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<mods:title id="F4F092A1AF7D408E571E56ED839D28D7">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>
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.— 
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<emphasis id="E5F89A11FF9DFF8F4FD84EAF48677ACE" box="[993,1326,1451,1477]" italics="true" pageId="65" pageNumber="178">Mantidactylus ambohimitombi</emphasis>
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is based on a series of 
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that include 
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from ‘ 
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. Blommers-Schl̂sser and Blanc (1991) noted that 
<typeStatus id="0837F8A1FF9DFF8F49724D0F48E2792E" box="[1355,1451,1547,1573]" pageId="65" pageNumber="178" type="syntype">syntypes</typeStatus>
<collectionCode id="B19DDEC6FF9DFF8F4F134D2F4E37794E" box="[810,894,1579,1605]" collectionName="United Kingdom, London, The Natural History Museum [formerly British Museum (Natural History)]" pageId="65" pageNumber="178">BMNH</collectionCode>
<specimenCode id="872AEE78FF9DFF8F4F994D2F4916794E" box="[928,1119,1579,1605]" pageId="65" pageNumber="178">
1947.2.26.31– 
<date id="A33260C3FF9DFF8F487A4D2F4916794E" box="[1091,1119,1579,1605]" pageId="65" pageNumber="178" value="1932-05">32</date>
</specimenCode>
may be referable to 
<taxonomicName id="108C3D80FF9DFF8F4F134D4F496D796E" box="[810,1060,1611,1637]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="65" pageNumber="178" phylum="Chordata" rank="species" species="curtus">
<emphasis id="E5F89A11FF9DFF8F4F134D4F496D796E" box="[810,1060,1611,1637]" italics="true" pageId="65" pageNumber="178">Mantidactylus curtus</emphasis>
</taxonomicName>
, without justification.
</materialsCitation>
We here designate the 
<materialsCitation id="67E44C5EFF9DFF8F4FC24D6F4EF479CE" ID-GBIF-Occurrence="4010830445" collectionCode="BMNH" pageId="65" pageNumber="178" specimenCode="1947.2.26.25" specimenCount="1" typeStatus="lectotype">
DNA-barcoded 
<specimenCount id="C18A8D8AFF9DFF8F48F24D6F487C798E" box="[1227,1333,1643,1669]" pageId="65" pageNumber="178" type="generic">specimen</specimenCount>
<collectionCode id="B19DDEC6FF9DFF8F496E4D6F48E2798E" box="[1367,1451,1643,1669]" collectionName="United Kingdom, London, The Natural History Museum [formerly British Museum (Natural History)]" pageId="65" pageNumber="178">BMNH</collectionCode>
<specimenCode id="872AEE78FF9DFF8F4F134D8F4EF679AD" box="[810,959,1675,1702]" pageId="65" pageNumber="178">1947.2.26.25</specimenCode>
, a large-sized individual of 65.0 mm SVL, as 
<typeStatus id="0837F8A1FF9DFF8F4F764DAF4EF079CE" box="[847,953,1707,1733]" pageId="65" pageNumber="178" type="lectotype">lectotype</typeStatus>
.
</materialsCitation>
<typeStatus id="0837F8A1FF9DFF8F4FF34DAF497379CE" box="[970,1082,1707,1733]" pageId="65" pageNumber="178" type="lectotype">Lectotype</typeStatus>
designation is justified by the need to stabilize this and other nomina in 
<taxonomicName id="108C3D80FF9DFF8F493E4DC848ED79EE" box="[1287,1444,1740,1765]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="65" pageNumber="309" phylum="Chordata" rank="subGenus" subGenus="Brygoomantis">
<emphasis id="E5F89A11FF9DFF8F493E4DC848ED79EE" box="[1287,1444,1740,1765]" italics="true" pageId="65" pageNumber="178">Brygoomantis</emphasis>
</taxonomicName>
, given the uncertain identity and morphological similarity of many taxa in the subgenus.
</paragraph>
</subSubSection>
<subSubSection id="9F961588FF9DFF8D4F634C2F484D792E" lastPageId="67" lastPageNumber="180" pageId="65" pageNumber="178" type="description">
<paragraph id="D7334603FF9DFF8F4F634C2F4946772E" blockId="65.[810,1452,1451,2085]" pageId="65" pageNumber="178">
<emphasis id="E5F89A11FF9DFF8F4F634C2F4EF9784E" box="[858,944,1835,1861]" italics="true" pageId="65" pageNumber="178">Identity</emphasis>
.— 
<taxonomicName id="108C3D80FF9DFF8F4FEB4C2F4E2B786E" authority="Boulenger, 1919" authorityName="Boulenger" authorityYear="1919" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="65" pageNumber="178" phylum="Chordata" rank="species" species="ambohimitombi">
<emphasis id="E5F89A11FF9DFF8F4FEB4C2F4863784E" box="[978,1322,1835,1861]" italics="true" pageId="65" pageNumber="178">Mantidactylus ambohimitombi</emphasis>
<bibRefCitation id="B31D3BF2FF9DFF8F49084C2F4E2B786E" author="Boulenger, G. A." pageId="65" pageNumber="178" pagination="257 - 261" refId="ref122966" refString="Boulenger, G. A. (1919 ' 1918 ') On the Madagascar frogs of the genus Mantidactylus. Proceedings of the Zoological Society of London, 1918, 257 - 261. https: // doi. org / 10.1111 / j. 1096 - 3642.1918. tb 02096. x" type="book chapter" year="1919">Boulenger, 1919</bibRefCitation>
</taxonomicName>
is typically considered a valid species (BlommersSchl̂sser &amp; Blanc 1991; 
<bibRefCitation id="B31D3BF2FF9DFF8F48674C6F4996788E" author="Frost, D. R." box="[1118,1247,1899,1925]" pageId="65" pageNumber="178" refId="ref125096" refString="Frost, D. R. (2021) Amphibian Species of the World: an Online Reference. Version 6.1 (Accessed 18 February 2021). Electronic Database accessible at http: // research. amnh. org / herpetology / amphibia / index. html. American Museum of Natural History, New York, USA" type="url" year="2021">Frost 2021</bibRefCitation>
