Maximum Likelihood tree inferred from an alignment of 533 nucleotides of the mitochondrial 16S rRNA gene for 146 samples of the Boophis marojezensis complex and one outgroup (B. picturatus). Main lineages A–H are as discussed in the text and candidate species numbers are assigned to lineages based on previous studies. Numbers at nodes are bootstrap proportions (in percent) and Bayesian posterior probabilities (not shown for some of the most shallow nodes).

Maximum Likelihood tree inferred from an alignment of 533 nucleotides of the mitochondrial 16S rRNA gene for 146 samples of the Boophis marojezensis complex and one outgroup (B. picturatus). Main lineages A–H are as discussed in the text and candidate species numbers are assigned to lineages based on previous studies. Numbers at nodes are bootstrap proportions (in percent) and Bayesian posterior probabilities (not shown for some of the most shallow nodes).

Maximum Likelihood tree inferred from an alignment of 6741 nucleotides of the mitochondrial 12S rRNA and 16S rRNA, ND1, COI and cyt b genes, and the nuclear KIAA1239, RAG-1 and SACS genes for the main lineages of the Boophis marojezensis group. The tree was rooted with B. picturatus (graphically removed to better illustrate branch lengths within the B. marojezensis group. Main lineages A–H of the B. marojezensis complex are as discussed in the text and in Figure 1. Values at nodes are bootstrap support in percent from a partitioned ML analysis with IQ-Tree, and posterior probabilities from an unpartitioned Bayesian Inference analysis with MrBayes.

Haplotype genealogies for three fragments of nuclear-encoded single-copy genes, RAG-1 (474 nucleotides; 37 samples), SACS (960 nt; 36 samples) and KIAA1239 (726 nt; 37 samples). Sequences were phased previous to the analysis, and each sample is therefore represented by two sequences in each network. Coloring of samples is according to their assignment to main mitochondrial lineages in Figure 1. Circles represent haplotypes, with size proportional to allele frequency. Dots on the branches indicate the number of mutational steps between haplotypes.

Map of Madagascar showing the distribution of species of the Boophis marojezensis complex. Only sites confirmed by molecular data are shown. Basemap shows vegetation across Madagascar from the Madagascar Vegetation Mapping Project (Moat and Smith 2007; formerly available at www.vegmad.org). Vegetation is colored as follows: green, humid forest (rainforest); red, western dry deciduous forest; bluish, western subhumid forest; orange, south western dry spiny forest-thicket; yellow, tapia forest.

Comparative spectrograms and oscillograms illustrating representative calls of various species in the Boophis marojezensis complex, all at 4000 ms time scale.

Preserved holotypes in dorsal and ventral views of the eight nominal species in the Boophis marojezensis complex described in this study.

Individuals of Boophis marojezensis in life. A Male holotype (ZFMK 57401) from Marojejy (low elevation). B Male paratype (ZSM 567/1999, previously ZFMK 57402). C Individual from Masoala probably assignable to this species (not sequenced). D Male from Masoala (ZSM 250/2016, FGZC 5439).

Individuals of Boophis kirki sp. nov. in life. A, D Male paratype KU 336967 (CRH 20) from Ranomafana in dorsolateral and ventral view. B, E Male paratype UADBA-CRH 15 (CRH 15) from Ranomafana in dorsolateral and ventral view. C Male paratype KU 336874 (CRH 17) from Ranomafana. F Male paratype KU 336875 (CRH 18) from Ranomafana. G Female (no type status) UADBA-CRH 430 (CRH 430) from Vohidrazana. H Male (no type status) KU 340727 (CRH 449) from Vohidrazana. I, J Male (no type status) KU 340730 (CRH 455) from Vohidrazana in dorsolateral and ventral view. Note the variation in iris color in both populations (Ranomafana and Vohidrazana), with some individual having red color in the outer iris area while others only have light orange color.

Individuals of Boophis picardi sp. nov. in life. A, B male paratype KU 340641 (CRH 275) from Vohimana in dorsolateral and ventral view. C, D male paratypes from An’Ala, not assignable to specific voucher specimens.

Individuals of Boophis pikei sp. nov. in life. A male paratype ZSM 394/2016 (ZCMV 15259) from Marojejy (Camp Simpona). B male holotype ZSM 393/2016 (ZCMV 15251) from Marojejy (Camp Simpona). C male from Marojejy (Camp Simpona), photo not assignable to a specific voucher specimen, photographed in 2016. D, E male paratype ZSM 1514/2012 (FGZC 3732) from Sorata.

Individuals (adult males) of Boophis siskoi sp. nov. in life. A paratype ZSM 10/2016 (MSZC 186) from Ampotsidy. B holotype ZSM 614/2001 (FGMV 2001.64) from the Tsaratanana Massif (Antsahamanara campsite).

Individuals of Boophis janewayae sp. nov. in life. A male holotype ZSM 472/2009 (ZCMV 11468) from Angozongahy campsite, western side of Makira plateau. B calling male (with deflated vocal sac, during a pause between calls) photographed in the field at night, at the type locality.

Individuals of Boophis archeri sp. nov. in life. A, B male holotype ZSM 12/2016 (MSZC 213) from Bevitagnono forest in dorsal and dorsolateral view. C male paratype ZSM 11/2016 (MSZC 198) from Andranonafindra forest. D female paratype, ZSM 13/2016 (MSZC 243) from Andranonafindra forest.

A paratype individual (probably a young male) of Boophis burnhamae sp. nov. in life: ZSM 392/2016 (ZCMV 15171) from Marojejia campsite in the Marojejy Massif.