohimitombi" status="ssp. nov." subSpecies="miloko">
<emphasis box="[136,562,1803,1829]" italics="true" pageId="72" pageNumber="185">Mantidactylus ambohimitombi miloko</emphasis>
</taxonomicName>
<emphasis bold="true" box="[569,670,1803,1829]" pageId="72" pageNumber="185">
<taxonomicNameLabel box="[569,670,1803,1829]" pageId="72" pageNumber="185" rank="subSpecies">ssp. nov.</taxonomicNameLabel>
</emphasis>
</paragraph>
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<subSubSection pageId="72" pageNumber="185" type="description">
<paragraph blockId="72.[136,778,1867,2085]" lastBlockId="72.[810,1452,171,2085]" pageId="72" pageNumber="185">
<emphasis box="[136,414,1867,1893]" italics="true" pageId="72" pageNumber="185">Identity and justification</emphasis>
.—This lineage of the
<taxonomicName box="[669,778,1868,1893]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="curtus">
<emphasis box="[669,778,1868,1893]" italics="true" pageId="72" pageNumber="185">M. curtus</emphasis>
</taxonomicName>
clade was considered as confirmed candidate species
<emphasis box="[747,778,1900,1925]" italics="true" pageId="72" pageNumber="185">M.</emphasis>
sp. 18 by
<bibRefCitation author="Vieites, D. R. &amp; Wollenberg, K. C. &amp; Andreone, F. &amp; Glaw, F. &amp; Vences, M." box="[256,489,1931,1957]" pageId="72" pageNumber="185" pagination="8267 - 8272" refId="ref134563" refString="Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. &amp; Vences, M. (2009) Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the USA, 106, 8267 - 8272. https: // doi. org / 10.1073 / pnas. 0810821106" type="journal article" year="2009">
Vieites
<emphasis box="[344,404,1931,1957]" italics="true" pageId="72" pageNumber="185">et al.</emphasis>
(2009)
</bibRefCitation>
due to its mitochondrial divergence in concert with slight but distinct differences in colour pattern, and as
<emphasis box="[449,480,1996,2021]" italics="true" pageId="72" pageNumber="185">M.</emphasis>
sp. Ca18 by
<bibRefCitation author="Perl, R. G. B. &amp; Nagy, Z. T. &amp; Sonet, G. &amp; Glaw, F. &amp; Wollenberg, K. C. &amp; Vences, M." pageId="72" pageNumber="185" pagination="197 - 206" refId="ref129051" refString="Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. &amp; Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942" type="journal article" year="2014">
Perl
<emphasis box="[714,778,1995,2021]" italics="true" pageId="72" pageNumber="185">et al.</emphasis>
(2014)
</bibRefCitation>
. It was depicted as
<taxonomicName box="[451,652,2027,2053]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="undetermined">
<emphasis box="[451,611,2027,2053]" italics="true" pageId="72" pageNumber="185">Mantidactylus</emphasis>
sp.
</taxonomicName>
<taxonomicName authority="Ambohitantely" authorityName="Ambohitantely" box="[661,778,2027,2053]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" isUncertain="true" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="curtus">
aff.
<emphasis box="[708,778,2028,2053]" italics="true" pageId="72" pageNumber="185">curtus</emphasis>
</taxonomicName>
“Ambohitantely” by
<bibRefCitation author="Glaw, F. &amp; Vences, M." box="[383,670,2059,2085]" pageId="72" pageNumber="185" refId="ref125641" refString="Glaw, F. &amp; Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences &amp; Glaw Verlags GbR, Cologne, Germany, 496 pp. Third Edition." type="book" year="2007">Glaw and Vences (2007)</bibRefCitation>
. We here consider this lineage provisionally as a subspecies of
<taxonomicName authorityName="Boulenger" authorityYear="1919" box="[810,1032,203,229]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="ambohimitombi">
<emphasis box="[810,1032,203,229]" italics="true" pageId="72" pageNumber="185">M. ambohimitombi</emphasis>
</taxonomicName>
based on the following rationale: (i) it belongs to the same general mitochondrial lineage as the nominal subspecies in the 16S tree, and the two are also closely related in the phylogenomic tree; (ii) morphologically, the two lineages are similar to each other, except for a somewhat more distinct dorsal pattern in
<emphasis box="[841,987,395,421]" italics="true" pageId="72" pageNumber="185">M. a. miloko</emphasis>
<emphasis bold="true" box="[997,1099,395,421]" pageId="72" pageNumber="185">ssp. nov.</emphasis>
; (iii) based on two samples in our mitochondrial tree, RJS 1877 and ACZC 4254, a very similar mitochondrial haplotype to the one from Ambohitantely also occurs at Ankaratra, suggesting the possibility of past or ongoing gene flow between localities or mitochondrial introgression;and (iv)based on sequences of specimens APR 10803, APR 10638, and APR 10663, the lineage also occurs in Angavokely and Ankazomivady, thus rather close to Ankaratra, suggesting the two lineages may be parapatric and could hypothetically have a hybrid zone. Since the status of the Ambohitantely population as a fully isolated evolutionary lineage is thus not fully verifiable with the data at hand, we consider the status as a subspecies of
<taxonomicName authorityName="Boulenger" authorityYear="1919" box="[986,1200,811,837]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="ambohimitombi">
<emphasis box="[986,1200,811,837]" italics="true" pageId="72" pageNumber="185">M. ambohimitombi</emphasis>
</taxonomicName>
to be adequate.