; 
<bibRefCitation id="B31D3BF2FF9DFF8F48C84C6F4E3878AE" author="Glaw, F. &amp; Vences, M." pageId="65" pageNumber="178" refId="ref125414" refString="Glaw, F. &amp; Vences, M. (1992 a) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences &amp; Glaw Verlags GbR, Cologne, Germany, 335 pp. First Edition." type="book" year="1992">Glaw &amp; Vences 1992a</bibRefCitation>
, 
<bibRefCitation id="B31D3BF2FF9DFF8F4FBB4C8F4EF778AE" author="Glaw, F. &amp; Vences, M." box="[898,958,1931,1957]" pageId="65" pageNumber="178" refId="ref125499" refString="Glaw, F. &amp; Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences &amp; Glaw Verlags GbR, Cologne, Germany, 478 pp. Second Edition." type="book" year="1994">1994</bibRefCitation>
), although it was considered of uncertain status by 
<bibRefCitation id="B31D3BF2FF9DFF8F4FAF4CAF49F878CE" author="Glaw, F. &amp; Vences, M." box="[918,1201,1963,1989]" pageId="65" pageNumber="178" 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>
. It is morphologically close to 
<taxonomicName id="108C3D80FF9DFF8F4FBC4CC84EA678EE" box="[901,1007,1996,2021]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="65" pageNumber="178" phylum="Chordata" rank="species" species="curtus">
<emphasis id="E5F89A11FF9DFF8F4FBC4CC84EA678EE" box="[901,1007,1996,2021]" italics="true" pageId="65" pageNumber="178">M. curtus</emphasis>
</taxonomicName>
but distinguished by its distinctly larger body size. It was not assigned to a genetic lineage by 
<bibRefCitation id="B31D3BF2FF9DFF8F4F13430F4942772E" author="Vieites, D. R. &amp; Wollenberg, K. C. &amp; Andreone, F. &amp; Glaw, F. &amp; Vences, M." box="[810,1035,2059,2085]" pageId="65" pageNumber="178" 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 id="E5F89A11FF9DFF8F4F4743084EFE772E" box="[894,951,2059,2085]" italics="true" pageId="65" pageNumber="178">et al.</emphasis>
(2009)
</bibRefCitation>
.
</paragraph>
<caption id="83F3168BFF9EFF8C4CB14D41499479B6" ID-DOI="http://doi.org/10.5281/zenodo.7441045" ID-Zenodo-Dep="7441045" httpUri="https://zenodo.org/record/7441045/files/figure.png" pageId="66" pageNumber="179" startId="66.[136,235,1605,1629]" targetBox="[151,1436,202,1584]" targetPageId="66">
<paragraph id="D7334603FF9EFF8C4CB14D41499479B6" blockId="66.[136,1452,1605,1725]" pageId="66" pageNumber="179">
<emphasis id="E5F89A11FF9EFF8C4CB14D414C5B7956" bold="true" box="[136,274,1605,1629]" pageId="66" pageNumber="179">FIGURE 11.</emphasis>
<taxonomicName id="108C3D80FF9EFF8C4D224D414CBA7956" box="[283,499,1605,1629]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="66" pageNumber="179" phylum="Chordata" rank="species" species="alutus">
<emphasis id="E5F89A11FF9EFF8C4D224D414CBA7956" box="[283,499,1605,1629]" italics="true" pageId="66" pageNumber="179">Mantidactylus alutus</emphasis>
</taxonomicName>
in life, in dorsolateral and ventral view. (a,b) Adult male from near Lake Mantasoa (ZSM 355/2000), photographed in 2000. (c,d) Adult male from near Antoetra (probably ZMA 19550 = FGMV 2002.50), photographed in 2003. (e) Adult male from Manjakatompo, photographed in 1991. (f) Specimen from Antakasina (this specimen has not been sequenced and its identification is therefore tentative; the locality is thus not included in the species account).
</paragraph>
</caption>
<caption id="83F3168BFF9FFF8D4CB14F79481E7BBE" ID-DOI="http://doi.org/10.5281/zenodo.7441047" ID-Zenodo-Dep="7441047" httpUri="https://zenodo.org/record/7441047/files/figure.png" pageId="67" pageNumber="180" startId="67.[136,235,1149,1173]" targetBox="[151,1435,186,1130]" targetPageId="67">
<paragraph id="D7334603FF9FFF8D4CB14F79481E7BBE" blockId="67.[136,1451,1149,1206]" pageId="67" pageNumber="180">
<emphasis id="E5F89A11FF9FFF8D4CB14F794C477B9E" bold="true" box="[136,270,1149,1173]" pageId="67" pageNumber="180">FIGURE 12.</emphasis>
Audiospectrogram and corresponding oscillogram of two advertisement calls from a regular call series of 
<taxonomicName id="108C3D80FF9FFF8D49234F7A4D8F7BBD" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="alutus">
<emphasis id="E5F89A11FF9FFF8D49234F7A4D8F7BBD" italics="true" pageId="67" pageNumber="180">Mantidactylus alutus</emphasis>
</taxonomicName>
, recorded on 21 January 2003 near Antoetra (20.5–21.0°C air temperature). Recording highpass-filtered at 375 Hz.
</paragraph>
</caption>
<paragraph id="D7334603FF9FFF8D4C814E2F48CC7A8E" blockId="67.[136,778,1323,2085]" lastBlockId="67.[810,1452,1323,2085]" pageId="67" pageNumber="180">
A 16S sequence of the 
<typeStatus id="0837F8A1FF9FFF8D4D834E2F4F687A4E" box="[442,545,1323,1349]" pageId="67" pageNumber="180" type="lectotype">lectotype</typeStatus>
surprisingly clusters with a lineage predominantly known from the Ankaratra Massif. We therefore redefine this lineage provisionally as corresponding to 
<emphasis id="E5F89A11FF9FFF8D4DB04E884F397AAE" box="[393,624,1419,1445]" italics="true" pageId="67" pageNumber="180">
<taxonomicName id="108C3D80FF9FFF8D4DB04E884F257AAE" authorityName="Boulenger" authorityYear="1919" box="[393,620,1419,1445]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="ambohimitombi">M. ambohimitombi</taxonomicName>
.
