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Mantidactylus kortei sp. nov. Identity and justification.—This lineage has only been found at high elevations of the Andohahela Massif, in a swamp and along streams. In previous DNA barcoding assessments, it has been considered as M. sp. 30 by Vieites et al. (2009) and M. sp. Ca30 by Perl et al. (2014). It is sister to M. noralottae from Isalo in the phylogenomic analysis (but according to 16S data may also be closely related to M. riparius sp. nov. which is not represented in the phylogenomic analysis; see below). However, it differs from M. noralottae in advertisement call and morphology (e.g. smaller body size; Table 4), and from its two siblings by a high genetic divergence (≥4.3% from M. noralottae, and ≥5.9% from M. riparius sp. nov.), supporting its species status. Some individuals of M. kortei sp. nov. are characterized by a relatively short snout, but this is apparently not the case in all individuals. Holotype.— ZSM 205/2005 (FGZC 2376), adult male, collected by P. Bora, F. Glaw, and M. Vences on 26 January 2005 near camp, Andohahela (24.5440°S, 046.7141°E, 1548 m a.s.l.), Anosy Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratypes. —A total of three paratypes: ZSM 203/2005 (FGZC 2377) and ZSM 204/2005 (FGZC 2375), two adult females with the same collection data as the holotype; ZSM 195/2005 (FGZC 2480), an adult female, collected by F. Glaw, M. Vences, and P. Bora on 28 January 2005 in a stream at high elevation, Andohahela (ca 24.544°S, ca 046.714°E, ca 1650 m a.s.l.). Diagnosis.— Mantidactylus kortei sp.nov. is a member of the M. betsileanus clade and related to M. noralottae in the phylogenomic analysis. See Table 4 for a list of diagnostic morphological characters. The combination of a relatively small body size (male SVL 27 mm), slightly tubercular dorsal skin with distinct continuous dorsolateral ridges, relatively large tympanum (13% of SVL in males), presence of a white marking on the snout tip in most specimens, and advertisement call consisting of a single pulsed note not repeated in regular series distinguishes M. kortei sp. nov. from species of all other clades. Species of the M. fergusoni clade are larger and have typically a more tubercular dorsum, while species of the M. curtus clade are often larger and most have a smaller tympanum. Some specimens of the new species have whitish dots on the flanks and only an indistinct white marking on the snout tip, which impedes their distinction from some species of the M. biporus, M. stelliger and M. inaudax clades where advertisement calls are unknown. However, the usually more pointed snout, larger tympanum, longer limbs, and overall different appearance of M. kortei sp. nov. should make a distinction straightforward (Table 4). Within the M. betsileanus clade, the new species can be distinguished from M. betsileanus, M. noralottae and M. tripunctatus by having fewer pulses per note in advertisement calls; furthermore from M. noralottae by smaller body size. Mantidactylus katae has a different advertisement call structure and larger femoral glands; M. jonasi has typically more pulses per note in advertisement calls, a slower pulse repetition rate, and a more tubercular dorsum; and M. incognitus has more expressed dorsal and dorsolateral ridges and supraocular tubercles (Table 4). For a detailed distinction from other new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. kortei sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype. —Adult male in good state of preservation (Fig. 33). Tissue sample taken ventrally from right thigh.Femoral gland partly detached to examine their structure internally. Body rather slender. Head wider than body. Snout rounded in dorsal view, truncate in lateral view. Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis weak, slightly concave. Loreal region slightly concave. Tympanum distinct, large, elliptical, wider than high, its diameter 88% of eye diameter. Supratympanic fold distinct, beginning straight, with a distinct, angular 90° bend midway towards insertion of forelimb. Tongue ovoid, distinctly posteriorly bifid. Maxillary teeth present. Vomerine teeth distinct in elliptical aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Outer metacarpal tubercle present, inner metacarpal tubercle present.Fingers without webbing. Relative length of fingers: I≤II Colour in preservative: dorsum brown, with indistinct irregular darker markings and some white spotting on flanks and with poorly contrasted crossbands on limbs. Loreal region light brown with dark markings. Snout tip with a distinct light dot. Venter beige, throat and chest with brown mottling. Lower lip ventrally with alternating light and brown spots. Colour in life similar to that in preservative, but more contrasted. Variation.—Variation in measurements is given in Table7.See Fig.44 for colouration in life and its variation. There may be pronounced sexual size dimorphism, but sample sizes are low (confirmed male SVL 27.4 mm [n = 1] vs confirmed female SVL 34.6–37.1 mm [n = 3]). Males may have a slightly larger tympanum than females (HTD/ED ratio is 73–80% in females, 88% in the male). Femoral glands in males are relatively prominent, but not prominently coloured in life, at least not in the only male available for examination. Natural history.—Calls of a large number of males were heard during the day, after heavy cyclonic rainfall from a sun-exposed swamp area in grassland directly next to primary rainforest. Several specimens were also found next to small streams in rainforest. Calls. —The advertisement call of M. kortei, recorded on 27 January2005 atAndohahela National Park (specimens not seen calling, therefore attribution of calls not completely certain, but very likely), 17.6°C air temperature (Vences et al. 2006: CD2, track 78), consists of a pulsed note (Fig. 45), emitted in series at irregular intervals and slow succession. Notes exhibited slight amplitude modulation, with maximum call energy occurring either during the first third of the note’s length or at the centre of the note, and the terminal pulse of the note always containing the lowest energy. In some calls, initial pulses were separated by longer inter-pulse intervals, whereas the 3-5 terminal pulses of all calls are narrowly spaced and have very short interpulse intervals. Numerical parameters of nine analysed calls, corresponding to at least two different individuals, are as follows: call duration (= note duration) 254–511 ms (356.9 ± 94.5 ms); 12–27 pulses per note (18.9 ± 4.6); pulse duration 3–6 ms (4.4 ± 0.8 ms); pulse repetition rate within notes (excluding narrowly spaced terminal pulses) 29.0–69.0 pulses/s (47.9 ± 10.1); dominant frequency 1123–1399 Hz (1224 ± 124 Hz); prevalent bandwidth 750–3200 Hz; call repetition rate (= note repetition rate) ca 4–11 calls/min. Tadpoles.— The tadpole of this species has not been described. Distribution.— Apparently microendemic to high elevations in Andohahela National Park (Fig. 7). Elevation range: ~ 1548 m a.s.l. Etymology.—We dedicate this species to Martin Korte, cellular neurobiologist of Braunschweig University of Technology, in recognition of his continued support of our research activities over the past 15 years.

Mantidactylus augustini sp. nov. Identity and justification.—This lineage is a member of the M. biporus clade from the lowlands of the North East of Madagascar and has previously been considered as confirmed candidate species M. sp. 22 by Vieites et al. (2009) and M. sp.Ca22by Perl et al. (2014).It was depicted as ‘ Mantidactylus sp. aff. biporus “Andranofotsy”’ by Glaw and Vences (2007). This lineage is sister to the true M. biporus according to our phylogenomic analysis, but is characterized by a high uncorrected pairwise-distance in the 16S rRNA marker (4.9–5.8%). It also is concordantly differentiated in the nuclear Rag-1 gene, not sharing its haplotype with M. biporus (Fig. 4). Moreover, the two lineages differ distinctly in male advertisement call. Based on the concordance of high mitochondrial divergence with nuclear and bioacoustic differentiation, we are convinced it represents a distinct species. A deep conspecific lineage of M. augustini sp. nov. co-occurs with the main lineage in Masoala. As we have only limited data on this lineage, we tentatively include it in our circumscription of this species, but note that it may later transpire to represent another distinct species. Holotype.— ZSM 122/2002 (MV 2001.1388), adult male, collected by M. Vences on 17 December 2001 at Andranofotsy (wood nearby, 15.4353°S, 049.8439°E, 85 m a.s.l.), Analanjirofo Region, Madagascar. 16S and cox1 barcode sequences of the holotype are available from GenBank (accessions AY848225 and JN133225). Paratypes.—A total of six paratypes: ZSM 740/2009 (ZCMV 11170), possibly female, collected by J.E. Randrianirina on 15 May 2009 at Melivinany ‘S 0I’, Manompana, Forêt de Befanjana (precise coordinates unavailable); MRSN A3600 (FN 7678 = ACZC 4904), adult female, collected by F. Andreone, and J.E. Randrianirina on 1 December 1998 in Beanjada ‘Corridor 1’, Ambatoledama Corridor, Masoala (ca 15.267°S, ca 049.983°E, ca 1000 m a.s.l.); MRSN A2905 (FN 7238 = ACZC 4897), possibly female, collected by J.E. Randrainirina on 6November 1998 at Andranobe, Masoala National Park (coordinates unavailable); MRSN A3540 (ACZC 4898), possibly female, collected by R. Nincheri on 24 July 1993 in Masomihenija forest, Ambodilalono, Masoala peninsula (coordinates unavailable); MRSN A6740 (FAZC 14292 = ACZC 4906), presumed subadult female, collected by J.E. Randrianirina on 27 April 2008 at Farankaraina (coordinates unavailable); MRSN A3737 (FAZC 10009 = ACZC 4905), juvenile, collected by F. Andreone and J.E. Randrianirina on 1 December 1999 at ‘Camp 4’, Antsarahan’Ambarato in the Ilampy Corridor, Masoala peninsula (15.3920°S, 050.0470°E, ca 550 m a.s.l.). Additional material. — ZFMK 70481 from Masoala probably belongs to this species but is not included in the paratype series due to the lack of genetic data. Diagnosis.— M. augustini sp. nov. is a member of the M. biporus clade, sister to M. biporus according to our phylogenomic analysis. See Table 4 for a list of diagnostic morphological characters. The combination of a small body size (male SVL 24 mm, female SVL 21–25 mm), rather smooth dorsal skin with weakly expressed dorsolateral ridges in some individuals, large tympanum size in males (13% of SVL), presence of white spots on flanks, absence of a white marking on the snout tip, and a short, pulsed advertisement call emitted in regular series, distinguishes M. augustini sp. nov. from species of the M. betsileanus, M. curtus, M. fergusoni, M. tricinctus, and M. ulcerosus clades. Mantidactylus inaudax (M. inaudax clade) is morphologically similar but appears to have shorter hindlimbs, less pulses per note, and higher pulse repetition rate in advertisement calls. M. augustini sp. nov. is distinguished from its sister species, M. biporus, by smaller body size, larger tympanum, longer hindlimbs, more pulses per note and a lower pulse rate in advertisement calls (Table 4), as well as a higher dominant frequency (1263–1356 Hz vs 832–997 Hz), and a lower call repetition rate (200–230 vs 300–360 calls/ min). For a distinction from the other (all new) species in the M. biporus, M. stelliger and M. inaudax clades, see the diagnoses in the respective species accounts below. A full list of molecular diagnostic sites in the 16S gene of M. augustini sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype. Adult male in moderate state of preservation (Fig. 62). Left foot missing (taken as tissue sample), femoral glands partly detached for examination in internal view. Body rather stout. Head wider than body. Snout rounded. Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis weak, slightly concave. Loreal region weakly concave.Tympanum distinct, large, rounded, diameter 82% of eye diameter. Supratympanic fold closely following outer edge of tympanum, not clearly recognisable as separate structure in preserved specimen. Tongue ovoid, distinctly posteriorly bifid. Maxillary teeth present. Vomerine teeth present in two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Outer metacarpal tubercle recognisable, inner metacarpal tubercle present. Fingers without webbing. Relative length of fingers: IVariation.—Variation in measurements is given in Table 10. See Fig. 65 for colouration in life and its variation. Data on sexual size dimorphism is inconclusive, and our sample size is small (confirmed male SVL 23.9 mm [n = 1] vs confirmed female SVL 20.9–25.0 mm [n = 3]). There does not seem to be a clear size dimorphism in tympanum diameter (HTD/ED ratio is 68–92% in females, 82% in the male). Femoral glands of the male in life were not very strongly expressed; small gland rudiments are recognisable also in a female (Fig. 65c). Natural history.—At the type locality, calling males were observed at night from the shore of a very shallow running water of a spring in rainforest. ...Continued on the next page ...Continued on the next page ) Calls.—The advertisement call of M. augustini, recorded on 16 December 2001 near Andranofotsy, 25.4°C air temperature (Vences et al. 2006: CD2, track 72), consisted of a short, pulsed note, emitted in regular series at fast succession (Fig. 66). Pulse repetition rate was distinctly lower at the beginning of calls and increased after approximately one half of the call’s duration. Amplitude modulation was present, with relative amplitude increasing from the beginning of the call, reaching its maximum at the last quarter of the call’s duration. Numerical parameters of seven analysed calls were as follows: call duration (= note duration) 154–236 ms (178.8 ± 26.6 ms); 9–15 pulses per note (10.4 ± 2.1); pulse duration 7–12 ms (9.1 ± 1.6); pulse repetition rate within notes 44.4–71.4 pulses/s (59.0 ± 10.3); dominant frequency 1263–1356 Hz (1304 ± 39 Hz); prevalent bandwidth 700–3150 Hz; call repetition rate (= note repetition rate) within regular series ca 200– 230 calls/min. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to the North East of Madagascar (Fig. 7). This species is known from Masoala, Manompana (Befanjana), Antsahataloka, and Andranofotsy. Elevation range: Ranging from near sea level (85 m a.s.l.) to ~ 1000 m a.s.l. Etymology.—We dedicate this species to Augustin Sarovy, an excellent musician, guide, and ecologist from Maroantsetra, whose help was crucial to collect the holotype of this new species and to record its call., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 265-271, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 496 pp. Third Edition.","Vences, M., Glaw, F. & Marquez, R. (2006) The Calls of the Frogs of Madagascar. 3 Audio CD's and booklet. Madrid, Spain, Fonoteca Zoologica, 44 pp."]}

Mantidactylus bletzae sp. nov. Identity and justification.—This lineage, known from two high-elevation sites in the Southern Central East and South East of Madagascar, was newly identified in this study. It resembles species of the M. biporus and M. inaudax clades phenotypically. By the presence of dorsolateral ridges, it resembles some species of the M. inaudax clade, but in the 16S tree, it is placed more closely to species of the M. biporus clade, although its relationships are not reliably resolved. We here consider it tentatively as a member of the M. biporus clade, pending its inclusion in a future phylogenomic analysis. The new species is characterized by a high mitochondrial divergence (≥5.7% from all other species; closest species M. inaudax bona species and M. madecassus), and an isolated position in the mitochondrial tree, without obvious close relationships to any other species. It also is concordantly differentiated in the nuclear Rag-1 gene with a unique haplotype (Fig. 4). Based on the concordance of high mitochondrial divergence with nuclear differentiation, we are convinced it represents a distinct species. Holotype.— ZSM 829/2014 (ZCMV 14771), adult female, collected by A. Rakotoarison, M. Bletz, D. Edmonds, and F. Randrianasolo on 11 November 2014 at Pic d’Ivohibe Special Reserve, Camp 3 (22.49710°S, 046.95758°E, 1566 m a.s.l.), Ihorombe Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratypes.—A total of two paratypes: ZSM 827/2014 (ZCMV 14763) and ZSM 828/2014 (ZCMV 14768), two adult females, with the same collection data as the holotype. Specimens from Ranomafana are not included in the paratype series as they were not examined morphologically and are genetically divergent. Diagnosis.— Mantidactylus bletzae sp. nov. is considered to be a member of the M. biporus clade based on affinities in the 16S tree. Its precise relationships remain unclarified, as it was missing from our phylogenomic analysis. See Table 4 for a list of diagnostic morphological characters. The combination of a small body size (female SVL 26–27 mm), slightly granular dorsal skin with weakly expressed but distinctly recognisable dorsolateral ridges, presence of white spots on flanks, and absence of a white marking on the snout tip distinguishes M. bletzae sp. nov. from species of the M. betsileanus, M. curtus, M. fergusoni, M. tricinctus, and M. ulcerosus clades. Mantidactylus inaudax (M. inaudax clade) is morphologically similar but appears to have a somewhat larger body size, no dorsolateral ridges, shorter hindlimbs, and less developed foot webbing; M. biporus has a larger body size, lacks dorsolateral ridges, and has less developed foot webbing; M. augustini has somewhat longer hindlimbs, somewhat less granular dorsal skin and less clearly expressed dorsolateral ridges, and less developed foot webbing (Table 4). For a distinction from the other new species in the M. biporus, M. stelliger and M. inaudax clades, see the diagnoses in the respective species accounts below. A full list of molecular diagnostic sites in the 16S gene of M. bletzae sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype.—Adult female in excellent state of preservation (Fig. 62). Tongue has been excised as tissue sample.Head as wide as body.Snout rounded in dorsal view. Nostrils directed laterally, not protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis weakly expressed, slightly concave. Loreal region concave. Tympanum distinct, small, rounded, its horizontal diameter about 71% of eye diameter. Supratympanic fold present, beginning straight, bending abruptly midway towards jaw / forelimb insertion. Maxillary teeth present. Vomerine teeth present in two small rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: I Colour in preservative: dorsum light brown with a quite distinct dark brown pattern, including dark longitudinal stripes running ventral of the dorsolateral folds, some dark patches in the vertebral region, and a broad dark band between the eyes. Fore- and hindlimbs with distinct dark brown crossbands. Ventrally whitish on belly and beige on limbs, with weak dark mottling on throat and chest, and lower jaw ventrally bordered by an interrupted dark line. The colour in life was not recorded. Variation.—Variation in measurements is given in Table 10. See Fig. 67 for colouration in life and its variation. No male specimens available to assess sexual dimorphism. Natural history.—Poorly known. Specimens were found in a swampy area near a small pool. Calls.— The call of this species has not been recorded. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to Southern Central East (Fig. 7). This species is known from Pic d’Ivohibe Special Reserve (Camp 3, at high elevation), and Maharira in Ranomafana National Park. Elevation range: 1248–1575 m a.s.l. Etymology.—We dedicate this species to Molly C. Bletz, who contributed to collecting the type specimens from Pic d’Ivohibe, in recognition of her substantial contributions to amphibian conservation and research in Madagascar.

Mantidactylus curtus clade This clade comprises several small to large-sized species (23.4–65.0 mm adult SVL) typically characterized by a relatively short snout and short hindlimbs, sometimes a weakly developed frenal stripe (very rare in other Brygoomantis), and occurrences either in the Central Region or in the western regions of Madagascar (South, West and North-West), including several montane species: Mantidactylus curtus, M. alutus, M. ambohimitombi, M. bourgati, M. madecassus, M. pauliani, as well as one new species and two new subspecies named herein (holotypes are depicted in Fig.9). The M.curtus clade was particularly difficult to revise because the calls of most species are— even by the standards of Brygoomantis —highly inconspicuous and rarely heard, and differences between males and females are not easy to recognise externally (males often have rather indistinct femoral glands, not too different from the gland rudiments of females). Two exceptions to this are M. alutus, a small-sized species with distinct femoral glands in males and distinct calls, and M. mahery sp. nov., a widespread species in the West, reaching into the North West and into the South (Isalo). Furthermore, our analysis with the Phylonetworks package suggested two instances of reticulated evolution in this clade (see above), which further complicates taxonomic inference., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on page 166, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023

Mantidactylus riparius sp. nov. Identity and justification.—This lineage of the M. betsileanus clade was first discovered by Cocca et al. (2018) and named ‘ Mantidactylus sp. aff. multiplicatus Ca 65 “Isalo”’. It was not included in earlier DNA barcoding assessments of Madagascar’s anuran diversity. It represents the third Brygoomantis species occurring in the Isalo massif, besides M. mahery and M. noralottae. Both this lineage and M. noralottae have so far only been recorded from Isalo and belong to the M. betsileanus clade according to the 16S tree. Mantidactylus kortei appears also to belong to this clade, and is morphologically and bioacoustically similar to this lineage. However, we here consider the Isalo lineage as a separate species from M. kortei due to its high genetic divergence of 5.9–6.8% in the 16S gene, absence of Rag-1 haplotype sharing and ecological divergence (found in canyons in the dry Isalo sandstone massif, vs M. kortei occurring only on high elevations in humid rainforest of Andohahela). Holotype.— ZSM 2403/2007 (ZCMV 5766), adult male, collected by L. du Preez, C. Weldon, O. Verneau, and L. Raharivololoniaina on 16 February 2007 at Isalo (Cascade des Nymphes), Ihorombe Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratypes.—A total of eight paratypes: ZSM 186/2021 (ACZCV 281, extraction ACP 2294, tissue ACZC 6908) and ZSM 187/2021 (ACZCV 283, ACP 2296, ACZC 6911), two probable females, collected on 25 November 2014 by A. Crottini, G.M. Rosa and F. Andreone at the Isalo Massif (Andriamanero: Antsifotra canyon); UADBA uncatalogued (ZCMV 5541 – 5544, ZCMV 5749, ZCMV 5775), six specimens of unkonwn sex and maturity, collected by L. du Preez, C. Weldon, O. Verneau, and L. Raharivololoniaina in February 2007 in the Isalo Massif. Diagnosis.— Mantidactylus riparius sp. nov. is a member of the M. betsileanus clade and related to M. noralottae and M.kortei based on the 16S tree (not included in the phylogenomic analysis). See Table 4 for a list of diagnostic morphological characters. The combination of a relatively small body size (male SVL 27 mm), slightly tubercular dorsal skin, relatively large tympanum (13% of SVLin males), and advertisement call consisting of a single pulsed note not repeated in regular series distinguishes M. riparius sp. nov. from species of all other clades. Species of the M. fergusoni clade are larger and have typically a more tubercular dorsum, while species of the M. curtus clade are often larger and most have a smaller tympanum. Some specimens of the new species have whitish dots on the flanks and most have only an indistinct white marking on the snout tip, which impedes their distinction from some species of the M. biporus, M. stelliger and M. inaudax clades where advertisement calls are unknown. However, the usually more pointed snout, larger tympanum, longer limbs, and overall different appearance of M. riparius sp. nov. should make a distinction straightforward (Table 4). Within the M. betsileanus clade, the new species can be distinguished from M. betsileanus, M. noralottae and M. tripunctatus by having fewer pulses per note in advertisement calls; furthermore from M. noralottae by smaller body size. Mantidactylus katae has a different advertisement call structure and larger femoral glands; M. jonasi has typically more pulses per note in advertisement calls, a lower pulse repetition rate, and a more tubercular dorsum; M. incognitus has more expressed dorsal and dorsolateral ridges and supraocular tubercles (Table 4). The new species is most similar to the allopatric M. kortei from which it cannot be reliably distinguished by morphology or calls, despite a tendency of a faster pulse rate in advertisement calls which however might be influenced by temperature (Table 4). For a detailed distinction from other new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. riparius sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype.—Adult male, in good state of preservation except for a large part of the right thigh excised for tissue sampling, and belly cut open (with some inner organs including bladder removed for parasite sampling) (Fig. 33). Body slender. Head slightly wider than body. Snout rounded in lateral view, slightly pointed in dorsal view. Nostrils directed laterally, slightly protuberant, nearer to tip of snout than to eye. Canthus rostralis straight; loreal region concave. Tympanum distinct and rather large, wider than high, horizontal diameter of tympanum 82% of horizontal eye diameter. Supratympanic fold distinct, following the outer edge of the tympanum, regularly curved. Tongue ovoid, bifid. Maxillary teeth present. Vomerine teeth form two small roundedaggregations,positionedposterolateraltochoanae. Choanae small and rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: IVariation.—Variation in measurements is given in Table 7. See Fig. 46 for colouration in life and its variation. Given the small sample sizes of measured individuals, an assessment of sexual dimorphism is not possible. Femoral glands in males are relatively weakly expressed and not conspicuously coloured in life. Natural history.—Found along semi-permanent streams and in natural pools of oasis at Isalo sandstone massif. It is a relatively shy species that hides in the crevices of the rocks. The species is found in syntopy with both Mantidactylus mahery and M. noralottae. Calls.— The advertisement call of M. riparius (FAZC 14746; ACP4528), recorded on 12 February 2011, at Isalo (Andriamanero), unknown air temperature, consisted of a pulsed note (Fig. 47) of variable duration, emitted in somewhat irregular series. Notes exhibited amplitude modulation, with call energy rapidly increasing from the beginning of the note, reaching its maximum after approximately one tenth of the note’s duration, continuously decreasing afterwards. Pulse repetition rate within notes was highest at the beginning and decreases towards the note’s end. Call energy was distributed in a wide frequency band. Numerical parameters of 12 analysed calls were as follows: call duration (= note duration) 249–697 ms (348.4 ± 139.2 ms); 15–41 pulses per note (21.0 ± 8.1); pulse duration 2–5 ms (3.8 ± 1.3 ms); pulse repetition rate within notes 49.2–114.3 pulses/s (74.0 ± 27.5); dominant frequency 1518–1574 Hz (1549 ± 28 Hz); prevalent bandwidth 950–7400 Hz; call repetition rate (= note repetition rate) within series ca 5–9 calls/ min. Tadpoles.— The tadpole of this species has not been described. Distribution.— Apparently microendemic to the Isalo massif (Fig. 7). Elevation range: 640–920 m a.s.l. Etymology.—The Latin adjective riparius, meaning ‘inhabiting the banks of rivers’, making reference to the preferred microhabitat of this (and other) Brygoomantis species., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 235-237, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Cocca, W., Rosa, G. M., Andreone, F., Aprea, G., Bergo, P. E., Mattioli, F., Mercurio, V., Randrianirina, J. E., Rosado, D., Vences, M. & Crottini, A. (2018) The herpetofauna (Amphibia, Crocodylia, Squamata, Testudines) of the Isalo Massif, Southwest Madagascar: combining morphological, molecular and museum data. Salamandra, 54, 178 - 200."]}