</paragraph>
</subSubSection>
<subSubSection pageId="72" pageNumber="185" type="materials_examined">
<paragraph blockId="72.[810,1452,171,2085]" pageId="72" pageNumber="185">
<materialsCitation collectingDate="2005-01-18" collectionCode="ZSM" collectorName="M. Vences; L. du Preez; P. Bora; L. Raharivololoniaina; R. D. Randrianiaina; T. Razafindraibe &amp; E. Randriamitso" country="Madagascar" elevation="1580" latitude="-18.1725" location="Ambohitantely Special Reserve" longLatPrecision="7" longitude="47.2768" municipality="Analamanga Region" pageId="72" pageNumber="185" specimenCode="2143" specimenCount="1" typeStatus="holotype">
<emphasis box="[858,961,843,869]" italics="true" pageId="72" pageNumber="185">
<typeStatus box="[858,961,843,869]" pageId="72" pageNumber="185">Holotype</typeStatus>
</emphasis>
.—
<collectionCode box="[995,1053,843,869]" collectionName="Germany, Muenchen [= Munich], Zoologische Staatssammlung" pageId="72" pageNumber="185">ZSM</collectionCode>
<specimenCode box="[1074,1180,843,870]" pageId="72" pageNumber="185">219/2005</specimenCode>
(FGZC 2143), adult male, collected by
<collectorName box="[1032,1156,875,901]" pageId="72" pageNumber="185">M. Vences</collectorName>
,
<collectorName box="[1171,1310,875,901]" pageId="72" pageNumber="185">L. du Preez</collectorName>
,
<collectorName box="[1325,1412,875,901]" pageId="72" pageNumber="185">P. Bora</collectorName>
,
<collectorName pageId="72" pageNumber="185">L. Raharivololoniaina</collectorName>
,
<collectorName box="[1036,1253,907,933]" pageId="72" pageNumber="185">R.D. Randrianiaina</collectorName>
,
<collectorName box="[1261,1447,907,933]" pageId="72" pageNumber="185">T. Razafindraibe</collectorName>
,
<collectorName box="[810,996,939,965]" pageId="72" pageNumber="185">E. Randriamitso</collectorName>
on
<date box="[1047,1242,939,966]" pageId="72" pageNumber="185" value="2005-01-18">
<collectingDate box="[1047,1242,939,966]" pageId="72" pageNumber="185" value="2005-01-18">18 January 2005</collectingDate>
</date>
at
<location LSID="urn:lsid:plazi:treatment:5F25F715FF94FF874CB14C0F49827CAE:D25310D8FF94FF86493A48AF4E967CEE" country="Madagascar" latitude="-18.1725" longLatPrecision="7" longitude="47.2768" municipality="Analamanga Region" name="Ambohitantely Special Reserve" pageId="72" pageNumber="185">Ambohitantely Special Reserve</location>
,
<locationDeviation box="[1002,1213,971,997]" pageId="72" pageNumber="185">Jardin Botanique</locationDeviation>
, at a site about 500 m fromthegeographicalcoordinates
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,
<geoCoordinate box="[1311,1448,1003,1029]" degrees="047.2768" direction="east" orientation="longitude" pageId="72" pageNumber="185" precision="5" value="47.2768">047.2768°E</geoCoordinate>
,
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<quantity box="[810,898,1035,1062]" metricMagnitude="3" metricUnit="m" metricValue="1.58" pageId="72" pageNumber="185" unit="m" value="1580.0">1580 m</quantity>
</elevation>
a.s.l.,
<collectingMunicipality box="[977,1214,1035,1061]" pageId="72" pageNumber="185">Analamanga Region</collectingMunicipality>
,
<collectingCountry box="[1228,1365,1035,1061]" name="Madagascar" pageId="72" pageNumber="185">Madagascar</collectingCountry>
.
