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<document ID-DOI="http://doi.org/10.5281/zenodo.5689491" ID-GBIF-Dataset="c3128d24-5293-4125-bc85-0678753fc483" ID-GBIF-Taxon="154520572" ID-Zenodo-Dep="5689491" LSID="urn:lsid:zoobank.org:act:1A79607B-E9D4-4214-BF47-9BC0B4320E5F" checkinTime="1553770082828" checkinUser="plazi" docAuthor="Scherz, Mark D., Hutter, Carl R., Rakotoarison, Andolalao, Riemann, Jana C., Rödel, Mark-Oliver, Ndriantsoa, Serge H., Glos, Julian, Roberts, Sam Hyde, Crottini, Angelica, Vences, Miguel &amp; Glaw, Frank" docDate="2019" docId="03F887F6FF880A68FD8B07A2D9046A60" docLanguage="en" docName="journal.pone.0213314.pdf" docOrigin="PLoS ONE 213314" docStyle="DocumentStyle{}" docTitle="Rhombophryne proportionalis Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw 2019, sp. nov." docType="treatment" docUuid="1A79607B-E9D4-4214-BF47-9BC0B4320E5F" docUuidSource="ZooBank" docVersion="8" lastPageId="32" lastPageNumber="33" masterDocId="FFC1FF8EFF930A48FF9E0658DC6D6955" masterDocTitle="Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera" masterLastPageNumber="45" masterPageNumber="1" pageId="27" pageNumber="28" updateTime="1636786376000" updateUser="ExternalLinkService">
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<mods:titleInfo>
<mods:title>Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera</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>Hutter, Carl R.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Rakotoarison, Andolalao</mods:namePart>
</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>Ndriantsoa, Serge H.</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Glos, Julian</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Roberts, Sam Hyde</mods:namePart>
</mods:name>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Crottini, Angelica</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>
<mods:name type="personal">
<mods:role>
<mods:roleTerm>Author</mods:roleTerm>
</mods:role>
<mods:namePart>Glaw, Frank</mods:namePart>
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<mods:title>PLoS ONE</mods:title>
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<mods:part>
<mods:date>2019</mods:date>
<mods:detail type="pubDate">
<mods:number>2019-03-27</mods:number>
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<mods:detail type="volume">
<mods:number>213314</mods:number>
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<mods:start>1</mods:start>
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<mods:classification>journal article</mods:classification>
<mods:identifier type="DOI">10.1371/journal.pone.0213314</mods:identifier>
<mods:identifier type="GBIF-Dataset">c3128d24-5293-4125-bc85-0678753fc483</mods:identifier>
<mods:identifier type="Zenodo-Dep">2614083</mods:identifier>
<mods:identifier type="ZooBank">91E597C8-7A80-46F9-B0E8-61F2524400F7</mods:identifier>
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<treatment ID-DOI="http://doi.org/10.5281/zenodo.5689491" ID-GBIF-Taxon="154520572" ID-Zenodo-Dep="5689491" LSID="urn:lsid:zoobank.org:act:1A79607B-E9D4-4214-BF47-9BC0B4320E5F" httpUri="http://treatment.plazi.org/id/03F887F6FF880A68FD8B07A2D9046A60" lastPageId="32" lastPageNumber="33" pageId="27" pageNumber="28">
<subSubSection pageId="27" pageNumber="28" type="nomenclature">
<paragraph blockId="27.[533,1038,506,535]" box="[533,1038,506,535]" pageId="27" pageNumber="28">
<heading bold="true" box="[533,1038,506,535]" fontSize="12" level="2" pageId="27" pageNumber="28" reason="0">
<emphasis bold="true" box="[533,1038,506,535]" pageId="27" pageNumber="28">
<taxonomicName authority="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw, 2019" authorityName="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw" authorityYear="2019" box="[533,929,506,535]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="28" phylum="Chordata" rank="species" species="proportionalis" status="sp. nov.">
<emphasis bold="true" box="[533,929,506,535]" italics="true" pageId="27" pageNumber="28">Rhombophryne proportionalis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[936,1038,507,535]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
<paragraph blockId="27.[533,1536,555,1897]" box="[533,1276,555,579]" pageId="27" pageNumber="28">
<uri box="[533,1276,555,579]" pageId="27" pageNumber="28">
urn:lsid:zoobank.org:act:
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1
<accessionNumber box="[807,894,555,579]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/A79607" pageId="27" pageNumber="28">A79607</accessionNumber>
B-E9D4-4214-BF47-9BC0B4320E5F
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</uri>
</paragraph>
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<subSubSection box="[565,997,590,614]" pageId="27" pageNumber="28" type="description">
<paragraph blockId="27.[533,1536,555,1897]" box="[565,997,590,614]" pageId="27" pageNumber="28">
(
<figureCitation box="[572,636,590,614]" captionStart="Fig 1" captionStartId="2.[192,220,1354,1373]" captionTargetBox="[192,1536,208,1332]" captionTargetId="figure@2.[192,1536,208,1332]" captionTargetPageId="2" captionText="Fig 1. Stylised representation of the latest phylogeny of the microhylid subfamily Cophylinae. Based on the phylogenetic tree of Tu et al. [20] (reconstructed from a concatenated multi-gene DNA sequence data set) with numerous nomenclatural updates. See that paper for support values and phylogenetic context. The sister species of Rhombophryne proportionalis sp. nov. is an undescribed candidate species (R. sp. Ca7) that is not included in this phylogeny; its taxonomy will be treated elsewhere. Taxa highlighted in red are those described herein. Inset photos show a selection of the new species described herein, and representatives of the other small-bodied genera, Stumpffia and Anilany. Inset photos are not to scale. https://doi.org/10.1371/journal.pone.0213314.g001" httpUri="https://zenodo.org/record/2614085/files/figure.png" pageId="27" pageNumber="28">Figs 1</figureCitation>
,
<figureCitation box="[648,661,590,614]" captionStart="Fig 2" captionStartId="7.[192,220,1818,1837]" captionTargetBox="[192,1536,851,1796]" captionTargetId="figure@7.[192,1536,851,1796]" captionTargetPageId="7" captionText="Fig 2. Holotypes of the new species described in this paper and their hands. Whole specimens in dorsal (left) and ventral (right) view. Hand images not to scale. https://doi.org/10.1371/journal.pone.0213314.g002" httpUri="https://zenodo.org/record/2614087/files/figure.png" pageId="27" pageNumber="28">2</figureCitation>
,
<figureCitation box="[673,686,590,614]" captionStart="Fig 3" captionStartId="8.[461,489,1766,1785]" captionTargetBox="[461,1535,208,1744]" captionTargetId="figure@8.[461,1535,208,1744]" captionTargetPageId="8" captionText="Fig 3. Currently known localities of the new taxa described in this paper. The base map is USGS SRTM 1-Arc second digital elevation model. https://doi.org/10.1371/journal.pone.0213314.g003" httpUri="https://zenodo.org/record/2614089/files/figure.png" pageId="27" pageNumber="28">3</figureCitation>
,