Mantidactylus noralottae Mercurio & Andreone, 2007 Type material.— The species is based on holotype (by original designation) MRSN A5317 from ‘ Ambovo, Parc National de l’Isalo, Fianarantsoa Faritany, Ranohira Fivondronana, 22°30.48′S, 45°21.15′E, 996 m a.s.l. ’. A total of 12 paratypes were defined in the original description: MRSN A5036 (FAZC 13021), MRSN A5035 (FAZC 13020), MRSN A5254 (FAZC 13008), MRSN A5318 (FAZC 13024), SMF 85861 (ex MRSN A5253 / FAZC 13007), SMF 85862–85864 (ex MRSN A5255– 5257 / FAZC 13011–13013), MRSN A5252 (FAZC 13005), PBZT-FAZC 12996, PBZT-FAZC 12998, and ZSM 49/2011 (ex MRSN A5319 / FAZC 13022), all with the same locality, date and collector as the holotype. Identity.—This species is genetically defined by the sequences of various paratypes published in the original description (MRSN A5252 and A5254; SMF 85861‒SMF 85864 corresponding to previous MRSN A5253 and A5255‒A5257; Mercurio & Andreone 2007). Unfortunately, no genetic data are available for the holotype or the call voucher paratype, MRSN A5317. However, both the holotype and the call voucher are comparatively large-sized individuals of 34.8 and 33.4 mm SVL, respectively, and thus distinctly larger than the second species of the M. betsileanus clade occurring at Isalo (described below as M. riparius sp. nov.). Furthermore, according to measurements in Mercurio & Andreone (2007) and here reproduced in Table 7, the male holotype of M. noralottae has a smaller relative tympanum size and smaller femoral glands than the third species of Brygoomantis at Isalo, M. mahery (see above and Tables 4‒5), confirming the holotype of M. noralottae is conspecific with the paratypes and other individuals usually assigned to this taxon. Diagnosis.—A member of the M. betsileanus clade as revealed by the phylogenomic analysis, sister to M. kortei sp. nov. described below. See Table 4 for a list of diagnostic morphological characters. The combination of a moderate body size in males (SVL 33–36 mm) and distinctly larger size in females (SVL 36–40 mm), rather smooth to slightly tubercular dorsal skin with distinct continuous dorsolateral ridges, relatively large tympanum (10–12% of SVL), absence of white spots on flanks, absence of a white marking on snout tip, and advertisement call consisting of a single, long note composed of ≥90 pulses distinguishes M. noralottae from species of all other clades (Table 4); M. noralottae may appear superficially similar to some species of the M. curtus clade but these have a smaller tympanum and less pulses in advertisement calls. Within the M. betsileanus clade, the species differs from all species except M. betsileanus (for M. incognitus sp. nov. calls are unknown) by a higher number of pulses in advertisement calls (Table 4); it differs from these two species by larger body size of males, and from M. betsileanus also by fewer pulses in advertisement calls (Table 4). For a detailed distinction from its sister species M. kortei sp. nov., from the sympatric M. riparius sp. nov., and from all other new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. noralottae in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table 7. See Fig. 36 for colouration in life and its variation. A light vertebral line can be present. There is moderate sexual size dimorphism (confirmed male SVL 32.8–35.5 mm [n = 4] vs confirmed female SVL 35.5– 40.0 mm [n = 7]). Tympanum size is somewhat larger in males compared to females (HTD/ED ratio is 56–63% in females, 63–95% in males). Femoral glands in males are not very prominent and not conspicuously coloured. Natural history.—According to Mercurio and Andreone (2007) the species inhabits canyons in the Isalo limestone massif, and can be found from the initial openings all the way to their deep end (eg. Anjofo waterfall). Individuals can climb on rocks and cling at 150– 200 cm height above the water or the ground. Mercurio and Andreone (2007) also provide some information on stomach content, according to which the species feeds on different groups of insects. Although the species cooccurs with M. noralottae at Isalo, the two species have so far not been found in the same streams or at exactly the same sites in this massif. Calls.—The advertisement call of M. noralottae, recorded on 18 December 2004, 20:00 h, at Ambovo, Isalo National Park, 20°C air temperature (from paratype MRSN A5319), consisted of a very long, regularly pulsed note (Fig. 37), emitted in irregular series. Each note showed some significant amplitude modulation with call energy slowly increasing to approximately the middle of the note’s duration and then slowly decreasing towards its end. Numerical parameters of five analysed calls were as follows: call duration (= note duration) 2054–2705 ms (2411.4 ± 273.9 ms); ca 92–108 pulses per note (100.6 ± 7.1); pulse duration 9–15 ms (10.9 ± 1.6 ms); pulse repetition rate within notes 37.4–41.2 pulses/s (39.7 ± 1.3); dominant frequency 1345–1405 Hz (1370 ± 26 Hz); prevalent bandwidth 1200–2100 Hz; call repetition rate (= note repetition rate) ca 12 calls/min. Tadpoles.— The tadpole of this species has not been described. ...Continued on the next page ...Continued on the next page ...Continued on the next page Distribution.— Apparently microendemic to the Isalo massif (Fig. 7). Elevation range: 640–996 m a.s.l. Etymology.—Eponym for Nora Lotta Mercurio née Fr̂hder, wife of V. Mercurio, one of the authors of the original description., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 217-223, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Mercurio, V. & Andreone, F. (2007) Two new canyon-dwelling frogs from the arid sandstone Isalo Massif, central-southern Madagascar (Mantellidae, Mantellinae). Zootaxa, 1574, 31 - 47. https: // doi. org / 10.11646 / zootaxa. 1574.1.2"]}

Mantidactylus eulenbergeri sp. nov. Identity and justification.—This lineage is a member of the M. biporus clade and has been considered as confirmed candidate species M. sp. 23 by Vieites et al. (2009), and M. sp. Ca23 by Perl et al. (2014). It was depicted as ‘ Mantidactylus sp. aff. biporus “Andasibe”’ by Glaw and Vences (2007). It shows a rather distinctive morphology with a very short snout in at least some specimens, and differs from other lineages of the M. biporus clade by concordant divergence in 16S and Rag-1.According to the phylogenomic analysis, it represents the sister taxon of M. brevirostris but differs from that lineage by a 16S distance of 8.6–9.1%, and possibly by a difference in foot webbing (Table 4). We consider the available evidence sufficient to assign a status of separate species to this lineage. Holotype.— ZSM 85/2002 (field number MV 2001.1092), adult male, collected by M. Vences on 23–25 November 2001 at Andasibe (18.9333°S, 048.4167°E, 915 m a.s.l.), Alaotra-Mangoro Region, Madagascar. 16S and cox1 barcode sequences of the holotype are available from GenBank (accessions AY848239 and JN133224). Paratypes.—A total of seven paratypes: ZSM 84/2002 (MV 2001.1090), adult female, with same collection data as the holotype; ZSM 919/2003 (FGMV 2002.949), putative female, collected by G. Aprea and collaborators on 20 February 2003 in Vohidrazana; ZSM 198/2021 (FAZC 15509, extraction ACP 3664, tissue ACZC 8596), ZSM 199/2021 (FAZC 15516, ACP 3671, ACZC 8603), MRSN A7047 (FAZC 15517, ACP 3672, ACZC 8604), MRSN A7048 (FAZC 15549, ACP 3702, ACZC 8636), MRSN A7049 (FAZC 15540, ACP 3693, ACZC 8627), all collected in January 2017 at Maromizaha (18.9713°S, 048.4642°E) by E. Coppola. Additional material.—The following specimens (without genetic data) are tentatively assigned to this species: ZFMK 52674–52675, collected by F. Glaw and M. Vences in February 1991 and ZFMK 62214 collected by F. Glaw on 1 February 1996 (all from Andasibe). Diagnosis.— Mantidactylus eulenbergeri sp. nov. is a member of the M. biporus clade, sister to M. brevirostris according to our phylogenomic analysis. See Table 4 for a list of diagnostic morphological characters. The combination of small body size (male SVL 20–23 mm, female SVL 25–28 mm), smooth dorsal skin with weakly expressed dorsolateral ridges sometimes recognisable, large tympanum size in males (12–14% of SVL), presence of (sometimes only few) white spots on flanks, and absence of a white marking on the snout tip, distinguishes M. eulenbergeri sp. nov. from species of the M. betsileanus, M. curtus, M. fergusoni, M. tricinctus, and M. ulcerosus clades. M. inaudax (M. inaudax clade) is morphologically similar but differs by larger body size; M. biporus has a larger body size; M. augustini has longer hindlimbs; M. bletzae has a more granular dorsal skin with dorsolateral ridges; M. brevirostris has a less developed foot webbing (Table 4). For a distinction from other new species in the M. biporus, M. stelliger and M. inaudax clades, see the diagnoses in the respective species accounts below. A full list of molecular diagnostic sites in the 16S gene of M. eulenbergeri sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype. —Adult male in moderate state of preservation (Fig. 62). Tissue sample removed from right thigh. Body stout. Head as wide as body. Snout very short and rounded in dorsal and lateral views. Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis weak, slightly concave. Loreal region weakly concave. Tympanum distinct, large, rounded, diameter 91% of eye diameter. Supratympanic fold indistinct, following exactly the outline of the large tympanum. Tongue ovoid, distinctly posteriorly bifid. Maxillary teeth present. Vomerine teeth present in two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Outer metacarpal tubercle recognisable, inner metacarpal tubercle present. Fingers without webbing. Relative length of fingers: I Nuptial pads absent. Foot longer than tibia (114%). Lateral metatarsalia separated. Inner metatarsal tubercle present. Outer metatarsal tubercle not present. Webbing formula: 1(1), 2i(1.5), 2e(1), 3i(2), 3e(1), 4i(2.5), 4e(2.5), 5(1). Relative length of toes: I Colour in preservative: dorsum brown, with distinct irregular darker markings. A dark brown band between eyes present. Forelimbs brown with distinct darker markings. Hindlimbs brown with distinct darker crossbands. Inguinal region without whitish spots. Snout tip without a light dot. Venter beige, throat darker than belly. Lower lip with distinct alternating light and brown spots. Toe discs dark. Toes light and dark striped. Colour in life as in preservative but more vibrant (Fig. 69). Variation.—Variation in measurements is given in Table 10. There is pronounced sexual size dimorphism (confirmed male SVL 20.0– 23.3 mm [n = 6] vs confirmed female SVL25.0–28.0mm [n = 4]). Horizontal tympanum diameter is 73–86% of eye diameter in males and 60–78% of eye diameter in females. Skin on the back is smooth. Colour on the back brown with few indistinct markings (e.g. ZSM 84/2002). Few white spots on the flanks are always present. Two dark spots on the back at level of forelimb insertion are present only in ZSM 84/2002. A light interrupted vertebral line is present in ZFMK 62214, ending on the snout tip with a distinct white dot. A light vertebral band is never present. A dark brown and more or less triangular band between eyes is always present. Lower lip with more (e.g. ZSM 85/2002) or less (e.g. ZSM 84/2002) distinct alternating light and brown spots. Venter and throat uniformly beige, in ZFMK 62214 with little white spots. A longitudinal white median line on thorax and throat is present in ZFMK 62214 and very faintly in ZSM 84/2002. Forelimbs brown with irregular darker markings and stripes. Femoral glands of adult males are large and prominent with one indistinct spot on the femoral gland as a small side structure proximal to the cloaca in ZSM 85/2002, with two distinct spots in ZFMK 62214. In females femoral glands are small but can be recognised (e.g. ZSM 84/2002), with two gland rudiments of the same size on each shank. Natural history.—A species found along small and shallow running water bodies in rainforest. Calls.—The call of this species has not been recorded Tadpoles.—The tadpole of this species has not been described. Distribution.— Endemic to the Northern Central East (Fig. 7). This species is known from Anivorano Est, Sahafina, Andasibe, Maromizaha, and Vohidrazana. Elevation range: 60–1100 m a.s.l. Etymology.—We dedicate this species to Klaus Eulenberger, former chief veterinary of Leipzig Zoo, in recognition of his contributions to knowledge on husbandry and veterinary care of captive amphibians and reptiles.

Mantidactylus ambohimitombi miloko ssp. nov. Identity and justification.—This lineage of the M. curtus clade was considered as confirmed candidate species M. sp. 18 by Vieites et al. (2009) due to its mitochondrial divergence in concert with slight but distinct differences in colour pattern, and as M. sp. Ca18 by Perl et al. (2014). It was depicted as ‘ Mantidactylus sp. aff. curtus “Ambohitantely”’ by Glaw and Vences (2007). We here consider this lineage provisionally as a subspecies of M. ambohimitombi based on the following rationale: (i) it belongs to the same general mitochondrial lineage as the nominal subspecies in the 16S tree, and the two are also closely related in the phylogenomic tree; (ii) morphologically, the two lineages are similar to each other, except for a somewhat more distinct dorsal pattern in M. a. miloko ssp. nov.; (iii) based on two samples in our mitochondrial tree, RJS 1877 and ACZC 4254, a very similar mitochondrial haplotype to the one from Ambohitantely also occurs at Ankaratra, suggesting the possibility of past or ongoing gene flow between localities or mitochondrial introgression;and (iv)based on sequences of specimens APR 10803, APR 10638, and APR 10663, the lineage also occurs in Angavokely and Ankazomivady, thus rather close to Ankaratra, suggesting the two lineages may be parapatric and could hypothetically have a hybrid zone. Since the status of the Ambohitantely population as a fully isolated evolutionary lineage is thus not fully verifiable with the data at hand, we consider the status as a subspecies of M. ambohimitombi to be adequate. Holotype.— ZSM 219/2005 (FGZC 2143), adult male, collected by M. Vences, L. du Preez, P. Bora, L. Raharivololoniaina, R.D. Randrianiaina, T. Razafindraibe, E. Randriamitso on 18 January 2005 at Ambohitantely Special Reserve, ‘Jardin Botanique’, at a site about 500 m fromthegeographicalcoordinates 18.1725°S, 047.2768°E, 1580 m a.s.l., Analamanga Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratype.—A single paratype: ZSM 237/2005 (FGZC 2172), adult female, with the same collection data as holotype. Additional material. —The following specimens probably belong to this taxon but are only assigned tentatively and not designated as paratypes because no molecular data is available: ZMA 6859 (two specimens with field numbers 675 and 676) and ZMA 6860 (six specimens with field numbers 910–914, 917), collected by R.M.A. Blommers-Schl ̂sser in 1972 at 1500 m (ZMA 6859) and 2200 m a.s.l. (ZMA 6860) at Tampoketsa d’Ankazobe. Diagnosis.— Mantidactylus ambohimitombi miloko is a member of the M. curtus clade, and the sister group of M. a. ambohimitombi + M. a. marefo. Morphologically, it is very similar to M. a. ambohimitombi. See Table 4 for a list of diagnostic morphological characters. The combination of relatively large body size of up to 49 mm, slightly granular skin without clearly defined dorsolateral ridges, and small tympanum diameter of a maximum of 10% of SVL in males, distinguishes M. a. miloko ssp. nov. from species of the other clades. Within the M. curtus clade, M. alutus, M. madecassus and M. pauliani have smaller body sizes and are distinguished by either a usually shorter snout (M. madecassus, M. pauliani, M. a. marefo), or presence of rather distinct dorsolateral ridges (M. alutus) (Table 4); M. curtus usually has a smoother skin and a somewhat shorter snout; M. bourgati is morphologically very similar but appears to occur only on the Andringitra Massif. Compared to other subspecies of M. ambohimitombi, the new subspecies differs from M. a. marefo by its distinct dorsal pattern, lack of bluish colour around the eye, and more pointed snout; and from M. a. ambohimitombi by the usually more distinct dorsal pattern. For detailed distinction from other new species and subspecies described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. a. miloko in pairwise comparisons to all other Brygoomantis species and subspecies is provided as Supplementary appendix. Description of the holotype.—Adult male in good state of preservation (Fig. 9). Tissue sample taken ventrally from right thigh. Femoral gland partly detached to examine their structure internally. Body rather slender. Head slightly wider than body. Snout rounded in dorsal and lateral view. Nostrils directed dorsolaterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis almost not recognisable, slightly concave. Loreal region slightly concave. Tympanum distinct, elliptical, wider than high, its diameter 78% of eye diameter. Supratympanic fold distinct, beginning straight, with a distinct, angular 90° bend at the posterior edge of tympanum towards insertion of forelimb. Tongue ovoid, distinctly posteriorly bifid. Maxillary teeth present. Vomerine teeth distinct in rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Outer metacarpal tubercle present, inner metacarpal tubercle present. Fingers without webbing. Relative length of fingers: I=IIVariation. —Variation in measurements is given in Table 5. Too few specimens have been sexed to assess the degree of sexual size dimorphism. Femoral glands distinct and large (but not differing in colour from surrounding ventral skin of thigh) in the male holotype (Fig. 16). In contrast, specimens from the ZMA collection are difficult to sex externally as femoral glands are often indistinct, as seems to be typical for several taxa in the M. curtus clade, possibly due to seasonal effects. Natural history.— Specimens were collected around clean highland streams running in open areas between forest fragments. Specimens at Angavokely and Ankazomivady that appear to belong to this taxon based on mitochondrial DNA were found on wet rocks along slow-moving parts of streams. Calls.—The call of this subspecies has not been recorded. Tadpoles.— Probably reported from Angavokely by Blommers-Schl̂sser (1979).. Distribution.— Endemic to the central highlands of Madagascar, north of the distribution of the nominal subspecies (Fig. 7). This subspecies is currently known from Ambohitantely, and probably also from Angavokely, and Ankazomivady. A mitochondrial haplotype corresponding to this subspecies has also been detected at Ankaratra, but due to the limited information on the sampling event and the absence of specimens (only tissue sampels were collected) this record (which might also represent mitochondrial introgression) requires confirmation. Elevation range: 1520–1735 m a.s.l. Etymology.—The subspecies name is derived from Malagasy word miloko, meaning ‘painted’, referring to the rather distinct dorsal pattern of well-delimited dark blotches characterizing this subspecies. The subspecies name is used as a noun in apposition., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 185-186, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 496 pp. Third Edition."]}

Mantidactylus gudrunae sp. nov. Identity and justification.—This lineage has been considered as confirmed candidate species M. sp. 7 by Vieites et al. (2009) and M. sp. Ca7 by Perl et al. (2014). It is a member of the M. tricinctus clade, and strongly differs from the two other lineages in the clade (M. tricinctus and M. grubenmanni) by concordant strong divergence in 16S and Rag-1 sequences. Furthermore, it also differs in various morphological features (see Diagnosis below). We are therefore confident that this lineage represents a distinct, evolutionarily isolated separate species. Holotype.— ZSM 146/2004 (field number FGZC 274), adult male, collected by F. Glaw, M. Puente, R.D. Randrianiaina, and M. Teschke (née Thomas) on 7 February 2004 at Manantantely (24.983°S, 046.917°E, 20–150 m a.s.l.), Anosy Region, Madagascar. 16S and cox1 barcode sequences of the holotype are available from GenBank (accessions AY848141 and JN133257). Paratypes.—A total of six paratypes: ZSM 136/2004 (FGZC 250), ZSM 138/2004 (FGZC 259), two adult males, and ZSM 154/2004 (FGZC 286), adult female, with the same collection data as the holotype (7–8 February 2004); ZSM 68/2004 (FGZC 115), adult female, collected by F. Glaw, M. Puente, M. Teschke (née Thomas), and R. Randrianiaina on 29–31 January 2004 at ‘Camp 1’, between Isaka and Eminiminy, Andohahela National Park (24.7586°S, 046.8542°E, 247 m a.s.l.); ZSM 95/2004 (FGZC 167), adult male, and ZSM 96/2004 (FGZC 168), adult female, collected by F. Glaw, M. Puente, M. Teschke (née Thomas), and R. Randrianiaina on 31 January 2004 above ‘Camp 1’, between Isaka and Eminiminy, Andohahela National Park (ca 24.750°S, ca 046.850°E, ca 350 m a.s.l.). Additional material.— The following specimens belong to genetically divergent populations and therefore are not included in the paratype series: ZSM 196/2005 (FGZC 2594), adult female, collected by F. Glaw, and P. Bora on 4 February 2005 in the forest at the QMM Climate Station, Sainte Luce (24.7798°S, 047.1713°E, 23 m a.s.l.); ZSM 181/2021 (ACZCV 375, extraction ACP 3589, tissue ACZC 8514), ZSM 182/2021 (ACZCV 376, ACP 3590, ACZC 8515), ZSM 183/2021 (ACZCV 377, ACP 3591, ACZC 8516), collected by S. Hyde Roberts at Sainte Luce (S9) on 10 October 2016; MRSN A7044 (FAZC 15282, ACP 0997, ACZC 4429), collected by F. Andreone and G.M. Rosa on 21 February 2012 at Sainte Luce; MRSN A7045 (FAZC 15419, ACP 1053, ACZC 4485) and MRSN A7046 (FAZC 15427, ACP 1057, ACZC 4489), one male and one female, collected by F. Andreone and G.M. Rosa on 29 February 2012 at Tsitongambarika, Ivohibe. Diagnosis.— Mantidactylus gudrunae sp. nov. is a member of the M. tricinctus clade as revealed by the phylogenomic analysis, and sister to a monophyletic group comprising M. tricinctus and M. grubenmanni. See Table 4 for a list of diagnostic morphological characters. The combination of small body size (male SVL 20–25 mm, female SVL 23–29 mm), presence of a whitish marking on snout tip and of a yellow inguinal marking, and absence of white spots on flanks, distinguishes M. gudrunae sp. nov. from members of other Brygoomantis clades (Table 4). Within the M. tricinctus clade, it differs from both M. tricinctus and M. grubenmanni by a slightly larger body size (male SVL 20–25 mm vs M. gudrunae sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. ...Continued on the next page Description of the holotype. —Adult male in good to moderate state of preservation (Fig. 56). Tongue removed as tissue sample; femoral glands partly detached for examination in internal view. Body relatively slender. Head as wide as body. Snout rounded in dorsal view, truncate in lateral view. Nostrils directed laterally, slightly protuberant, nearer to tip of snout than to eye. Canthus rostralis weakly recognisable, slightly concave; loreal region slightly concave. Tympanum distinct and large, rounded, horizontal diameter of tympanum 88% of horizontal eye diameter. Supratympanic fold in its first part almost identical to tympanum edge, thereafter distinct, running rather straight from behind eye and bending about 70° close to posterior edge of tympanum towards forelimb insertion. Maxillary teeth present. Vomerine teeth form two elongated aggregations, directed posteromedially from choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: I Colour in preservative: dorsally almost uniformly brown, with a dark band between the eyes bordering on a lighter colour on the anterior head surface. Some white spots and markings laterally on head. Limbs with dark crossbands. Ventrally, beige on limbs, brown with distinct white pattern on throat, chest and anterior belly. Larger white spots arranged to form a median intrreupted white line on throat. Lower lip ventrally with alternating white/ dark brown pattern. In life, colourarion was similar but more contrasted. A small yellowish marking was present in the inguinal region. The light ventral pattern was bright silvery white. Variation.—Variation in measurements is given in Table 9. See Fig. 61 for colouration in life and its variation. There is weak sexual size dimorphism (confirmed male SVL 20.2–24.4 mm [n = 7] vs confirmed female SVL 22.6–28.6 mm [n = 6]). Males have a larger tympanum than females (HTD/ED ratio is 62–79% in females, 75– 96% in males). Femoral glands of males in life distinct and coloured with a conspicuous yellowish shade; a large and distinct distal ulcerous macrogland is clearly visible, as is a smaller proximal granular gland field. Natural history.—Specimens have been found along slow running water bodies in coastal rainforest. They are active during the night and call from water. Their call is rarely heard. The colouration of this species is quite variable, with some specimens showing an orangish colouration on the arms or on the dorsal stripe. Sometimes reminiscent of the colouration of species in the subgenus Ochthomantis (e.g. Fig. 61f). Calls.—The calls of this species have not yet been recorded. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to the South East of Madagascar (Fig. 7). This species is known from Andohahela, Manantantely, Sainte Luce, and Tsitongambarika. Elevation range: 23–415 m a.s.l. Etymology.—We dedicate this species to Gudrun Grubenmann from Z̧rich. Together with her husband Moritz, she has been travelling in Madagascar for many decades and has supported our research with important observations of Malagasy amphibians and reptiles.