</materialsCitation>
A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis.
</paragraph>
<paragraph blockId="72.[810,1452,171,2085]" pageId="72" pageNumber="185">
<emphasis box="[858,961,1132,1157]" italics="true" pageId="72" pageNumber="185">Paratype</emphasis>
.—A single
<materialsCitation collectionCode="ZSM" collectorName="M. Vences; L. du Preez; P. Bora; L. Raharivololoniaina; R. D. Randrianiaina; T. Razafindraibe &amp; E. Randriamitso" pageId="72" pageNumber="185" specimenCode="237/2005" specimenCount="1" typeStatus="paratype">
<typeStatus box="[1132,1233,1131,1157]" pageId="72" pageNumber="185">paratype</typeStatus>
:
<collectionCode box="[1262,1320,1131,1157]" collectionName="Germany, Muenchen [= Munich], Zoologische Staatssammlung" pageId="72" pageNumber="185">ZSM</collectionCode>
<specimenCode box="[1346,1452,1131,1158]" pageId="72" pageNumber="185">237/2005</specimenCode>
(FGZC 2172), adult
<specimenCount box="[1038,1117,1163,1189]" pageId="72" pageNumber="185" type="female">female</specimenCount>
, with the same collection data as holotype.
</materialsCitation>
</paragraph>
<paragraph blockId="72.[810,1452,171,2085]" pageId="72" pageNumber="185">
<emphasis box="[858,1104,1227,1253]" italics="true" pageId="72" pageNumber="185">Additional material.</emphasis>
—The following specimens probably belong to this taxon but are only assigned tentatively and not designated as paratypes because no molecular data is available:
<materialsCitation collectionCode="ZMA" collectorName="R. M. A. Blommers-Schl" elevation="1500" location="Tampoketsa d'Ankazobe" pageId="72" pageNumber="185" specimenCode="6859" specimenCount="2">
<collectionCode box="[1136,1197,1323,1349]" pageId="72" pageNumber="185">ZMA</collectionCode>
<specimenCode box="[1207,1263,1323,1350]" pageId="72" pageNumber="185">6859</specimenCode>
(
<specimenCount box="[1281,1452,1323,1349]" pageId="72" pageNumber="185" type="generic">two specimens</specimenCount>
with field numbers 675 and 676) and
</materialsCitation>
<materialsCitation collectionCode="ZMA" collectorName="R. M. A. Blommers-Schl" elevation="2200" location="Tampoketsa d'Ankazobe" pageId="72" pageNumber="185" specimenCode="6860" specimenCount="6">
ZMA 6860 (
<specimenCount pageId="72" pageNumber="185" type="generic">six specimens</specimenCount>
with field numbers 910914, 917), collected by
<collectorName box="[844,1116,1419,1447]" pageId="72" pageNumber="185">R.M.A. Blommers-Schl</collectorName>
̂sser in 1972 at 1500 m (
<collectionCode box="[1391,1452,1419,1445]" pageId="72" pageNumber="185">ZMA</collectionCode>
<specimenCode box="[810,868,1451,1478]" pageId="72" pageNumber="185">6859</specimenCode>
) and
<quantity box="[944,1037,1451,1477]" metricMagnitude="3" metricUnit="m" metricValue="2.2" pageId="72" pageNumber="185" unit="m" value="2200.0">
<elevation box="[944,1037,1451,1477]" metricMagnitude="3" metricUnit="m" metricValue="2.2" pageId="72" pageNumber="185" unit="m" value="2200.0">2200 m</elevation>
</quantity>
a.s.l. (
<collectionCode box="[1127,1188,1451,1477]" pageId="72" pageNumber="185">ZMA</collectionCode>
<specimenCode box="[1202,1260,1451,1477]" pageId="72" pageNumber="185">6860</specimenCode>
) at
<location LSID="urn:lsid:plazi:treatment:5F25F715FF94FF874CB14C0F49827CAE:D25310D8FF94FF86491D4EAF4EFF7AEE" name="Tampoketsa d'Ankazobe" pageId="72" pageNumber="185">Tampoketsa dAnkazobe</location>
.