</emphasis>
In previous studies the populations from the Ankaratra Massif have been considered as confirmed candidate species 
<emphasis id="E5F89A11FF9FFF8D4EF94EC84F967AEE" box="[704,735,1484,1509]" italics="true" pageId="67" pageNumber="180">M.</emphasis>
sp. 19 by 
<bibRefCitation id="B31D3BF2FF9FFF8D4CED4EEF4CF7790E" author="Vieites, D. R. &amp; Wollenberg, K. C. &amp; Andreone, F. &amp; Glaw, F. &amp; Vences, M." box="[212,446,1515,1541]" pageId="67" pageNumber="180" 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 id="E5F89A11FF9FFF8D4D154EE84C2E790E" box="[300,359,1515,1541]" italics="true" pageId="67" pageNumber="180">et al.</emphasis>
(2009)
</bibRefCitation>
, and 
<emphasis id="E5F89A11FF9FFF8D4E394EE84F56790E" box="[512,543,1516,1541]" italics="true" pageId="67" pageNumber="180">M.</emphasis>
sp. Ca19 by 
<bibRefCitation id="B31D3BF2FF9FFF8D4E874EEF4DB7792E" author="Perl, R. G. B. &amp; Nagy, Z. T. &amp; Sonet, G. &amp; Glaw, F. &amp; Wollenberg, K. C. &amp; Vences, M." pageId="67" pageNumber="180" 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 id="E5F89A11FF9FFF8D4ECC4EE84DEC792E" italics="true" pageId="67" pageNumber="180">et al.</emphasis>
(2014)
</bibRefCitation>
. They were referred to as ‘ 
<emphasis id="E5F89A11FF9FFF8D4E794D084F16792E" box="[576,607,1548,1573]" italics="true" pageId="67" pageNumber="180">M.</emphasis>
sp. 
<taxonomicName id="108C3D80FF9FFF8D4EAA4D0F4E43792E" authority="Ankaratra" authorityName="Ankaratra" box="[659,778,1547,1573]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" isUncertain="true" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="curtus">
aff. 
<emphasis id="E5F89A11FF9FFF8D4EFD4D084E43792E" box="[708,778,1548,1573]" italics="true" pageId="67" pageNumber="180">curtus</emphasis>
</taxonomicName>
“Ankaratra”’ by 
<bibRefCitation id="B31D3BF2FF9FFF8D4D764D2F4F1B794E" author="Schmidt, H. &amp; Strauss, A. &amp; Glaw, F. &amp; Teschke, M. &amp; Vences, M." box="[335,594,1579,1605]" pageId="67" pageNumber="180" pagination="48 - 60" refId="ref132292" refString="Schmidt, H., Strauss, A., Glaw, F., Teschke, M. &amp; Vences, M. (2009) Description of tadpoles of five frog species in the subgenus Brygoomantis from Madagascar (Mantellidae: Mantidactylus). Zootaxa, 1988, 48 - 60. https: // doi. org / 10.11646 / zootaxa. 1988.1.4" type="journal article" year="2009">
Schmidt 
<emphasis id="E5F89A11FF9FFF8D4D804D284CBE794E" box="[441,503,1579,1605]" italics="true" pageId="67" pageNumber="180">et al.</emphasis>
(2009)
</bibRefCitation>
. We emphasize that this attribution is preliminary; since only mtDNA data (no genomic information) are available from the 
<typeStatus id="0837F8A1FF9FFF8D4CB14D8F4DBD79AE" box="[136,244,1675,1701]" pageId="67" pageNumber="180" type="lectotype">lectotype</typeStatus>
, and no fresh samples are available from Ambohimitombo forest where the original 
<typeStatus id="0837F8A1FF9FFF8D4E534DAF4F8A79CE" box="[618,707,1707,1733]" pageId="67" pageNumber="180" type="syntype">syntype</typeStatus>
series was collected, we cannot exclude that mitochondrial introgression has taken place, potentially blurring a hypothetical differentiation between the Ankaratra and Ambohimitombo populations. Our Phylonetworks analysis provided evidence for gene flow of syntopic 
<taxonomicName id="108C3D80FF9FFF8D4CB14C684DBC788E" box="[136,245,1900,1925]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="curtus">
<emphasis id="E5F89A11FF9FFF8D4CB14C684DBC788E" box="[136,245,1900,1925]" italics="true" pageId="67" pageNumber="180">M. curtus</emphasis>
</taxonomicName>
into 
<taxonomicName id="108C3D80FF9FFF8D4D164C684F16788E" 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="[303,607,1899,1925]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="subSpecies" species="ambohimitombi" subSpecies="marefo">
<emphasis id="E5F89A11FF9FFF8D4D164C684F16788E" box="[303,607,1899,1925]" italics="true" pageId="67" pageNumber="180">M. ambohimitombi marefo</emphasis>
</taxonomicName>
<emphasis id="E5F89A11FF9FFF8D4E5C4C6F4F82788E" bold="true" box="[613,715,1899,1925]" pageId="67" pageNumber="180">ssp. nov.</emphasis>
from Itremo (
<figureCitation id="4FB75A86FF9FFF8D4CD04C8F4C7D78AD" box="[233,308,1931,1958]" captionStart="FIGURE 5" captionStartId="46.[136,235,1559,1583]" captionTargetBox="[151,1436,185,1539]" captionTargetId="figure-35@46.[151,1436,185,1539]" captionTargetPageId="46" captionText="FIGURE 5. Maximum-Likelihood tree based on a partitioned analysis of 12,818 nuclear-encoded markers obtained via the FrogCap strategy, calculated with IQ-tree, for 58 representative individuals of species-level lineages in the subgenus Mantidactylus (Brygoomantis). Note that three species are missing from this analysis (M. bletzae sp. nov., M. marintsoai sp. nov., and M. riparius sp. nov.) and the identity of the two samples of M. katae sp. nov. in this tree is uncertain. For those lineages that previously (Perl et al. 2014; Vieites et al. 2009) had candidate species numbers assigned, these are reported in parenthesis after the name used in the classification proposed herein. All branches were fully supported by SH-like approximate likelihood ratio tests with 1000 pseudoreplicates (100% support, symbolized by black dots at nodes).The tree was rooted with Mantidactylus grandidieri (subgenus Mantidactylus) as outgroup (removed from graphical representation for better visualization of ingroup relationships), with the inclusion of M. melanopleura (subgenus Chonomantis) as hierarchical outgroup. Red arrows in the M. curtus clade indicate the two reticulation events detected by a Phylonetworks analysis performed separately for this clade (note that the topology recovered by this analysis in the Phylonetworks analysis differs in the position of M. bourgati; see original results of the Phylonetworks analysis in Zenodo repository, DOI 10.5281/zenodo.668741" figureDoi="http://doi.org/10.5281/zenodo.7441033" httpUri="https://zenodo.org/record/7441033/files/figure.png" pageId="67" pageNumber="180">Fig. 5</figureCitation>
), providing a first hint that reticulated evolution may have played a role in the origin of the various morphologically divergent frogs that we here subsume in the species 
<emphasis id="E5F89A11FF9FFF8D4D8F4CE84FD6770E" box="[438,671,2027,2053]" italics="true" pageId="67" pageNumber="180">
<taxonomicName id="108C3D80FF9FFF8D4D8F4CE84FD2770E" authorityName="Boulenger" authorityYear="1919" box="[438,667,2027,2053]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="ambohimitombi">M. ambohimitombi</taxonomicName>
.