Mantidactylus manerana fotaka ssp. nov. Identity and justification.—This lineage was newly identified in this study. It is characterized by a high divergence in 16S but haplotype sharing in Rag-1 with the nominal form, M. manerana manerana, which occurs allopatrically and lacks strong morphological differentiation, and is therefore described here as a new subspecies. Holotype.— ZSM 1588/2012 (FGZC 3776), adult male, collected by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F. M. Ratsoavina, and A. Razafimanantsoa on 1 December 2012 at a campsite in the Sorata Massif (13.6851°S, 049.4417°E, 1279 m a.s.l.), Sava Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratypes.—A total of four paratypes: ZSM 1587/2012 (FGZC 3770), adult female, and UADBA uncatalogued (FGZC 3777), unsexed, with same collection data as holotype; ZSM 1586/2012 (FGZC 3753), male, and UADBA uncatalogued (FGZC 3639), adult female, collected by the same collectors as the holotype on 28–30 November 2012 in bamboo forest above Sorata campsite (ca 13.6752°S, ca 49.4410°E, ca 1485 m a.s.l.). Diagnosis.— Mantidactylus manerana fotaka ssp. nov. is a lineage here considered as subspecies of M. manerana due to its high morphological similarity. It is the direct sister group of M. m. manerana according to our phylogenomic analysis. See Table 4 and the diagnosis of M. manerana above for for a list of diagnostic morphological characters and of differences to other species of Brygoomantis. Morphologically, this poorly known taxon appears to differ from the nominal subspecies, M. m. manerana, by shorter hindlimbs (Table 4). A full list of molecular diagnostic sites in the 16S gene of M. manerana fotaka ssp. nov. in pairwise comparisons to all other Brygoomantis species and subspecies is provided as Supplementary appendix. Description of the holotype.—Adult male in excellent state of preservation (Fig. 73). Tongue removed as tissue sample. Body stout. Head as wide as body. Snout rounded in dorsal and lateral views. Nostrils directed laterally, slightly protuberant, nearer to tip of snout than to eye. Canthus rostralis weakly recognisable, slightly concave; loreal region slightly concave. Tympanum distinct, large, wider than high, horizontal diameter of tympanum 92% of horizontal eye diameter. Supratympanic fold distinct, beginning straight above, with gentle ca 60° bend midway, following edge of tympanum. Tongue removed. Maxillary teeth present. Vomerine teeth form two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: I Colour in preservative: dorsally almost uniformly brown. Only a few white spots along flanks and laterally on head. A small white patch on tip of snout. Reatively distinct dark crossbands on limbs. Fingers and toes with alternating pattern of light and dark colour. Ventrally light beige, with rather contrasted brown pigmentation on throat and chest with light spots and vermiculations, including a median light line on throat, and light-dark pattern ventrally on lower lip. Colour of holotype in life not documented. Variation.—Variation in measurements is given in Table 11. See Fig. 78 for colouration in life. There may be some sexual size dimorphism, but our sample size is small (confirmed male SVL 26.2 mm [n = 1] vs confirmed female SVL 29.2 mm [n = 1]). In females, small, yellowish gland rudiments are visible. Natural history.—Largely unknown. Specimens were collected from an area of disturbed rainforest. Calls.— The call of this subspecies has not been recorded. Tadpoles.— The tadpole of this subspecies has not been described. Distribution.— Apparently microendemic to the Sorata massif (Fig. 7). Elevation range: 1398–1599 m a.s.l. Etymology.—The subspecies name is derived from the Malagasy word fotaka, meaning ‘mud’, in reference to the microhabitat in which this and other Brygoomantis dwell. The name is used as a noun in apposition.

Mantidactylus tripunctatus Angel, 1930 bona species Type material.— Mantidactylus tripunctatus Angel, 1930 is based on syntypes: MNHN 1931.21, 1931.23–25 and MCZ 14280 [formerly MNHN 1931.22] (Barbour & Loveridge 1946; Guibé 1978), from ‘ Pic St. Louis, province de Fort-Dauphin’ and ‘ Befotaka, province de Farafangana... à l’altitude de 700 mètres, au bord d’un torrent, en forêt’. We here designate MNHN 1931.24, probably a subadult/ juvenile specimen from Pic St. Louis, as lectotype because we could obtain genetic data from this specimen. Lectotype designation is justified by the need to stabilize this and other nomina in Brygoomantis, given the uncertain identity and morphological similarity of many taxa in the subgenus. Identity.—This nomen has been considered a nomen dubium by Guibé (1978), Blommers-Schl̂sser and Blanc (1991) and Glaw and Vences (1992a), and as a junior synonym of M. betsileanus by Frost (2021). Using barcode fishing we obtained a 16S sequence of the lectotype which firmly clusters among sequences of a lineage morphologically similar to M. betsileanus that is widespread and common in the Tolagnaro (= Fort Dauphin) area, including the environments of the Pic St. Louis, and considered as M. sp. 29 or M. sp. Ca29 by Vieites et al. (2009) and Perl et al. (2014), and depicted as ‘ Mantidactylus sp. aff. betsileanus “Tolagnaro”’ by Glaw andVences(2007).In the phylogenomic tree, a specimen of M. tripunctatus is placed in a subclade with M. noralottae and two other new species described below as M. katae sp. nov. and M. kortei sp. nov., and relationships between these species are also supported by the 16S tree; all of the species differ from each other in their advertisement calls, and M. noralottae also is characterized by larger body size (Table 4), confirming their species-level distinctness and justifying elevation of M. tricinctus to species status. Diagnosis.—A member of the M. betsileanus clade as revealed by the phylogenomic analysis, probably sister to M. katae sp. nov. described below (but see below for uncertainties regarding the samples of M. katae included in the phylogenomic analysis). See Table 4 for a list of diagnostic morphological characters. The combination of a relatively small body size in males (SVL 26–27 mm) and distinctly larger size in females (SVL 33–35 mm), slightly tubercular dorsal skin with distinct continuous dorsolateral ridges, reduced webbing (one phalanx of fifth toe free of web), absence of white spots on flanks, presence of a white marking on snout tip, and advertisement call consisting of a single, long note composed of ≥70 pulses distinguishes M. tripunctatus from species of all other clades (Table 4). Within the M. betsileanus clade, the species differs from M. betsileanus by a lower number of pulses in advertisement calls and a lower pulse repetition rate; and from M. noralottae by smaller body size and presence of a distinct white marking on snout tip (Table 4). For a distinction from the new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. tripunctatus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table 7. See Fig. 38 for colouration in life and its variation. A light vertebral line can be present. There is pronounced sexual size dimorphism (confirmed male SVL 26.4–27.0 mm [n = 3] vs confirmed female SVL 32.9–34.7 mm [n = 5]). Tympanum size is quite variable but does not seem to differ consistently and strongly between sexes (HTD/ED ratio is 53–93% in females, 93–95% in males). Natural history.—At the base of Pic St. Louis (Tolagnaro) we observed calling males at night, sitting at the edge of shallow puddles in a small, very slowly running stream surrounded by remains of rainforest. Calls.—Advertisement calls of individuals probably belonging to M. tripunctatus (but not DNA barcoded), recorded in February 1991 at a site near Tolagnaro, air temperature unknown (Vences et al. 2006: CD2, track 68, cut 1), consists of a long, regularly pulsed note (Fig. 39), emitted in series. The available recording was of relatively poor quality and partly suffered from the overlap of calls of different individuals, making it difficult to assess and measure all parameters precisely. Numerical parameters of five analysed calls were as follows: call duration (= note duration) 1380–1870 ms (1612.8 ± 197.8 ms); ca 70–80 pulses per note (estimate according to overlap of calls); pulse duration 11–19 ms (14.2 ± 2.6 ms); pulse repetition rate within notes 41.7–47.6 pulses/s (45.6 ± 2.3); dominant frequency 1383–1556 Hz (1460 ± 75 Hz); prevalent bandwidth 1200–3400 Hz; call repetition rate (= note repetition rate) ca 8–9 calls/min. Calls recorded on 1 January 1992 at Pic St. Louis near Tolagnaro, 23°C air temperature (Vences et al. 2006: CD2, track 68, cuts 2 & 3), generally agree in character with the other calls from Tolagnaro described above. Although difficult to evaluate due to overlapping calls of multiple individuals, call duration seems to be longer, roughly ranging from 2100–2600 ms. Pulse repetition rate in these calls is slightly lower and ranges from ca 30–39 pulses/s. Tadpoles.— The tadpole of this species has not been described. Distribution.— Apparently microendemic to a small area in far South East of Madagascar (Fig. 7). This species is known from Andohahela, Manantantely, Mandena, Nahampoana, Pic St. Louis, and Tsitongambarika. Elevation range: 8–415 m a.s.l. Etymology.—Latin adjective meaning ‘having three spots’, presumably in reference to some feature of the colouration., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 223-224, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Angel, F. (1930) Description d'un Batracien nouveau de Madagascar, appartenant au genre Mantidactylus (Materiaux des Missions de M. R. Decary). Bulletin du Museum National d'Histoire Naturelle, Paris, Serie 2, 2, 619 - 620.","Barbour, T. & Loveridge, A. (1946) First supplement to typical reptiles and amphibians. Bulletin of the Museum of Comparative Zoology, 96, 59 - 214.","Guibe, J. (1978) Les batraciens de Madagascar. Bonner zoologische Monographien, 11, 1 - 140.","Glaw, F. & Vences, M. (1992 a) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 335 pp. First Edition.","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","Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Vences, M., Glaw, F. & Marquez, R. (2006) The Calls of the Frogs of Madagascar. 3 Audio CD's and booklet. Madrid, Spain, Fonoteca Zoologica, 44 pp."]}

Mantidactylus manerana manerana ssp. nov. Identity and justification.—The nominal subspecies was previously considered as unconfirmed candidate species M. sp. 16 by Vieites et al. (2009), and M. sp. Ca16 by Perl et al. (2014). It was depicted as ‘ Mantidactylus sp. aff. biporus “Andranofotsy”’ by Glaw and Vences (2007). Holotype.— ZSM 500/2016 (field number ZCMV 15162), adult male, collected by M.D. Scherz, A. Rakotoarison, M. Bletz, M. Vences, and J. Razafindraibe on 17 November 2016 at Camp 3 ‘Simpona’, Marojejy National Park (14.43661°S, 049.74335°E, 1325 m a.s.l.), Sava Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratypes.—A total of 16 paratypes: ZSM 209/2005 (FGZC 2881), adult male, ZSM 206/2005 (FGZC 2837) and ZSM 5064/2005 (ZCMV 2019), two adult females, collected by F. Glaw, M. Vences, and R.D. Randrianiaina on 16–17 February 2005 at the type locality; ZSM 208/2005 (FGZC 2847) and ZSM 207/2005 (FGZC 2846), two adult males, collected by F. Glaw, M. Vences, and R.D. Randrianiaina on 16 February 2005 above Camp 3 ‘Simpona’, Marojejy National Park; UADBA uncatalogued (ZCMV 2017–2018, FGZC 2854), three unsexed specimens, UADBA uncatalogued (FGZC 2835), subadult, and UADBA uncatalogued (FGZC 2836), female, collected by F. Glaw, M. Vences, and R.D. Randrianiaina on 16–17 February 2005 around the type locality; UADBA uncatalogued (ZCMV 2087), male, collected by F.Glaw, M.Vences, and R.D. Randrianiaina on 18 February 2005 on the trail between Camp 1 ‘Mantella’ and Camp 2 ‘Marojejia’, Marojejy National Park (precise coordinates unavailable); UADBA uncatalogued (ZCMV 15190, ZCMV 15301, ZCMV 15302), three females, UADBA uncatalogued (ZCMV 15297), unsexed adult, UADBA uncatalogued (ZCMV 15303), subadult, with the same collection data as the holotype. Diagnosis.—See diagnosis for the species, M. manerana, above; for distinction from the other two subspecies, see their diagnoses below. . ...Continued on the next page Description of the holotype.—Adult male in excellent state of preservation (Fig. 73). Tissue from left thigh removed. Body rather stout. Head as wide as body. Snout rounded in dorsal view. Nostrils directed laterally, not protuberant, nearer to tip of snout than to eye. Canthus rostralis weakly recognisable, slightly concave; loreal region slightly concave. Tympanum distinct, large, wider than high, horizontal diameter of tympanum 85% of horizontal eye diameter. Supratympanic fold distinct, beginning straight above, with gentle 45° bend midway towards insertion of forelimb. Tongue ovoid, distinctly bifid posteriorly. Maxillary teeth present. Vomerine teeth form two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: IVariation.—Variation in measurements is given in Table 11. See Fig. 76 for colouration in life and its variation. There is weak sexual size dimorphism (confirmed male SVL 25.0– 27.5 mm [n = 4] vs confirmed female SVL 28.0– 29.4 mm [n = 2]). Males appear to have only slightly larger tympanum sizes than females (HTD/ED ratio is 60–78% in females, 67–86% in males). Femoral glands of males are relatively small but distinct in life, with a yellowish or orange shade, and with both a distal ulcerous macrogland and a proximal granular gland field recognizable. Natural history.—Poorly known. Calling specimens were collected at dusk from a stream in primary rainforest. 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). Distribution.— Microendemic to the Marojejy massif (Fig. 7). This subspecies is known exclusively from high elevation in Marojejy National Park (from Camp 2 ‘Marojejia’ to above Camp 3 ‘Simpona’). Elevation range: 615–1576 m a.s.l., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 287-292, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 496 pp. Third Edition.","Schmidt, H., Strauss, A., Glaw, F., Teschke, M. & 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"]}

Mantidactylus stelliger sp. nov. Identity and justification.—This lineage, phenotypically similar to species in the M. biporus clade, occupies an isolated position in the phylogenomic tree and in the 16S tree. It was newly discovered in this study and therefore has not been included in previous DNA barcoding assessments of Madagascar’s amphibians.We here consider it as distinct species due to its isolated phylogenetic position, high mitochondrial divergences of at least 6.8% uncorrected pairwise 16S distance to all other Brygoomantis species, and morphological differences (see Diagnosis below). Holotype.— ZSM 2381/2007 (ZCMV 5932), adult male, collected by M. Vences, K.C. Wollenberg, and E. Rajeriarison on 3 March 2007 at Ambohitsara (21.3572°S, 047.8157°E), Vatovavy-Fitovinany Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratypes.—A total of four paratypes: ZSM 2379/2007 (ZCMV 5870), adult female, and ZSM 2380/2007 (ZCMV 5922), adult male, with the same collection data as the holotype; UADBA uncatalogued (ZCMV 5923, ZCMV 5931), two specimes of unknown sex and maturity, with the same collection data as the holotype. Diagnosis.— Mantidactylus stelliger sp. nov. is the sole member of the M. stelliger clade according to our phylogenomic analysis. See Table 4 for a list of diagnostic morphological characters. The combination of small body size (male SVL 21–23 mm, female SVL 31 mm), slightly granular dorsal skin without dorsolateral ridges, large tympanum size in males (12% of SVL), presence of white spots on flanks in at least some individuals, and absence of a white marking on the snout tip, distinguishes M. stelliger sp. nov. from species of the M. betsileanus, M. curtus, M. fergusoni, M. tricinctus, and M. ulcerosus clades. M. inaudax (M. inaudax clade) differs by larger body size and more developed foot webbing. Species of the M. biporus clade differ as follows: M. biporus by larger body size and and more developed foot webbing; M. augustini by longer hindlimbs and more developed foot webbing; M. bletzae by more developed foot webbing and presence of dorsolateral ridges; M. brevirostris possibly by somewhat smoother dorsal skin and smaller femoral glands; M. eulenbergeri by more developed foot webbing and smoother dorsal skin; M. glosi by shorter hindlimbs and more developed foot webbing (Table 4). For a distinction from new species in the M. inaudax clade, see the diagnoses in the respective species accounts below. A full list of molecular diagnostic sites in the 16S gene of M. stelliger sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype. —Adult male in good state of preservation (Fig. 62). Part of right thigh muscle removed as tissue sample. Femoral glands partly detached for examination in internal view. Body rather stout. Head wider than body. Snout rounded in dorsal view. Nostrils directed laterally, not protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis not clearly recognisable. Loreal region very weakly concave. Tympanum distinct, rounded, its horizontal diameter about 67% of eye diameter. Supratympanic fold present, beginning straight, and gently curving midway towards jaw/forelimb insertion, following the rounded form of the tympanum. Tongue ovoid and bifid. Maxillary teeth present. Vomerine teeth present in two small rounded aggregations, positioned posterolateral to choanae. Choanae more or less rounded, somewhat elliptical/ slit-like. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: IVariation.—Variation in measurements is given in Table10. See Fig. 72 for colouration in life and its variation. There may be pronounced sexual size dimorphism, but our sample size is small (confirmed male SVL 21.6–22.6 mm [n = 2] vs confirmed female SVL 31.1 mm [n = 1]). Male and female relative tympanum sizes do not seem to differ (HTD/ED ratio is 74% in the female, 67–74% in the males). Femoral glands of males in life relatively distinct, with a yellowish tone, and mostly consisting of a distinct distal ulcerous macrogland, with only small remnants of a proximal granular gland field. Natural history.—Poorly known. Specimens were collected from near a stream in a remnant of primary rainforest. At Ranomafana National Park specimens were found during day and night at two streams in primary forest (elevational range between 777–835 m a.s.l.) sitting in shallow parts of the streams. Two females with visible eggs were detected at Sahalavakely on 22 February 2011, another female with visible eggs was found at Sahalavabe on 23 March 2011. Calls.— The call of this species has not been recorded. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to the Southern Central East (Fig. 7). This species is known from Ambohitsara and Ranomafana National Park (Sahalavabe and Sahalavakely near Beremby). Elevation range: 294–860 m a.s.l. Etymology.—The Latin adjective in the masculine nominative singular stelliger, meaning ‘starry’, in reference to the white spots that are often present on the flanks of this species., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 279-282, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023

Mantidactylus biporus clade A species-rich, diverse and probably not monophyletic group, containing species characterized by a mostly rather small body size (20.5–35.8 mm adult SVL), short and rounded snout, and typically stout body shape with short hindlimbs. Species in this group often have scattered white spots, especially on flanks and laterally on the head. Contains: M. biporus and five new species, described based on holotypes depicted in Fig. 62. Note that several other species previously thought to be related to M. biporus (or reported to be morphologically similar), are assigned to the M. ulcerosus clade (M. schulzi and M. steinfartzi; see above), and to the M. inaudax and M. stelliger clades (see respective accounts below). One of the new species named in the following (M. bletzae sp. nov.) is not included in the phylogenomic tree, and its relationships are not reliably resolved in the 16S tree. We here assign it tentatively to the M. biporus clade, but it might also be related to the M. inaudax clade., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on page 262, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023

Mantidactylus curtus (Boulenger, 1882) Type material.— Rana curta Boulenger, 1882 is based on seven syntypes, four of which are still available: BMNH 1947.2.10.28–31 from ‘ East Betsileo’ and ‘ Ankafana, Betsileo’. We here designate the DNA-barcoded specimen BMNH 1947.2.10.30, probably an adult male (sex not unambiguously confirmed), as lectotype. In consequence, the type locality is now ‘ East Betsileo’. Lectotype designation is justified by the need to stabilize this and other nomina in Brygoomantis, given the uncertain identity and morphological similarity of many taxa in the subgenus. ...Continued on the next page ) ...Continued on the next page Identity.—The name Mantidactylus curtus has been applied to a complex of genetically divergent lineages inhabiting various mountain ranges and areas of the central high plateau of Madagascar (e.g. Blommers-Schl̂sser & Blanc 1991; Glaw & Vences 1992a). Glaw and Vences (2006) revalidated M. bourgati to refer to the lineage of the Andringitra Massif, but the identity of M. curtus remained uncertain. We here provide a 16S sequence of the lectotype that clusters with a lineage from various localities not far from the type locality (e.g. Antoetra, Itremo, Col des Tapias), providing definitive evidence of the assignment of the nomen curtus to this lineage. Evidence of introgression of genomic material from this species (M. curtus) into a syntopic lineage (M. ambohimitombi marefo ssp. nov., described below) was found in the Phylonetworks analysis (Fig. 5). Since the latter taxon appears to have a limited distribution range in the Itremo Massif, and the observed reticulation only concerned one M. curtus specimen from the same site, it is likely that this inter-species gene flow is localized and does not compromise the identity of M. curtus as independently evolving lineage. Synonyms.—Boulenger (1895) considered Rana inaudax Peracca, 1893 to be a synonym of M. curtus, but that species name is revalidated below. Diagnosis.— A member of the M. curtus clade and sister to M. bourgati. See Table 4 for a list of diagnostic morphological characters. The combination of relatively large body size of up to 39 mm, smooth skin, absence of dorsolateral ridges, strongly developed foot webbing with fully webbed fifth toe, and relatively short snout distinguishes this species from species of the other clades. Within the M. curtus clade, M. alutus, M. madecassus and M. pauliani have smaller body sizes (Table 4). Mantidactylus curtus has smooth dorsal skin, constituting a difference to many specimens of M. ambohimitombi and M. bourgati where the skin is somewhat granular.As far as known, M. curtus and its sister species M. bourgati occur allopatrically and therefore can be distinguished based on localities. 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 M. curtus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table 5. See Fig. 10 for colouration in life and its variation. Evidence for sexual size dimorphism is inconclusive (confirmed male SVL 33.7 mm [n = 1] vs confirmed female SVL 32.5–39.8 mm [n = 8]). In the male specimen MRSN A6757, FGL x FGW is 5.6 mm x 2.6 mm. In many other individuals, femoral glands are less distinct, and some of them cannot be reliably sexed by external examination. Natural history.— Specimens were found in the vicinity of highland streams, usually quite close to the water. Calls.—The call of this species has not been recorded. Tadpoles. —A tadpole of M. curtus (ZSM 943/2004) was described by Schmidt et al. (2009). The tadpole description by Blommers-Schl̂sser (1979), based on materialfromManjakatompo(Ankaratra) andAngavokely, probably refers to different species, as M. curtus is not among the species we have recorded from Ankaratra here. We have not re-sampled Angavokely, but M. alutus and M. ambohimitombi miloko ssp. nov. occur at sites nearby, the latter of which closely resembles M. curtus. Distribution.— Endemic to a small area of the central highlands of Madagascar (Fig. 7). This species is known from Ambositra, Ankazomivady, Antoetra, Antsirakambiaty forest, Col des Tapias, Ibity, Itremo, and Vatolampy. Elevation range: 1300–2090 m a.s.l. Etymology.— Latin adjective meaning ‘shortened’ or ‘short’, presumably in reference to the short snout of the species., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 166-170, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Boulenger, G. A. (1882) Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum. Taylor and Francis, London, UK. 2 nd Edition.","Glaw, F. & Vences, M. (1992 a) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 335 pp. First Edition.","Vences, M., Glaw, F. & Marquez, R. (2006) The Calls of the Frogs of Madagascar. 3 Audio CD's and booklet. Madrid, Spain, Fonoteca Zoologica, 44 pp.","Peracca, M. G. (1893) Descrizione di nuove specie di rettili e anfibi di Madagascar. Nota II (1). Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Universita di Torino, 8, 1 - 16. https: // doi. org / 10.5962 / bhl. part. 27224","Schmidt, H., Strauss, A., Glaw, F., Teschke, M. & 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"]}