</materialsCitation>
</paragraph>
</subSubSection>
<subSubSection lastPageId="73" lastPageNumber="186" pageId="72" pageNumber="185" type="diagnosis">
<paragraph blockId="72.[810,1452,171,2085]" lastBlockId="73.[136,778,171,2085]" lastPageId="73" lastPageNumber="186" pageId="72" pageNumber="185">
<emphasis box="[858,1427,1515,1541]" italics="true" pageId="72" pageNumber="185">
Diagnosis.—
<taxonomicName authorityName="Scherz &amp; Crottini &amp; Hutter &amp; Hildenbrand &amp; Andreone &amp; Fulgence &amp; Köhler &amp; Ndriantsoa &amp; Ohler &amp; Preick &amp; Rakotoarison &amp; Rancilhac &amp; Raselimanana &amp; Riemann &amp; Rödel &amp; Rosa &amp; Streicher &amp; Vieites &amp; Köhler &amp; Hofreiter &amp; Glaw &amp; Vences" authorityYear="2022" box="[1002,1427,1515,1541]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="subSpecies" species="ambohimitombi" subSpecies="miloko">Mantidactylus ambohimitombi miloko</taxonomicName>
</emphasis>
is a member of the
<taxonomicName box="[1005,1113,1548,1573]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="curtus">
<emphasis box="[1005,1113,1548,1573]" italics="true" pageId="72" pageNumber="185">M. curtus</emphasis>
</taxonomicName>
clade, and the sister group of
<emphasis box="[810,1249,1579,1605]" italics="true" pageId="72" pageNumber="185">
M. a.
<taxonomicName authorityName="Boulenger" authorityYear="1919" box="[880,1056,1579,1605]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="ambohimitombi">ambohimitombi</taxonomicName>
+ M. a. marefo.
</emphasis>
Morphologically, it is very similar to
<emphasis box="[1047,1296,1611,1637]" italics="true" pageId="72" pageNumber="185">
M. a.
<taxonomicName authorityName="Boulenger" authorityYear="1919" box="[1120,1296,1611,1637]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="ambohimitombi">ambohimitombi</taxonomicName>
</emphasis>
. See
<tableCitation box="[1365,1451,1611,1637]" captionStart="TABLE 4" captionStartId="54.[134,222,490,516]" captionTargetPageId="54" captionText="TABLE 4. Summary of the most important diagnostic characters of species in the Mantidactylus subgenus Brygoomantis. SVL is given only based on reliably sexed specimens, summarised and rounded to the nearest mm from original measurements in Tables 511. Dorsolateral ridges are always weakly expressed in Brygoomantis, but clearly visible in some species while completely absent in others. Dorsal skin rugosity is estimated from pictures in life and can appear different in preservative: 0, smooth; 1, finely granular; 2, coarsely granular / tubercular. Webbing at toe 5 is given as 0, half or less than half of one phalanx free of web, not counting terminal phalanx (corresponding to numbers 00.5 in webbing formula); 1, about one phalanx free of web (0.751.25); 2, more than one phalanx free of web (1.52). (Yes) in parentheses indicates that the respective character state is found in some individuals of the respective species, and often only weakly expressed, while it is missing in others. Lat.met., state of the lateral metatarsalia, either separated (sep.) by webbing or connected (conn.) by tissue. NM, no measurements available. *For the advertisement calls of M. katae sp. nov. and M. fergusoni sp. nov., homology of call components remains unclear. Here, each unit is interpreted as singlepulse call (= single-pulse note) arranged in series (under this definition the provided pulse rate reflects call rate), but it would also be possible to consider the entire vocalization as single-note multi-pulsed call with very long duration, containing very widely spaced pulses. Values for pulse rate (pulses/second) are rounded. We here evaluate the presence versus absence of regular call series (last column), where regular series are considered as such, if calls are emitted at really regular intervals in series of similar duration containing a more or less defined number of calls. # In M. ambohimitombi, several unsexed individuals of the type series, probably females, reach sizes of up to 65 mm, being the largest of all Brygoomantis known. †One female of M. biporus of 19.3 mm SVL is probably a not fully mature specimen; other females of the species measure 3136 mm SVL." httpUri="http://table.plazi.org/id/83F3168BFFEAFFF84CBF4AEE4C717C8F" pageId="72" pageNumber="185" tableUuid="83F3168BFFEAFFF84CBF4AEE4C717C8F">Table 4</tableCitation>