<figureCitation box="[697,710,590,614]" captionStart="Fig 5" captionStartId="10.[200,228,1285,1304]" captionTargetBox="[200,1536,208,1263]" captionTargetId="figure@10.[200,1536,208,1263]" captionTargetPageId="10" captionText="Fig 5. One-second spectrograms and oscillograms of the calls of the new species described here. Insets represent the respective species but not the calling specimens. (a) Mini mum gen. et sp. nov., paratype ZMB 81993 from Manombo, (b) Mini scule gen. et sp. nov., ZSM 265/2018 from Sainte Luce, (c) Rhombophryne proportionalis sp. nov., part of a call (note series) of a specimen from Bepia campsite, Tsaratanana (not collected), (d) Anodonthyla eximia sp. nov., specimen not collected, from Maharira (Ranomafana). https://doi.org/10.1371/journal.pone.0213314.g005" httpUri="https://zenodo.org/record/2614093/files/figure.png" pageId="27" pageNumber="28">5</figureCitation>
,
<figureCitation box="[722,747,590,614]" captionStart="Fig 10" captionStartId="28.[115,143,1722,1741]" captionTargetBox="[115,1536,208,1700]" captionTargetId="figure@28.[115,1536,208,1700]" captionTargetPageId="28" captionText="Fig 10. Rhombophryne proportionalis sp. nov., holotype ZSM 1826/2010, in life. (a) dorsolateral, (b) dorsal, and (c) ventral view. https://doi.org/10.1371/journal.pone.0213314.g010" httpUri="https://zenodo.org/record/2614103/files/figure.png" pageId="27" pageNumber="28">10</figureCitation>
and
<figureCitation box="[799,824,590,614]" captionStart="Fig 11" captionStartId="29.[192,220,1159,1178]" captionTargetBox="[192,1536,208,1137]" captionTargetId="figure@29.[192,1536,208,1137]" captionTargetPageId="29" captionText="Fig 11. Osteology of Rhombophryne proportionalis sp. nov. holotype (ZSM 1826/2010). (a, b) Whole skeleton in (a) dorsal and (b) ventral view. (c-f) Skull in (c) lateral, (d) ventral, (e) anterior, and (f) dorsal view. (g) Foot in ventral view. (h) Hand in ventral view. For abbreviations, see Fig 6. https://doi.org/10.1371/journal.pone.0213314.g011" httpUri="https://zenodo.org/record/2614105/files/figure.png" pageId="27" pageNumber="28">11</figureCitation>
,
<tableCitation box="[836,924,590,614]" captionStart="Table 1" captionStartId="12.[96,146,518,537]" captionText="Table 1. Morphological measurements (in mm) of new species of miniaturised cophyline microhylids. Measurement abbreviations are listed in the Materials and Methods. A = Adult, M = Male, F = Female (lowercase indicates probable subadult), p = paratype, h = holotype (holotypes are also bolded), na = not available." pageId="27" pageNumber="28">Tables 1</tableCitation>
and
<tableCitation box="[976,989,590,614]" captionStart="Table 2" captionStartId="13.[533,583,194,213]" captionTargetBox="[533,1512,335,774]" captionTargetId="graphics@13.[533,1536,323,778]" captionTargetPageId="13" captionText="Table 2. Bioacoustic parameters of new species of miniaturised cophyline microhylids. Data on S. miery from Rakotoarison et al. [14], provided for comparison with the sympatric A. eximia. Values are presented as mean ± standard deviation, with range in brackets. na = not applicable. In all species except R. proportionalis calls consist of a single note according to the definition herein, and in these species call duration is therefore synonymous with note duration." httpUri="http://table.plazi.org/id/DF2E6668FF9E0A45FD8B069ADEFD686E" pageId="27" pageNumber="28" tableUuid="DF2E6668FF9E0A45FD8B069ADEFD686E">2</tableCitation>
)
</paragraph>
</subSubSection>
<subSubSection pageId="27" pageNumber="28" type="discussion">
<paragraph blockId="27.[533,1536,555,1897]" pageId="27" pageNumber="28">
<emphasis bold="true" box="[565,659,625,649]" pageId="27" pageNumber="28">Remark.</emphasis>
This species was previously referred to as
<taxonomicName box="[1106,1209,625,649]" class="Amphibia" family="Microhylidae" genus="Stumpffia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis box="[1106,1209,625,649]" italics="true" pageId="27" pageNumber="28">Stumpffia</emphasis>
</taxonomicName>
sp. Ca34 [
<bibRefCitation author="Scherz MD &amp; Vences M &amp; Rakotoarison A &amp; Andreone F &amp; Kohler J &amp; Glaw F" box="[1319,1344,625,649]" journalOrPublisher="Molecular Phylogenetics and Evolution." pageId="27" pageNumber="28" pagination="372 - 81" part="100" refId="ref23908" refString="15. Scherz MD, Vences M, Rakotoarison A, Andreone F, Kohler J, Glaw F, et al. Reconciling molecular phylogeny, morphological divergence and classification of Madagascan narrow-mouthed frogs (Amphibia: Microhylidae). Molecular Phylogenetics and Evolution. 2016; 100: 372 - 81. https: // doi. org / 10.1016 / j. ympev. 20 16.04. 0 19 PMID: 270856 71" title="et al. Reconciling molecular phylogeny, morphological divergence and classification of Madagascan narrow-mouthed frogs (Amphibia: Microhylidae)" type="journal article" year="2016">15</bibRefCitation>
,
<bibRefCitation author="Perl RGB &amp; Nagy ZT &amp; Sonet G &amp; Glaw F &amp; Wollenberg KC &amp; Vences M." box="[1356,1381,625,649]" journalOrPublisher="Amphibia-Reptilia." pageId="27" pageNumber="28" pagination="197 - 206" part="35" refId="ref25709" refString="46. Perl RGB, Nagy ZT, Sonet G, Glaw F, Wollenberg KC, Vences M. DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia. 2014; 35: 197 - 206. https: // doi. org / 10.1163 / 15685381 - 0 0 0 0 2942" title="DNA barcoding Madagascar's amphibian fauna" type="journal article" year="2014">46</bibRefCitation>
] and as
<taxonomicName box="[533,636,660,684]" class="Amphibia" family="Microhylidae" genus="Stumpffia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis box="[533,636,660,684]" italics="true" pageId="27" pageNumber="28">Stumpffia</emphasis>
</taxonomicName>
sp. 39 MV-2012 (
<accessionNumber box="[824,940,659,683]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/KC351481" pageId="27" pageNumber="28">
<collectionCode box="[824,861,659,683]" pageId="27" pageNumber="28">KC</collectionCode>
351481
</accessionNumber>
) [
<bibRefCitation author="Tu N &amp; Yang M &amp; Liang D &amp; Zhang P." box="[961,986,659,683]" journalOrPublisher="Molecular Phylogenetics and Evolution." pageId="27" pageNumber="28" pagination="85 - 91" part="126" refId="ref24267" refString="20. Tu N, Yang M, Liang D, Zhang P. A large-scale phylogeny of Microhylidae inferred from a combined dataset of 121 genes and 427 taxa. Molecular Phylogenetics and Evolution. 2018; 126: 85 - 91. https: // doi. org / 10.1016 / j. ympev. 201 8.03. 0 36 PMID: 2964958 1" title="A large-scale phylogeny of Microhylidae inferred from a combined dataset of 121 genes and 427 taxa" type="journal article" year="2018">20</bibRefCitation>
].
</paragraph>
</subSubSection>
<subSubSection pageId="27" pageNumber="28" type="materials_examined">
<paragraph blockId="27.[533,1536,555,1897]" pageId="27" pageNumber="28">
<materialsCitation ID-GBIF-Occurrence="2090010997" collectingDate="2010-06-16" collectionCode="KU" collectorName="M. Vences &amp; D. R. Vieites &amp; R. D. Randrianiaina &amp; S. Rasamison &amp; E. Rajeriarison." country="Madagascar" county="Diana Region" elevation="2294" location="Bepia Campsite" municipality="Tsaratanana" pageId="27" pageNumber="28" specimenCode="ZSM 1826, ZCMV 12404" specimenCount="1" stateProvince="Antsiranana" typeStatus="holotype">
<emphasis bold="true" box="[565,675,694,718]" pageId="27" pageNumber="28">
<typeStatus box="[565,670,694,718]" pageId="27" pageNumber="28">Holotype</typeStatus>
.