</emphasis>
A more in-depth analysis of ranges, and of gene flow among various lineages of the 
<taxonomicName id="108C3D80FF9FFF8D48794E2849F97A4E" box="[1088,1200,1324,1349]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="curtus">
<emphasis id="E5F89A11FF9FFF8D48794E2849F97A4E" box="[1088,1200,1324,1349]" italics="true" pageId="67" pageNumber="180">M. curtus</emphasis>
</taxonomicName>
clade (
<emphasis id="E5F89A11FF9FFF8D49324E2849387A6E" italics="true" pageId="67" pageNumber="180">
<taxonomicName id="108C3D80FF9FFF8D49324E2848377A4E" box="[1291,1406,1324,1349]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="curtus">M. curtus</taxonomicName>
, 
<taxonomicName id="108C3D80FF9FFF8D49B44E284EC67A6E" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="bourgati">M. bourgati</taxonomicName>
, 
<taxonomicName id="108C3D80FF9FFF8D4FA04E4849387A6E" authorityName="Boulenger" authorityYear="1919" box="[921,1137,1355,1381]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="ambohimitombi">M. ambohimitombi</taxonomicName>
</emphasis>
) is necessary to understand their evolutionary history and verify their taxonomy.
</paragraph>
<paragraph id="D7334603FF9FFF8D4F634E8F484D792E" blockId="67.[810,1452,1323,2085]" pageId="67" pageNumber="180">
In our phylogenomic tree, two other lineages form a monophyletic group with specimens of 
<taxonomicName id="108C3D80FF9FFF8D48E04EA848E57ACE" authorityName="Boulenger" authorityYear="1919" box="[1241,1452,1451,1477]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="ambohimitombi">
<emphasis id="E5F89A11FF9FFF8D48E04EA848E57ACE" box="[1241,1452,1451,1477]" italics="true" pageId="67" pageNumber="180">M. ambohimitombi</emphasis>
</taxonomicName>
from Ankaratra, and these are in subsequent accounts described as subspecies of 
<taxonomicName id="108C3D80FF9FFF8D48524EE8480F790E" authorityName="Boulenger" authorityYear="1919" box="[1131,1350,1515,1541]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="ambohimitombi">
<emphasis id="E5F89A11FF9FFF8D48524EE8480F790E" box="[1131,1350,1515,1541]" italics="true" pageId="67" pageNumber="180">M. ambohimitombi</emphasis>
</taxonomicName>
; see the rationale in the respective accounts below.
</paragraph>
</subSubSection>
<subSubSection id="9F961588FF9FFF8B4F634D284F037D4E" lastPageId="69" lastPageNumber="182" pageId="67" pageNumber="180" type="diagnosis">
<paragraph id="D7334603FF9FFF8B4F634D284F037D4E" blockId="67.[810,1452,1323,2085]" lastBlockId="69.[136,778,171,2085]" lastPageId="69" lastPageNumber="182" pageId="67" pageNumber="180">
<emphasis id="E5F89A11FF9FFF8D4F634D284E85794E" box="[858,972,1580,1605]" italics="true" pageId="67" pageNumber="180">Diagnosis</emphasis>
.— 
<taxonomicName id="108C3D80FF9FFF8D4FD74D2F4825794E" authorityName="Boulenger" authorityYear="1919" box="[1006,1388,1579,1605]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="ambohimitombi">
<emphasis id="E5F89A11FF9FFF8D4FD74D2F4825794E" box="[1006,1388,1579,1605]" italics="true" pageId="67" pageNumber="180">Mantidactylus ambohimitombi</emphasis>
</taxonomicName>
is a member of the 
<taxonomicName id="108C3D80FF9FFF8D4FC14D484921796E" box="[1016,1128,1612,1637]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="curtus">
<emphasis id="E5F89A11FF9FFF8D4FC14D484921796E" box="[1016,1128,1612,1637]" italics="true" pageId="67" pageNumber="180">M. curtus</emphasis>
</taxonomicName>
clade; it is here defined as containing three deep genetic lineages considered as subspecies, and is sister to the morphologically very distinct 
<taxonomicName id="108C3D80FF9FFF8D4FBB4DA8496279CE" box="[898,1067,1707,1733]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="madecassus">
<emphasis id="E5F89A11FF9FFF8D4FBB4DA8496279CE" box="[898,1067,1707,1733]" italics="true" pageId="67" pageNumber="180">M. madecassus</emphasis>
</taxonomicName>
. See 
<tableCitation id="9A0E73B8FF9FFF8D485C4DAF49FF79CE" box="[1125,1206,1707,1733]" 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 5‒11. 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 0–0.5 in webbing formula); 1, about one phalanx free of web (0.75–1.25); 2, more than one phalanx free of web (1.5–2). (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 single–pulse 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 31–36 mm SVL." httpUri="http://table.plazi.org/id/83F3168BFFEAFFF84CBF4AEE4C717C8F" pageId="67" pageNumber="180" tableUuid="83F3168BFFEAFFF84CBF4AEE4C717C8F">Table 4</tableCitation>
for a list of diagnostic morphological characters. The following account only diagnoses the nominal subspecies 
<emphasis id="E5F89A11FF9FFF8D488C4DE848E5780E" box="[1205,1452,1771,1797]" italics="true" pageId="67" pageNumber="180">