for a list of diagnostic morphological characters. The combination of relatively large body size of up to
<quantity box="[1366,1447,1675,1701]" metricMagnitude="-2" metricUnit="m" metricValue="4.9" pageId="72" pageNumber="185" unit="mm" value="49.0">49 mm</quantity>
, slightly granular skin without clearly defined dorsolateral ridges, and small tympanum diameter of a maximum of 10% of SVL in males, distinguishes
<emphasis box="[1245,1395,1771,1797]" italics="true" pageId="72" pageNumber="185">M. a. miloko</emphasis>
<emphasis bold="true" pageId="72" pageNumber="185">ssp. nov.</emphasis>
from species of the other clades. Within the
<taxonomicName class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="curtus">
<emphasis italics="true" pageId="72" pageNumber="185">M. curtus</emphasis>
</taxonomicName>
clade,
<emphasis box="[964,1261,1835,1861]" italics="true" pageId="72" pageNumber="185">
<taxonomicName box="[964,1074,1835,1861]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="alutus">M. alutus</taxonomicName>
,
<taxonomicName box="[1087,1261,1835,1861]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="madecassus">M. madecassus</taxonomicName>
</emphasis>
and
<taxonomicName box="[1319,1451,1835,1861]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="pauliani">
<emphasis box="[1319,1451,1835,1861]" italics="true" pageId="72" pageNumber="185">M. pauliani</emphasis>
</taxonomicName>
have smaller body sizes and are distinguished by either a usually shorter snout (
<emphasis italics="true" pageId="72" pageNumber="185">
<taxonomicName box="[1107,1289,1899,1925]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="madecassus">M. madecassus</taxonomicName>
,
<taxonomicName box="[1307,1447,1899,1925]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="pauliani">M. pauliani</taxonomicName>
, M. a. marefo
</emphasis>
), or presence of rather distinct dorsolateral ridges (
<taxonomicName box="[901,1013,1963,1989]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="alutus">
<emphasis box="[901,1013,1963,1989]" italics="true" pageId="72" pageNumber="185">M. alutus</emphasis>
</taxonomicName>
) (
<tableCitation box="[1046,1138,1963,1989]" captionStart="TABLE 4" captionStartId="54.[134,222,490,516]" captionTargetPageId="54" captionText="TABLE 4. Summary of the most important diagnostic characters of species in the Mantidactylus subgenus Brygoomantis. SVL is given only based on reliably sexed specimens, summarised and rounded to the nearest mm from original measurements in Tables 511. Dorsolateral ridges are always weakly expressed in Brygoomantis, but clearly visible in some species while completely absent in others. Dorsal skin rugosity is estimated from pictures in life and can appear different in preservative: 0, smooth; 1, finely granular; 2, coarsely granular / tubercular. Webbing at toe 5 is given as 0, half or less than half of one phalanx free of web, not counting terminal phalanx (corresponding to numbers 00.5 in webbing formula); 1, about one phalanx free of web (0.751.25); 2, more than one phalanx free of web (1.52). (Yes) in parentheses indicates that the respective character state is found in some individuals of the respective species, and often only weakly expressed, while it is missing in others. Lat.met., state of the lateral metatarsalia, either separated (sep.) by webbing or connected (conn.) by tissue. NM, no measurements available. *For the advertisement calls of M. katae sp. nov. and M. fergusoni sp. nov., homology of call components remains unclear. Here, each unit is interpreted as singlepulse call (= single-pulse note) arranged in series (under this definition the provided pulse rate reflects call rate), but it would also be possible to consider the entire vocalization as single-note multi-pulsed call with very long duration, containing very widely spaced pulses. Values