</emphasis>
<specimenCode box="[681,790,694,718]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="27" pageNumber="28" type="Museum">ZSM 1826</specimenCode>
/2010 (
<specimenCode box="[863,1012,694,718]" collectionCode="ZCMV" pageId="27" pageNumber="28">ZCMV 12404</specimenCode>
, GenBank accession number
<accessionNumber isEnumeration="true" pageId="27" pageNumber="28">
KC351380 and
<collectionCode box="[533,570,729,753]" country="USA" name="Biodiversity Institute, University of Kansas" pageId="27" pageNumber="28" type="University or college">KU</collectionCode>
937808
</accessionNumber>
for 5 and 3 fragments of the
<emphasis box="[964,1069,729,753]" italics="true" pageId="27" pageNumber="28">16S rRNA</emphasis>
gene, respectively, and
<accessionNumber box="[1319,1427,729,753]" httpUri="http://www.ncbi.nlm.nih.gov/nucleotide/KF611640" pageId="27" pageNumber="28">KF611640</accessionNumber>
for the
<emphasis box="[533,581,764,788]" italics="true" pageId="27" pageNumber="28">cox1</emphasis>
gene), an adult male specimen (seen calling, not recorded) collected at
<location LSID="urn:lsid:plazi:treatment:03F887F6FF880A68FD8B07A2D9046A60:8E8E603BFF880A53FAA904A3D9B06A46" box="[1335,1501,763,787]" country="Madagascar" county="Diana Region" municipality="Tsaratanana" name="Bepia Campsite" pageId="27" pageNumber="28" stateProvince="Antsiranana">Bepia Campsite</location>
on the
<collectingMunicipality box="[573,704,798,822]" pageId="27" pageNumber="28">Tsaratanana</collectingMunicipality>
massif (Camp 3; 14.1182˚S, 48.9782˚E,
<elevation box="[1124,1254,798,822]" metricMagnitude="3" metricUnit="m" metricValue="2.294" pageId="27" pageNumber="28" unit="m" value="2294.0">
<quantity box="[1124,1202,798,822]" metricMagnitude="3" metricUnit="m" metricValue="2.294" pageId="27" pageNumber="28" unit="m" value="2294.0">2294 m</quantity>
a.s.l.
</elevation>
),
<collectingCounty box="[1275,1423,798,822]" pageId="27" pageNumber="28">Diana Region</collectingCounty>
, former
<collectingRegion box="[533,767,833,857]" country="Madagascar" name="Antsiranana" pageId="27" pageNumber="28">Antsiranana province</collectingRegion>
, northern
<collectingCountry box="[878,1005,833,857]" name="Madagascar" pageId="27" pageNumber="28">Madagascar</collectingCountry>
on
<collectingDate box="[1045,1182,833,857]" pageId="27" pageNumber="28" value="2010-06-16">
<date box="[1045,1182,833,857]" pageId="27" pageNumber="28" value="2010-06-16">16 June 2010</date>
</collectingDate>
by
<collectorName box="[1219,1334,833,857]" pageId="27" pageNumber="28">M. Vences</collectorName>
,
<collectorName box="[1343,1473,833,857]" pageId="27" pageNumber="28">D.R. Vieites</collectorName>
,
<collectorName pageId="27" pageNumber="28">R.D. Randrianiaina</collectorName>
,
<collectorName box="[697,840,867,891]" pageId="27" pageNumber="28">S. Rasamison</collectorName>
, and
<collectorName box="[896,1060,867,891]" pageId="27" pageNumber="28">E. Rajeriarison.</collectorName>
</materialsCitation>
</paragraph>
<paragraph blockId="27.[533,1536,555,1897]" pageId="27" pageNumber="28">
<materialsCitation ID-GBIF-Occurrence="2090013495" collectingDate="2010-06-16" collectionCode="MK" collectorName="M. Vences &amp; D. R. Vieites &amp; R. D. Randrianiaina &amp; S. Rasamison &amp; E. Rajeriarison." country="Madagascar" county="Diana Region" elevation="2294" location="Bepia Campsite" municipality="Tsaratanana" pageId="27" pageNumber="28" specimenCode="ZSM 1840, ZCMV 12405" specimenCount="1" stateProvince="Antsiranana" typeStatus="paratype">
<emphasis bold="true" box="[565,681,902,926]" pageId="27" pageNumber="28">
<typeStatus box="[565,676,902,926]" pageId="27" pageNumber="28">Paratypes</typeStatus>
.
</emphasis>
<specimenCode box="[687,796,902,926]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="27" pageNumber="28" type="Museum">ZSM 1840</specimenCode>
/2010 (
<specimenCode box="[869,1018,902,926]" collectionCode="ZCMV" pageId="27" pageNumber="28">ZCMV 12405</specimenCode>
, GenBank accession number
<accessionNumber isEnumeration="true" pageId="27" pageNumber="28">
KC351481 and
<collectionCode box="[533,574,937,961]" country="Kenya" name="National Museum of Kenya" pageId="27" pageNumber="28">MK</collectionCode>
459317
</accessionNumber>
for 5 and 3 fragments of the
<emphasis box="[968,1073,937,961]" italics="true" pageId="27" pageNumber="28">16S rRNA</emphasis>
gene, respectively), adult male specimen with the same collection data as the holotype; and
</materialsCitation>
<materialsCitation ID-GBIF-Occurrence="2090007783" collectingDate="2010-06-16" collectionCode="ZSM, DRV" country="Madagascar" elevation="2503" location="Andranomadio Campsite in Tsaratanana" pageId="27" pageNumber="28" specimenCode="ZSM 636, DRV 6224" specimenCount="1" stateProvince="Antsiranana" typeStatus="paratype">
<specimenCode box="[1064,1160,971,995]" collectionCode="ZSM" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="27" pageNumber="28" type="Museum">ZSM 636</specimenCode>
/2014 (
<specimenCode box="[1233,1350,971,995]" collectionCode="DRV" pageId="27" pageNumber="28">DRV 6224</specimenCode>
), an adult presumed male specimen collected at
<location LSID="urn:lsid:plazi:treatment:03F887F6FF880A68FD8B07A2D9046A60:8E8E603BFF880A53FC1D05B6D95A6D53" box="[899,1335,1006,1030]" country="Madagascar" name="Andranomadio Campsite in Tsaratanana" pageId="27" pageNumber="28" stateProvince="Antsiranana">Andranomadio Campsite in Tsaratanana</location>
(Camp 4; 14.0801˚S, 48.9854˚E,
<elevation box="[770,900,1041,1065]" metricMagnitude="3" metricUnit="m" metricValue="2.503" pageId="27" pageNumber="28" unit="m" value="2503.0">
<quantity box="[770,849,1041,1065]" metricMagnitude="3" metricUnit="m" metricValue="2.503" pageId="27" pageNumber="28" unit="m" value="2503.0">2503 m</quantity>
a.s.l.
</elevation>
) on
<collectingDate box="[949,1086,1041,1065]" pageId="27" pageNumber="28" value="2010-06-16">
<date box="[949,1086,1041,1065]" pageId="27" pageNumber="28" value="2010-06-16">16 June 2010</date>
</collectingDate>
by the same collectors
</materialsCitation>
.
</paragraph>
</subSubSection>
<subSubSection lastPageId="28" lastPageNumber="29" pageId="27" pageNumber="28" type="diagnosis">
<paragraph blockId="27.[533,1536,555,1897]" pageId="27" pageNumber="28">
<emphasis bold="true" box="[565,682,1075,1099]" pageId="27" pageNumber="28">Diagnosis.</emphasis>
A diminutive frog assigned to the genus
<taxonomicName box="[1115,1275,1076,1100]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis box="[1115,1275,1076,1100]" italics="true" pageId="27" pageNumber="28">Rhombophryne</emphasis>
</taxonomicName>
on the basis of absence of clavicles, presence of vomerine, maxillary, and premaxillary teeth, short and broad skull, and genetic affinities. It is separated by uncorrected p-distances of 7.012.9% in the analysed 3 fragment of the
<emphasis box="[703,808,1180,1204]" italics="true" pageId="27" pageNumber="28">16S rRNA</emphasis>
gene from other members of the genus
<taxonomicName box="[1230,1390,1180,1204]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis box="[1230,1390,1180,1204]" italics="true" pageId="27" pageNumber="28">Rhombophryne</emphasis>
</taxonomicName>
.