M. a. 
<taxonomicName id="108C3D80FF9FFF8D48C54DEF48E5780E" authorityName="Boulenger" authorityYear="1919" box="[1276,1452,1771,1797]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="ambohimitombi">ambohimitombi</taxonomicName>
</emphasis>
from other species in 
<taxonomicName id="108C3D80FF9FFF8D48714C0849AC782E" box="[1096,1253,1804,1829]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="309" phylum="Chordata" rank="subGenus" subGenus="Brygoomantis">
<emphasis id="E5F89A11FF9FFF8D48714C0849AC782E" box="[1096,1253,1804,1829]" italics="true" pageId="67" pageNumber="180">Brygoomantis</emphasis>
</taxonomicName>
; see below for diagnoses and comparisons of the two other subspecies. The combination of relatively large body size of up to 
<quantity id="1074EBE6FF9FFF8D4F704C6F4ED2788E" box="[841,923,1899,1926]" metricMagnitude="-2" metricUnit="m" metricValue="5.1" pageId="67" pageNumber="180" unit="mm" value="51.0">51 mm</quantity>
at Ankaratra (up to 
<quantity id="1074EBE6FF9FFF8D48B34C6F4995788E" box="[1162,1244,1899,1926]" metricMagnitude="-2" metricUnit="m" metricValue="6.5" pageId="67" pageNumber="180" unit="mm" value="65.0">65 mm</quantity>
in the 
<typeStatus id="0837F8A1FF9FFF8D49084C6F4828788E" box="[1329,1377,1899,1925]" pageId="67" pageNumber="180">type</typeStatus>
series from Ambohimitombo forest), slightly granular skin without dorsolateral ridges in most specimens, strongly developed foot webbing with fully webbed fifth toe, small tympanum diameter with a maximum of 11% of SVL in males, distinguishes 
<emphasis id="E5F89A11FF9FFF8D482143084859772E" box="[1048,1296,2059,2085]" italics="true" pageId="67" pageNumber="180">
M. a. 
<taxonomicName id="108C3D80FF9FFF8D4859430F4859772E" authorityName="Boulenger" authorityYear="1919" box="[1120,1296,2059,2085]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="67" pageNumber="180" phylum="Chordata" rank="species" species="ambohimitombi">ambohimitombi</taxonomicName>
</emphasis>
from species of the other clades. Within the 
<taxonomicName id="108C3D80FF99FF8B4DE64BA84F037FCE" box="[479,586,172,197]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="curtus">
<emphasis id="E5F89A11FF99FF8B4DE64BA84F037FCE" box="[479,586,172,197]" italics="true" pageId="69" pageNumber="182">M. curtus</emphasis>
</taxonomicName>
clade, 
<emphasis id="E5F89A11FF99FF8B4EA04BA84C7C7FEE" italics="true" pageId="69" pageNumber="182">
<taxonomicName id="108C3D80FF99FF8B4EA04BA84E4C7FCE" box="[665,773,171,197]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="alutus">M. alutus</taxonomicName>
, 
<taxonomicName id="108C3D80FF99FF8B4CB14BC84C7C7FEE" box="[136,309,203,229]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="madecassus">M. madecassus</taxonomicName>
</emphasis>
and 
<taxonomicName id="108C3D80FF99FF8B4D574BC84CBB7FEE" box="[366,498,203,229]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="pauliani">
<emphasis id="E5F89A11FF99FF8B4D574BC84CBB7FEE" box="[366,498,203,229]" italics="true" pageId="69" pageNumber="182">M. pauliani</emphasis>
</taxonomicName>
have smaller body sizes and are distinguished by either a usually shorter snout (
<emphasis id="E5F89A11FF99FF8B4ED24BE84CD67E2E" italics="true" pageId="69" pageNumber="182">
<taxonomicName id="108C3D80FF99FF8B4ED24BE84C587E2E" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="madecassus">M. madecassus</taxonomicName>
, 
<taxonomicName id="108C3D80FF99FF8B4D254A084CD67E2E" box="[284,415,267,293]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="pauliani">M. pauliani</taxonomicName>
</emphasis>
), or advertisement calls emitted in regular series (in 
<taxonomicName id="108C3D80FF99FF8B4D524A284C9C7E4E" box="[363,469,299,325]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="alutus">
<emphasis id="E5F89A11FF99FF8B4D524A284C9C7E4E" box="[363,469,299,325]" italics="true" pageId="69" pageNumber="182">M. alutus</emphasis>
</taxonomicName>
, vs single notes) (
<tableCitation id="9A0E73B8FF99FF8B4E9C4A2F4FB57E4E" box="[677,764,299,325]" 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 5‒11. 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 0–0.5 in webbing formula); 1, about one phalanx free of web (0.75–1.25); 2, more than one phalanx free of web (1.5–2). (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 single–pulse 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 31–36 mm SVL." httpUri="http://table.plazi.org/id/83F3168BFFEAFFF84CBF4AEE4C717C8F" pageId="69" pageNumber="182" tableUuid="83F3168BFFEAFFF84CBF4AEE4C717C8F">Table 4</tableCitation>
); 
<taxonomicName id="108C3D80FF99FF8B4CB14A484DBE7E6E" box="[136,247,332,357]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="curtus">
<emphasis id="E5F89A11FF99FF8B4CB14A484DBE7E6E" box="[136,247,332,357]" italics="true" pageId="69" pageNumber="182">M. curtus</emphasis>
</taxonomicName>
usually has a smoother skin and a somewhat shorter snout; 
<taxonomicName id="108C3D80FF99FF8B4D1C4A684CE17E8E" box="[293,424,363,389]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="bourgati">
<emphasis id="E5F89A11FF99FF8B4D1C4A684CE17E8E" box="[293,424,363,389]" italics="true" pageId="69" pageNumber="182">M. bourgati</emphasis>
</taxonomicName>
is morphologically very similar but appears to occur only on the Andringitra Massif. For detailed distinction from new species described herein, see the respective species accounts.A full list of molecular diagnostic sites in the 16S gene of 
<emphasis id="E5F89A11FF99FF8B4E2F4AE84E437D0E" box="[534,778,491,517]" italics="true" pageId="69" pageNumber="182">
M. a. 
<taxonomicName id="108C3D80FF99FF8B4E634AEF4E437D0E" authorityName="Boulenger" authorityYear="1919" box="[602,778,491,517]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="ambohimitombi">ambohimitombi</taxonomicName>
</emphasis>
in pairwise comparisons to all other 
<taxonomicName id="108C3D80FF99FF8B4E2F49084FFA7D2E" box="[534,691,524,549]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="309" phylum="Chordata" rank="subGenus" subGenus="Brygoomantis">
<emphasis id="E5F89A11FF99FF8B4E2F49084FFA7D2E" box="[534,691,524,549]" italics="true" pageId="69" pageNumber="182">Brygoomantis</emphasis>
</taxonomicName>
species is provided as Supplementary appendix.