for pulse rate (pulses/second) are rounded. We here evaluate the presence versus absence of regular call series (last column), where regular series are considered as such, if calls are emitted at really regular intervals in series of similar duration containing a more or less defined number of calls. # In M. ambohimitombi, several unsexed individuals of the type series, probably females, reach sizes of up to 65 mm, being the largest of all Brygoomantis known. †One female of M. biporus of 19.3 mm SVL is probably a not fully mature specimen; other females of the species measure 3136 mm SVL." httpUri="http://table.plazi.org/id/83F3168BFFEAFFF84CBF4AEE4C717C8F" pageId="72" pageNumber="185" tableUuid="83F3168BFFEAFFF84CBF4AEE4C717C8F">Table 4</tableCitation>
);
<taxonomicName box="[1166,1280,1964,1989]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="curtus">
<emphasis box="[1166,1280,1964,1989]" italics="true" pageId="72" pageNumber="185">M. curtus</emphasis>
</taxonomicName>
usually has a smoother skin and a somewhat shorter snout;
<taxonomicName box="[1319,1451,1995,2021]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="72" pageNumber="185" phylum="Chordata" rank="species" species="bourgati">
<emphasis box="[1319,1451,1995,2021]" italics="true" pageId="72" pageNumber="185">M. bourgati</emphasis>
</taxonomicName>
is morphologically very similar but appears to occur only on the Andringitra Massif. Compared to other subspecies of
<taxonomicName authorityName="Boulenger" authorityYear="1919" box="[170,388,171,197]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="73" pageNumber="186" phylum="Chordata" rank="species" species="ambohimitombi">
<emphasis box="[170,388,171,197]" italics="true" pageId="73" pageNumber="186">M. ambohimitombi</emphasis>
</taxonomicName>
, the new subspecies differs from
<emphasis box="[136,282,203,229]" italics="true" pageId="73" pageNumber="186">M. a. marefo</emphasis>
by its distinct dorsal pattern, lack of bluish colour around the eye, and more pointed snout; and from
<emphasis box="[136,379,267,293]" italics="true" pageId="73" pageNumber="186">
M. a.
<taxonomicName authorityName="Boulenger" authorityYear="1919" box="[203,379,267,293]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="73" pageNumber="186" phylum="Chordata" rank="species" species="ambohimitombi">ambohimitombi</taxonomicName>
</emphasis>
by the usually more distinct dorsal pattern. For detailed distinction from other new species and subspecies described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of
<emphasis box="[282,426,395,421]" italics="true" pageId="73" pageNumber="186">M. a. miloko</emphasis>
in pairwise comparisons to all other
<taxonomicName box="[200,357,428,453]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="73" pageNumber="309" phylum="Chordata" rank="subGenus" subGenus="Brygoomantis">
<emphasis box="[200,357,428,453]" italics="true" pageId="73" pageNumber="186">Brygoomantis</emphasis>
</taxonomicName>
species and subspecies is provided as Supplementary appendix.
</paragraph>
</subSubSection>
<subSubSection pageId="73" pageNumber="186" type="description">
<paragraph blockId="73.[136,778,171,2085]" pageId="73" pageNumber="186">
<emphasis box="[184,507,491,517]" italics="true" pageId="73" pageNumber="186">
Description of the
<typeStatus box="[411,507,491,517]" pageId="73" pageNumber="186">holotype</typeStatus>
</emphasis>
.—Adult male in good state of preservation (
<figureCitation box="[414,495,523,549]" captionStart="FIGURE 9" captionStartId="52.[136,235,1772,1796]" captionTargetBox="[155,1429,206,1743]" captionTargetId="figure-35@52.[151,1436,198,1753]" captionTargetPageId="52" captionText="FIGURE 9. Preserved holotypes of newly named species and subspecies in the M. curtus clade. Scale bars equal 5 mm." figureDoi="http://doi.org/10.5281/zenodo.7441041" httpUri="https://zenodo.org/record/7441041/files/figure.png" pageId="73" pageNumber="186">Fig. 9</figureCitation>
). Tissue sample taken ventrally from right thigh. Femoral gland partly detached to examine their structure internally. Body rather slender.