</paragraph>
<paragraph blockId="27.[533,1536,555,1897]" pageId="27" pageNumber="28">
<taxonomicName authority="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw, 2019" authorityName="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw" authorityYear="2019" box="[565,878,1214,1238]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="28" phylum="Chordata" rank="genus" species="proportionalis" status="sp. nov.">
<emphasis box="[565,878,1214,1238]" italics="true" pageId="27" pageNumber="28">Rhombophryne proportionalis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[884,966,1214,1238]" pageId="27" pageNumber="28" rank="species">sp. nov.</taxonomicNameLabel>
is characterised by the unique combination of the following characters (n =
<specimenCount box="[777,847,1249,1273]" pageId="27" pageNumber="28" type="male">3 male</specimenCount>
specimens): (1) male SVL 11.0
<quantity box="[1188,1283,1249,1273]" metricMagnitude="-2" metricUnit="m" metricValue="1.23" pageId="27" pageNumber="28" unit="mm" value="12.3">12.3 mm</quantity>
; (2) ED/HL 0.400.48; (3) HW/SVL 0.330.37; (4)
<collectionCode box="[827,891,1283,1307]" pageId="27" pageNumber="28">FARL</collectionCode>
/SVL 0.330.35; (5) TIBL/SVL 0.340.36; (6) HIL/SVL 0.21 1.32; (7) finger 1 reduced, 2 and 4 short; (8) toe 1 highly reduced, 2 somewhat reduced; (9) maxillary and premaxillary teeth present; (10) vomerine teeth present; (11) lateral colour bor- der absent; (12) black inguinal spots sometimes present; (13) postchoanal vomer present, spatulate, medially fused to parasphenoid; (14) nasal broad and not laterally displaced; (15) quadratojugal-maxilla contact strong and broad; (16) zygomatic ramus of squamosal long, thick, curved, and horizontal; (17) clavicles absent; (18) prepollex short and triangular; (19) carpal 2 present; (20) finger phalangeal formula 2-2-3-3; (21) toe phalangeal formula 2-2-3-4- 3; (22) unpulsed calls emitted in series of 917 calls at irregular intervals; (23) non-frequency modulated calls; (24) call dominant frequency 5460 ± 117 Hz (n = 79); (25) call duration
</paragraph>
<paragraph blockId="27.[533,1536,555,1897]" box="[533,1252,1630,1654]" pageId="27" pageNumber="28">45.4 ± 8.2 ms (n = 79); (26) inter-call interval 63.0 ± 9.0 ms (n = 73).</paragraph>
<paragraph blockId="27.[533,1536,555,1897]" pageId="27" pageNumber="28">
Among extremely miniaturized cophylines, this species is unique in possessing vomerine teeth. It can be distinguished from almost all other miniaturised species in lacking clavicles (also absent in
<emphasis box="[692,704,1735,1758]" italics="true" pageId="27" pageNumber="28">S</emphasis>
.
<emphasis box="[716,833,1734,1758]" italics="true" pageId="27" pageNumber="28">contumelia</emphasis>
,
<emphasis box="[845,857,1735,1758]" italics="true" pageId="27" pageNumber="28">S</emphasis>
.
<emphasis box="[869,963,1734,1758]" italics="true" pageId="27" pageNumber="28">obscoena</emphasis>
,
<emphasis box="[975,987,1735,1758]" italics="true" pageId="27" pageNumber="28">S</emphasis>
.
<emphasis box="[999,1205,1734,1758]" italics="true" pageId="27" pageNumber="28">davidattenboroughi</emphasis>
,
<emphasis box="[1217,1229,1735,1758]" italics="true" pageId="27" pageNumber="28">S</emphasis>
.
<emphasis box="[1241,1317,1734,1758]" italics="true" pageId="27" pageNumber="28">makira</emphasis>
,
<emphasis box="[1329,1341,1735,1758]" italics="true" pageId="27" pageNumber="28">S</emphasis>
.
<emphasis box="[1353,1428,1734,1758]" italics="true" pageId="27" pageNumber="28">achillei</emphasis>
, and
<emphasis box="[1486,1498,1735,1758]" italics="true" pageId="27" pageNumber="28">S</emphasis>
.
<emphasis box="[533,662,1769,1793]" italics="true" pageId="27" pageNumber="28">analanjirofo</emphasis>
, and some specimens of
<emphasis box="[926,938,1769,1792]" italics="true" pageId="27" pageNumber="28">S</emphasis>
.
<emphasis box="[951,1052,1769,1793]" italics="true" pageId="27" pageNumber="28">tridactyla</emphasis>
, unpublished data). It is also characterised by a dark colouration of the lateral surface of the head with a distinct colour border, and less reduced fingers and toes. Confusion with juvenile
<taxonomicName box="[1066,1226,1838,1862]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="27" pageNumber="28" phylum="Chordata" rank="genus">
<emphasis box="[1066,1226,1838,1862]" italics="true" pageId="27" pageNumber="28">Rhombophryne</emphasis>
</taxonomicName>
species is still possible, but these have much larger teeth proportional to their skull size, and most lack the distinct lateral
</paragraph>
<caption httpUri="https://zenodo.org/record/2614103/files/figure.png" pageId="28" pageNumber="29" startId="28.[115,143,1722,1741]" targetBox="[115,1536,208,1700]" targetPageId="28">
<paragraph blockId="28.[115,1219,1722,1782]" pageId="28" pageNumber="29">
<emphasis bold="true" box="[115,812,1722,1741]" pageId="28" pageNumber="29">
Fig 10.
<taxonomicName authority="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw, 2019" authorityName="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw" authorityYear="2019" box="[181,444,1722,1741]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="28" pageNumber="29" phylum="Chordata" rank="genus" species="proportionalis" status="sp. nov.">
<emphasis bold="true" box="[181,444,1722,1741]" italics="true" pageId="28" pageNumber="29">Rhombophryne proportionalis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[449,517,1722,1741]" pageId="28" pageNumber="29" rank="species">sp. nov.</taxonomicNameLabel>
, holotype ZSM 1826/2010, in life.
</emphasis>
(a) dorsolateral, (b) dorsal, and (c) ventral view. https://doi.org/10.1371/journal.pone.0213314.g01 0
</paragraph>
</caption>
<paragraph blockId="28.[533,1519,1817,1875]" pageId="28" pageNumber="29">
head colouration and possess clavicles. The call is unique among the frogs of
<collectingCountry box="[1346,1473,1817,1841]" name="Madagascar" pageId="28" pageNumber="29">Madagascar</collectingCountry>
and is instantly distinctive in being emitted as a rapid, high-pitched series of tonal notes.
</paragraph>
</subSubSection>
<caption httpUri="https://zenodo.org/record/2614105/files/figure.png" pageId="29" pageNumber="30" startId="29.[192,220,1159,1178]" subCaptionStartIDs="29.[1264,1291,1184,1203]" subCaptionStarts="Fig 6" targetBox="[192,1536,208,1137]" targetPageId="29">
<paragraph blockId="29.[192,1534,1159,1245]" pageId="29" pageNumber="30">
<emphasis bold="true" box="[192,954,1159,1178]" pageId="29" pageNumber="30">
Fig 11. Osteology of
<taxonomicName authority="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw, 2019" authorityName="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw" authorityYear="2019" box="[374,637,1159,1178]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="29" pageNumber="30" phylum="Chordata" rank="genus" species="proportionalis" status="sp. nov.">
<emphasis bold="true" box="[374,637,1159,1178]" italics="true" pageId="29" pageNumber="30">Rhombophryne proportionalis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[641,709,1159,1178]" pageId="29" pageNumber="30" rank="species">sp. nov.</taxonomicNameLabel>
holotype (ZSM 1826/2010).
</emphasis>
(a, b) Whole skeleton in (a) dorsal and (b) ventral view. (c-f) Skull in (c) lateral, (d) ventral, (e) anterior, and (f) dorsal view. (g) Foot in ventral view. (h) Hand in ventral view. For abbreviations, see Fig 6. https://doi.org/10.1371/journal.pone.0213314.g01 1
</paragraph>
</caption>
<subSubSection lastPageId="32" lastPageNumber="33" pageId="29" pageNumber="30" type="description">
<paragraph blockId="29.[533,1536,1279,1858]" pageId="29" pageNumber="30">
<emphasis bold="true" box="[565,808,1279,1303]" pageId="29" pageNumber="30">
<typeStatus box="[565,668,1279,1303]" pageId="29" pageNumber="30">Holotype</typeStatus>
description.