</paragraph>
</subSubSection>
<caption id="83F3168BFF98FF8A4CB14DAE4F3A7889" ID-DOI="http://doi.org/10.5281/zenodo.7441049" ID-Zenodo-Dep="7441049" httpUri="https://zenodo.org/record/7441049/files/figure.png" pageId="68" pageNumber="181" startId="68.[136,235,1706,1730]" targetBox="[241,1348,202,1685]" targetPageId="68">
<paragraph id="D7334603FF98FF8A4CB14DAE4F3A7889" blockId="68.[136,1452,1706,1922]" pageId="68" pageNumber="181">
<emphasis id="E5F89A11FF98FF8A4CB14DAE4C4679C9" bold="true" box="[136,271,1706,1730]" pageId="68" pageNumber="181">FIGURE 13.</emphasis>
<emphasis id="E5F89A11FF98FF8A4D2D4DAE4F2879C9" box="[276,609,1706,1730]" italics="true" pageId="68" pageNumber="181">
<taxonomicName id="108C3D80FF98FF8A4D2D4DAE4CEC79C9" authorityName="Boulenger" authorityYear="1895" box="[276,421,1706,1730]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="68" pageNumber="181" phylum="Chordata" rank="genus">Mantidactylus</taxonomicName>
a. 
<taxonomicName id="108C3D80FF98FF8A4DFB4DAE4F2879C9" authorityName="Boulenger" authorityYear="1919" box="[450,609,1706,1730]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="68" pageNumber="181" phylum="Chordata" rank="species" species="ambohimitombi">ambohimitombi</taxonomicName>
</emphasis>
in life, in dorsolateral and ventral view, and their habitat. (a) Adult specimen (ACZC 6937; unsexed) from Ankaratra. (b,c) Adult specimen, possibly an adult male due to relatively distinct femoral glands, photographed in 1992. (d,e) Adult male (ACZC 8376; note the well-developed, prominent femoral glands and relatively large tympanum) from Ankaratra. (f) Adult specimen (ACZC 8377; unsexed) from Ankaratra. (g) Adult specimen (ACZC 8109; unsexed) with dorsal stripe from Ankaratra. (h) Pond at high-elevation on the Ankaratra Massif. (i) Landscape at high elevation in the Ankaratra Massif; streams in the valleys are habitat of 
<emphasis id="E5F89A11FF98FF8A4DC14C4F4F897869" box="[504,704,1866,1890]" italics="true" pageId="68" pageNumber="181">
<taxonomicName id="108C3D80FF98FF8A4DC14C4F4FF57869" authorityName="Boulenger" authorityYear="1919" box="[504,700,1866,1890]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="68" pageNumber="181" phylum="Chordata" rank="species" species="ambohimitombi">M. ambohimitombi</taxonomicName>
,
</emphasis>
and in area of rapids, of 
<taxonomicName id="108C3D80FF98FF8A4FF94C4F497F7869" box="[960,1078,1866,1890]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="68" pageNumber="181" phylum="Chordata" rank="species" species="pauliani">
<emphasis id="E5F89A11FF98FF8A4FF94C4F497F7869" box="[960,1078,1866,1890]" italics="true" pageId="68" pageNumber="181">M. pauliani</emphasis>
</taxonomicName>
. (j) Stream in Manjakatompo forest, Ankaratra Massif, habitat of 
<taxonomicName id="108C3D80FF98FF8A4D954C6F4F247889" authorityName="Boulenger" authorityYear="1919" box="[428,621,1898,1922]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="68" pageNumber="181" phylum="Chordata" rank="species" species="ambohimitombi">
<emphasis id="E5F89A11FF98FF8A4D954C6F4F247889" box="[428,621,1898,1922]" italics="true" pageId="68" pageNumber="181">M. ambohimitombi</emphasis>
</taxonomicName>
.
</paragraph>
</caption>
<subSubSection id="9F961588FF99FF8B4C8149484C307CCE" pageId="69" pageNumber="182" type="description">
<paragraph id="D7334603FF99FF8B4C8149484E437D8E" blockId="69.[136,778,171,2085]" pageId="69" pageNumber="182">
<emphasis id="E5F89A11FF99FF8B4C8149484C697D6E" box="[184,288,588,613]" italics="true" pageId="69" pageNumber="182">Variation</emphasis>
.—Variation in measurements is given in 
<tableCitation id="9A0E73B8FF99FF8B4CB1496F4D927D8D" box="[136,219,619,646]" 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="69" pageNumber="182" tableUuid="83F3168BFFE7FFF54CBF4AD74A627D52">Table 5</tableCitation>
. See 
<figureCitation id="4FB75A86FF99FF8B4D2B496F4C167D8E" box="[274,351,619,645]" captionStart="FIGURE 13" captionStartId="68.[136,235,1706,1730]" captionTargetBox="[241,1348,202,1685]" captionTargetId="figure-35@68.[239,1348,202,1685]" captionTargetPageId="68" captionText="FIGURE 13. Mantidactylus a. ambohimitombi in life, in dorsolateral and ventral view, and their habitat. (a)Adult specimen (ACZC 6937; unsexed) fromAnkaratra.(b,c)Adult specimen, possibly an adult male due to relatively distinct femoral glands, photographed in 1992. (d,e) Adult male (ACZC 8376; note the well-developed, prominent femoral glands and relatively large tympanum) from Ankaratra. (f) Adult specimen (ACZC 8377; unsexed) from Ankaratra. (g) Adult specimen (ACZC 8109; unsexed) with dorsal stripe from Ankaratra. (h) Pond at high-elevation on the Ankaratra Massif. (i) Landscape at high elevation in the Ankaratra Massif; streams in the valleys are habitat of M. ambohimitombi, and in area of rapids, of M. pauliani. (j) Stream in Manjakatompo forest, Ankaratra Massif, habitat of M. ambohimitombi." figureDoi="http://doi.org/10.5281/zenodo.7441049" httpUri="https://zenodo.org/record/7441049/files/figure.png" pageId="69" pageNumber="182">Fig. 13</figureCitation>
for colouration in life and its variation.
