Mantidactylus manerana, Scherz & Crottini & Hutter & Hildenbrand & Andreone & Fulgence & Köhler & Ndriantsoa & Ohler & Preick & Rakotoarison & Rancilhac & Raselimanana & Riemann & Rödel & Rosa & Streicher & Vieites & Köhler & Hofreiter & Glaw & Vences, 2022

Scherz, Mark D., Crottini, Angelica, Hutter, Carl R., Hildenbrand, Andrea, Andreone, Franco, Fulgence, Thio Rosin, Köhler, Gunther, Ndriantsoa, Serge Herilala, Ohler, Annemarie, Preick, Michaela, Rakotoarison, Andolalao, Rancilhac, Loïs, Raselimanana, Achille P., Riemann, Jana C., Rödel, Mark-Oliver, Rosa, Gonçalo M., Streicher, Jeffrey W., Vieites, David R., Köhler, Jörn, Hofreiter, Michael, Glaw, Frank & Vences, Miguel, 2022, 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, Megataxa 7 (2), pp. 113-311 : 287

Mantidactylus manerana sp. nov.

Identity and justification.—A lineage of the M. inaudax clade, differing from M. inaudax by ≥3.5% uncorrected distance in the 16S gene ( Table 2 ), and here considered to represent a distinct species due to the deep divergence in the phylogenomic tree ( Fig. 5 ), and bioacoustics differences ( Table 4 ). Mantidactylus manerana sp. nov. is comprised of three deep mitochondrial lineages, one from Marojejy (Ca16), one from Sorata, and one from Befanjana. These three lineages differ by a strong mitochondrial divergence of 3.1–4.1% uncorrected distance in 16S, but exhibit haplotype sharing in Rag-1, and do not differ substantially morphologically. Bioacoustic data are not available from the Sorata and Befanjana populations to assess whether these lineages are different in call characters.The presence of mitochondrial divergence among these allopatric lineages without substantial differentiation in available phenotypic data and with haplotype sharing in a nuclearencoded gene leads us to recognise them as subspecies, and we in the following assign them the subspecies names M. manerana manerana ssp. nov., M. manerana fotaka ssp. nov., and M. manerana antsanga ssp. nov.

Diagnosis.— Mantidactylus manerana sp. nov. belongs to the M. inaudax clade and, together with two related lineages here considered as subspecies (described below), forms the sister species of M. inaudax , according to our phylogenomic analysis. See Table 4 for a list of diagnostic morphological characters. The combination of small to moderate body size (male SVL 25–38 mm, female SVL 28–38 mm), slightly granular dorsal skin with weakly expressed but clearly recognisable dorsolateral ridges, presence of white spots on flanks, and absence of a white marking on the snout tip, distinguishes M. manerana from species of the M. betsileanus , M. curtus , M. fergusoni , M. tricinctus , and M. ulcerosus clades. M. stelliger ( M. stelliger clade) differs by smaller body size and less developed foot webbing. Species of the M. biporus clade differ as follows: M. biporus by larger body size, absence of dorsolateral ridges, fewer pulses per note and and higher pulse repetition rate in advertisement calls; M. augustini by longer hindlimbs and fewer pulses per note in advertisement calls; M. bletzae by a somewhat smaller body size and more developed foot webbing; M. brevirostris by less developed foot webbing and absence of dorsolateral ridges, and possibly by smaller body size; M. eulenbergeri by slightly smaller body size and smoother dorsal skin; M. glosi by smaller body size and shorter hindlimbs in comparison with most individuals of M. manerana sp. nov., as well as shorter duration of advertisement calls ( Table 4 ). The species is very similar morphologically to its sister species M. inaudax which, however, has less distinct dorsolateral ridges and fewer pulses per note in advertisement calls. For a distinction of the nominal subspecies, M. m. manerana from the other two subspecies recognised, see the diagnoses in the respective subspecies accounts below. A full list of molecular diagnostic sites in the 16S gene of M. manerana sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix.

Distribution.—Occurs on mountains of the North East and in forests in the Northern Central East. This species is known from Ambatobe, Befanjana, Marojejy (from camp 2 upwards, i.e. above 700 m a.s.l.), Sahavontsira, and Sorata (from 1279 m a.s.l. upwards). Elevation range: 14–1599 m a.s.l.

Etymology.—The species name is derived from the Malagasy adjective manerana, meaning ‘observed everywhere’ (in a specific place), and refers to the rather common occurrence of this species on the Marojejy Massif. The species name is used as a noun in apposition.

Figures

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