</emphasis>
Specimen in a good state of preservation, part of the right thigh removed as a tissue sample. Body somewhat rhomboid; head wider than long, narrower than body; snout rounded in dorsal view, squared in lateral view; nostrils directed laterally, not protuberant, closer to eye than to tip of snout; canthus rostralis rounded, concave; loreal region flat, vertical; tympanum indistinct, round, about 57% of eye diameter; supratympanic fold distinct, weakly raised, curving slightly from posterior corner of eye over tympanum toward axilla; tongue very broad, disc-like, posteriorly free, unlobed; maxillary teeth present; vomerine teeth present in two tiny patches either side of the midline; choanae rounded. Forelimbs slen- der; subarticular tubercles faint, single; outer metacarpal tubercle faint, paired; inner metacarpal tubercle distinct, elongated; hand without webbing; first finger reduced, second and fourth short; relative length of fingers 1
<emphasis box="[875,896,1626,1649]" italics="true" pageId="29" pageNumber="30">&lt;</emphasis>
2 = 4
<emphasis box="[948,969,1626,1649]" italics="true" pageId="29" pageNumber="30">&lt;</emphasis>
3; finger tips not expanded. Hind limbs robust; TIBL 34% of SVL; lateral metatarsalia strongly connected; inner metatarsal tubercle thin and indistinct; outer metatarsal tubercle small and indistinct; no webbing between toes; first toe highly reduced, second toe short; relative length of toes 1
<emphasis box="[1062,1083,1730,1753]" italics="true" pageId="29" pageNumber="30">&lt;</emphasis>
2
<emphasis box="[1095,1116,1730,1753]" italics="true" pageId="29" pageNumber="30">&lt;</emphasis>
5
<emphasis box="[1128,1149,1730,1753]" italics="true" pageId="29" pageNumber="30">&lt;</emphasis>
3
<emphasis box="[1161,1182,1730,1753]" italics="true" pageId="29" pageNumber="30">&lt;</emphasis>
4; fifth toe distinctly shorter than third. Skin on dorsum smooth in preservative, without distinct dorsolateral folds. In life, the dorsal skin was smooth was scattered tubercles, and distinct ridges above the scapular region (
<figureCitation box="[542,607,1834,1858]" captionStart="Fig 10" captionStartId="28.[115,143,1722,1741]" captionTargetBox="[115,1536,208,1700]" captionTargetId="figure@28.[115,1536,208,1700]" captionTargetPageId="28" captionText="Fig 10. Rhombophryne proportionalis sp. nov., holotype ZSM 1826/2010, in life. (a) dorsolateral, (b) dorsal, and (c) ventral view. https://doi.org/10.1371/journal.pone.0213314.g010" httpUri="https://zenodo.org/record/2614103/files/figure.png" pageId="29" pageNumber="30">Fig 10</figureCitation>
). Ventral skin smooth in preservative, granular in life.
</paragraph>
<paragraph blockId="30.[533,1536,208,1896]" pageId="30" pageNumber="31">After eight years in 70% ethanol, the dorsum is chocolate brown in colour, darker over the head, with a faint dark brown line running from the inguinal region anteriorly toward the eye. The dorsal leg has a dark brown crossband on the shank. The lateral head has a distinct colour border to the dorsum, being a much darker brown, its border defined by the supratympanic fold. The flank has an indistinct colour border to the venter. The venter is brown with cream flecks, slightly darker and less flecked on the chin. The ventral legs and arms are as the ventral abdomen in colour. The bottom of the foot is dark brown along the medial half. Colour in life as in preservative but more vibrant.</paragraph>
<paragraph blockId="30.[533,1536,208,1896]" pageId="30" pageNumber="31">
<emphasis bold="true" box="[565,680,485,509]" pageId="30" pageNumber="31">Variation.</emphasis>
For measurements, see
<tableCitation box="[934,1012,485,509]" captionStart="Table 1" captionStartId="12.[96,146,518,537]" captionText="Table 1. Morphological measurements (in mm) of new species of miniaturised cophyline microhylids. Measurement abbreviations are listed in the Materials and Methods. A = Adult, M = Male, F = Female (lowercase indicates probable subadult), p = paratype, h = holotype (holotypes are also bolded), na = not available." pageId="30" pageNumber="31">Table 1</tableCitation>
. The
<typeStatus box="[1071,1172,485,509]" pageId="30" pageNumber="31">paratypes</typeStatus>
are in general very similar to the
<typeStatus box="[533,626,520,544]" pageId="30" pageNumber="31">holotype</typeStatus>
in morphology.
<collectionCode box="[803,855,520,544]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
636/2014 is slightly smaller than the other specimens, and has a slightly shorter, more rounded head. Its supratympanic fold is also less curved than those of the other specimens, being rather more straight from the eye to above the arm.
<collectionCode box="[1370,1422,589,613]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
1840/ 2010 has a more massive body than the others. The colouration of the specimens is relatively consistent, with the whole venter of
<collectionCode box="[915,967,658,682]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
1840/2010 being darker than those of the other two specimens. The dorsolateral lines of the
<typeStatus box="[957,1050,693,717]" pageId="30" pageNumber="31">holotype</typeStatus>
are present in
<collectionCode box="[1206,1258,693,717]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
1840/2010, but not in
<collectionCode box="[533,585,728,752]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
636/2014, instead being broken in that specimen into spots above the suprascapular region and lines in the inguinal region. The crossbands of the shanks are less distinct in
<collectionCode box="[1461,1513,762,786]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
1840/2010 than the other two specimens.