</paragraph>
<paragraph id="D7334603FF99FF8B4CB1498F4E437C2E" blockId="69.[136,778,171,2085]" pageId="69" pageNumber="182">
A light vertebral stripe occurs in few individuals. There is pronounced sexual size dimorphism (at Ankaratra, largest confirmed male SVL 37.7 vs female SVL 
<quantity id="1074EBE6FF99FF8B4EE049CF4DF17C0E" metricMagnitude="-2" metricUnit="m" metricValue="5.06" pageId="69" pageNumber="182" unit="mm" value="50.6">50.6 mm</quantity>
). As discussed above, the 
<typeStatus id="0837F8A1FF99FF8B4DE749EF4F477C0E" box="[478,526,747,773]" pageId="69" pageNumber="182">type</typeStatus>
series is comprised of particularly large-sized animals reaching 65.0 mm in SVL.
</paragraph>
<paragraph id="D7334603FF99FF8B4CB1482F4C307CCE" blockId="69.[136,778,171,2085]" pageId="69" pageNumber="182">
Femoral glands are only distinct in some male specimens, possibly due to seasonal effects; in 
<specimenCount id="C18A8D8AFF99FF8B4E23484F4F8D7C6E" box="[538,708,843,869]" pageId="69" pageNumber="182" type="generic">two specimens</specimenCount>
, ZSM 190/2021 (ACZCV 334) and ZSM 195/2021 (ACZCV 342), FGL and FGW are 
<quantity id="1074EBE6FF99FF8B4D9B488F4CB17CAE" box="[418,504,907,933]" metricMagnitude="-3" metricUnit="m" metricValue="4.7" pageId="69" pageNumber="182" unit="mm" value="4.7">4.7 mm</quantity>
x 
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and 4.0 mm x 3.0 mm, respectively.
</paragraph>
</subSubSection>
<subSubSection id="9F961588FF99FF8B4C8148CF4C457B8E" pageId="69" pageNumber="182" type="biology_ecology">
<paragraph id="D7334603FF99FF8B4C8148CF4C457B8E" blockId="69.[136,778,171,2085]" pageId="69" pageNumber="182">
<emphasis id="E5F89A11FF99FF8B4C8148CF4C2B7CEE" box="[184,354,971,997]" italics="true" pageId="69" pageNumber="182">Natural history</emphasis>
. 
<emphasis id="E5F89A11FF99FF8B4D5148C84CC87CEE" box="[360,385,972,997]" italics="true" pageId="69" pageNumber="182">—</emphasis>
A common species on theAnkaratra Massif, especially above the tree line along streams and swamp in montane savanna and heatland, but also in rainforest (reported by 
<bibRefCitation id="B31D3BF2FF99FF8B4DB34F2F4F1E7B4E" author="Vences, M. &amp; Andreone, F. &amp; Glaw, F. &amp; Raminosoa, N. &amp; Randrianirina, J. E. &amp; Vieites, D. R." box="[394,599,1067,1093]" pageId="69" pageNumber="182" pagination="263 - 284" refId="ref133786" refString="Vences, M., Andreone, F., Glaw, F., Raminosoa, N., Randrianirina, J. E. &amp; Vieites, D. R. (2002) Amphibians and reptiles of the Ankaratra Massif: reproductive diversity, biogeography and conservation of a montane fauna in Madagascar. Italian Journal of Zoology, 69, 263 - 284. https: // doi. org / 10.1080 / 11250000209356469" type="journal article" year="2002">
Vences 
<emphasis id="E5F89A11FF99FF8B4DD94F284F517B4E" box="[480,536,1067,1093]" italics="true" pageId="69" pageNumber="182">et al.</emphasis>
2002
</bibRefCitation>
under the name 
<taxonomicName id="108C3D80FF99FF8B4CB14F484DB37B6E" box="[136,250,1100,1125]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="curtus">
<emphasis id="E5F89A11FF99FF8B4CB14F484DB37B6E" box="[136,250,1100,1125]" italics="true" pageId="69" pageNumber="182">M. curtus</emphasis>
</taxonomicName>
). Males were found calling during the day underwater.
</paragraph>
</subSubSection>
<subSubSection id="9F961588FF99FF8B4C844F8F4CE978AE" pageId="69" pageNumber="182" type="description">
<paragraph id="D7334603FF99FF8B4C844F8F4DF5784E" blockId="69.[136,778,171,2085]" pageId="69" pageNumber="182">
<emphasis id="E5F89A11FF99FF8B4C844F8F4C537BAE" box="[189,282,1163,1189]" italics="true" pageId="69" pageNumber="182">Calls.—</emphasis>
The advertisement call of 
<emphasis id="E5F89A11FF99FF8B4E8A4F884C727BCE" italics="true" pageId="69" pageNumber="182">
M. a. 
<taxonomicName id="108C3D80FF99FF8B4CB14FAF4C727BCE" authorityName="Boulenger" authorityYear="1919" box="[136,315,1195,1221]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="ambohimitombi">ambohimitombi</taxonomicName>
</emphasis>
, recorded on 
<date id="A33260C3FF99FF8B4DDD4FAF4FF37BCE" box="[484,698,1195,1221]" pageId="69" pageNumber="182" value="2006-02-16">16 February 2006</date>
, 14:50 h, at Ankaratra, consists of a short, pulsed note (
<figureCitation id="4FB75A86FF99FF8B4EE24FCF4DEE7A0E" captionStart="FIGURE 14" captionStartId="70.[136,235,1149,1173]" captionTargetBox="[151,1435,186,1130]" captionTargetId="figure-35@70.[151,1435,185,1130]" captionTargetPageId="70" captionText="FIGURE 14. Audiospectrogram and corresponding oscillogram of one advertisement call of Mantidactylus a. ambohimitombi, recorded on 16 February 2006 at Ankaratra. Recording bandpass-filtered at 220–6000 Hz." figureDoi="http://doi.org/10.5281/zenodo.7441051" httpUri="https://zenodo.org/record/7441051/files/figure.png" pageId="69" pageNumber="182">Fig. 14</figureCitation>
), emitted in series at slow succession and somewhat irregular intervals. The calls were emitted from several shy animals at the border of a mountain stream, partly underwater, and identification of the calling specimen was therefore impossible. Notes exhibit slight amplitude modulation, with maximum call energy occurring either at first third of the note’s length or at the centre of the note, and the terminal pulse of the note always being of lowest energy. The initial pulse is sometimes separated from the second pulse by a slightly longer inter-pulse interval. Numerical parameters of 20 analysed calls are as follows: call duration (= note duration) 136– 218 ms (180.2 ± 18.7 ms); 9–14 pulses per note (10.9 ± 1.5); pulse duration 5–8 ms (6.3 ± 1.0 ms); pulse repetition rate within notes 56.1–69.3 pulses/s (62.3 ± 5.1); dominant frequency 925–1012 Hz (959 ± 30 Hz); prevalent bandwidth 500–3100 Hz; call repetition rate (= note repetition rate) in regular series ca 11–20 calls/ min.