Mantidactylus pauliani Guibé, 1974 Type material.— Mantidactylus pauliani Guibé, 1974 is based on the holotype (by original designation) MNHN 1972.1508 from ‘ Nosiarivo (massif d’ l’Ankaratra)’. There are eight paratypes (Vences & Glaw 1999): MNHN 1972.1509–1516. Identity.— Mantidactylus pauliani is a morphologically distinct and apparently microendemic species restricted to high elevations on the Ankaratra Massif. Its identity has been assessed by Vences and Glaw (1999) and is unambiguous due to its microendemic distribution and typical short-snouted appearance. Therefore, no genetic data from the name-bearing type were collected. Diagnosis.—A member of the M. curtus clade and sister to M. mahery sp. nov. (described below), from which it strongly differs morphologically. See Table 4 for a list of diagnostic morphological characters. The combination of a body size of 25–34 mm (Table 5), small tympanum size in males (8–9% of SVL), smooth dorsal skin without dorsolateral ridges, absence of vomerine teeth, and strongly expressed foot webbing with fully webbed fifth toe distinguishes M. pauliani from species of the other clades in Brygoomantis. Within the M. curtus clade, this high-elevation endemic differs from all species exept M. curtus, M. madecassus, and M. ambohimitombi marefo, by a conspicuously short snout in most specimens, from M. madecassus by the single (vs bilobed) subarticular tubercles, and from M. a. marefo by absence of a bluish ring around the eye. Mantidactylus pauliani is endemic to high elevations at the Ankaratra Massif, where it is sympatric with M. a. ambohimitombi, which differs by larger body size, more pointed snout,and more contrasted dorsal pattern. 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 M. madecassus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table 5. See Fig. 20 for colouration in life and its variation. There is weak sexual size dimorphism (confirmed male SVL 29.5–31.0 mm [n = 2] vs confirmed female SVL 31.1–33.7 mm [n = 4]), and males have a slightly larger tympanum diameter than females (Vences & Glaw 1999). This is consistent with weak sexual size dimorphism reported by Andreone et al. (2014). Based on formalin-fixed and well-preserved voucher specimens of the MNHN collection, Vences and Glaw (1999) illustrate femoral glands in internal view, and document that females have weakly developed glands that are reminiscent in structure of those of males but with overall smaller gland granules. This same phenomenon of relatively well-developed gland rudiments in females, may also apply to several other species in the M. curtus clade and could make it difficult to sex preserved individuals. Future studies should assess whether femoral gland prominence in these frogs might also be influenced by seasonal effects. Natural history. —Specimens were found sitting in the water or on exposed rocks in montane streams both inside and outside of forest (see Vences et al. 2002 for more information). Mantidactylus pauliani is rarely encountered and considered highly threatened (Andreone et al. 2005). Age structure, population estimate, and status of infection with Batrachochytrium dendrobatidis were studied by Andreone et al. (2014). They found adult specimens ranging 3–8 years old, with no significant difference in age between males and females. Specimens reach sexual maturity in the second year in males and third year in females. Chytrid was not identified in these frogs. Calls.—The call of this species has not been recorded. Tadpoles.— The tadpole of this species has not yet been described in detail. Distribution.— Apparently microendemic to high elevations on the Ankaratra massif (Fig. 7). Elevation range: all verified sites are from> 2000 m a.s.l. (up to at least 2200 m a.s.l.) (see Vences et al. 2002 for more information). Etymology.— Eponym for R. Paulian, who initiated and directed the CNRS programme ‘Study of montane ecosystems in the Malagasy region’ (RCP 225) (loosely translated from Guibé 1973b)., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 190-193, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Vences, M. & Glaw, F. (1999) Variation in Mantidactylus madecassus Millot & Guibe, 1950, a little known Malagasy frog, with resurrection of Mantidactylus pauliani Guibe, 1974. Herpetological Journal, 9, 101 - 110.","Andreone, F., Crottini, A., Rabemananjara, F. C. E., Randrianirina, J. E., Razafindrabe, T. & Tessa, G. (2014) Age structure, population estimate and Bd-status of two Critically Endangered frogs from the Ankaratra Massif (Madagascar), Boophis willliamsi and Mantidactylus pauliani (Amphibia: Mantellidae). Scripta Herpetologica, Studies on Amphibians and Reptiles in honour of Benedetto Lanza, 17 - 29.","Vences, M., Andreone, F., Glaw, F., Raminosoa, N., Randrianirina, J. E. & 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","Andreone, F., Cadle, J. E., Cox, N., Glaw, F., Nussbaum, R. A., Raxworthy, C. J., Stuart, S. N., Vallan, D. & Vences, M. (2005) Species review of amphibian extinction risks in Madagascar: conclusions from the Global Amphibian Assessment. Conservation Biology, 19, 1790 - 1802. https: // doi. org / 10.1111 / j. 1523 - 1739.2005.00249. x","Guibe, J. (1973 b) Batraciens nouveaux de Madagascar. Bulletin du Museum National d'Histoire Naturelle, Paris, Serie 3, 171, 1169 - 1192."]}

Mantidactylus inaudax (Peracca, 1893) bona species Type material.— The nomen is based on a series of three syntypes, originally under the single number MZUT An727, but now divided into An727.1 – An727.3, from ‘ dintorni di Andrangoloaka e dalla vicina valle dell’Umbi’ (Gavetti &Andreone1993; Peracca1893). Andrangoloaka is at ca 19.00°S, ca 047.95°E. These specimens were thought lost by Guibé (1978) and Blommers-Schl̂sser and Blanc (1991) but were rediscovered by Gavetti and Andreone (1993). We here designate as lectotype MZUT An727.1, an adult female individual of 32.1 mm SVL (Fig. 73). Lectotype designation is justified by the need to stabilize this and other nomina in Brygoomantis, given the uncertain identity and morphological similarity of many taxa in the subgenus. Identity.— The name Rana inaudax Peracca, 1893 was placed in the synonymy of M. curtus by BlommersSchl̂sser and Blanc (1991) although this synonymy was doubted by Gavetti and Andreone (1993) because it was not based on specimen examination (see also Frost 2021). We here provide a 16S sequence of the lectotype that clusters with a lineage from various localities not far from the type locality of Andrangoloaka (e.g. Fierenana and the road between Moramanga and Anosibe An’Ala). We thereby provide genetic evidence of the assignment of the nomen inaudax to this lineage, which was previously considered as the true M. biporus (Vieites et al. 2009), and we therefore revalidate the name M. inaudax for this lineage. The lineage to which the lectotype belongs is closely related to another from Ambohitantely and other nearby sites that has been assigned to candidate species M. sp. 17 by Vieites et al. (2009), and M. sp. Ca17 by Perl et al. (2014). It was depicted as ‘ M. sp. aff. biporus “Ambohitantely”’ by Glaw and Vences (2007). In the course of this study, we discovered several other deep lineages from the western slope of the Makira reserve, from Ampotsidy, and from Fierenana and the road Moramanga to Anosibe An’Ala (previously considered as the true M. biporus) that all form a mitochondrial clade with Ca17 and partly share Rag-1 haplotypes with it. Our phylogenomic analysis of this clade recovers these different lineages as a monophyletic group, but with a different topology of relationships among them than recovered in our mitochondrial tree. We conservatively treat all of these deep lineages as conspecific. Additional material. — The following specimens are all tentatively assigned to M. inaudax, despite substantial genetic variation among several populations (visualized in Fig. 2): ZSM 180/2005 (FGZC 2146), adult male, and ZSM 181/2005 (FGZC 2147), adult female, collected by M. Vences, L. du Preez, P. Bora, L. Raharivololoniaina, R.D. Randrianiaina, T. Razafindraibe, and E. Randriamitso on 18 January 2005 at Ambohitantely Jardin Botanique (ca 18.17°S, ca 047.27°E, ca 1580 m a.s.l.); ZFMK 60141–60142, adult male and female, collected by F. Glaw and D. Vallan on 6 April 1995 at Ambohitantely (ca 18.17°S, ca 047.27°E, ca 1580 m a.s.l.); ZMA 19310 (FGMV 2002.2435), ZMA 19311 (FGMV 2002.2439), ZMA 19314 (FGMV 2002.2429), ZMA 19331 (FGMV 2002.2441), four adult females, ZMA 19341 (FGMV 2002.2252), putative female, and ZMA 19343 (FGMV 2002.2423) and ZMA 19345 (FGMV 2002.2251), two adult males, collected by M. Vences, D.R. Vieites, and collaborators on 19 February 2003 in Fierenana (18.5299°S, 048.5901°E); ZSM 1768/2008 (ZCMV 8869), adult male, collected by D.R. Vieites, J. Patton, P. Bora, and M. Vences on 22 February 2008 in the Andranogorika forest fragment, near the road to Brieville (17.76781°S, 047.98415°E); ZSM 363/2010 (FGZC 4358), adult male, collected by F. Glaw, J. K̂hler, P.-S. Gehring, M. Pabijan, K. Mebert, E. Rajeriarison, F. Randrianasolo, and S. Rasamison on 8 April 2010 in Fanamby forest (18.4214°S, 047.9383°E, 1315 m a.s.l.); ZSM 398/2006 (ZCMV 3259), adult male, collected by M. Vences, R.D. Randrianiaina, and E. Edwards on 25 March 2006 at the crossroad between Moramanga–Anosibe An’Ala and Besariaka (19.0959°S, 048.2402°E); ZSM 548/2009 (ZCMV 11213), adult male, collected by M. Vences, D.R. Vieites, F.M. Ratsoavina, R.D. Randrianiaina, E. Rajeriarison, T. Rajofiarison, and J. Patton on 23 June 2009 at Hevirina (pandanus swamp), Makira (15.4490°S, 049.1119°E, 1093 m a.s.l.); ZSM 85/2016 (MSZC 0161), adult male, collected by M.D. Scherz, J. Borrell, L. Ball, T. Starnes, E. Razafimandimby, D.H. Nomenjanahary, and J. Rabearivony on 8 January 2016 beside a muddy spring in Ampotsidy, Bealanana District (14.4194°S, 048.7194°E, 1340 m a.s.l.); ZSM 88/2016 (MSZC 0178), adult male, collected by M.D. Scherz, J. Borrell, L. Ball, T. Starnes, E. Razafimandimby, D.H. Nomenjanahary, and J. Rabearivony on 8 January 2016 in a pandanus swamp in Ampotsidy, Bealanana District (14.4169°S, 048.7144°E, 1371 m a.s.l.). Diagnosis.— Mantidactylus inaudax belongs to the M. inaudax clade according to our phylogenomic analysis.See Table4 for a list of diagnostic morphological characters. The combination of small to moderate body size (male SVL 22–30 mm, female SVL 27–33 mm), rather smooth dorsal skin without dorsolateral ridges, large tympanum size in males (12–16% of SVL), presence of white spots on flanks in at least some individuals, and absence of a white marking on the snout tip, distinguishes M. inaudax 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 smaller tympanum in males, and a higher pulse repetition rate in advertisement calls; M. augustini by longer hindlimbs, fewer pulses per note, and lower pulse repetition rate in advertisement calls; M. bletzae by a somewhat smaller body size, presence of dorsolateral ridges, longer hindlimbs, and more developed foot webbing; M. brevirostris by less developed foot webbing, possibly smaller body size, and differences in colour pattern; M. eulenbergeri by smaller body size; M. glosi by smaller body size, more granular dorsal skin and shorter hindlimbs (Table 4). For a distinction from new species and subspecies in the M. inaudax clade, see the diagnoses in the respective taxa accounts below. A full list of molecular diagnostic sites in the 16S gene of M. inaudax in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Translation of original description.— To facilitate a revised treatment of this nomen, we here provide a translation of Peracca’s detailed description of Rana inaudax from the original Italian: Vomerine teeth in two oblique groups behind the posterior margin of the choanae. Cordiform tongue, rather small, bifurcated in two posteriorly short rounded appendages. Moderate head, subacute snout protruding about 1 mm over the lower jaw; rounded canthus rostralis; slightly concave loreal region; inter-orbital space equal in width to the upper eyelid, equaling the distance between the antero-internal eye and nostril. Tympanum clearly visible, about ¾ of the eye diameter, equal in diameter to the distance between the antero-internal corner of the eye and the nostril, surmounted by a small skin fold starting from the posterior-external corner of the eye and disappearing at the origin of the arm. Digits of the feet terminated by discs almost twice as large as those of the hands. Internal metatarsal tubercle oval, protruding, very large. A small external metatarsal tubercle, conical. Almost entirely webbed toes. By pulling the posterior extremities forward along the body, the tibio-tarsal joint barely reaches the posterior corner of the eye. The skin of the head, the back, the hips, the upper surface of the posterior extremities, and the posterior surface of the thighs is finely granular; in other regions it is smooth. On the lower surface of the thighs a small circular glandular relief can be observed on each side, presenting a median depression in which 5 or 6 point-like pores (femoral pores) are visible. Colouration—Basic colour of the upper parts grey-brown or slate grey, more or less light. A black spot connects the eyes, preceded by a lighter band. On the back there is an irregular dark spot, sometimes shaped like a V. The posterior extremities have narrow black bands. The lips and cheeks are dotted with white. Lower face of a dirty yellowish white, turning to flesh-grey on the throat, irregularly speckled with white. Dimensions: ♀ ♀ ♁ Length from snout to vent mm 33 29.5 22 of the arm 18.5 15.5 13.5 of the leg 47.5 38 32 of the shank 14.5 12 10 of the foot 15 13 10.5 Head width 13 11 9 Inner metatarsal tubercle 2 2 1.5 Three specimens. Description of referred specimen ZSM 180/2005 (FGZC 2146).— Adult male in good state of preservation. Tissue removed from right thigh; femoral glands partly detached for examination in internal view. Body rather stout. Head as wide as body. Snout rounded in dorsal and lateral views. Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis weak, slightly concave. Loreal region weakly concave. Tympanum distinct, large, elliptical, diameter 78% of eye diameter. Supratympanic fold distinct, beginning straight, with a rather distinct bend midway towards insertion of forelimb, following the outer edge of the tympanum. Tongue ovoid, distinctly posteriorly bifid. Maxillary teeth present. Vomerine teeth present in two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Outer and inner metacarpal tubercles present. Fingers without webbing. Relative length of fingers: IVariation.—Variation in measurements is given in Table 11. See Fig. 74 for colouration in life and its variation. There is pronounced sexual size dimorphism (confirmed male SVL 21.7–25.1 mm [n = 10] vs confirmed female SVL 26.9–32.1 mm [n = 7]). Males have larger tympanum sizes than females (HTD/ED ratio is 63–82% in females, 84–111% in males). Skin on the back is smooth (ZSM 180/2005) or granular (ZFMK 60141, ZFMK 60142). Colour on the back is light brown with few indistinct markings in ZFMK 60141 and ZFMK 60142, darker brown with few indistinct markings in ZSM 180/2005. A dark brown more or less triangular band between eyes is always present. Two dark spots on the back at the level of the forelimb insertion are always present (not visible for ZFMK 60141 and ZFMK 60142 because their skin is in pieces). A light vertebral line or a light vertebral band are never present. A small but distinct light dot on the snout tip is present in the two males. Lower lip with more (e.g. ZFMK 60141) or less (e.g. ZFMK 60142) distinct alternating light and brown spots. Venter and throat from uniformly beige in ZFMK 60141 and ZFMK 60142 or brown with distinct mottling in ZSM 180/2005. A longitudinal white median line on abdomen and throat is never present. Hindlimbs more or less distinctly striped. Forelimbs brown with irregular darker markings and stripes. Femoral glands of adult males are consistently large and prominent. The femoral glands of ZSM 180/2005 have an extensive proximal granular gland field, while such a structure is not recognisable in ZFMK 60141. In external view an external central depression in the distal ulcerous macrogland component of the femoral glands can be seen, and in life, the glands are of yellowish colour. In females femoral glands cannot be recognised in preservative, but in life, rudimentary glands consisting of two or three small structures of similar size are recognisable (Fig. 74e, l). Natural history.—Regularly found in clean, running waters, often in areas of highly disturbed and degraded rainforest, but also in springs and swamps within undisturbed rainforest. Near Moramanga calling males were heard and collected during the day, from a slowly moving small stream near degraded rainforest and with just some remaining trees close to the water. Calls.—The advertisement call of M. inaudax, recorded on 23 June 2009, at Makira West, at an estimated air temperature of 20–25°C, consisted of a pulsed note, emitted isolated or in short series containing 5–7 calls (Fig. 75). Notes exhibit amplitude modulation, with call energy rapidly 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 decreases towards the note’s end. Numerical parameters of 14 analysed calls were as follows: call duration (= note duration) 196–367 ms (268.6 ± 66.4 ms); 19–28 pulses per note (22.6 ± 3.9); pulse duration 3–6 ms (4.3 ± 0.9 ms); pulse repetition rate within notes 55.6–130.4 pulses/s (85.8 ± 23.9); dominant frequency 1146–1276 Hz (1200 ± 41 Hz); prevalent bandwidth 800–3400 Hz; call repetition rate (= note repetition rate) within series ca 75 calls/min. Calls recorded on 25 March 2006 at the crossing of the Moramanga-Anosibe An’Ala and Besariaka roads, at an estimated air temperature of 20–25°C, were emitted in regular series and agreed perfectly in character with the calls described from Makira West above and differd only slightly in numerical parameters (n = 9): call duration (= note duration) 199–312 ms (254.9 ± 35.9 ms); 17–23 pulses per note (20.3 ± 2.2); pulse duration 3–7 ms (4.7 ± 1.1 ms); pulse repetition rate within notes 61.2–142.9 pulses/s (87.1 ± 28.9); dominant frequency 738–961 Hz (814 ± 72 Hz); prevalent bandwidth 500–3500 Hz; call repetition rate (= note repetition rate) within series ca 60–90 calls/min. The most significant difference was the lower dominant frequency which was possibly due to a larger SVL of the calling male. Tadpoles.— The tadpole of M. inaudax was described under the name ‘ M. biporus ’ by Knoll et al. (2007). Distribution.— Endemic to the highlands and rainforests of the Northern Central East and Central Madagascar, also occurring at sites in the North West and Ambirano Regions (Fig. 7). This species is known from Ambohitantely, Ampotsidy, Andrangoloaka, Anjozorobe, Fierenana, the vicinity of Lake Alaotra, the western slope of Makira, and the Moramanga-Anosibe An’Ala/ Besariaka crossroad. Elevation range: 948–1580 m a.s.l. Etymology.— Latin adjective meaning ‘shy’ or ‘hesitant’. It is a third-declension one-termination adjective, and thus is effectively invariable with respect to the gender of the genus., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 282-287, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Peracca, M. G. (1893) Descrizione di nuove specie di rettili e anfibi di Madagascar. Nota II (1). Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Universita di Torino, 8, 1 - 16. https: // doi. org / 10.5962 / bhl. part. 27224","Gavetti, E. & Andreone, F. (1993) Revised catalogue of the herpetological collection in Turin University. 1. Amphibia. Cataloghi. Museo Regionale di Scienze Naturali. Torino 10: 1 - 187. Cataloghi X, Museo Regionale di Scienze Naturali, Torino, Italy.","Guibe, J. (1978) Les batraciens de Madagascar. Bonner zoologische Monographien, 11, 1 - 140.","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","Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 496 pp. Third Edition.","Knoll, A., K ˆ hler, J., Glaw, F., Teschke, M. & Vences, M. (2007) Larval morphology in four species of Madagascan frogs of the subgenus Brygoomantis (Mantellidae: Mantidactylus). Zootaxa, 59, 49 - 59. https: // doi. org / 10.11646 / zootaxa. 1616.1.4"]}

Mantidactylus mahery sp. nov. Identity and justification.—This lineage has been considered as confirmed candidate species M. sp. 14 by Vieites et al. (2009) and M. sp. Ca14 by Perl et al. (2014). This is a relatively large-sized species of Brygoomantis fromwesternMadagascarwithamorphologysuperficially similar to M. ulcerosus which, however, is not its closest relative and concordantly differs in mitochondrial and nuclear genes and in advertisement call structure. Both species occur syntopically at least at one site (Makira). Although M. mahery sp. nov. mainly occurs in western Madagascar unlike the other species of the M. curtus clade which live in the central highlands, the phylogenomic data unambiguously support its inclusion in the group. Even more surprising, the phylogenomic tree places the species sister to M. pauliani, which is a montane endemic from the Ankaratra Massif that differs in numerous morphological characters. The status of M. mahery sp. nov. as a separate species is well supported by multiple lines of evidence. Holotype. — ZSM 23/2004 (field number FGZC 37), adult male collected by F. Glaw, M. Puente, R. Randrianiaina, and M. Teschke (née Thomas) on 21 January 2004 in Isalo at a creek near Ranohira (22.5856°S, 045.3997°E, 813 m a.s.l.), Ihorombe Region, Madagascar. A 16S barcode sequence of the holotype is available from GenBank (accession AY848286). Paratypes.—A total of 12 paratypes: ZSM 25/2004 (FGZC 39), adult male, and ZSM 26/2004 (FGZC 42), adult female, with the same collection data as the holotype; ZSM 567/2009 (ZCMV 11457) and ZSM 569/2009 (ZCMV 11484), two adult females, collected by M. Vences, D.R. Vieites, F.M. Ratsoavina, R.D. Randrianiaina, E. Rajeriarison, T. Rajofiarison, and J. Patton on 20 June 2009 in Sahaovy (‘Camp 0’), Makira (15.4889°S, 049.0785°E, 607 m a.s.l.); ZSM 9/2006 (FGZC 682), adult male, and ZSM 61/2006 (FGZC 793), adult female, collected by F. Glaw, J. K̂hler, P. Bora, and H. Enting on 18 and 23 March 2006, respectively, at Antranopasazy (‘Camp 1’), Tsingy de Bemaraha National Park (18.7086°S, 044.7189°E, 146 m a.s.l.); ZSM 134/2006 (FGZC 938), ZSM 135/2006 (FGZC 939), two adult males, and ZSM 136/2006 (FGZC 940), adult female, collected by F. Glaw, J. K̂hler, P. Bora, and H. Enting on 31 March 2006 at Andafiabe on the Beboka River (‘Camp 2’), Tsingy de Bemaraha National Park (18.7842°S, 044.7794°E, 177 m a.s.l.); ZSM 927/2003 (FG / MV 2002.1421), putative male, ZSM 941/2003 (FG / MV 2002-1485), adult male, and ZSM 942/2003 (FG / MV 2002-1486), adult female, collected by G. Aprea, M. Puente, L. Raharivololoniaina, M. Teschke (née Thomas), and D.R. Vieites between 29 January and 1 February 2003 at Hotel Isalo Ranch (22.5929°S, 045.3928°E, ca 800 m a.s.l.). Diagnosis.— Mantidactylus mahery sp. nov. is a member of the M. curtus clade as revealed by the phylogenomic analysis, and sister to the morphologically strongly different M. pauliani. While all other species in the M. curtus clade occur on the central plateau of Madagascar, M. mahery is distributed in western Madagascar, including some rather arid areas where it appears to be the only Brygoomantis present, and it is also present in one locality in the North West (western slope of Makira). See Table 4 for a list of diagnostic morphological characters. The combination of a large body size of up to 49 mm, slightly granular skin with (weakly expressed) dorsolateral ridges, absence of white spots on flanks and of white marking on snout tip, and short pulsed advertisement calls emitted in regular series distinguishes the new species from species of the other clades. In the North West it can occur syntopically with the similarly sized M. ulcerosus which however has a distinctly more tubercular dorsal skin, and more pulses per note in advertisement calls. Within the M. curtus clade, the new species differs by its larger tympanum diameter in males (11–13% of SVL) from all other species (Table 4). Mantidactylus alutus, M. madecassus and M. pauliani furthermore are smaller and have a shorter snout (Table 4). For detailed distinction from other new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. alutus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype. —Adult male in good state of preservation (Fig. 9). Tongue excised as tissue sample (tongue no longer present); femoral glands partly detached for examination in internal view. Body rather slender (in this, differing from many other specimens of this species which are stouter). Head as wide as body. Snout rounded in dorsal and lateral views. Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis weak, slightly concave. Loreal region weakly concave. Tympanum distinct, large, elliptical, diameter about 93% of eye diameter. Supratympanic fold present, beginning straight,with a rather distinct bend midway towards forelimb insertion. Maxillary teeth present. Vomerine teeth present in two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticalur tubercles single. Outer metacarpal tubercle present, inner metacarpal tubercle present. Fingers without webbing. Relative length of fingers: I=II Colour in preservative: dorsum dark brown. Forelimbs brown with poorly defined darker markings. Hindlimbs brown with poorly defined darker crossbands. Inguinal region without whitish spots. Snout tip without a light dot. Venter beige with brown mottling, throat darker than belly. Lower lip with alternating light and brown spots. Toe discs dark. Colour in life of holotype unknown. Variation. —Variation in measurements is given in Table 5. See Fig. 21 for colouration in life and its variation. In life, dorsum is brown with distinct darker markings. Dark band between eyes is present. Forelimbs brown with very indistinct darker markings; hindlimbs with indistinct darker crossbands. Belly beige; throat with distinct white and brown mottling. A longitudinal white line on abdomen and throat is present. Femoral glands distinctly orange. There is moderate sexual size dimorphism(confirmed male SVL 29.2–37.2mm [n =7] vs confirmed female SVL 34.2–48.6 mm [n = 6]). Horizontal tympanum diameter is 80–100% of eye diameter in males and 69–86% of eye diameter in females. Skin on the back with very few indistinct tubercles on the flanks. Colour on the back varies from light brown with distinct darker markings (e.g. ZSM 927/2003) to uniformly dark brown. Two dark spots on the back at the level of forelimb insertion always more or less distinctly present, except in the holotype (ZSM 23/2004), whose colour is too dark to see any markings.A dark brown more or less triangular band between eyes is always present. A light vertebral band or line is not present. An indistinct light dot on the snout tip is never present except in ZSM 25/2004. Lower lip with more (e.g. ZSM 23/2004) or less (e.g. ZSM 25/2004) distinct alternating light and brown spots. Venter and throat from uniformly beige with faint markings (e.g. ZSM 26/2004) to dark brown mottled (e.g. ZSM 23/2004). A longitudinal light median line on abdomen and throat is present in ZSM 25/2004. Hindlimbs always distinctly striped (e.g. ZSM 941/2003) except in ZSM 23/2004 where hindlimbs are striped indistinctly dark-brown. Forelimbs brown with irregular darker markings and stripes. Femoral glands of males large and prominent with a clear proximal granular gland field in ZSM 25/2004 and ZSM 941/2003, in ZSM 23/2004 and ZSM 927/2003 less prominent with indistinct proximal granular gland field. In external view a central depression in the middle of the femoral gland can be seen, thus indicating a distal ulcerous macrogland. In females femoral glands are always small but distinctly present (e.g. ZSM 942/2003), but a proximal granular gland field is never present. In life, males in reproductive state have femoral glands orange coloured (Fig. 21b, h), and sometimes (Fig. 21b) the proximal granular gland field is larger and more prominent than the distal ulcerous macrogland, which is uncommon among mantellines; the granular gland fields on the two opposite thighs contact each other medially. Natural history.—The species is known from various sites in the West and North West of Madagascar, reaching into the South at Isalo. It has been found along running water (including very slowly running streams) in and outside of forest. At the western slope of the Makira Reserve it was found with M. ulcerosus and M. jonasi sp. nov. (see below) along a relatively large stream in degraded remnants of rainforest. Calls.— The advertisement call of M. mahery, recorded on 28 January 1994 at Isalo National Park, near Ranohira, 23.4°C air temperature (Vences et al. 2006: CD 2, track 76), consists of a short, pulsed note, emitted in regular series at fast succession (Fig. 22). Pulse repetition rate is distinctly higher at the beginning of calls and significantly reduces after approximately one quarter of the call’s duration. Amplitude modulation is present, with highest call energy occurring at the beginning of the call and continuously decreasing towards its end. Numerical parameters of eight analysed calls are as follows: call duration (= note duration) 120– 144 ms (131.8 ± 8.7 ms); 21–26 pulses per note (24.1 ± 2.3); pulse duration 1–2 ms (1.6 ± 0.5); pulse repetition rate within notes 107.8–667.0 pulses/s (331.4 ± 205.3); dominant frequency 1004–1270 Hz (1151 ± 95 Hz); prevalent bandwidth 800–4500 Hz; call repetition rate (= note repetition rate) within regular series ca 211–218 calls/min. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to an eclectic collection of disparate localities, mostly in the West of Madagascar, but reaching the eastern rainforest escarpment at Makira in the North West, and Tsaranoro in the Central region (Fig. 7). This species is known from Isalo (various localities), Forêt de Beanka, Makay, Makira West (Sahaovy, Camp 0), Tsaranoro, and Tsingy de Bemaraha (various localities). Elevation range: 120–960 m a.s.l. Etymology. —The species name is derived from the Malagasy adjective mahery, meaning ‘big’ or ‘strong’, and refers to the rather stout body shape of this species. The name is used as a noun in apposition.