</paragraph>
<paragraph blockId="30.[533,1536,208,1896]" pageId="30" pageNumber="31">
<emphasis bold="true" box="[565,710,832,856]" pageId="30" pageNumber="31">Bioacoustics.</emphasis>
Specimens called only during the day, from open, shrubby landscape between dense vegetation, on the ground. In some areas, numerous specimens could be heard calling in a chorus. Calls were recorded from an uncollected specimen by M. Vences (
<figureCitation box="[1336,1405,901,925]" captionStart="Fig 5" captionStartId="10.[200,228,1285,1304]" captionTargetBox="[200,1536,208,1263]" captionTargetId="figure@10.[200,1536,208,1263]" captionTargetPageId="10" captionText="Fig 5. One-second spectrograms and oscillograms of the calls of the new species described here. Insets represent the respective species but not the calling specimens. (a) Mini mum gen. et sp. nov., paratype ZMB 81993 from Manombo, (b) Mini scule gen. et sp. nov., ZSM 265/2018 from Sainte Luce, (c) Rhombophryne proportionalis sp. nov., part of a call (note series) of a specimen from Bepia campsite, Tsaratanana (not collected), (d) Anodonthyla eximia sp. nov., specimen not collected, from Maharira (Ranomafana). https://doi.org/10.1371/journal.pone.0213314.g005" httpUri="https://zenodo.org/record/2614093/files/figure.png" pageId="30" pageNumber="31">Fig 5C</figureCitation>
,
<tableCitation box="[1417,1495,901,925]" captionStart="Table 2" captionStartId="13.[533,583,194,213]" captionTargetBox="[533,1512,335,774]" captionTargetId="graphics@13.[533,1536,323,778]" captionTargetPageId="13" captionText="Table 2. Bioacoustic parameters of new species of miniaturised cophyline microhylids. Data on S. miery from Rakotoarison et al. [14], provided for comparison with the sympatric A. eximia. Values are presented as mean ± standard deviation, with range in brackets. na = not applicable. In all species except R. proportionalis calls consist of a single note according to the definition herein, and in these species call duration is therefore synonymous with note duration." httpUri="http://table.plazi.org/id/DF2E6668FF9E0A45FD8B069ADEFD686E" pageId="30" pageNumber="31" tableUuid="DF2E6668FF9E0A45FD8B069ADEFD686E">Table 2</tableCitation>
) at around 11h40 on
<date box="[722,859,936,960]" pageId="30" pageNumber="31" value="2010-06-15">
15 June
<quantity box="[808,886,936,960]" metricMagnitude="1" metricUnit="m" metricValue="5.1053999999999995" pageId="30" pageNumber="31" unit="in" value="2010.0">2010 in</quantity>
</date>
Camp Bepia (14.11822˚S, 048.97822˚E,
<quantity box="[1311,1390,936,960]" metricMagnitude="3" metricUnit="m" metricValue="2.294" pageId="30" pageNumber="31" unit="m" value="2294.0">2294 m</quantity>
a.s.l.), and further calls were heard but not recorded at Camp Andranomadio (14.0801˚S, 048.9854˚E,
<quantity box="[533,612,1005,1029]" metricMagnitude="3" metricUnit="m" metricValue="2.503" pageId="30" pageNumber="31" unit="m" value="2503.0">2503 m</quantity>
a.s.l.). A precise description of the call structure is difficult; the vocalization is a series of short tonal units that cannot readily be assigned to units. In past studies, we have described roughly comparable structures differently: in
<emphasis box="[1014,1237,1075,1099]" italics="true" pageId="30" pageNumber="31">
<taxonomicName box="[1014,1117,1075,1099]" class="Amphibia" family="Microhylidae" genus="Stumpffia" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">Stumpffia</taxonomicName>
psologlossa
</emphasis>
, where the units have a very short duration, as a single call composed of pulses [
<bibRefCitation author="Rakotoarison A &amp; Scherz MD &amp; Glaw F &amp; Kohler J &amp; Andreone F &amp; Franzen M" box="[1075,1100,1109,1133]" journalOrPublisher="Vertebrate Zoology." pageId="30" pageNumber="31" pagination="271 - 398" part="67" refId="ref23849" refString="14. Rakotoarison A, Scherz MD, Glaw F, Kohler J, Andreone F, Franzen M, et al. Describing the smaller majority: Integrative taxonomy reveals twenty-six new species of tiny microhylid frogs (genus Stumpffia) from Madagascar. Vertebrate Zoology. 2017; 67 (3): 271 - 398." title="et al. Describing the smaller majority: Integrative taxonomy reveals twenty-six new species of tiny microhylid frogs (genus Stumpffia) from Madagascar" type="journal article" year="2017">14</bibRefCitation>
], in
<emphasis box="[1149,1449,1109,1133]" italics="true" pageId="30" pageNumber="31">
<taxonomicName box="[1149,1309,1109,1133]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">Rhombophryne</taxonomicName>
mangabensis
</emphasis>
as a series of notes [
<bibRefCitation author="Glaw F &amp; Kohler J &amp; Vences M." box="[697,722,1144,1168]" journalOrPublisher="Zoosystematics and Evolution." pageId="30" pageNumber="31" pagination="235 - 43" part="86" refId="ref25994" refString="52. Glaw F, Kohler J, Vences M. A new fossorial frog, genus Rhombophryne, from Nosy Mangabe Special Reserve, Madagascar. Zoosystematics and Evolution. 2010; 86 (2): 235 - 43. https: // doi. org / 10.1002 / zoos. 20100 0 0 0 6" title="A new fossorial frog, genus Rhombophryne, from Nosy Mangabe Special Reserve, Madagascar" type="journal article" year="2010">52</bibRefCitation>
], and in
<emphasis box="[817,833,1145,1168]" italics="true" pageId="30" pageNumber="31">R</emphasis>
.
<emphasis box="[845,922,1145,1168]" italics="true" pageId="30" pageNumber="31">minuta</emphasis>
two closely spaced units were seen as components of a single note [
<bibRefCitation author="Glaw F &amp; Kohler J &amp; Vences M." box="[663,688,1178,1202]" journalOrPublisher="Zoosystematics and Evolution." pageId="30" pageNumber="31" pagination="235 - 43" part="86" refId="ref25994" refString="52. Glaw F, Kohler J, Vences M. A new fossorial frog, genus Rhombophryne, from Nosy Mangabe Special Reserve, Madagascar. Zoosystematics and Evolution. 2010; 86 (2): 235 - 43. https: // doi. org / 10.1002 / zoos. 20100 0 0 0 6" title="A new fossorial frog, genus Rhombophryne, from Nosy Mangabe Special Reserve, Madagascar" type="journal article" year="2010">52</bibRefCitation>
]. To allow comparison within
<taxonomicName box="[1015,1175,1179,1203]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis box="[1015,1175,1179,1203]" italics="true" pageId="30" pageNumber="31">Rhombophryne</emphasis>
</taxonomicName>
we here define the tonal units in the vocalizations of the new species as notes, and the entire series as a call, but emphasize that it also would be possible to define each unit as call and the entire series as call series, or to consider each unit as a pulse as their duration falls within the 550 ms range for which the pulse category was recommended by Köhler et al. [
<bibRefCitation author="Kohler J &amp; Jansen M &amp; Rodriguez A &amp; Kok PJR &amp; Toledo LF &amp; Emmrich M" box="[1010,1035,1317,1341]" journalOrPublisher="Zootaxa." pageId="30" pageNumber="31" pagination="1 - 124" part="4251" refId="ref25028" refString="33. Kohler J, Jansen M, Rodriguez A, Kok PJR, Toledo LF, Emmrich M, et al. The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice. Zootaxa. 2017; 4251 (1): 1 - 124. https: // doi. org / 10.11646 / zootaxa. 425 1.1. 1 PMID: 28609991" title="et al. The use of bioacoustics in anuran taxonomy: theory, terminology, methods and recommendations for best practice" type="journal article" year="2017">33</bibRefCitation>
].
</paragraph>
<paragraph blockId="30.[533,1536,208,1896]" pageId="30" pageNumber="31">
Calls are rapid series of high-pitched slightly frequency modulated notes. Each call consists of a series of 917 notes, and calls are repeated at long and irregular intervals. For detailed call parameters, see
<tableCitation box="[702,780,1421,1445]" captionStart="Table 2" captionStartId="13.[533,583,194,213]" captionTargetBox="[533,1512,335,774]" captionTargetId="graphics@13.[533,1536,323,778]" captionTargetPageId="13" captionText="Table 2. Bioacoustic parameters of new species of miniaturised cophyline microhylids. Data on S. miery from Rakotoarison et al. [14], provided for comparison with the sympatric A. eximia. Values are presented as mean ± standard deviation, with range in brackets. na = not applicable. In all species except R. proportionalis calls consist of a single note according to the definition herein, and in these species call duration is therefore synonymous with note duration." httpUri="http://table.plazi.org/id/DF2E6668FF9E0A45FD8B069ADEFD686E" pageId="30" pageNumber="31" tableUuid="DF2E6668FF9E0A45FD8B069ADEFD686E">Table 2</tableCitation>
.
</paragraph>
<paragraph blockId="30.[533,1536,208,1896]" box="[565,1510,1456,1480]" pageId="30" pageNumber="31">
<emphasis bold="true" box="[565,775,1456,1480]" pageId="30" pageNumber="31">
Osteology (
<figureCitation box="[692,759,1456,1480]" captionStart="Fig 11" captionStartId="29.[192,220,1159,1178]" captionTargetBox="[192,1536,208,1137]" captionTargetId="figure@29.[192,1536,208,1137]" captionTargetPageId="29" captionText="Fig 11. Osteology of Rhombophryne proportionalis sp. nov. holotype (ZSM 1826/2010). (a, b) Whole skeleton in (a) dorsal and (b) ventral view. (c-f) Skull in (c) lateral, (d) ventral, (e) anterior, and (f) dorsal view. (g) Foot in ventral view. (h) Hand in ventral view. For abbreviations, see Fig 6. https://doi.org/10.1371/journal.pone.0213314.g011" httpUri="https://zenodo.org/record/2614105/files/figure.png" pageId="30" pageNumber="31">Fig 11</figureCitation>
).