</paragraph>
<paragraph id="D7334603FF99FF8B4C814C4F4CE978AE" blockId="69.[136,778,171,2085]" pageId="69" pageNumber="182">
<emphasis id="E5F89A11FF99FF8B4C814C4F4C6D786E" box="[184,292,1867,1893]" italics="true" pageId="69" pageNumber="182">Tadpoles.</emphasis>
—A tadpole of 
<emphasis id="E5F89A11FF99FF8B4DE24C484F9D786E" box="[475,724,1867,1893]" italics="true" pageId="69" pageNumber="182">
M. a. 
<taxonomicName id="108C3D80FF99FF8B4E1D4C4F4F9D786E" authorityName="Boulenger" authorityYear="1919" box="[548,724,1867,1893]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="ambohimitombi">ambohimitombi</taxonomicName>
</emphasis>
was described under the name ‘ 
<emphasis id="E5F89A11FF99FF8B4D8F4C684C9C788E" box="[438,469,1900,1925]" italics="true" pageId="69" pageNumber="182">M.</emphasis>
sp. 
<taxonomicName id="108C3D80FF99FF8B4E384C6F4F3A788E" authority="Ankaratra" authorityName="Ankaratra" box="[513,627,1899,1925]" class="Amphibia" family="Mantellidae" genus="Mantidactylus" isUncertain="true" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="curtus">
aff. 
<emphasis id="E5F89A11FF99FF8B4E144C684F3A788E" box="[557,627,1900,1925]" italics="true" pageId="69" pageNumber="182">curtus</emphasis>
</taxonomicName>
“Ankaratra”’ by 
<bibRefCitation id="B31D3BF2FF99FF8B4C924C8F4CD578AE" author="Schmidt, H. &amp; Strauss, A. &amp; Glaw, F. &amp; Teschke, M. &amp; Vences, M." box="[171,412,1931,1957]" pageId="69" pageNumber="182" pagination="48 - 60" refId="ref132292" refString="Schmidt, H., Strauss, A., Glaw, F., Teschke, M. &amp; Vences, M. (2009) Description of tadpoles of five frog species in the subgenus Brygoomantis from Madagascar (Mantellidae: Mantidactylus). Zootaxa, 1988, 48 - 60. https: // doi. org / 10.11646 / zootaxa. 1988.1.4" type="journal article" year="2009">
Schmidt 
<emphasis id="E5F89A11FF99FF8B4D364C884C0178AE" box="[271,328,1931,1957]" italics="true" pageId="69" pageNumber="182">et al.</emphasis>
(2009)
</bibRefCitation>
.
</paragraph>
</subSubSection>
<subSubSection id="9F961588FF99FF8B4C814CAF49057E6E" pageId="69" pageNumber="182" type="distribution">
<paragraph id="D7334603FF99FF8B4C814CAF49057E6E" blockId="69.[136,778,171,2085]" lastBlockId="69.[810,1452,171,421]" pageId="69" pageNumber="182">
<emphasis id="E5F89A11FF99FF8B4C814CAF4C2878CE" box="[184,353,1963,1989]" italics="true" pageId="69" pageNumber="182">Distribution.—</emphasis>
Apparently endemic to a small area of the central highlands of 
<collectingCountry id="AF9B0693FF99FF8B4DA84CCF4F5178EE" box="[401,536,1995,2021]" name="Madagascar" pageId="69" pageNumber="182">Madagascar</collectingCountry>
(
<figureCitation id="4FB75A86FF99FF8B4E1F4CCF4F2078EE" box="[550,617,1995,2021]" 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="69" pageNumber="182">Fig. 7</figureCitation>
). The nominal form is known from Ankaratra. Mitochondrial sequences assignable to this lineage have also been recorded from Analafohy and Antoetra; however, confirmation is needed, especially at Antoetra, as to whether this is evidence of true co-occurrence, or is a result of introgression with 
<emphasis id="E5F89A11FF99FF8B49B44BE84E3F7E2E" italics="true" pageId="69" pageNumber="182">
<taxonomicName id="108C3D80FF99FF8B49B44BE84E3A7E2E" class="Amphibia" family="Mantellidae" genus="Mantidactylus" kingdom="Animalia" order="Anura" pageId="69" pageNumber="182" phylum="Chordata" rank="species" species="curtus">M. curtus</taxonomicName>
,
</emphasis>
which is common in this locality. The 
<typeStatus id="0837F8A1FF99FF8B49184A0F48187E2E" box="[1313,1361,267,293]" pageId="69" pageNumber="182">type</typeStatus>
locality Ambohimitombo forest is close to Antoetra. Elevation range: 
<quantity id="1074EBE6FF99FF8B4F4E4A4F49587E6E" box="[887,1041,331,358]" metricMagnitude="3" metricUnit="m" metricValue="1.765" metricValueMax="2.38" metricValueMin="1.15" pageId="69" pageNumber="182" unit="m" value="1765.0" valueMax="2380.0" valueMin="1150.0">1150–2380 m</quantity>
a.s.l.
</paragraph>
</subSubSection>
<subSubSection id="9F961588FF99FF8B4F634A6F491B7EAE" pageId="69" pageNumber="182" type="etymology">
<paragraph id="D7334603FF99FF8B4F634A6F491B7EAE" blockId="69.[810,1452,171,421]" pageId="69" pageNumber="182">
<emphasis id="E5F89A11FF99FF8B4F634A6F4E9B7E8E" box="[858,978,363,389]" italics="true" pageId="69" pageNumber="182">Etymology</emphasis>
.—Formulated from the 
<typeStatus id="0837F8A1FF99FF8B49344A6F48747E8E" box="[1293,1341,363,389]" pageId="69" pageNumber="182">type</typeStatus>
locality, ‘Ambohimitombo Forest’.
</paragraph>
</subSubSection>
</treatment>
</document>