Mantidactylus steinfartzi sp. nov. Identity and justification.—This lineage was considered as M. sp. 33 by Vieites et al. (2009) and M. sp. Ca33 by Perl et al. (2014) and Vences et al. (2018). It was depicted as ‘ Mantidactylus sp. aff. biporus “Tsaratanàna Antsahamanara”’ by Glaw and Vences (2007). It occurs in sympatry(although not strict syntopy)with its sister species M. schulzi but is distinguished from it by advertisement calls, morphology, and concordant differentiation of 16S and Rag-1. It was previously thought (e.g. Glaw & Vences 2007) to be related to M. biporus, but our phylogenomic tree firmly places it in the M. ulcerosus clade. Holotype.— ZSM 658/2001 (FGMV 2001.107), adult male (seen calling), collected by F. Andreone, F. Mattioli, J.E. Randrianirina, and M. Vences between 4–9 February 2001 at Antsahamanara ‘Camp 1’ (14.0450°S, 048.7844°E, ca 1000 m a.s.l.), Manarikoba forest, Tsaratanàna Massif, Diana Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratypes.—A total of 11 paratypes: ZSM 659/2001 (FG / MV 2001.110)and ZSM 663/2001 (FG / MV 2001.118), two adult males, and ZSM 655/2001 (FG / MV 2001.68), ZSM 657/2001 (FG / MV 2001.98), ZSM 660/2001 (FG / MV 2001.113), ZSM 661/2001 (FG / MV 2001.116), four adult females, with the same collection data as the holotype; ZSM 843/2003 (FG / MV 2002.0810), and ZMA 19567 (FG / MV 2002.2315), two adult males, and ZSM 844/2003 (FG / MV 2002.0811) and ZMA 19568 (FG / MV 2002.2317), two adult females, collected by F. Glaw, R.D. Randrianiaina, and M. Vences on 3 February 2003 at ‘Camp 1’ on the Manongarivo Massif (13.9770°S, 048.4220°E, 751 m a.s.l.); UADBA-A uncatalogued (FGZC 3791), specimen of unknown age and sex, collected by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F.M. Ratsoavina, and A. Razafimanantsoa on 3 December 2012 near Ambodimandresy (13.7133°S, 049.4911°E, 778 m a.s.l.). Diagnosis.— Mantidactylus steinfartzi sp. nov. is a member of the M. ulcerosus clade as revealed by the phylogenomic analysis, and sister to the sympatric M. schulzi. See Table 4 for a list of diagnostic morphological characters. The combination of a small body size of up to 28 mm, smooth to slightly tubercular dorsal skin, absence of dorsolateral ridges, large tympanum size in males (15–17% of SVL), presence of white spots on flanks, and absence or weak expression of a white marking on snout tip, distinguishes M. steinfartzi sp. nov. from most other species of Brygoomantis from other clades: members of the M. betsileanus clade typically have a distinct white marking on the snout tip and no white spots on the flanks, and (except for M. betsileanus and M. riparius sp. nov. which differ in the number of pulses per note; Table 4) a lower pulse rate in advertisement calls; members of the M. fergusoni clade are larger, have a more granular dorsal skin, no white spots on the flanks, and lower pulse rate in advertisement calls; and members of the M. biporus and M. inaudax clades have, as far as known, fewer pulses per note in advertisement calls (Table 4). Within the M. ulcerosus clade, the new species differs by a distinctly smaller body size and several other characters from M. bellyi and M. ulcerosus. It is morphologically rather similar to its sister species M. schulzi which, however, usually has no white dots on the flanks and a more distinctly expressed white marking on the snout, more granular dorsal skin, a smaller tympanum in males, a slightly larger male body size, and also differs in details of advertisement calls: a rather irregularly emitted note of quite variable number of pulses in M. schulzi vs less variability in pulse number and emission of short series of usually two calls in M. steinfartzi sp. nov. 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 M. steinfartzi sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype.—Adult male in a mediocre state of preservation (softly fixed, similar to all other available specimens of this species; Fig. 23); muscle tissue from left thigh removed, femoral glands partly detached for examination in internal view. Body rather stout. Head as wide as body. Snout rather pointed. Nostrils directed laterally, slightly protuberant, nearer to tip of snout than to eye. Canthus rostralis weakly recognisable, slightly concave; loreal region slightly concave. Tympanum distinct, large, wider than high, horizontal diameter of tympanum 106% of horizontal eye diameter. Supratympanic fold distinct, beginning straight above, with a rather distinct 45° bend midway towards insertion of forelimb, following the outline of the large tympanum. Tongue ovoid, distinctly bifid posteriorly. Vomerine teeth form two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: I Colour in preservative: dorsally almost uniformly brown, with a few indistinct and irregular large markings. A somewhat darker patch is present between the eyes. Limbs with poorly contrasted dark crossbands. Flanks and sides of head with scattered whitish spots. Snout tip with a poorly contrasted light spot. Venter beige, throat and chest with brown pigment and a light medial line on the throat. Lower lip ventrally with distinct alternating light and brown spots. Colouration in life not recorded for holotype specimen. Variation.—Variation in measurements is given in Table 6. See Fig. 31 for colouration in life. There is moderate sexual size dimorphism (confirmed male SVL 17.3–22.3 mm [n = 5] vs confirmed female SVL 21.9–28.2 mm [n = 6]). Males have a distinctly larger tympanum than females (HTD/ED ratio is 59–73% in females, 94– 114% in males). Compared to the holotype, in other male specimens the femoral glands were smaller (Fig. 31), and smaller than in the sister species M. schulzi. In life the glands have a slightly yellowish tone (Fig. 31). Natural history.—All specimens were observed in small streams and brooks in primary rainforest. Calling males were observed from one headwater pool, calling from positions directly next to the water during the day. Calls.— The advertisement call recorded on 4 February 2001 at Antsahamanara Campsite, Manarikoba forest, Tsaratanàna Strict Nature Reserve, 20°C air temperature (Vences et al. 2006: CD2, track 73), consists of a pulsed note (Fig. 32), emitted in groups containing two calls. Notes exhibit slight amplitude modulation, with maximum call energy occurring at approximately the middle of the note’s duration. Pulse repetition rate within notes is higher at the beginning of the note and slightly decreases after approximately the first quarter of the note’s duration. Numerical parameters of six analysed calls were as follows: call duration (= note duration) 516–721 ms (615.3 ± 87.8 ms); 40–54 pulses per note (47.2 ± 5.1); pulse duration 2–5 ms (2.9 ± 1.0 ms); pulse repetition rate within notes 69.8– 115.4 pulses/s (87.2 ± 17.5); dominant frequency 3193– 3716 Hz (3416 ± 184 Hz); prevalent bandwidth 2700–4300 Hz; call repetition rate (= note repetition rate) within call groups ca 24–36 calls/min. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to the Sambirano Region in northern Madagascar (Fig. 7). This species is known from Tsaratanàna (Manarikoba, type locality), Manongarivo, and Ambodimandresy. Elevation range: 751–1000 m a.s.l. Etymology.—We dedicate this species with an apparent ecological (elevational) component in species formation to our colleague Sebastian Steinfartz, in recognition of his contributions to the field of ecologydriven population differentiation and speciation

Mantidactylus biporus (Boulenger, 1889) Type material.—According to Blommers-Schl̂sser and Blanc (1991), Rana biporus Boulenger, 1889 is based on syntypes BMNH 1947.2.26.46–52 from ‘ Madagascar’. We here designate BMNH 1947.2.26.47, an adult female from which we could obtain a DNA sequence, as lectotype. Lectotype designation is justified by the need to stabilize this and other nomina in Brygoomantis, given the uncertain identity and morphological similarity of many taxa in the subgenus. Identity.—Consideringthelargenumberofgenetically highly divergent lineages conforming at least roughly to the morphology of M. biporus, and the lack of a precise type locality, the identity of this nomen has long remained obscure. Using barcode fishing we obtained a sequence of the lectotype which clusters with specimens of a previously (Perl et al. 2014; Vieites et al. 2009) unreported lineage found in Betampona as well as in An’Ala, allowing us herein to newly ascribe this name to that lineage. The lineage previously (Vieites et al. 2009) assigned to M. biporus is reassigned to M. inaudax below. It is worth mentioning that in the original description (Boulenger 1889), the species was described as ‘ Rana biporus ’, and the species epithet may have been meant as a noun in apposition, making the emendation ‘ Rana bipora ’ (e.g. Blommers-Schl̂sser & Blanc 1991; Guibé 1978) unjustified (see also Frost 2021). However, given the species is now in the genus Mantidactylus (of masculine gender) this has no bearing on the name Mantidactylus biporus as currently used. The species has been previously referred to as Mantidactylus sp. aff. biporus [Ca HM364733] (sp. Ca76) by Rosa et al. (2011, 2012). Synonyms.—Previously Mantidactylus brauni Ahl, 1929 was considered a synonym, but we have shown here that that nomen is a junior synonym of M. ulcerosus. Reference specimens. — ZSM 1982/2006 (ZCMV 2425), adult male, and ZSM 396/2006 (ZCMV 1483), adult female, collected by D.R. Vieites, M. Vences, F. Rabemananjara, P. Bora, C. Weldon, and J. Patton on 07– 19 February 2006 at An’Ala (forest camp) (18.91926°S, 048.48796°E, 889 m a.s.l.); MRSN A6180 (FAZC 13480), adult male, collected by G.M. Rosa on 4 February 2007 at Sahabefoza in Betampona (17.9142°S, 049.2077°E, 349 m a.s.l.); MRSN A6266 (FAZC 13675), adult male, collected by G.M. Rosa on 27 February 2007 at Vohitsivalana in Betampona (17.8862°S, 049.2025°E, 517 m a.s.l.); MRSN A6374 (FAZC 13835), adult male, collected by G.M. Rosa on 29 October 2007 at Sahabefoza in Betampona (at geographical coordinates 17.91438°S, 49.20778°E, 325 m a.s.l); ZSM 184/2021 (ACZCV 201 = ACZC 5694), putative female, collected on 14 November 2013 at Vohitsivalana, Betampona (17.8862°S, 049.2026°E), by A. Crottini, D. Salvi, E. Scanarini, Georges, and Jean Noël. Diagnosis.—A member of the M. biporus clade, sister to the new species M. augustini sp. nov. (described below) according to our phylogenomic analysis. See Table 4 for a list of diagnostic morphological characters. The combination of a moderate body size (male SVL 28- 32 mm, female SVL 31–36 mm [one probably immature female 19.3 mm]), rather smooth dorsal skin without dorsolateral ridges, moderate to relatively large tympanum size in males (10–12% of SVL), presence of white spots on flanks, absence of a white marking on the snout tip, and a short, pulsed advertisement call emitted in rapid succession in regular series, distinguishes M. biporus from species of the M. betsileanus, M. curtus, M. fergusoni, M. tricinctus, and M. ulcerosus clades. Mantidactylus inaudax (M. inaudax clade) is morphologically similar but appears to have a larger tympanum in males, and a lower pulse repetition rate in advertisement calls. Mantidactylus biporus is distinguished from its sister species M. augustini sp. nov. by larger body size, smaller tympanum, shorter hindlimbs, less pulses per note and higher pulse rate in advertisement calls. For a distinction from the other (all new) species in the M. biporus, M. stelliger and M. inaudax clades, see the diagnoses in the respective species accounts below. A full list of molecular diagnostic sites in the 16S gene of M. biporus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table10. See Fig. 63 for colouration in life and its variation. A light vertebral line can be present. There is weak sexual size dimorphism (confirmed male SVL 28.0– 31.6 mm [n = 5] vs confirmed female SVL 30.5–35.8 mm [n = 2]; one further female, ZSM 184/2021, measures only 19.3 mm but we hypothesize it is an immature specimen). Males have a slightly larger tympanum than females (HTD/ED ratio is 60–70% in females, 69–79% in males). Femoral glands of males in life are not documented. Females, both in preservative as in life (Fig. 63c) have a distinct pattern of two distinct gland rudiments next to each other which almost certainly explains the species name. Natural history.—Males call during the day and night from flooding zones of small forest streams. Calls.— The advertisement call of M. biporus recorded at Betampona on 31 October 2007 at 20:00 h, 19°C air temperature (Rosa et al. 2011: track 34), consisted of a short, regularly pulsed note (Fig. 64), emitted in regular series at very fast succession. Slight amplitude modulation was present, with relative amplitude increasing from the beginning of the call, reaching its maximum approximately at the last third of the note’s duration. Calls in call series tended to become louder and longer from the beginning to the end of a series. Numerical parameters of 45 analysed calls of two individuals were as follows: call duration (= note duration) 93–132 ms (109.7 ± 11.2 ms); 15–21 pulses per note (17.4 ± 1.8); pulse duration 4–5 ms (4.7 ± 0.5); pulse repetition rate within notes 130.4–210.5 pulses/s (165.8 ± 27.9); dominant frequency 832–997 Hz (884 ± 61 Hz); prevalent bandwidth 400–3000 Hz; call repetition rate (= note repetition rate) within regular series ca 300–360 calls/min. Call series (n = 4) had a duration of 2290–5875 ms. Tadpoles.— The tadpole of this species has not been described. The tadpole described under this name by Knoll et al. (2007) refers to M. inaudax bona species, see below. Distribution.— Endemic to low-elevation rainforest in the Northern Central East (Fig. 7). This species is known from An’Ala and Betampona. The type locality cannot currently be narrowed down. Elevation range: 190–840 m a.s.l. Etymology.— Latin noun in apposition meaning ‘double pores’, presumably in reference to the femoral glands of this species where especially in females, two separate gland rudiments are visible., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 263-265, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Boulenger, G. A. (1889) Descriptions of new reptiles and batrachians from Madagascar. Annals and Magazine of Natural History, Series 6, 4, 244 - 248. https: // doi. org / 10.1080 / 00222938909460511","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Guibe, J. (1978) Les batraciens de Madagascar. Bonner zoologische Monographien, 11, 1 - 140.","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","Rosa, G. M., Marquez, R. & Andreone, F. (2011) The astonishing calls of the frogs of Betampona. Museo Regionale di Scienze Naturali and Fonoteca Zoologica, Torino, Italy.","Rosa, G. M., Andreone, F., Crottini, A., Hauswaldt, J. S., Noel, J., Rabibisoa, N. H., Randriambahiniarime, M. O., Rebelo, R. & Raxworthy, C. J. (2012) The amphibians of the relict Betampona low-elevation rainforest, eastern Madagascar: an application of the integrative taxonomy approach to biodiversity assessments. Biodiversity and Conservation, 21, 1531 - 1559. https: // doi. org / 10.1007 / s 10531 - 012 - 0262 - x","Ahl, E. (1929 ' 1928 ') Beschreibung neuer Fr ˆ sche aus Madagascar. Mitteilungen aus dem Zoologischen Museum in Berlin, 14, 469 - 484.","Knoll, A., K ˆ hler, J., Glaw, F., Teschke, M. & Vences, M. (2007) Larval morphology in four species of Madagascan frogs of the subgenus Brygoomantis (Mantellidae: Mantidactylus). Zootaxa, 59, 49 - 59. https: // doi. org / 10.11646 / zootaxa. 1616.1.4"]}

Mantidactylus incognitus sp. nov. Identity and justification.—This enigmatic lineage is phenotypically assigned to the M. betsileanus clade and has been considered as unconfirmed candidate species M. sp. 34 by Vieites et al. (2009), and M. sp. Ca34 by Perl et al. (2014). Only a minimal amount of information on this lineage is available. It is phylogenetically sister to M. betsileanus in our FrogCap analysis (Fig. 5) with which it also shares Rag-1 haplotypes, and it occurs parapatrically but in close geographic proximity with this species. We here consider it as distinct species due to its highly divergent mitochondrial DNA sequences (6.2–7.6% 16S divergence to M. betsileanus), and the distinct dorsal ridges, and tubercles above the eye seen in the holotype (absent or more weakly expressed in M. betsileanus). Furthermore, also the FrogCap analysis supports a substantial genomic divergence, given the long branch length separating this lineage from M. betsileanus. Mantidactylus incognitus sp. nov. is the only species of Brygoomantis for which no photos in life are available (same applies also to one subspecies, M. manerana antsanga ssp. nov.; see below). Holotype.— ZSM 669/2009 (ZCMV 8945), probably a subadult female, collected by P.-S. Gehring, F.M. Ratsoavina, and E. Rajeriarison on 22 April 2009 at Mahanoro (19.6536°S, 048.7780°E), Antsinanana Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratypes.—A total of two paratypes: UADBA uncatalogued (MV 2001.1156 = 2002.G9), specimen of unknown sex and maturity, collected by M. Vences on 26– 27 November 2001 in Vohidrazana (precise coordinates unavailable); MRSN A6694 (RJS 1801 = ACZC 4189), adult male, collected by J.E. Randrianirina at Anivorano Est (18.7638°S, 048.9468°E, 60 m a.s.l.). Additional material.—Two series of tadpoles, ZSM 1042/2004 and ZSM 1043/2004, from Vohidrazana. Diagnosis.— Mantidactylus incognitus sp. nov. is a member of the M. betsileanus clade as revealed by the phylogenomic analysis, representing the sister species of M. betsileanus. See Table 4 for a list of diagnostic morphological characters. The combination of a relatively small body size in males (SVL 27 mm), dorsal skin with several distinct dorsolateral ridges and supraocular tubercles, reduced webbing (one phalanx of fifth toe free of web), absence of white spots on flanks, and presence of a white marking on snout tip, distinguishes M. incognitus sp. nov. from species of all other clades (Table 4). Within the M. betsileanus clade, the new species differs from M. noralottae by smaller body size and presence of a distinct white marking on snout tip; and from M. betsileanus, M. noralottae, and M. tripunctatus by distinct dorsal ridges (in addition to the dorsolateral ridges) and strongly expressed supraocular tubercles. For a distinction from the other new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. incognitus sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype.—Specimen in moderate state of preservation (soft fixation); probably a subadult female with rudimentary ovary based on gonad examination. Some muscle tissue removed from left thigh (Fig. 33), and a longitudinal lateroventral cut made for gonad examination. Some skin missing from left thigh and shank dorsally. Body relatively slender. Head as wide as body. Snout rather truncate in dorsal and lateral views which might be a preservation artefact. Nostrils directed laterally, slightly protuberant, nearer to tip of snout than to eye. Canthus rostralis weakly recognisable, slightly concave; loreal region slightly concave. Tympanum distinct, rather small, rounded, horizontal diameter of tympanum61%of horizontal eye diameter.Supratympanic fold distinct, beginning straight behind eye, with gentle ca 45° bending midway towards forelimb insertion. Tongue ovoid, distinctly bifid. Maxillary teeth present. Vomerine teeth form two small rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: IVariation.—Variation in measurements is given in Table 7. Colouration in life unknown. Insufficient number of reliably sexed, adult specimens available to assess sexual dimorphism. Natural history.—Very little is known on the habits of this species. Tadpoles at Vohidrazana were collected from a stream in degraded rainforest. Calls.—The call of this species is unknown. Tadpoles.— The tadpole of M. incognitus was described under the name ‘ Mantidactylus sp. aff. betsileanus “Vohidrazana”’ by Knoll et al. (2007). Distribution.— Distributed in a rather small area of the Northern Central East (Fig. 7). This species is known from Anivorano Est, Bemandrevo, Andekaleka, Mahanoro, Sahafina, and Vohidrazana. Elevation range: 10–810 m a.s.l. Etymology.—The Latin adjective incognitus, meaning ‘unknown’, referring to the extremely poor knowledge we have on this genetically distinct species., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 224-226, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Knoll, A., K ˆ hler, J., Glaw, F., Teschke, M. & Vences, M. (2007) Larval morphology in four species of Madagascan frogs of the subgenus Brygoomantis (Mantellidae: Mantidactylus). Zootaxa, 59, 49 - 59. https: // doi. org / 10.11646 / zootaxa. 1616.1.4"]}