</emphasis>
Based on
<collectionCode box="[883,935,1456,1480]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
1826/2010 (figured) and
<collectionCode box="[1205,1257,1456,1480]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
636/2014 (not figured).
</paragraph>
<paragraph blockId="30.[533,1536,208,1896]" pageId="30" pageNumber="31">
<emphasis bold="true" box="[565,837,1490,1514]" pageId="30" pageNumber="31">
Cranium (
<figureCitation box="[681,747,1490,1514]" captionStart="Fig 11" captionStartId="29.[192,220,1159,1178]" captionTargetBox="[192,1536,208,1137]" captionTargetId="figure@29.[192,1536,208,1137]" captionTargetPageId="29" captionText="Fig 11. Osteology of Rhombophryne proportionalis sp. nov. holotype (ZSM 1826/2010). (a, b) Whole skeleton in (a) dorsal and (b) ventral view. (c-f) Skull in (c) lateral, (d) ventral, (e) anterior, and (f) dorsal view. (g) Foot in ventral view. (h) Hand in ventral view. For abbreviations, see Fig 6. https://doi.org/10.1371/journal.pone.0213314.g011" httpUri="https://zenodo.org/record/2614105/files/figure.png" pageId="30" pageNumber="31">Fig 11</figureCitation>
C11F).
</emphasis>
Shape and proportions. Skull short and stout, relatively wide, widest at quadratojugal anterior to the otic region. Short rostrum. Braincase broad.
</paragraph>
<paragraph blockId="30.[533,1536,208,1896]" lastBlockId="31.[533,1535,208,1861]" lastPageId="31" lastPageNumber="32" pageId="30" pageNumber="31">
<emphasis bold="true" box="[565,736,1560,1584]" pageId="30" pageNumber="31">Neurocranium.</emphasis>
Well ossified except the weakly calcified otic capsules. Sphenethmoid well ossified, in contact with the frontoparietal in
<collectionCode box="[1010,1062,1594,1618]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
1826/2010 but not in
<collectionCode box="[1295,1347,1594,1618]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="30" pageNumber="31" type="Museum">ZSM</collectionCode>
636/2014, with a calcified anterior medial cone; broadly separated from prootic. Prootic in dorsal contact with lateral flange of frontoparietal, ventral contact with parasphenoid alae, not approaching contralateral. Septomaxilla miniscule, roughly spiralled, similar in shape to those of other
<taxonomicName class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="30" pageNumber="31" phylum="Chordata" rank="genus">
<emphasis italics="true" pageId="30" pageNumber="31">Rhombophryne</emphasis>
</taxonomicName>
(see [
<bibRefCitation author="Scherz MD &amp; Hawlitschek O &amp; Andreone F &amp; Rakotoarison A &amp; Vences M &amp; Glaw F." box="[668,693,1733,1757]" journalOrPublisher="Zootaxa." pageId="30" pageNumber="31" pagination="301 - 40" part="4273" refId="ref24597" refString="25. Scherz MD, Hawlitschek O, Andreone F, Rakotoarison A, Vences M, Glaw F. A review of the taxonomy and osteology of the Rhombophryne serratopalpebrosa species group (Anura: Microhylidae) from Madagascar, with comments on the value of volume rendering of micro-CT data to taxonomists. Zootaxa. 2017; 4273 (3): 301 - 40. https: // doi. org / 10.11646 / zootaxa. 427 3.3. 1 PMID: 28610237" title="A review of the taxonomy and osteology of the Rhombophryne serratopalpebrosa species group (Anura: Microhylidae) from Madagascar, with comments on the value of volume rendering of micro-CT data to taxonomists" type="journal article" year="2017">25</bibRefCitation>
]). Columella (stapes) well ossified, pars media plectra (stylus) steeply sloping upwards proximally to its broad, bilobed, flattened pars interna plectra (baseplate). Nasal broad, curved over the nasal capsule, triangular, acuminate maxillary process not closely approaching maxillary pars facialis, broadly separated from contralateral, situated anterior to frontoparietal. Frontoparietal with slanted anterior edge, laterally bulging, with short lateral flange covering prootic, posteriorly weakly connected to exoccipital, anteroventrally contacting sphenethmoid, lacking any dorsal process (suggestions of such processes in
<collectionCode box="[1374,1426,242,266]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="31" pageNumber="32" type="Museum">ZSM</collectionCode>
1826/ 2010).
</paragraph>
<paragraph blockId="31.[533,1535,208,1861]" pageId="31" pageNumber="32">Parasphenoid with broad, squared cultriform process and broad perpendicular alae, shorter than frontoparietals, posteromedial process not participating in foramen magnum, anteroventrally in contact with postchoanal vomer and not in contact with neopalatine. Vomer divided into pre- and postchoanal portions; prechoanal portion narrow, arcuate, triradiate, with a short lateral and anterior ramus and long posterior ramus; postchoanal portion spatulate bearing a single vomerine tooth or a pair thereof, separated from contralateral by a narrow space, in dorsal contact with the parasphenoid proximally and the neopalatine distally, lacking an anterior projection. Neopalatine laminar, broader than lateral postchoanal portion of vomer, laterally approaching but not contacting the maxilla, exceeding the lateral-most point of the postchoanal vomer to approach the anterior tip of the parasphenoid but not in contact with it, such that the vomer contacts the neopalatine near its midpoint and not at its terminus.</paragraph>
<paragraph blockId="31.[533,1535,208,1861]" pageId="31" pageNumber="32">Maxillary arcade quite robust, maxilla and premaxilla bearing numerous diminutive teeth, premaxilla and maxilla in narrow contact anteriorly. Premaxilla with a broad acuminate dorsal alary process rising vertically, pars palatina shallowly divided into a narrow palatine process and broad lateral process. Maxilla with a low triangular pars facialis and a narrow pars palatina, its posterior tip acuminate and broadly overlapping the quadratojugal, the lingual surface of the pars palatina in broad contact with the anterior ramus of the pterygoid.</paragraph>
<paragraph blockId="31.[533,1535,208,1861]" pageId="31" pageNumber="32">Pterygoid with a short medial ramus, long anterior ramus with broad contact with the maxilla, leaving a channel for the pterygoid cartilage, posterior ramus broad and posterolaterally calcified to the quadratojugal complex. Quadratojugal bowed laterally, broadly connected to the ventral ramus of the squamosal, and with a broad articular surface with the maxilla, bearing a large posteroventral knob. Squamosal with a broadened ventral ramus and narrow otic and zygomatic rami, the otic ramus long and thin and oriented dorsally, the zygomatic ramus short and thin, oriented anteriorly.</paragraph>
<paragraph blockId="31.[533,1535,208,1861]" pageId="31" pageNumber="32">Mandible slim and edentate, largely unremarkable, with a low coronoid process on the angulosplenial. Mentomeckelians strongly connected to the dentary, with unusual, flat ventrolateral projections.</paragraph>
<paragraph blockId="31.[533,1535,208,1861]" box="[565,1415,1248,1272]" pageId="31" pageNumber="32">Posteromedial processes of hyoid proximally pointed with a broad medial crista.</paragraph>
<paragraph blockId="31.[533,1535,208,1861]" pageId="31" pageNumber="32">
<emphasis bold="true" box="[565,1121,1282,1306]" pageId="31" pageNumber="32">
Postcranial skeleton (
<figureCitation box="[804,1106,1282,1306]" captionStart="Fig 11" captionStartId="29.[192,220,1159,1178]" captionTargetBox="[192,1536,208,1137]" captionTargetId="figure@29.[192,1536,208,1137]" captionTargetPageId="29" captionText="Fig 11. Osteology of Rhombophryne proportionalis sp. nov. holotype (ZSM 1826/2010). (a, b) Whole skeleton in (a) dorsal and (b) ventral view. (c-f) Skull in (c) lateral, (d) ventral, (e) anterior, and (f) dorsal view. (g) Foot in ventral view. (h) Hand in ventral view. For abbreviations, see Fig 6. https://doi.org/10.1371/journal.pone.0213314.g011" httpUri="https://zenodo.org/record/2614105/files/figure.png" pageId="31" pageNumber="32">Fig 11A, 11B, 11G and 11H</figureCitation>
).