</paragraph>
<paragraph blockId="73.[136,778,171,2085]" pageId="73" pageNumber="186">Head slightly wider than body. Snout rounded in dorsal and lateral view. Nostrils directed dorsolaterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis almost not recognisable, slightly concave. Loreal region slightly concave. Tympanum distinct, elliptical, wider than high, its diameter 78% of eye diameter. Supratympanic fold distinct, beginning straight, with a distinct, angular 90° bend at the posterior edge of tympanum towards insertion of forelimb. Tongue ovoid, distinctly posteriorly bifid. Maxillary teeth present. Vomerine teeth distinct in rounded aggregations, positioned posterolateral to choanae. Choanae rounded.</paragraph>
<paragraph blockId="73.[136,778,171,2085]" pageId="73" pageNumber="186">Subarticular tubercles single. Outer metacarpal tubercle present, inner metacarpal tubercle present. Fingers without webbing. Relative length of fingers: I=II&lt;IV&lt;III. Finger discs minimally enlarged. Nuptial pads absent. Foot longer than tibia (113%). Lateral metatarsalia separated.</paragraph>
<paragraph blockId="73.[136,778,171,2085]" pageId="73" pageNumber="186">Inner metatarsal tubercle present. Outer metatarsal tubercle small but recognisable. Webbing formula: 1(0.25), 2i(1), 2e(0.5), 3i(1.5), 3e(1), 4i(2), 4e(2), 5(0.75).</paragraph>
<paragraph blockId="73.[136,778,171,2085]" pageId="73" pageNumber="186">Relative length of toes: I&lt;II&lt;V=III&lt;IV. Skin on the upper surface smooth, with some granules on flanks, and some longitudinal tubercles forming interrupted and weakly expressed dorsolateral folds.Ventral side smooth. Femoral glands distinct but relatively small, with a distal ulcerous macrogland consisting of at least eight large granules and an external central depression, and a weakly expressed proximal granular gland field visible in internal view.</paragraph>
<paragraph blockId="73.[136,778,171,2085]" pageId="73" pageNumber="186">Colour in preservative: light brown dorsally with large and contrasted dark brown patches all over the dorsal surface, distinct dark crossbands on limbs, and a light frenal stripe with some dark markings on the upper lip.</paragraph>
<paragraph blockId="73.[136,778,171,2085]" pageId="73" pageNumber="186">
Ventrally uniformly light grey with alternating dark-light pattern ventrally on the lower lip. Colour in life similar to that in preservative, but more contrasted (
<figureCitation box="[602,686,1707,1733]" captionStart="FIGURE 16" captionStartId="74.[136,235,1645,1669]" captionTargetBox="[151,1436,185,1625]" captionTargetId="figure-35@74.[151,1436,185,1625]" captionTargetPageId="74" captionText="FIGURE 16. Mantidactylus ambohimitombi miloko ssp. nov. in life, in dorsolateral and ventral view. (a,b,c) Adult male (holotype ZSM 219/2005 = FGZC 2143; note distinct and large femoral glands), from Ambohitantely Special Reserve." figureDoi="http://doi.org/10.5281/zenodo.7441055" httpUri="https://zenodo.org/record/7441055/files/figure.png" pageId="73" pageNumber="186">Fig. 16</figureCitation>
).
</paragraph>
<paragraph blockId="73.[136,778,171,2085]" pageId="73" pageNumber="186">
<emphasis box="[184,296,1740,1765]" italics="true" pageId="73" pageNumber="186">Variation.</emphasis>
—Variation in measurements is given in
<tableCitation box="[136,220,1771,1798]" captionStart="TABLE 5" captionStartId="59.[134,222,467,493]" captionTargetPageId="59" captionText="TABLE 5. Morphometric measurements (all in mm) of voucher specimens of the Mantidactylus curtus clade. Type status is given in square brackets after voucher number: HT, holotype; PT, paratype; LT, lectotype; PLT, paralectotype. An asterisk (*) marks lectotypes designated in the current paper. For abbreviations of measurements, see Materials and Methods. All specimens are probable adults, but sex is difficult to ascertain externally in many species of this clade due to weak expression of femoral glands in male specimens, and is therefore only given for some specimens where gonad examination was possible. NM, not measured. # Measurements taken by Gavetti and Andreone (1993)" httpUri="http://table.plazi.org/id/83F3168BFFE7FFF54CBF4AD74A627D52" pageId="73" pageNumber="186" tableUuid="83F3168BFFE7FFF54CBF4AD74A627D52">Table 5</tableCitation>
. Too few specimens have been sexed to assess the degree of sexual size dimorphism. Femoral glands distinct and large (but not differing in colour from surrounding ventral skin of thigh) in the male
<typeStatus box="[529,627,1867,1893]" pageId="73" pageNumber="186">holotype</typeStatus>
(
<figureCitation box="[645,731,1867,1893]" captionStart="FIGURE 16" captionStartId="74.[136,235,1645,1669]" captionTargetBox="[151,1436,185,1625]" captionTargetId="figure-35@74.[151,1436,185,1625]" captionTargetPageId="74" captionText="FIGURE 16. Mantidactylus ambohimitombi miloko ssp. nov. in life, in dorsolateral and ventral view. (a,b,c) Adult male (holotype ZSM 219/2005 = FGZC 2143; note distinct and large femoral glands), from Ambohitantely Special Reserve." figureDoi="http://doi.org/10.5281/zenodo.7441055" httpUri="https://zenodo.org/record/7441055/files/figure.png" pageId="73" pageNumber="186">Fig. 16</figureCitation>
). In contrast, specimens from the ZMA collection are difficult to sex externally as femoral glands are often indistinct, as seems to be typical for several taxa in the
<taxonomicName box="[599,705,1964,1989]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="73" pageNumber="186" phylum="Chordata" rank="species" species="curtus">
<emphasis box="[599,705,1964,1989]" italics="true" pageId="73" pageNumber="186">M. curtus</emphasis>
</taxonomicName>
clade, possibly due to seasonal effects.