Mantidactylus bellyi Mocquard, 1895 Type material.— Mantidactylus bellyi Mocquard, 1895 is based on the holotype MNHN 1893.240 (by monotypy) from ‘ Montagne d’Ambre’ (according to Guibé 1978), and this number had been given incorrectly as 1893.420 by Guibé (1950). According to the original description, there are no paratypes. Identity.—This species was previously considered a junior synonym of M. curtus (e.g. Blommers-Schl̂sser & Blanc 1991) and was resurrected as separate species by Glaw and Vences (2006). It is the sister species of M. ulcerosus. A very extensive Brygoomantis sampling at its type locality Montagne d’Ambre, reflected by 150 sequences in our 16S data set, found only two lineages at this Massif in northernmost Madagascar: one corresponding to a lineage morphologically similar to M. betsileanus and described below as M. jonasi sp. nov., and one corresponding to specimens of a lineage typically (Glaw & Vences 2006, 2007) assigned to M. bellyi. We obtained barcode fishing 16S data of the holotype MNHN 1893.240 and confirm this nomen has been correctly assigned. Diagnosis.—A member of the M. ulcerosus clade as revealed by the phylogenomic analysis, and sister to the morphologically very similar M. ulcerosus. See Table 4 for a list of diagnostic morphological characters. The combination of a large body size of up to 46 mm, strongly tubercular dorsal skin in most individuals, absence of dorsolateral ridges, large tympanum size in males (10– 17% of SVL) and absence of white spots on flanks and of white marking on snout tip distinguishes M. bellyi from species of the other clades. Some species in the M. fergusoni clade can be morphologically similar, but they occur in eastern Madagascar (vs Sambirano and North West regions), and have strongly different advertisement calls (Table 4). Within the M. ulcerosus clade, the species differs by its large body size and tubercular dorsal skin from M. schulzi, and by its advertisement call consisting of single calls (vs note series) from M. schulzi and M. ulcerosus. 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 M. bellyi in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Re-description of the holotype. Adult male in poor state of preservation. Skin on various places lacerated and completely discoloured. Body rather stout. Head wider than body. Snout rounded. Nostrils directed laterally, slightly protuberant. Position of nostrils not recognisable due to the bad state of preservation. Canthus rostralis weak, slightly concave. Loreal region weakly concave. Tympanum distinct, large, rounded, diameter about 75 % of eye diameter. Supratympanic fold distinct, beginning straight, with a rather distinct bend midway towards jaw. Tongue ovoid, distinctly posteriorly bifid. Maxillary teeth present. Vomerine teeth present in two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Outer metacarpal tubercle present, inner metacarpal tubercle present. Fingers without webbing. Relative length of fingers: IVariation.—Variation in measurements is given in Table 6. See Figs 26–27 for colouration in life and its variation. Specimens with a light vertebral stripe occur. There is moderate sexual size dimorphism (confirmed male SVL 31.7–41.0 mm [n = 11] vs confirmed female SVL 36.7–46.4 mm [n = 12]). Males have a larger tympanum than females (HTD/ED ratio is 53–77% in females, 71–113% in males).Femoral glands in males large and distinct, orange coloured in life, with the proximal granular gland field sometimes occupying a larger surface than the distal ulcerous macrogland (Fig. 26o) which is relatively unusual in mantellines; the proximal granular gland field on the two opposite thighs contact each other medially. Rasolonjatovo et al. (2022) describe the genetic diversity and phylogeographic structure of M. bellyi on Montagne d’Ambre. Natural history.—Similar to M. ulcerosus, M. bellyi is found in and along small shallow streams in primary as well as degraded rainforest and dry forest. Males were heard calling at night from the water or at the edge of the water. Rasolonjatovo et al. (2018) report on the attempted predation of a Boophis tadpole by an adult M. bellyi and reported an adult female M. bellyi repeatedly emitting a series of rapid scratch-like vocalisations from a hidden place on the rough magmatic rock forming the edge of the pool. Rasolonjatovo et al. (2020) report on thermal ecology of M. bellyi on Montagne d’Ambre across its elevational range (467–1394 m a.s.l.), and Rasolonjatovo et al. (2022) described its genetic diversity and phylogeographic structure on Montagne d’Ambre. Calls.— The advertisement call of M. bellyi, recorded on 17 March 2000 at the entrance of Montagne d’Ambre National Park, 21.6°C air temperature (Vences et al. 2006: CD 2, track 75), consists of a regularly pulsed note (Fig. 28), emitted at irregular intervals, but not regular call series. Notes exhibited distinct amplitude modulation, with call energy being highest at the beginning of the note, followed by continuous decrease of energy towards the note’s end. Pulses were very narrowly spaced. Numerical parameters of three analysed calls were as follows: call duration (= note duration) 594–682 ms (630.0 ± 46.1 ms); 41–46 pulses per note (43.7 ± 2.5); pulse duration 7–11 ms (8.3 ± 1.2 ms); pulse repetition rate within notes 63.8– 77.8 pulses/s (69.9 ± 4.9); dominant frequency 2497–2583 Hz (2532 ± 32 Hz), with a second peak of almost identical energy at around 880–920 Hz; prevalent bandwidth 660– 3200 Hz; call repetition rate (= note repetition rate) not identifiable with available recordings. Variation in the advertisement call over populations of M. bellyi on Montagne d’Ambre was investigated by Rasolonjatovo et al. (2022) for 23 male specimens. In all cases, the calls typically consisted of a single note, with minor variation of call duration between individuals, and without significant differences in call parameters between sites. Dominant frequency was negatively correlated with body size. Tadpoles. —The tadpole of this species has not been described. Distribution. —Distributed in the North and North East of Madagascar; apparently allopatrically distributed with respect to its sister species, M. ulcerosus (Fig. 7). In Montagne d’Ambre National Park (the type locality) it is well documented from across the whole elevational range of the mountain. Additionally, our genetic results confirm its presence in Ankarana, Fanambana forest, Andrakata, Marojejy, Montagne des Français, Belambo, Andapa, and Andrafainkona/Ambarata. Elevation range: 53–1372 m a.s.l. Etymology.— Eponym for a ‘Mr Belly’ according to the original description, who collected the type specimens. This probably refers to Édouard Belly, to whom Charles A.Alluaud referred as ‘adjoint à ma mission par le Muséum de Paris’ for his travels in northern Madagascar (Alluaud 1893)., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 205-206, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Guibe, J. (1978) Les batraciens de Madagascar. Bonner zoologische Monographien, 11, 1 - 140.","Guibe, J. (1950) Catalogue des Types d'Amphibiens du Museum National d'Histoire Naturelle. Imprimerie Nationale, Paris, France.","Vences, M., Glaw, F. & Marquez, R. (2006) The Calls of the Frogs of Madagascar. 3 Audio CD's and booklet. Madrid, Spain, Fonoteca Zoologica, 44 pp.","Rasolonjatovo, S. M., Scherz, M. D., Schmidt, R., Glos, J., Rakotoarison, A., Raselimanana, A. P. & Vences, M. (2022) Population diversification in the frog Mantidactylus bellyi on an isolated massif in northern Madagascar: genetic, morphological, bioacoustic and ecological evidence. PLoS One, 17, e 0263764. https: // doi. org / 10.1371 / journal. pone. 0263764","Rasolonjatovo, S. M., Scherz, M. D., Raselimanana, A. P. & Vences, M. (2018) Tadpole predation by Mantidactylus bellyi Mocquard, 1895 with brief description of the site and morphological measurements of the specimen. Herpetology Notes, 11, 747 - 750.","Rasolonjatovo, S. M., Scherz, M. D., Rakotoarison, A., Glos, J., Raselimanana, A. P. & Vences, M. (2020) Field body temperatures in the rainforest frog Mantidactylus (Brygoomantis) bellyi from northern Madagascar: Variance and predictors. Malagasy Nature, 14, 57 - 68.","Alluaud, C. A. (1893) Correspondance. Compte rendu des Seances de la Societe de Geographie et de la Commission centrale, 14 (Numero Supplementaire), 355."]}

Mantidactylus tricinctus (Guibé, 1947) Type material.— Gephyromantis tricinctus Guibé, 1947 was originally described based on two syntypes: MNHNP 1931.26‒27. As discussed by Glaw and Vences (1999), Guibé (1947) considered the female MNHN 1931.26 as ‘gynétype’ and the male MNHN 1931.27 as ‘androtype’ (see Frizzell 1933) whereas the other four specimens of the original series of G. tricinctus, containing the following specimens: MNHN 1931.26A (relabelled MNHN 1994.611), and MNHN 1931.26B (relabelled MNHN 1994.612) from Befotaka; and MNHN 1931.27A (relabelled MNHN 1994.613), and MNHN 1931.27B (relabelled MNHN 1994.614)from Vondrozo, all collected by R. Decary, were expressly indicated as ‘paratypes’ and thus are not name-bearing specimens. Guibé (1950) considered the ‘gynétype’ and the ‘androtype’ each as holotype, but as the nomen then was based on two specimens, these have to be considered syntypes. The nomen is currently based on lectotype MNHN 1931.26 designated by Blommers-Schl̂sser and Blanc (1991). Identity.— Glaw and Vences (1999) resurrected M. tricinctus which was previously considered a synonym of M. biporus (see Blommers-Schl̂sser & Blanc 1991) based on new material from An’Ala. The genetic data presented herein suggest three morphologically similar deeply divergent mitochondrial lineages that conform with the lectotype (and paralectotypes) of tricinctus in their morphological characters (e.g. small body size, reduced webbing). Our attempts of barcode fishing from the lectotype was unsuccessful and the few 16S reads obtained were inconclusive, probably contaminated with Homo sapiens reads. However, we succeeded in PCRamplifying and sequencing 16S from a specimen (ZSM 176/2006 = MVTIS 16559) from the Midongy du Sud National Park (Bora et al. 2007) (also known as BefotakaMidongy), and thus presumably from very close to the type locality. Based on the phylogenetic position of this sample, we circumscribe M. tricinctus to a lineage known from Midongy, Manombo Special Reserve, and Ambahavala in the Anosy Mountains, and consider the lineages from the southernmost Southern Central East, and from the Northern Central East, as two new species named below. Additional material. —ZSM 176/2006 (BOR 1066), adult male, and ZSM 177/2006, adult female, collected by P. Bora between September and October 2005 in BefotakaMidongy National Park (precise coordinates unavailable); ZSM 2377/2007 (ZCMV 5444), adult female, and ZSM 2415/2007 (ZCMV 5420), adult male, collected by M. Vences, G. Safarek, E. Rajeriarison, and T. Rajeriarison on 23 February 2007 1 km south of ‘site 2’, Manombo Special Reserve (precise coordinates not taken). Diagnosis.—A member of the M. tricinctus clade as revealed by the phylogenomic analysis, and sister to M. grubenmanni sp. nov. described below. See Table 4 for a list of diagnostic morphological characters. The combination of very small body size (below 20 mm male SVL), connected lateral metatarsalia, reduced webbing, presence of a light (often yellowish) marking on snout tip, a yellow inguinal marking, and a short, pulsed advertisement call emitted in rapid succession in regular series, readily distinguishes M. tricinctus from all other nominal species of Brygoomantis. Most similar to its sister species M. grubenmanni sp. nov. and to a lesser degree, to M. gudrunae sp. nov.; for comparisons, see the diagnoses of these species below. For detailed distinction from other new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. tricinctus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table 9. See Fig. 57 for colouration in life. There is weak sexual size dimorphism (confirmed male SVL 16.8–19.2 mm [n = 6] vs confirmed female SVL 18.0– 23.4 mm [n = 3]). Males have a larger tympanum than females (HTD/ED ratio is 59–73% in females, 73–105% in males). Femoral glands in males not very prominent, with a yellowish tone in life. Natural history.—At Manombo, males were heard emitting their advertisement calls from the shore of slowmoving streams in degraded rainforest. Calls.—The advertisement call of M. tricinctus, recorded on 23 February 2007 at Manombo, at an estimated air temperature of 25°C, consisted of a short, pulsed note, emitted in regular series at very fast succession (Fig. 58). Amplitude modulation was present, with relative amplitude increasing from the beginning of the call, reaching its maximum with the terminal pulse. Numerical parameters of 64 analysed calls were as follows: call duration (= note duration) 45–113 ms (83.4 ± 19.0 ms); 8–19 pulses per note (15.5 ± 3.6); pulse duration 2–5 ms (2.8 ± 1.1); pulse repetition rate within notes 147.1–214.3 pulses/s (177.2 ± 27.7); dominant frequency 2787–2906 Hz (2851 ± 59 Hz), with a second peak of almost equal energy at ca 2100–2200 Hz; prevalent bandwidth 1400–5300 Hz; call repetition rate (= note repetition rate) within regular series ca 470– 580 calls/min. Call series consist of 35–42 calls (n = 3), but calls were also emitted isolated (possibly territorial function) or in short groups containing 2–3 calls. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to the South East (Fig. 7). This species is known from Ambahavala, Midongy du Sud/Befotaka-Midongy National Park (type locality), and Manombo. Elevation range: 30–900 m a.s.l. Etymology.—Latin adjective formed from the stems ‘tri’ meaning ‘three’ and ‘cinctus’ meaning ‘crowned’ or ‘girded’, presumably in reference to the colour pattern., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 253-255, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Guibe, J. (1947) Trois Gephyromantis nouveaux de Madagascar (Batraciens). Bulletin du Museum National d'Histoire Naturelle, Paris, Serie 2, 19, 151 - 155.","Vences, M. & Glaw, F. (1999) Variation in Mantidactylus madecassus Millot & Guibe, 1950, a little known Malagasy frog, with resurrection of Mantidactylus pauliani Guibe, 1974. Herpetological Journal, 9, 101 - 110.","Frizzell, D. L. (1933) Terminology of types. The American Midland Naturalist, 14, 637 - 668. https: // doi. org / 10.2307 / 2420124","Guibe, J. (1950) Catalogue des Types d'Amphibiens du Museum National d'Histoire Naturelle. Imprimerie Nationale, Paris, France.","Bora, P., Otisitraka Randriambahiniarime, M., Rabemananjara, F. C. E., Ravoahangimalala Ramilijaona, O., Glaw, F. & Vences, M. (2007) A rapid assessment survey of the herpetofauna at Befotaka-Midongy National Park, south-eastern Madagascar. Mitteilungen aus dem Museum f ¸ r Naturkunde in Berlin - Zoologische Reihe, 83, 170 - 178. https: // doi. org / 10.1002 / mmnz. 200700007"]}

Mantidactylus fergusoni sp. nov. Identity and justification.—This lineage has been previously considered as confirmed candidate species M. sp. 26 by Vieites et al. (2009) and M. sp. Ca26 by Perl et al. (2014). It was depicted as ‘ Mantidactylus sp. aff. betsileanus “Andranofotsy”’ by Glaw and Vences (2007). Our phylogenomic analysis confirms that this lineage is in a major clade (here named the M. fergusoni clade) containing only scientifically unnamed lineages, and its status as distinct species is therefore out of question. However, deciding how many species exist in this clade is complicated. Mantidactylus sp. Ca26 forms a clade with several other divergent mitochondrial lineages of uncertain status; here we consider the lineage previously named M. sp. 21 (Vieites et al. 2009) or M. sp. Ca21 (Perl et al. 2014) from Nosy Mangabe, Makira, Masoala, and Cap Est/Ambato as conspecific with M. fergusoni (note that M. sp. 21 was wrongly placed in the tree of Vieites et al. [2009], probably due to a sequence confusion in the alignment used for phylogenetic analysis). As a further deep conspecific lineage we consider samples from Marojejy and additional sites in the North East. Samples from Nosy Boraha (M. sp. Ca27) according to our phylogenomic analysis (Fig. 5) form a clade with the lineage M. sp. Ca25 (see Vieites et al. 2009) from different sites on the mainland adjacent to Nosy Boraha, and together are sister to M. fergusoni sp. nov.; the Nosy Boraha population is described below as M. jahnarum sp. nov. based on its bioacoustic differentiation. Holotype.— ZSM 126/2002 (MV 2001.1389), adult male, collected by M. Vences on 17 December 2001 at Andranofotsy (15.4353°S, 049.8439°E, 85 m a.s.l.), Analanjirofo Region, Madagascar. A 16S barcode sequence of the holotype is available from GenBank (accession AY848214). Paratypes.— A total of six paratypes: ZSM 124/2002 (MV 2001.1433), female, ZSM 125/2002 (MV 2001.1434), adult male, and ZSM 176/2002 (FGMV 2001.1383), specimen of unknown sex and maturity, all with the same collection data as the holotype; ZSM 5050/2005 (ZCMV 2125), adult female, and ZSM 5051/2005 (ZCMV 2136), adult male, collected by F. Glaw, M. Vences, and R.D. Randrianiaina on 22 February 2005 on Nosy Mangabe (ca 15.50°S, 049.77°E, 50–100 m a.s.l.); ZSM 355/2010 (FGZC 4277), adult female, collected by F. Glaw, J. K̂hler, P.-S. Gehring, M. Pabijan, F.M. Ratsoavina on 3 April 2010 at Ambodivoahangy (15.2899°S, 049.6203°E, ca 100 m a.s.l.). Additional material.— The following three specimens are assigned to M. fergusoni sp. nov. but come from a genetically divergent population and have in part (ZFMK specimens) not been sequenced, and therefore they are not included in the paratype series: ZSM 201/2005 (FGZC 2746), adult male, collected by F. Glaw, M. Vences, and R.D. Randrianiaina on 14 February 2005 at Marojejy, Camp 1 ‘Mantella’ (14.4377°S, 049.7756°E, 481 m a.s.l.); ZFMK 59938, adult male, and ZFMK 59939, adult female, collected by F. Glaw and O. Ramilison on 22 February 1995 at Marojejy (near Camp Mantella). Diagnosis.—A member of the M. fergusoni clade as revealed by the phylogenomic analysis, and sister to M. jahnarum sp. nov. described below. See Table 4 for a list of diagnostic morphological characters. The combination of a small to moderate body size in males (SVL up to 25– 30 mm) and distinctly larger body size in females (36– 42 mm), tubercular dorsal skin, large tympanum size in males (10–14% of SVL), absence of white spots on flanks and of white marking on snout tip, and advertisement call consisting of a single-pulse note distinguishes M. fergusoni sp. nov. from most species of the other clades. Two species from the M. ulcerosus clade (M. ulcerosus and M. bellyi) can be morphologically similar, but they occur in the Sambirano and North West regions, and have strongly differing advertisement calls (Table 4). One species of the M. betsileanus clade (M. katae) has an advertisement call of similar general structure, but has a faster and more regular call repetition rate, less tubercular dorsum, larger femoral glands, and a distinct white marking on snout tip (Table 4). M. fergusoni sp. nov. may also show superficial similarities to other species of the M. betsileanus clade but does not appear to occur sympatrically with any of them; in general it has a more tubercular dorsum and differs from all of these by advertisement call structure (Table 4). The M. fergusoni clade contains only species newly named herein; for a distinction from these other species, see below. A full list of molecular diagnostic sites in the 16S gene of M. fergusoni sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype.—Adult male in good state of preservation (Fig. 48). Part of left shank muscle removed as tissue sample, femoral glands partly detached for examination in internal view. Body slender. Head wider than body. Snout rounded.Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis very weak, slightly concave. Loreal region concave. Tympanum distinct, large, slightly wider than high, its horizontal diameter about 91% of eye diameter. Supratympanic fold present, beginning straight, with a rather distinct bend midway towards jaw / forelimb insertion. Tongue ovoid, distinctly bifid. Maxillary teeth present. Vomerine teeth present in two rounded to ovoid aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: I≤II Relative length of toes: I Colour in preservative: dorsum brown with a minimal reddish shade. Forelimbs and hindlimbs with distinct brown crossbands. Venter beige, chest and throat rather dark brown with distinct light markings and a central median line on throat. Colour in life was similar to preservative, but with a distinct reddish-brown colour dorsally. Variation.—Variation in measurements is given in Table 8. See Fig. 49 for colouration in life and its variation. There is pronounced sexual size dimorphism (confirmed male SVL 24.9–29.9 mm [n = 5] vs confirmed female SVL 35.7–42.2 mm [n = 4]). Males have a somewhat larger tympanum than females (HTD/ED ratio is 69–67% in females, 67–96% in males). Femoral glands are distinct but not particularly prominent in males, and only with a slight yellowish tone in life; very small, rudimentary glands recognisable in females. Natural history.—Habitat and habits of this species are poorly known, but so far it has been found in primary or somewhat degraded rainforest. At Andranofotsy, calling males were observed at night from the shore of a shallow puddle next to a small spring in rainforest. Calls.—The advertisement call of M. fergusoni, recorded on 16 December 2001 near Andranofotsy, 25.4°C air temperature (Vences et al. 2006, CD 2, track 69), consists of a simple, short, single pulse ‘click’ note, emitted in series (Fig. 50). Inter-call intervals were irregular with some calls (= notes) occurring in pairs with lower inter-call interval inbetween. Call energy was distributed in a wide frequency band. Numerical parameters of 13 analysed calls were as follows: call duration (= note duration) 9–12 ms (10.5 ± 0.8 ms); 1 pulse per note (1.0 ± 0.0); pulse duration = note duration = call duration; dominant frequency 1626–1690 Hz (1669 ± 26 Hz); prevalent bandwidth 700–5700 Hz; call repetition rate (= note repetition rate) within series ca 210–290 calls/min. Calls recorded on 31 March 2010 at Ambodivohangy, Makira area (from specimen ZSM 361/2010 = FGZC 4219), 24°C estimated air temperature, agree with those described above from Andranofotsy, apart from somewhat lower call repetition rate within series. Calls consisted of a simple short ‘click’ note emitted isolated or more often in series at irregular intervals. Numerical parameters of 28 analysed calls were as follows: call duration (= note duration) 7–13 ms (8.9 ± 1.4 ms); 1 pulse per note (1.0 ± 0.0); pulse duration = note duration = call duration; dominant frequency 1571–1722 Hz (1638 ± 54 Hz); prevalent bandwidth 700–5500 Hz; call repetition rate (= note repetition rate) within series ca 100–170 calls/min. Tadpoles.— The tadpole of this species has not been described. Voucher Field number Sex Locality SVL HW HL ED HTD END NSD NND FORL HAL HIL FOTL FOL TIBL FGL FGW M. fergusoni sp. nov. (Ca26) ZSM 126/2002 [HT]FGMV 2001.1389MAndranofotsy27.810.311.64.44.03.01.92.817.58.748.722.515.114.53.62.6ZFMK 59938 #NAMMarojejy, Camp Mantella26.010.69.73.92.62.91.82.98.98.0NMNM13.414.52.82.2ZSM 125/2002 [PT]FGMV 2001.1434MAndranofotsy26.210.111.24.23.12.42.03.115.87.741.819.113.313.43.92.4ZSM 201/2005 #FGZC 2746MMarojejy, Camp Mantella24.99.910.04.13.32.71.82.710.27.9NMNM12.314.03.42.3ZSM 5051/2005 [PT]ZCMV 2136MNosy Mangabe29.911.412.84.13.63.52.43.317.68.545.621.014.014.83.72.5ZFMK 59939 #NAFMarojejy, Camp Mantella35.714.713.45.42.93.82.14.012.710.4NMNM16.319.6NANAZSM 124/2002 [PT]FGMV 2001.1433FAndranofotsy36.113.614.55.63.63.92.33.122.510.260.228.319.418.9NANAZSM 355/2010 [PT]FGZC 4277FMakira40.315.216.35.53.84.12.43.924.611.866.530.021.221.2NANAZSM 5050/2005 [PT]ZCMV 2125FNosy Mangabe42.215.416.76.04.04.22.73.824.411.969.430.621.021.7NANA M. georgei sp. nov. (Ca35/Ca36) ZSM 455/2005 [HT]ZCMV 806MToamasina30.711.912.44.94.73.21.93.517.48.046.220.614.113.74.03.3ZSM 454/2005ZCMV 803MMaroantsetra29.010.912.04.63.73.31.83.417.48.646.920.313.913.84.02.4ZSM 456/2005 [PT]ZCMV 807MToamasina28.410.511.94.44.22.91.83.516.38.446.320.413.813.54.63.3 M. jahnarum sp. nov. (Ca27) ZSM 423/2006 [HT]ZCMV 3390MNosy Boraha30.311.112.64.73.33.11.83.217.28.946.121.114.014.44.83.0... Continued on the next page Distribution.— Endemic to low-elevations of Northern Central East and North East (Fig. 7). This species is known from Ambato, Ambinanifaho, Ambodirafia Tokana, Ambodiriana, Ambodivohangy (Makira), Ampasimazava, Andranofotsy (type locality), Andrantambe, Antanambe, Antsahanoro, Befanjana, Belambo, various sites within Marojejy National Park, Masoala, Nosy Mangabe, and the Sambava region. Elevation range: 12–1326 m a.s.l. Etymology.—We dedicate this species to Barry Ferguson, an outspoken worker on environmental justice for Madagascar, in acknowledgement of his support of the first author’s work on Madagascar’s herpetofauna.

Mantidactylus jahnarum sp. nov. Identity and justification.—This lineage of the M. fergusoni clade has previously considered as confirmed candidate species M. sp. 27 by Vieites et al. (2009), and M. sp. Ca27 by Perl et al. (2014). It was depicted as ‘ Mantidactylus sp. aff. betsileanus “Nosy Boraha”’ by Glaw and Vences (2007). The phylogenomic data place it sister to M. fergusoni, and its mitochondrial divergence from M. fergusoni (2.0–3.1%) is comparatively low, compared with other divergences observed among species of Brygoomantis. However, it distinctly differs by its advertisement calls (heard in multiple years from many individuals at the type locality, the islet Nosy Boraha) that consists of several short, pulsed notes, somewhat reminiscent of the call of species of the M. ulcerosus clade. We therefore here consider this lineage as the separate species M. jahnarum sp. nov. which, according to genomic data, may also include populations from the mainland adjacent to Nosy Boraha which were named M. sp. Ca25 by Vieites et al. (2009) and which we here provisionally include in M. jahnarum sp. nov. as deep conspecific lineage. Holotype.— ZSM 423/2006 (ZCMV 3390), adult male (call voucher), collected by M. Vences and J.E. Randrianirina on 7–8 March 2006 at Maromandia village on Nosy Boraha (16.9089°S, 049.8678°E, 20 m a.s.l.), Analanjirofo Region, Madagascar. 16S and cox1 barcode sequences of the holotype are available from GenBank (accessions FJ559258 and JN133220). Paratypes.—A total of three paratypes: ZSM 424/2006 (ZCMV 3393), adult male, with same collection data as holotype; ZSM 520/2006 (ZCMV 3218) and ZSM 521/2006 (ZCMV 3219), two adult females, collected by M. Vences and J.E. Randrianirina on 7–8 March 2006 in a forest several km from Maromandia village (coordinates not taken), Nosy Boraha (= Île Sainte Marie). Diagnosis.— Mantidactylus jahnarum sp. nov. is a member of the M. fergusoni clade as revealed by the phylogenomic analysis, and is the sister species of M. fergusoni. See Table 4 for a list of diagnostic morphological characters.The combination of a moderate body size (male SVL 29–30 mm, female SVL 30–34 mm), moderately tubercular dorsal skin, moderate tympanum size in males (11% of SVL), absence of white spots on flanks, presence of a white marking on the snout tip of many individuals (especially males), and regularly pulsed advertisement call emitted in regular series distinguishes M. jahnarum sp. nov. from most species of the other clades (Table 4). Two species from the M. ulcerosus clade (M. ulcerosus and M. bellyi) can be morphologically similar, but they occur in the Sambirano and North West regions, and have higher pulse repetition rates in advertisement calls (Table 4). Mantidactylus jahnarum sp. nov. shows similarities to species of the M. betsileanus clade but does not appear to occur sympatrically with any of them; in general, it has a more tubercular dorsum and differs from most of these by advertisement call structure (Table 4). Within the M. fergusoni clade, it differs from M. fergusoni and M. georgei in advertisement call structure: M. fergusoni has single-pulse calls repeated 2–5 times per second, whereas M. georgei does not arrange calls in regular series. For a distinction from other new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. jahnarum sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype.—Adult male in good state of preservation (Fig. 48). Part of right thigh muscle removed as tissue sample; femoral glands partly detached for examination in internal view. Body relatively slender. Head as wide as body. Snout rounded. Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis weak, rather straight. Loreal region concave. Tympanum distinct, large, slightly wider than high, its horizontal diameter about 75% of eye diameter. Supratympanic fold present, beginning straight, with a rather distinct bend midway towards jaw / forelimb insertion. Tongue ovoid, distinctly bifid. Maxillary teeth present. Vomerine teeth present in two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: I≤IIVariation.—Variation in measurements is given in Table 8. See Fig. 53 for colouration in life and its variation. There is moderate sexual size dimorphism (confirmed male SVL 28.5–30.3 mm [n = 2] vs confirmed female SVL 30.4–33.6 mm [n = 2]). Also, sexual dimorphism in relative tympanum size seems to be absent (HTD/ED ratio is 62– 73% in females, 69–70% in males). Femoral glands in life are rather distinct and prominent in males, with a large distal ulcerous macrogland, often with a somewhat yellowish colour but still with some of the dark pigmentation found elsewhere on the ventral side of the thigh. Natural history.—Similar to M. georgei, this species appears to tolerate heavy habitat degradation. At Nosy Boraha, it can be found next to streams in rainforest, but calling males can also be heard at night from slowly running, shallow water bodies in highly degraded forest, including plantations and swamps next to villages shaded by some larger trees. Calls.— The advertisement call of M. jahnarum, recorded on 7 March 2006 at Maromandia village, Nosy Boraha, from the holotype, at an at 25°C, consisted of a regularly pulsed note of variable duration (Fig. 54), emitted in series at regular intervals. Pulses were very short in duration. Regular call series seemed to consist of 4–6 calls, with the first call of the series being of longest duration. Notes exhibited slight amplitude modulation, with maximum call energy occurring shortly after the beginning of the note, continuously decreasing towards the note’s end. Numerical parameters of 20 analysed calls were as follows: call duration (= note duration) 568–1558 ms (976.9 ± 277.9 ms); 16–45 pulses per note (29.1 ± 7.9); pulse duration 1–2 ms (1.1 ± 0.3 ms); pulse repetition rate within notes 25.0–35.4 pulses/s (29.3 ± 3.9); dominant frequency 2971–3120 Hz (2999 ± 60 Hz); prevalent bandwidth 1000–3400 Hz; call repetition rate (= note repetition rate) within regular series ca 36-47 calls/min. Tadpoles.— The tadpole of this species has not been described. Distribution.— Apparently microendemic to the islet of Nosy Boraha (Fig. 7), although a related lineage provisionally assigned to this species occurs on the adjacent mainland. Elevation range: ~ 20 m a.s.l. Etymology.—We dedicate this species to our microbiologist colleagues Martina and Dieter Jahn, in recognition of their support of biodiversity research at the University of Technology in Braunschweig. We intentionally deviate from Article 31.1.2 of the Code and form the name using the feminine plural ending -arum, and not the masculine plural ending, -orum, with the intention of drawing attention to the already overly maledominated taxonomic nomenclature, and the desire for a more egalitarian declension. To stabilise the nomen as coined here, we define it as a noun in apposition., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 247-250, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 496 pp. Third Edition."]}