</emphasis>
Eight procoelous unfused presacrals, much broader than long, lacking neural spines, with round posterior articular processes, presacral I with a mostly complete neural arch, presacrals IIIV with thicker and longer transverse processes than VVIII. Sacrum with expanded diapophyses, the leading and trailing edges roughly equally angled, the articulation
<typeStatus box="[954,999,1421,1445]" pageId="31" pageNumber="32">type</typeStatus>
IIB
<emphasis box="[1045,1102,1422,1445]" italics="true" pageId="31" pageNumber="32">sensu</emphasis>
Emerson [
<bibRefCitation author="Emerson SB" box="[1218,1243,1421,1445]" journalOrPublisher="Biological Journal of the Linnean Society." pageId="31" pageNumber="32" pagination="153 - 68" part="11" refId="ref25518" refString="42. Emerson SB. The ilio-sacral articulation in frogs: form and function. Biological Journal of the Linnean Society. 1979; 11: 153 - 68. https: // doi. org / 10.1111 / j. 1095 - 8312.1979. tb 00032. x" title="The ilio-sacral articulation in frogs: form and function" type="journal article" year="1979">42</bibRefCitation>
]. Urostyle bicondylar, long, broadening posteriorly, without lateral processes and with a low dorsal ridge.
</paragraph>
<paragraph blockId="31.[533,1535,208,1861]" pageId="31" pageNumber="32">Pectoral girdle without ossified prezonal or postzonal elements, lacking ossified clavicles. Coracoid broad, strongly flared with a large medial articular surface with the contralateral. Scapula also robust, with a broad pars acromialis distinct from the pars glenoidalis. Cleithrum ossified for half the width of the scapular border, acuminate, thickened anteriorly. Suprascapula unossified.</paragraph>
<paragraph blockId="31.[533,1535,208,1861]" pageId="31" pageNumber="32">
Humerus with a well-developed crista ventralis and no medial or lateral cristae. Radioulna broad with a distinct sulcus intermedius. Carpals well ossified in
<collectionCode box="[1219,1271,1698,1722]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="31" pageNumber="32" type="Museum">ZSM</collectionCode>
1826/2010 and poorly ossified in
<collectionCode box="[647,699,1733,1757]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="31" pageNumber="32" type="Museum">ZSM</collectionCode>
636/2014, composed of radiale, ulnare, prepollical element, element Y, carpal 2, and large post-axial element formed by carpals 35. Finger phalangeal formula is standard (2-2-3-3). Small distal knobs on terminal phalanges of the fingers. A very small prepollex is present in
<collectionCode box="[645,697,1837,1861]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="31" pageNumber="32" type="Museum">ZSM</collectionCode>
1826/2010 but is not visible in
<collectionCode box="[1027,1079,1837,1861]" country="Germany" httpUri="http://grbio.org/cool/4b7r-7sec" name="Bavarian State Collection of Zoology" pageId="31" pageNumber="32" type="Museum">ZSM</collectionCode>
636/2014.
</paragraph>
<paragraph blockId="32.[533,1537,208,821]" pageId="32" pageNumber="33">Pubis partly calcified, iliac shafts passing ventral to and beyond sacrum, nearly cylindrical, without a dorsal crest and with a weak dorsal prominence and shallow oblique groove. Femur weakly sigmoid with a low posterior crest. Tibiofibula shorter than femur, with a sulcus intermedius. Tibiale and fibulare weakly fused proximally and distally. T1 and T2+3 tarsals present, T1 considerably smaller than T2+3. Centrale present but not large. Prehallux unossified. Phalangeal formula standard (2-2-3-4-3).</paragraph>
</subSubSection>
<subSubSection pageId="32" pageNumber="33" type="distribution">
<paragraph blockId="32.[533,1537,208,821]" pageId="32" pageNumber="33">
<emphasis bold="true" box="[565,1165,416,440]" pageId="32" pageNumber="33">Distribution, natural history, and conservation status.</emphasis>
<taxonomicName authority="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw, 2019" authorityName="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw" authorityYear="2019" box="[1170,1483,416,440]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="32" pageNumber="33" phylum="Chordata" rank="genus" species="proportionalis" status="sp. nov.">
<emphasis box="[1170,1483,416,440]" italics="true" pageId="32" pageNumber="33">Rhombophryne proportionalis</emphasis>
</taxonomicName>
<taxonomicNameLabel pageId="32" pageNumber="33" rank="species">sp. nov.</taxonomicNameLabel>
is known from two localities (Bepia and Andranomadio) on the Tsaratanana massif in northern
<collectingCountry box="[635,765,485,509]" name="Madagascar" pageId="32" pageNumber="33">Madagascar</collectingCountry>
. Both of these sites fall within the Tsaratanana National Park (formerly a Strict Nature Reserve). It is a terrestrial species that lives among the leaf litter. Two other cophyline frogs,
<emphasis box="[709,891,555,579]" italics="true" pageId="32" pageNumber="33">
<taxonomicName authorityName="Boulenger" authorityYear="1882" box="[709,809,555,579]" class="Amphibia" family="Microhylidae" genus="Platypelis" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="32" pageNumber="33" phylum="Chordata" rank="genus">Platypelis</taxonomicName>
alticola
</emphasis>
and
<emphasis box="[943,958,555,578]" italics="true" pageId="32" pageNumber="33">P</emphasis>
.
<emphasis box="[970,1147,555,578]" italics="true" pageId="32" pageNumber="33">tsaratananaensis</emphasis>
, are also known from these locations, and both are currently listed as Endangered, due to their small distribution
<emphasis box="[1395,1416,590,613]" italics="true" pageId="32" pageNumber="33">&lt;</emphasis>
<quantity box="[1422,1513,589,613]" metricMagnitude="6" metricUnit="m" metricValue="2.0" pageId="32" pageNumber="33" unit="km" value="2000.0">2000 km</quantity>
<superScript attach="left" box="[1513,1522,583,599]" fontSize="7" pageId="32" pageNumber="33">2</superScript>
, presence in a single threat-defined location, and potentially on-going decline in habitat quality. We therefore suggest
<taxonomicName authority="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw, 2019" authorityName="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw" authorityYear="2019" box="[762,937,658,683]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="proportionalis" status="sp. nov.">
<emphasis box="[762,778,659,682]" italics="true" pageId="32" pageNumber="33">R</emphasis>
.
<emphasis box="[790,937,659,683]" italics="true" pageId="32" pageNumber="33">proportionalis</emphasis>
</taxonomicName>
<taxonomicNameLabel box="[943,1025,658,682]" pageId="32" pageNumber="33" rank="species">sp. nov.</taxonomicNameLabel>
also to be Endangered, following the same rationale.
</paragraph>
</subSubSection>
<subSubSection pageId="32" pageNumber="33" type="etymology">
<paragraph blockId="32.[533,1537,208,821]" pageId="32" pageNumber="33">
<emphasis bold="true" box="[565,692,728,752]" pageId="32" pageNumber="33">Etymology.</emphasis>
The species epithet is the Latin adjective
<taxonomicName authorityName="Scherz &amp; Hutter &amp; Rakotoarison &amp; Riemann &amp; Rödel &amp; Ndriantsoa &amp; Glos &amp; Roberts &amp; Crottini &amp; Vences &amp; Glaw" authorityYear="2019" box="[1127,1274,728,752]" class="Amphibia" family="Microhylidae" genus="Rhombophryne" higherTaxonomySource="GBIF" kingdom="Animalia" order="Anura" pageId="32" pageNumber="33" phylum="Chordata" rank="species" species="proportionalis">
<emphasis box="[1127,1274,728,752]" italics="true" pageId="32" pageNumber="33">proportionalis</emphasis>
</taxonomicName>
meaning proportional, in reference to the comparatively proportional dwarfism that this species has apparently undergone (see discussion). It is a feminine adjective in the nominative singular.
</paragraph>
</subSubSection>
</treatment>
</document>