</paragraph>
<paragraph blockId="73.[136,778,171,2085]" lastBlockId="73.[810,1452,171,933]" pageId="73" pageNumber="186">
<emphasis box="[184,390,2027,2053]" italics="true" pageId="73" pageNumber="186">Natural history.—</emphasis>
Specimens were collected around clean highland streams running in open areas between forest fragments. Specimens at Angavokely and Ankazomivady that appear to belong to this taxon based on mitochondrial DNA were found on wet rocks along slow-moving parts of streams.
</paragraph>
<paragraph blockId="73.[810,1452,171,933]" pageId="73" pageNumber="186">
<emphasis box="[858,916,299,325]" italics="true" pageId="73" pageNumber="186">Calls</emphasis>
.—The call of this subspecies has not been recorded.
</paragraph>
<paragraph blockId="73.[810,1452,171,933]" pageId="73" pageNumber="186">
<emphasis box="[858,989,363,389]" italics="true" pageId="73" pageNumber="186">Tadpoles.—</emphasis>
Probably reported from Angavokely by Blommers-Schl̂sser (1979)..
</paragraph>
</subSubSection>
<subSubSection pageId="73" pageNumber="186" type="distribution">
<paragraph blockId="73.[810,1452,171,933]" pageId="73" pageNumber="186">
<emphasis box="[858,1026,427,453]" italics="true" pageId="73" pageNumber="186">Distribution.—</emphasis>
Endemic to the central highlands of
<collectingCountry box="[843,980,459,485]" name="Madagascar" pageId="73" pageNumber="186">Madagascar</collectingCountry>
, north of the distribution of the nominal subspecies (
<figureCitation box="[948,1020,491,517]" captionStart="FIGURE 7" captionStartId="49.[136,235,1996,2020]" captionTargetBox="[252,1336,200,1976]" captionTargetId="figure-20@49.[252,1336,200,1976]" captionTargetPageId="49" captionText="FIGURE 7. Mantidactylus subgenus Brygoomantis species distribution based on verified records, divided by clade (Mantidactylus stelliger sp. nov. is presented alongside the inaudax clade for practicality). Colours for species correspond to those in Figs 2 and 4. Inset map shows the geographic regions of Madagascar identified by Boumans et al. (2007), referred to throughout the text." figureDoi="http://doi.org/10.5281/zenodo.7441037" httpUri="https://zenodo.org/record/7441037/files/figure.png" pageId="73" pageNumber="186">Fig. 7</figureCitation>
). This subspecies is currently known from Ambohitantely, and probably also from Angavokely, and Ankazomivady. A mitochondrial haplotype corresponding to this subspecies has also been detected at Ankaratra, but due to the limited information on the sampling event and the absence of specimens (only tissue sampels were collected) this record (which might also represent mitochondrial introgression) requires confirmation. Elevation range:
<quantity box="[1160,1315,747,774]" metricMagnitude="3" metricUnit="m" metricValue="1.6275" metricValueMax="1.735" metricValueMin="1.52" pageId="73" pageNumber="186" unit="m" value="1627.5" valueMax="1735.0" valueMin="1520.0">15201735 m</quantity>
a.s.l.
</paragraph>
</subSubSection>
<subSubSection pageId="73" pageNumber="186" type="etymology">
<paragraph blockId="73.[810,1452,171,933]" pageId="73" pageNumber="186">
<emphasis box="[858,978,779,805]" italics="true" pageId="73" pageNumber="186">Etymology</emphasis>
.—The subspecies name is derived from Malagasy word miloko, meaning painted, referring to the rather distinct dorsal pattern of well-delimited dark blotches characterizing this subspecies. The subspecies name is used as a noun in apposition.
</paragraph>
</subSubSection>
</treatment>
</document>