Mantidactylus ambohimitombi ambohimitombi Boulenger, 1919 Type material.— Mantidactylus ambohimitombi Boulenger, 1919 is based on a series of syntypes that include BMNH 1947.2.26.25–32 from ‘ Ambohimitombo Forest’. Blommers-Schl̂sser and Blanc (1991) noted that syntypes BMNH 1947.2.26.31– 32 may be referable to Mantidactylus curtus, without justification. We here designate the DNA-barcoded specimen BMNH 1947.2.26.25, a large-sized individual of 65.0 mm SVL, as lectotype. Lectotype designation is justified by the need to stabilize this and other nomina in Brygoomantis, given the uncertain identity and morphological similarity of many taxa in the subgenus. Identity.— Mantidactylus ambohimitombi Boulenger, 1919 is typically considered a valid species (BlommersSchl̂sser & Blanc 1991; Frost 2021; Glaw & Vences 1992a, 1994), although it was considered of uncertain status by Glaw and Vences (2007). It is morphologically close to M. curtus but distinguished by its distinctly larger body size. It was not assigned to a genetic lineage by Vieites et al. (2009). A 16S sequence of the lectotype surprisingly clusters with a lineage predominantly known from the Ankaratra Massif. We therefore redefine this lineage provisionally as corresponding to M. ambohimitombi. In previous studies the populations from the Ankaratra Massif have been considered as confirmed candidate species M. sp. 19 by Vieites et al. (2009), and M. sp. Ca19 by Perl et al. (2014). They were referred to as ‘ M. sp. aff. curtus “Ankaratra”’ by Schmidt et al. (2009). We emphasize that this attribution is preliminary; since only mtDNA data (no genomic information) are available from the lectotype, and no fresh samples are available from Ambohimitombo forest where the original syntype 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 M. curtus into M. ambohimitombi marefo ssp. nov. from Itremo (Fig. 5), 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 M. ambohimitombi. A more in-depth analysis of ranges, and of gene flow among various lineages of the M. curtus clade (M. curtus, M. bourgati, M. ambohimitombi) is necessary to understand their evolutionary history and verify their taxonomy. In our phylogenomic tree, two other lineages form a monophyletic group with specimens of M. ambohimitombi from Ankaratra, and these are in subsequent accounts described as subspecies of M. ambohimitombi; see the rationale in the respective accounts below. Diagnosis.— Mantidactylus ambohimitombi is a member of the M. curtus clade; it is here defined as containing three deep genetic lineages considered as subspecies, and is sister to the morphologically very distinct M. madecassus. See Table 4 for a list of diagnostic morphological characters. The following account only diagnoses the nominal subspecies M. a. ambohimitombi from other species in Brygoomantis; see below for diagnoses and comparisons of the two other subspecies. The combination of relatively large body size of up to 51 mm at Ankaratra (up to 65 mm in the type 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 M. a. ambohimitombi from species of the other clades. Within the M. curtus clade, M. alutus, M. madecassus and M. pauliani have smaller body sizes and are distinguished by either a usually shorter snout (M. madecassus, M. pauliani), or advertisement calls emitted in regular series (in M. alutus, vs single notes) (Table 4); M. curtus usually has a smoother skin and a somewhat shorter snout; M. bourgati 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 M. a. ambohimitombi in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table 5. See Fig. 13 for colouration in life and its variation. 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 50.6 mm). As discussed above, the type series is comprised of particularly large-sized animals reaching 65.0 mm in SVL. Femoral glands are only distinct in some male specimens, possibly due to seasonal effects; in two specimens, ZSM 190/2021 (ACZCV 334) and ZSM 195/2021 (ACZCV 342), FGL and FGW are 4.7 mm x 3.8 mm and 4.0 mm x 3.0 mm, respectively. Natural history. — 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 Vences et al. 2002 under the name M. curtus). Males were found calling during the day underwater. Calls.— The advertisement call of M. a. ambohimitombi, recorded on 16 February 2006, 14:50 h, at Ankaratra, consists of a short, pulsed note (Fig. 14), 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. Tadpoles. —A tadpole of M. a. ambohimitombi was described under the name ‘ M. sp. aff. curtus “Ankaratra”’ by Schmidt et al. (2009). Distribution.— Apparently endemic to a small area of the central highlands of Madagascar (Fig. 7). 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 M. curtus, which is common in this locality. The type locality Ambohimitombo forest is close to Antoetra. Elevation range: 1150–2380 m a.s.l. Etymology.—Formulated from the type locality, ‘Ambohimitombo Forest’., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 178-182, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["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","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","Glaw, F. & Vences, M. (1992 a) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 335 pp. First Edition.","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 478 pp. Second Edition.","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 496 pp. Third Edition.","Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Schmidt, H., Strauss, A., Glaw, F., Teschke, M. & 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","Vences, M., Andreone, F., Glaw, F., Raminosoa, N., Randrianirina, J. E. & 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"]}

Mantidactylus alutus (Peracca, 1893) Type material.— Rana aluta Peracca, 1893 is based on 25 syntypes according to the original description, 14 of the colour morph ‘forma’A and 11 of the ‘forma’ B.According to Frost (2021), Rana aluta Peracca, 1893 includes the following syntypes: MZUT An725 and An729, MNHN 1894.1–2, and specimens in BMNH, all from ‘dintorni di Andrangoloaka e dalla vicina valle dell’Umbi’. However, Gavetti and Andreone (1993) designated MZUT An725.1 as lectotype and redescribed this specimen including morphological measurements (summarised in Table 4). They listed 11 paralectotypes of colour morph A (MZUT An725.2–12) and 13 of colour morph B (MZUT An729), resulting in a total of 25 type specimens, although their numbers of individuals attributed to both colour morphs differ from the original description (Peracca 1893). However, since the total number of type specimens is in accordance with the number mentioned by Peracca (1893) the claim by Blommers-Schl̂sser and Blanc (1991) concerning the presence of two other syntypes at Paris (MNHN 1894.1 and 1894.2) does not seem to be correct (Gavetti & Andreone 1993: 106) and the same must be assumed for the specimens claimed by Boulenger (1895 ‘1894’) to have been received by the BMNH from Peracca. An additional non-type specimen is MZUT An917 from Andrangoloaka (Gavetti & Andreone 1993), demonstrating that more than the 25 types were available in the MZUT collection. The MNHN and BMNH specimens are therefore not paralectotypes. Identity.—In this study we obtained genetic data via barcode fishing from specimen MNHN 1894.1 from Andrangoloaka, marked as ‘type’ of M. alutus in the MNHN catalogue and provided by M.G. Peracca according to the MNHN catalogue, but, as discussed above, probably not representing one of the paralectotypes. The 16S sequence of this specimen clustered among specimens from the central highlands of Madagascar that are typically considered as M. alutus (BlommersSchl̂sser 1979; Blommers-Schl̂sser & Blanc 1991; Glaw & Vences 1992a, 1994, 2007). Although this information does not refer to the lectotype of the species, little doubts thus remain that the nomen M. alutus has been correctly applied to this small-sized lineage from the central highlands. ...Continued on thte next page ...Continued on thte next page ) ...Continued on thte next page ) ...Continued on thte next page ...Continued on thte next page ) Synonyms.— The taxon Mantidactylus laevis Angel, 1929 was listed as a dubious species by Guibé (1978) and considered a synonym of M. alutus by Glaw and Vences (1992a). The holotype MNHN 1929.208, collected by G. Petit from the vicinity of Antananarivo (type locality ‘environs de Tananarive’; SVL 32 mm according to the original description) was reported to be lost according to Guibé (1978), and this has been confirmed by S. Grosjean (pers. comm. to A. Ohler, 22 January 2022). Because M. alutus is the only species of Brygoomantis occurring in the Antananarivo area, little doubts remain that this synonymy is correct. In order to stabilize synonymies and as the holotype has been lost for more than 40 years, we here designate the lectotype of Rana aluta, MZUT An 725.1 as the neotype of Mantidactylus laevis Angel, 1929. Diagnosis.—A member of the M. curtus clade and sister to a monophyletic group of all other species of the clade. See Table 4 for a list of diagnostic morphological characters. The combination of relatively small body size of up to 31 mm, slightly granular skin with (weakly expressed) dorsolateral ridges, small tympanum diameter of ca 9% of SVL in males, absence of small white spots on flanks, presence of a light frenal stripe in most specimens, and advertisement calls as a regular series of short pulsed notes distinguishes M. alutus from species of the other clades. Within the M. curtus clade, M. ambohimitombi, M. bourgati and M. curtus have larger body sizes, and M. madecassus and M. pauliani are high-elevation endemics with usually a shorter snout and absence of dorsolateral ridges (Table 4). 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 M. alutus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table 5. See Fig. 11 for colouration in life and its variation. Some individuals have a light vertebral stripe. There is moderate sexual size dimorphism (confirmed male SVL 23.4–27.2 mm [n = 2] vs confirmed female SVL 31.0 mm [n = 2]). Males with large and distinct femoral glands (e.g. Fig. 11b); in MNHN 1894.1 and 1894.2, FGL and FGW are 2.4 mm x 1.4 mm and 1.7 mm x 1.4 mm, respectively. The glands can be of orange/yellow colour in some individuals in life (e.g. Fig. 11b), which may be related to the reproductive state. Natural history.— Typically found in slow-flowing parts of streams or associated swamps in Madagascar’s highlands. Often at the edge of forest or in streams devoid of forest, with some gallery vegetation only. Males call from the edge or very shallow parts of water, from concealed positions during the day or from more open positions at night. See Vences et al. (2002) for observations from Ankaratra. This species is also abundant in parks of Antananarivo, e.g. in the Tsimbazaza garden (Glaw & Vences 2007). Calls.— The advertisement call of M. alutus, recorded on 21 January 2003, 17:30 h, near Antoetra, 20.5–21.0°C air temperature (Vences et al. 2006: CD 2, track 61, cut 1), consists of a short, regularly pulsed note, emitted in series at regular intervals (Fig. 12). Notes exhibit distinct amplitude modulation, with amplitude continuously increasing from the beginning, reaching maximum call energy at the middle of the note, before continuously decreasing towards the note’s end. Numerical parameters of eight analysed calls are as follows: call duration (= note duration) 149–290 ms (234.3 ± 53.5 ms); 20–27 pulses per note (24.1 ± 2.7); pulse duration 5–8 ms (6.3 ± 1.0 ms); pulse repetition rate within notes 83.3–130.4 pulses/s (108.2 ± 17.3); dominant frequency 1040–1116 Hz (1081 ± 26 Hz); prevalent bandwidth 600–4800 Hz; call repetition rate (= note repetition rate) within regular series ca 110 calls/min. Calls recorded on 1 January 1994, 18:20 h, at Manjakatompo, 18°C air temperature (Vences et al. 2006: CD 2, track 61, cut 2) generally agree in character with the calls described above. Calls of a series containing nine calls have the following parameters: call duration (= note duration) 254–310 ms (278.5 ± 16.8 ms); 21–27 pulses per note (23.7 ± 2.2); pulse duration 3–6 ms (4.9 ± 0.7 ms); pulse repetition rate within notes 66.7–120.0 pulses/s (83.2 ± 19.0); dominant frequency 1270–1378 Hz (1326 ± 45 Hz); prevalent bandwidth 800–5600 Hz; call repetition rate (= note repetition rate) within series ca 115 calls/min. Tadpoles.—The tadpoles of M. alutus were described by Blommers-Schl̂sser (1979) and Schmidt et al. (2009). Distribution.— Endemic to the central highlands of Madagascar (Fig. 7). This species is known from Antananarivo, Ankaratra, Ibity, Ambohitantely, Ranomafanakely,Antoetra, a swamp in the Alaotra region, Mantasoa, Forêt de Tampina, and Tsinjoarivo. Elevation range: 933–2090 m a.s.l. Etymology.— Probably derived from the Greek adjective ἄλουτος, meaning ‘unwashed’ or ‘speckled’, presumably in reference to the dorsal colour pattern., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 170-178, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Peracca, M. G. (1893) Descrizione di nuove specie di rettili e anfibi di Madagascar. Nota II (1). Bollettino dei Musei di Zoologia ed Anatomia comparata della R. Universita di Torino, 8, 1 - 16. https: // doi. org / 10.5962 / bhl. part. 27224","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","Gavetti, E. & Andreone, F. (1993) Revised catalogue of the herpetological collection in Turin University. 1. Amphibia. Cataloghi. Museo Regionale di Scienze Naturali. Torino 10: 1 - 187. Cataloghi X, Museo Regionale di Scienze Naturali, Torino, Italy.","Glaw, F. & Vences, M. (1992 a) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 335 pp. First Edition.","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 478 pp. Second Edition.","Angel, F. (1929) Materiaux de la Mission G. Petit a Madagascar. Description de trois Batraciens nouveaux appartenant aux genres Mantidactylus et Gephyromantis. Bulletin du Museum National d'Histoire Naturelle, Paris, Serie 2, 1, 358 - 362.","Guibe, J. (1978) Les batraciens de Madagascar. Bonner zoologische Monographien, 11, 1 - 140.","Vences, M., Andreone, F., Glaw, F., Raminosoa, N., Randrianirina, J. E. & 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","Vences, M., Wahl-Boos, G., Hoegg, S., Glaw, F., Oliveira, E. S., Meyer, A. & Perry, S. (2007) Molecular systematics of mantelline frogs from Madagascar and the evolution of their femoral glands. Biological Journal of the Linnean Society, 92, 529 - 539. https: // doi. org / 10.1111 / j. 1095 - 8312.2007.00859. x","Vences, M., Glaw, F. & Marquez, R. (2006) The Calls of the Frogs of Madagascar. 3 Audio CD's and booklet. Madrid, Spain, Fonoteca Zoologica, 44 pp.","Schmidt, H., Strauss, A., Glaw, F., Teschke, M. & 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"]}

Mantidactylus madecassus (Millot & Guibé, 1950) Type material.—The taxon Racophorus [misspelling of Rhacophorus in the original description] (Philautus) madecassus Millot and Guibé, 1950 is based on the lectotype MNHN 1953.246 from ‘ Andringitra: Cirque Boby (altitude: 2.520 métres)’, designated by Vences and Glaw (1999). The eight paralectotypes, all from the same locality, are numbered MNHN 1989.3590–3597. Identity.— Mantidactylus madecassus is a morphologically distinct and apparently microendemic species restricted to high elevations on the Andringitra Massif. Its identity has been unambiguously assessed by Vences and Glaw (1999) based on morphology, and is here confirmed by a 16S sequence obtained from the lectotype, MNHN 1953.246. Diagnosis.—A member of the M. curtus clade and sister to M. ambohimitombi from which it strongly differs morphologically. See Table 4 for a list of diagnostic morphological characters. The combination of a body size of 27–34 mm (Table 5), smooth dorsal skin without dorsolateral ridges, strongly expressed foot webbing with almost fully webbed fifth toe, small tympanum size in males (7–10% of SVL), vomerine teeth absent, and especially, the bilobed subarticular tubercles (unique in Mantidactylus madecassus and illustrated by Vences & Glaw 1999) distinguishes this species from species of the other clades in Brygoomantis. Within the M. curtus clade, this high-elevation endemic differs from all species by its double or rather bilobed subarticular tubercles, and from all species exept M. curtus, M. pauliani, and M. ambohimitombi marefo, by a conspicuously short snout in many specimens. The species is an endemic to high elevations on the Andringitra Massif, where at slightly lower elevations, also M.bourgati occurs;a distinction from that species can readily be achieved by the combination of bilobed subarticular tubercles, shorter snout, usually smaller size and smoother skin, as well as more uniform silvery-whitish ventral colour in M. madecassus. 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 M. madecassus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table 5. See Fig. 19 for colouration in life and its variation. There is weak sexual size dimorphism (confirmed male SVL 27.0– 29.8 mm [n = 2] vs confirmed female SVL 29.3–33.7 mm [n = 5]). For a more detailed discussion of this species’ morphology and a morphometric comparison to M. pauliani, see Vences and Glaw (1999). Relative tympanum size is larger in males than in females (Vences & Glaw 1999). Femoral glands in males include a distal ulcerous macrogland only, and rudimentary glands are also present in females. Natural history. —Specimens were found in small, cold, clear mountain streams at high elevations of the Andringitra Massif. They were sitting in the water, especially in shallow side puddles next to the canal-like streams, at night. Calls.—The call of this species has not been recorded. Tadpoles.— The tadpole of M. madecassus was described by Thomas et al. (2005). Distribution.— Apparently microendemic to high elevations on the Andringitra Massif (Fig. 7). Elevation range: ~ 2488 m a.s.l. Etymology.—Latin adjective referring to the occurrence of the species in Madagascar., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 189-190, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Millot, J. & Guibe, J. (1950) Les batraciens du nord de l'Andringitra (Madagascar). Memoires de l'Institut Scientifique de Madagascar, 4, 197 - 206.","Vences, M. & Glaw, F. (1999) Variation in Mantidactylus madecassus Millot & Guibe, 1950, a little known Malagasy frog, with resurrection of Mantidactylus pauliani Guibe, 1974. Herpetological Journal, 9, 101 - 110.","Thomas, M., Raharivololoniaina, L., Glaw, F., Vences, M. & Vieites, D. R. (2005) Montane tadpoles in Madagascar: molecular identification and description of the larval stages of Mantidactylus elegans, Mantidactylus madecassus, and Boophis laurenti from the Andringitra Massif. Copeia, 2005, 174 - 183. https: // doi. org / 10.1643 / CH- 03 - 293 R 2"]}

Mantidactylus bourgati Guibé, 1974 Type material.— Mantidactylus bourgati Guibé, 1974 is based on the holotype MNHN 1972.437 (given in error as 1972.427 in the original publication but correct in the handwritten catalogue) by original designation from ‘ Ambalamarovandana’, a site in the Andringitra Mountains, and seven paratypes (MNHN 1972.440, 1972.449, 1972.464, 1972.472, 1972.476, 1972.479, and 1972.491) Identity.— Mantidactylus bourgati has long been considered a synonym of M. curtus but was resurrected as a separate species by Glaw and Vences (2006) based primarily on strong genetic divergences. We here provide a 16S sequence obtained by barcode fishing from the holotype MNHN 1972.437 confirming the attribution of the lineage containing all samples of curtus -like stream frogs from the Andringitra Massif to this nomen. Diagnosis.—A member of the M. curtus clade and sister to M. curtus. See Table 4 for a list of diagnostic morphological characters. The combination of relatively large body size of at least up to 41 mm (Table 5; larger specimens observed in the field), slightly granular skin, strongly expressed foot webbing with almost fully webbed fifth toe distinguishes this species from species of the other clades. Within the M. curtus clade, M. alutus, M. madecassus and M. pauliani have smaller body sizes (Table 4). M. curtus has a very smooth dorsal skin while in M. bourgati the skin is usually somewhat granular. Morphological distinction from M. a. ambohimitombi and M. a. miloko is most difficult, but as far as known, the distributions of these taxa do not overlap with that of M. bourgati. M. bourgati differs further from M. alutus by longer advertisement calls emitted at irregular intervals (vs regular series in M. alutus). 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 M. bourgati in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation. —Variation in measurements is given in Table 5. See Fig. 17 for colouration in life and its variation. From our sample, it is unclear whether sexual size dimorphism exists in this species; in the specimens measured, male SVl is 33.0– 38.9 mm (n=2) and female SVL is 32.3–40.0 mm (n=13). Femoral glands in adult males in life are often clearly visible and prominent (Fig. 17) but in preserved specimens, a clear distinction between males and females (which probably also have visible gland rudiments) is often not possible without examination of gonads. Natural history.—Specimens of M. bourgati are common in and around small slow-flowing montane streams at high elevations in the Andringitra Massif, e.g. on the Andohariana Plateau at 2000 m a.s.l. where they can be found at the edge of the water. They do not seem to occur higher up the massif, e.g. at Cuvette Boby we found only M. madecassus. However, at somewhat lower elevations (e.g. Imaitso forest at ca 1500 m a.s.l.), M. bourgati is also common along large streams running through rainforest. Our call recordings were made from a swamp area next to such a stream where specimens were observed during the day in the shallow water. Calls.— The advertisement call of M. bourgati, has been recorded on 16 January 1994, 10:00 h, at Andringitra National Park, in a forest swamp area close to Ambalamarina, 19°C air temperature, but the calling specimens could not be seen, and the attribution to M. bourgati is tentative (Vences et al. 2006: CD 2, track 77). The call consists of a regularly pulsed note(Fig.18),emitted in series at somewhat irregular intervals. Notes exhibit some amplitude modulation, with amplitude continuously increasing from the beginning, reaching maximum call energy at the middle of the note, before continuously decreasing towards the note’s end. Numerical parameters of four analysed calls are as follows: call duration (= note duration) 756–1020 ms (872.8 ± 112.6 ms); 36–50 pulses per note (43.8 ± 5.9); pulse duration 8–10 ms (8.7 ± 0.8 ms); pulse repetition rate within notes 45.8–54.1 pulses/s (50.8 ± 4.1); dominant frequency 1063–1106 Hz (1087 ± 18 Hz); prevalent bandwidth 880–1740 Hz; call repetition rate (= note repetition rate) within series ca 19–29 calls/min. Tadpoles.— The tadpole of this species has not been described. Distribution.— Apparently microendemic to Andringitra and nearby areas (Fig. 7). This species is known from Andringitra (various sites, type locality), Belambo, Fivahona, and Tsaranoro. Elevation range: 940–2488 m a.s.l. Etymology.—Eponym for ‘Professor [Robert M.] Bourgat of the University of Lomé (Togo), who collected several frogs during his voyages in Madagascar’(translated from Guibé 1973a)., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 186-189, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Vences, M., Glaw, F. & Marquez, R. (2006) The Calls of the Frogs of Madagascar. 3 Audio CD's and booklet. Madrid, Spain, Fonoteca Zoologica, 44 pp.","Guibe, J. (1973 a) Batraciens nouveaux de Madagascar. Bulletin du Museum National d'Histoire Naturelle, Paris, Serie 3, 145, 1009 - 1017."]}

