Bioacoustics of Mantidactylus manerana

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.

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.

Calls. —The advertisement call of M. manerana manerana , recorded from the holotype on 17 November 2016 , 17:20 h, at Camp Simpona, Marojejy National Park, ca 20°C air temperature, consisted of a pulsed note, emitted isolated or in short series ( Fig. 77 ). Notes exhibit amplitude modulation, with call energy constantly increasing from the beginning of the note, reaching its maximum after approximately one third of the note’s duration. Pulse repetition rate within notes was highest at the beginning and decreased towards the note’s end. Numerical parameters of eight analysed calls were as follows: call duration (= note duration) 260–363 ms (292.2 ± 37.4 ms); 26–31 pulses per note (27.9 ± 1.8); pulse duration 3–5 ms (4.5 ± 0.7 ms); pulse repetition rate within notes 62.5–115.9 pulses/s (93.9 ± 15.6); dominant frequency 1452–1539 Hz (1498 ± 30 Hz); prevalent bandwidth 720–5300 Hz; call repetition rate (= note repetition rate) within series ca 30 calls/min. Tadpoles.— The tadpole of M. manerana manerana was described under the name ‘ M. sp. aff. biporus “Marojejy”’ by Schmidt et al. (2009) .