Mantidactylus ulcerosus clade This clade contains two species characterized by a rather stout morphology and medium to large body sizes (28.5– 46.4 mm adult SVL), occurring in the North, Sambirano, and North West regions, and from some sites in the North East (M. ulcerosus, M. bellyi) as well as two small-sized species (17.3–28.9 mm adult SVL) apparently restricted to the Tsaratanàna and Manongarivo Massifs in the Sambirano Region (M. schulzi, and one new species named herein as M. steinfartzi sp. nov., based on the holotype depicted in Fig. 23). Mantidactylus ulcerosus (Boettger, 1880) Type material.—The taxon Limnodytes ulcerosus Boettger, 1880 is based on a syntype series from ‘ insula Nossi-Bé’ that according to Frost (2021) included SMF 1068.1a–b, MCZ 9331–9334 and 2164 (on exchange from SMF; Barbour & Loveridge 1929), and UMMZ 60296 (on exchange from MCZ). The name-bearing type has been considered to be SMF 6605, lectotype designated by implication as it was considered the holotype by Mertens (1967), Guibé (1978) and Blommers-Schl̂sser (1979). However, in the original description, Boettger (1880) mentioned explicitely a male and a female specimen, and provided morphometric data for these two specimens. Also, he stated ‘multa spec.’ after the locality information indicating that his description was based on more than one specimen. Mertens (1967) and subsequent authors therefore must have been aware of the fact that Limnodytes ulcerosus was based on syntypes. Furthermore, Article 74.5 of the Code (ICZN 1999) states unambiguously for lectotype designations before 2000, that ‘When the original work reveals that the taxon had been based on more than one specimen, a subsequent use of the term “ holotype ” does not constitute a valid lectotype designation unless the author, when wrongly using that term, explicitly indicated that he or she was selecting from the type series that particular specimen to serve as the name-bearing type.’ Since the lectotype designation by the authors listed above was not explicit in terms of the Code, we here stabilize it by expressly designating SMF 6605 (from which we obtained a 16S sequence) as lectotype of Limnodytes ulcerosus out of the available series of syntypes, in agreement with the choice of this specimen by previous authors. Identity.—Only one lineage of Brygoomantis has so far been reliably recorded from Nosy Be (Andreone et al. 2003; and the present study), and the nomen ulcerosus has been correctly assigned to this lineage in the recent literature (e.g. Glaw & Vences 1992a, 1994, 2007; Penny et al. 2017; Perl et al. 2014; Vieites et al. 2009). This assignment was confirmed by the 16S sequence of the lectotype, obtained by barcode fishing and included in our molecular analysis. Genetic results confirmed that this species occurs in the Sambirano region, and at Sahamalaza in the North West (Penny et al. 2017), across the northern mountain escarpment from Tsaratanàna to Makira. However, all records from eastern Madagascar (e.g. Blommers-Schl̂sser & Blanc 1991) belong to other, unrelated species of Brygoomantis. Synonyms.—The nomen Mantidactylus brauni Ahl, 1929 is usually considered a junior synonym of Mantidactylus biporus (e.g. Frost 2021; Guibé 1978) but has been considered doubtfully distinct from that species by Blommers-Schl̂sser and Blanc (1991), and as a nomen dubium by Glaw and Vences (1992a). It is based on two unnumbered syntypes from ‘Akkoraka (Central-Madagascar)’ according to the original description, and on a ‘holotype’ ZMB 31617 (a supposed lectotype designation ‘by implication’) by Guibé (1978). However, Blommers-Schl̂sser & Blanc (1991) doubted whether the syntypes of this species have been correctly identified, and furthermore, as discussed above, a lectotype designation by implication is not a valid nomenclatural act according to article 74.5 of the Code. We located the specimen ZMB 53737 (locality according to catalogue: ‘Akkorotha, Madagaskar’; collector / donor: [S.G.] Braun) at the Museum f̧r Naturkunde (Berlin) labelled as type of M. brauni and succeeded to obtain a 16S sequence by barcode fishing from this specimen. Consequently, we here deviate from Guibé (1978) and designate specimen ZMB 53737 as lectotype of Mantidactylus brauni Ahl, 1929, which is justified in order to clarify the identity of this nomen. According to the respective 16S sequence, the lectotype firmly cluster among sequences of M. ulcerosus, and based on this genetic information we include Mantidactylus brauni as junior synonym of M. ulcerosus. Diagnosis.—A member of the M. ulcerosus clade as revealed by the phylogenomic analysis, and sister to the morphologically very similar M. bellyi. See Table 4 for a list of diagnostic morphological characters. The combination of a large body size of up to 45 mm, strongly tubercular dorsal skin usually without welldefined dorsolateral ridges, large tympanum size in males (11–14% of SVL), absence of white spots on flanks and of white marking on snout tip, and pulsed advertisement calls emitted in regular series distinguishes M. ulcerosus from species of the other clades. Some species in the M. fergusoni clade can be morphologically similar, but they occur in eastern Madagascar (vs Sambirano and North West regions), and have highly different advertisement calls (Table 4). Within the M. ulcerosus clade, the new species differs by its large body size and tubercular dorsal skin from M. schulzi, and by its advertisement call emitted in regular call series from M. bellyi (single calls). M. ulcerosus can occur sympatrically with M. mahery in the North West of Madagascar, but that species differs by a smoother skin and fewer pulses per note in advertisement calls. 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 M. ulcerosus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Re-description of the lectotype (SMF 6605). Adult female in good state of preservation. Body stout. Head as wide as body. Snout rounded. Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis weak, slightly concave. Loreal region weakly concave. Tympanum distinct, large, rounded, diameter about 68% of eye diameter. Supratympanic fold distinct, beginning straight, with a rather distinct bend midway towards insertion of forelimb. Tongue ovoid, distinctly posteriorly bifid. Maxillary teeth present.Vomerine teeth present in two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Fingers without webbing. Relative length of fingers: IVariation.—Variation in measurements is given in Table 6. See Fig. 24 for colouration in life and its variation. There is moderate sexual size dimorphism (confirmed male SVL 28.5–35.9 mm [n = 8] vs confirmed female SVL 32.5–45.4 [n = 14]). Males have a larger tympanum than females (HTD/ED ratio is 53–79% in females, 74– 95% in males). Specimens with a light vertebral stripe, or with a broad light vertebral band, occur regularly. Dorsolateral ridges are absent in many individuals, but can be recognised in others (Fig. 24). Femoral glands in males are distinct, often orange-coloured in life, and with distinct distal ulcerous macrogland and proximal granular gland field; the granular gland fields on the two opposite thighs contact each other medially. Natural history.—Common along small streams with shallow water and associated swamps, where males emit their advertisement calls at night from the ground next to the water or sitting in shallow water. Males often begin calling in the late afternoon. Found in primary rainforest or transitional/dry forest and also in degraded areas and secondary forest, as long as some vegetation cover is present around the streams. Previous descriptions of eggs and embryos (Blommers-Schl̂sser 1979) refer to other species of Brygoomantis. Figure 24n shows a clutch of eggs probably assignable to this species. ...Continued on the next page ...Continued on the next page ...Continued on the next page ...Continued on the next page Calls.— The advertisement call of M. ulcerosus, recorded on 10 February 1992, 19:00 h, at Nosy Be (Vences et al. 2006: CD 2, track 74, cut 1), consists of a pulsed note (Fig. 25), emitted in series at regular intervals and very fast succession (short inter-call intervals). Notes exhibit some complexity in pulse structure, with the initial pulse of each note being distinctly longer in duration and containing highest call energy compared to subsequent pulses. This initial pulse seems to contain some substructure and amplitude modulation and sounds more distorted than subsequent pulses. Relative amplitude of pulses slightly decreases towards the end of the note. Within the recorded short call series of four calls, these become successively shorter from the beginning to the end of the series. Numerical parameters of four analysed calls were as follows: call duration (= note duration) 453– 779 ms (593.0 ± 143.2 ms); 45–65 pulses per note (54.8 ± 9.0); pulse duration 1–3 ms (2.3 ± 0.6 ms); pulse duration of initial pulses 27–37 ms (31.8 ± 4.1 ms); pulse repetition rate within notes (excluding initial pulse) 83.9–104.5 pulses/s (92.4 ± 7.6); dominant frequency 3101–3605 Hz (3308 ± 196 Hz); prevalent bandwidth 880–5500 Hz; call repetition rate (= note repetition rate) within regular call series ca 68 calls/min. Calls recorded from a chorus on 30 June 2009 from a site at km 27 of the road from Antsohihy to Mandritsara, 24°C air temperature, generally agreed in character with calls described from Nosy Be. They also exhibit the complex pulse structure described above, with initial pulses being very narrowly spaced and sometimes fused. Calls were emitted in series, containing 4–9 calls. The recording was difficult to analyse due to many overlapping calls, but the following call parameters could be measured (10 calls analysed): call duration (= note duration) 355– 565 ms (447.4 ± 63.0 ms); pulse duration 1–4 ms (1.8 ± 0.9 ms); pulse repetition rate within notes varied from ca 50–220 pulses/s; dominant frequency 2798–2885 Hz (2847 ± 37 Hz); prevalent bandwidth 1000–3800 Hz; call repetition rate (= note repetition rate) within regular call series ca 70–88 calls/min. Calls recorded on 24 February 2001 atAnkarafantsika, 28°C air temperature, also agreed with the calls described above, with very narrowly spaced initial pulses, partly fused. Calls were emitted in series containing 3–5 calls. Recordings were of poor quality, containing overlapping calls and background noise. However, the following parameters could be measured (8 calls analysed): call duration (= note duration) 551–1047 ms (667.3 ± 160.0 ms); pulse repetition rate within notes varied from ca 50–400 pulses/s (maximum values from initial pulses); prevalent bandwidth 900–3400 Hz; call repetition rate (= note repetition rate) within regular call series ca 66–87 calls/min. Distress calls of a female from Nosy Be were described by Glaw and Vences (1992b). Tadpoles.— Tadpoles assignable to this species (from the type locality Nosy Be) were briefly mentioned and their tooth formula described by Glaw & Vences (1994). The tadpole description by (Blommers-Schl̂sser 1979) was based on east coast specimens not identified by genetics that almost certainly belong to other species of Brygoomantis. Distribution.— Widespread in the North West (including the western slopes of the Makira Reserve) and Sambirano regions of Madagascar, over a wide range of elevations and habitat types (Fig. 7). This species is known from Angorony, Ankarafantsika, Antsatramidola, Benavony, Berara, the border of the Bealanana district (Bevitagnono and Irogno forest), between Antsohihy and Mandritsara, between Antoshihy and Port Berger, Makira West (Sahaovy, Camp 0), Nosy Be (type locality) including Lokobe National Park, Sahamalaza, and Tsaratanàna. It was also recorded from Nosy Komba (Hyde Roberts & Daly 2014). Records from the forests of Belambo and Anjiamangirana and in the Namoroka National Park (Raselimanana 2008) require genetic confirmation. Elevation range: 0–1093 m a.s.l. Etymology.— Latin adjective meaning ‘full of sores’ or ‘ulcerated’, presumably in reference to the granular dorsal skin or maybe to the femoral glands., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 195-205, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Boettger, O. (1880) Diagnoses reptilium et batrachiorum novorum a Carolo Ebenau in insula Nossi-Be madagascariensi lectorum. Zoologischer Anzeiger, 3, 279 - 283.","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","Barbour, T. & Loveridge, A. (1929) Typical reptiles and amphibians in the Museum of Comparative Zo ˆ logy. Bulletin of the Museum of Comparative Zoology, 69, 205 - 360.","Mertens, R. (1967) Die herpetologische Sektion des Natur- Museums und Forschungs-Institutes Senckenberg in Frankfurt a. M. nebst einem Verseichnis ihrer Typen. Senckenbergiana Biologica, 48, 1 - 106.","Guibe, J. (1978) Les batraciens de Madagascar. Bonner zoologische Monographien, 11, 1 - 140.","ICZN. (1999) International Code of Zoological Nomenclature. The International Trust for Zoological Nomenclature, London, 306 pp. 4 th Edition.","Andreone, F., Glaw, F., Nussbaum, R. A., Raxworthy, C. J., Vences, M. & Randrianirina, J. E. (2003) The amphibians and reptiles of Nosy Be (NW Madagascar) and nearby islands: a case study of diversity and conservation of an insular fauna. Journal of Natural History, 37, 2119 - 2149. https: // doi. org / 10.1080 / 00222930210130357","Glaw, F. & Vences, M. (1992 a) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 335 pp. First Edition.","Glaw, F. & Vences, M. (1994) A Fieldguide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 478 pp. Second Edition.","Penny, S. G., Crottini, A., Andreone, F., Bellati, A., Rakotozafy, L. M. S., Holderied, M. W., Schwitzer, C. & Rosa, G. M. (2017) Combining old and new evidence to increase the known biodiversity value of the Sahamalaza Peninsula, Northwest Madagascar. Contributions to Zoology, 86, 273 - 296. https: // doi. org / 10.1163 / 18759866 - 08604002","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Ahl, E. (1929 ' 1928 ') Beschreibung neuer Fr ˆ sche aus Madagascar. Mitteilungen aus dem Zoologischen Museum in Berlin, 14, 469 - 484.","Vences, M., Glaw, F. & Marquez, R. (2006) The Calls of the Frogs of Madagascar. 3 Audio CD's and booklet. Madrid, Spain, Fonoteca Zoologica, 44 pp.","Glaw, F. & Vences, M. (1992 b) Zur Kenntnis der Gattungen Boophis, Aglyptodactylus und Mantidactylus (Amphibia: Anura) aus Madagaskar, mit Beschreibung einer neuen Art. Bonner zoologische Beitrage, 43, 45 - 77.","Hyde Roberts, S. & Daly, C. (2014) A rapid herpetofaunal assessment of Nosy Komba Island, northwestern Madagascar, with new locality records for seventeen species. Salamandra, 50, 18 - 26.","Raselimanana, A. P. (2008) Herpetofaune des forets seches malgaches. Malagasy Nature, 1, 46 - 75."]}

Mantidactylus fergusoni sp. nov. Identity and justification.—This lineage has been previously considered as confirmed candidate species M. sp. 26 by Vieites et al. (2009) and M. sp. Ca26 by Perl et al. (2014). It was depicted as ‘ Mantidactylus sp. aff. betsileanus “Andranofotsy”’ by Glaw and Vences (2007). Our phylogenomic analysis confirms that this lineage is in a major clade (here named the M. fergusoni clade) containing only scientifically unnamed lineages, and its status as distinct species is therefore out of question. However, deciding how many species exist in this clade is complicated. Mantidactylus sp. Ca26 forms a clade with several other divergent mitochondrial lineages of uncertain status; here we consider the lineage previously named M. sp. 21 (Vieites et al. 2009) or M. sp. Ca21 (Perl et al. 2014) from Nosy Mangabe, Makira, Masoala, and Cap Est/Ambato as conspecific with M. fergusoni (note that M. sp. 21 was wrongly placed in the tree of Vieites et al. [2009], probably due to a sequence confusion in the alignment used for phylogenetic analysis). As a further deep conspecific lineage we consider samples from Marojejy and additional sites in the North East. Samples from Nosy Boraha (M. sp. Ca27) according to our phylogenomic analysis (Fig. 5) form a clade with the lineage M. sp. Ca25 (see Vieites et al. 2009) from different sites on the mainland adjacent to Nosy Boraha, and together are sister to M. fergusoni sp. nov.; the Nosy Boraha population is described below as M. jahnarum sp. nov. based on its bioacoustic differentiation. Holotype.— ZSM 126/2002 (MV 2001.1389), adult male, collected by M. Vences on 17 December 2001 at Andranofotsy (15.4353°S, 049.8439°E, 85 m a.s.l.), Analanjirofo Region, Madagascar. A 16S barcode sequence of the holotype is available from GenBank (accession AY848214). Paratypes.— A total of six paratypes: ZSM 124/2002 (MV 2001.1433), female, ZSM 125/2002 (MV 2001.1434), adult male, and ZSM 176/2002 (FGMV 2001.1383), specimen of unknown sex and maturity, all with the same collection data as the holotype; ZSM 5050/2005 (ZCMV 2125), adult female, and ZSM 5051/2005 (ZCMV 2136), adult male, collected by F. Glaw, M. Vences, and R.D. Randrianiaina on 22 February 2005 on Nosy Mangabe (ca 15.50°S, 049.77°E, 50–100 m a.s.l.); ZSM 355/2010 (FGZC 4277), adult female, collected by F. Glaw, J. K̂hler, P.-S. Gehring, M. Pabijan, F.M. Ratsoavina on 3 April 2010 at Ambodivoahangy (15.2899°S, 049.6203°E, ca 100 m a.s.l.). Additional material.— The following three specimens are assigned to M. fergusoni sp. nov. but come from a genetically divergent population and have in part (ZFMK specimens) not been sequenced, and therefore they are not included in the paratype series: ZSM 201/2005 (FGZC 2746), adult male, collected by F. Glaw, M. Vences, and R.D. Randrianiaina on 14 February 2005 at Marojejy, Camp 1 ‘Mantella’ (14.4377°S, 049.7756°E, 481 m a.s.l.); ZFMK 59938, adult male, and ZFMK 59939, adult female, collected by F. Glaw and O. Ramilison on 22 February 1995 at Marojejy (near Camp Mantella). Diagnosis.—A member of the M. fergusoni clade as revealed by the phylogenomic analysis, and sister to M. jahnarum sp. nov. described below. See Table 4 for a list of diagnostic morphological characters. The combination of a small to moderate body size in males (SVL up to 25– 30 mm) and distinctly larger body size in females (36– 42 mm), tubercular dorsal skin, large tympanum size in males (10–14% of SVL), absence of white spots on flanks and of white marking on snout tip, and advertisement call consisting of a single-pulse note distinguishes M. fergusoni sp. nov. from most species of the other clades. Two species from the M. ulcerosus clade (M. ulcerosus and M. bellyi) can be morphologically similar, but they occur in the Sambirano and North West regions, and have strongly differing advertisement calls (Table 4). One species of the M. betsileanus clade (M. katae) has an advertisement call of similar general structure, but has a faster and more regular call repetition rate, less tubercular dorsum, larger femoral glands, and a distinct white marking on snout tip (Table 4). M. fergusoni sp. nov. may also show superficial similarities to other species of the M. betsileanus clade but does not appear to occur sympatrically with any of them; in general it has a more tubercular dorsum and differs from all of these by advertisement call structure (Table 4). The M. fergusoni clade contains only species newly named herein; for a distinction from these other species, see below. A full list of molecular diagnostic sites in the 16S gene of M. fergusoni sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype.—Adult male in good state of preservation (Fig. 48). Part of left shank muscle removed as tissue sample, femoral glands partly detached for examination in internal view. Body slender. Head wider than body. Snout rounded.Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis very weak, slightly concave. Loreal region concave. Tympanum distinct, large, slightly wider than high, its horizontal diameter about 91% of eye diameter. Supratympanic fold present, beginning straight, with a rather distinct bend midway towards jaw / forelimb insertion. Tongue ovoid, distinctly bifid. Maxillary teeth present. Vomerine teeth present in two rounded to ovoid aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: I≤IIVariation.—Variation in measurements is given in Table 8. See Fig. 49 for colouration in life and its variation. There is pronounced sexual size dimorphism (confirmed male SVL 24.9–29.9 mm [n = 5] vs confirmed female SVL 35.7–42.2 mm [n = 4]). Males have a somewhat larger tympanum than females (HTD/ED ratio is 69–67% in females, 67–96% in males). Femoral glands are distinct but not particularly prominent in males, and only with a slight yellowish tone in life; very small, rudimentary glands recognisable in females. Natural history.—Habitat and habits of this species are poorly known, but so far it has been found in primary or somewhat degraded rainforest. At Andranofotsy, calling males were observed at night from the shore of a shallow puddle next to a small spring in rainforest. Calls.—The advertisement call of M. fergusoni, recorded on 16 December 2001 near Andranofotsy, 25.4°C air temperature (Vences et al. 2006, CD 2, track 69), consists of a simple, short, single pulse ‘click’ note, emitted in series (Fig. 50). Inter-call intervals were irregular with some calls (= notes) occurring in pairs with lower inter-call interval inbetween. Call energy was distributed in a wide frequency band. Numerical parameters of 13 analysed calls were as follows: call duration (= note duration) 9–12 ms (10.5 ± 0.8 ms); 1 pulse per note (1.0 ± 0.0); pulse duration = note duration = call duration; dominant frequency 1626–1690 Hz (1669 ± 26 Hz); prevalent bandwidth 700–5700 Hz; call repetition rate (= note repetition rate) within series ca 210–290 calls/min. Calls recorded on 31 March 2010 at Ambodivohangy, Makira area (from specimen ZSM 361/2010 = FGZC 4219), 24°C estimated air temperature, agree with those described above from Andranofotsy, apart from somewhat lower call repetition rate within series. Calls consisted of a simple short ‘click’ note emitted isolated or more often in series at irregular intervals. Numerical parameters of 28 analysed calls were as follows: call duration (= note duration) 7–13 ms (8.9 ± 1.4 ms); 1 pulse per note (1.0 ± 0.0); pulse duration = note duration = call duration; dominant frequency 1571–1722 Hz (1638 ± 54 Hz); prevalent bandwidth 700–5500 Hz; call repetition rate (= note repetition rate) within series ca 100–170 calls/min. Tadpoles.— The tadpole of this species has not been described. Voucher Field number Sex Locality SVL HW HL ED HTD END NSD NND FORL HAL HIL FOTL FOL TIBL FGL FGW M. fergusoni sp. nov. (Ca26) ZSM 126/2002 [HT]FGMV 2001.1389MAndranofotsy27.810.311.64.44.03.01.92.817.58.748.722.515.114.53.62.6ZFMK 59938 #NAMMarojejy, Camp Mantella26.010.69.73.92.62.91.82.98.98.0NMNM13.414.52.82.2ZSM 125/2002 [PT]FGMV 2001.1434MAndranofotsy26.210.111.24.23.12.42.03.115.87.741.819.113.313.43.92.4ZSM 201/2005 #FGZC 2746MMarojejy, Camp Mantella24.99.910.04.13.32.71.82.710.27.9NMNM12.314.03.42.3ZSM 5051/2005 [PT]ZCMV 2136MNosy Mangabe29.911.412.84.13.63.52.43.317.68.545.621.014.014.83.72.5ZFMK 59939 #NAFMarojejy, Camp Mantella35.714.713.45.42.93.82.14.012.710.4NMNM16.319.6NANAZSM 124/2002 [PT]FGMV 2001.1433FAndranofotsy36.113.614.55.63.63.92.33.122.510.260.228.319.418.9NANAZSM 355/2010 [PT]FGZC 4277FMakira40.315.216.35.53.84.12.43.924.611.866.530.021.221.2NANAZSM 5050/2005 [PT]ZCMV 2125FNosy Mangabe42.215.416.76.04.04.22.73.824.411.969.430.621.021.7NANA M. georgei sp. nov. (Ca35/Ca36) ZSM 455/2005 [HT]ZCMV 806MToamasina30.711.912.44.94.73.21.93.517.48.046.220.614.113.74.03.3ZSM 454/2005ZCMV 803MMaroantsetra29.010.912.04.63.73.31.83.417.48.646.920.313.913.84.02.4ZSM 456/2005 [PT]ZCMV 807MToamasina28.410.511.94.44.22.91.83.516.38.446.320.413.813.54.63.3 M. jahnarum sp. nov. (Ca27) ZSM 423/2006 [HT]ZCMV 3390MNosy Boraha30.311.112.64.73.33.11.83.217.28.946.121.114.014.44.83.0... Continued on the next page Distribution.— Endemic to low-elevations of Northern Central East and North East (Fig. 7). This species is known from Ambato, Ambinanifaho, Ambodirafia Tokana, Ambodiriana, Ambodivohangy (Makira), Ampasimazava, Andranofotsy (type locality), Andrantambe, Antanambe, Antsahanoro, Befanjana, Belambo, various sites within Marojejy National Park, Masoala, Nosy Mangabe, and the Sambava region. Elevation range: 12–1326 m a.s.l. Etymology.—We dedicate this species to Barry Ferguson, an outspoken worker on environmental justice for Madagascar, in acknowledgement of his support of the first author’s work on Madagascar’s herpetofauna., Published as part of 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, pp. 113-311 in Megataxa 7 (2) on pages 237-244, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/7441023, {"references":["Vieites, D. R., Wollenberg, K. C., Andreone, F., K ˆ hler, J., Glaw, F. & 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","Perl, R. G. B., Nagy, Z. T., Sonet, G., Glaw, F., Wollenberg, K. C. & Vences, M. (2014) DNA barcoding Madagascar's amphibian fauna. Amphibia-Reptilia, 35, 197 - 206. https: // doi. org / 10.1163 / 15685381 - 00002942","Glaw, F. & Vences, M. (2007) A Field Guide to the Amphibians and Reptiles of Madagascar. Vences & Glaw Verlags GbR, Cologne, Germany, 496 pp. Third Edition.","Vences, M., Glaw, F. & Marquez, R. (2006) The Calls of the Frogs of Madagascar. 3 Audio CD's and booklet. Madrid, Spain, Fonoteca Zoologica, 44 pp."]}

Mantidactylus manerana antsanga ssp. nov. Identity and justification.— This lineage was previously considered as unconfirmed candidate species M. sp. 15 by Vieites et al. (2009), and M. sp. Ca15 by Perl et al. (2014). It is characterized by a high divergence in 16S but haplotype sharing in Rag-1 with the nominal form, M. manerana manerana, which occurs allopatrically. Little is known about this form, with limited data on morphology and no data on bioacoustics or colour in life. We therefore describe it here as subspecies. Holotype.— MRSN A5633 (FAZC 13371, ACZC 10079), adult female, collected by F. Andreone, F. Mattioli, and J.E. Randrianirina on 24 January 2006 in Sahavontsira (ca 16.91°S, 049.22°E), Analanjirofo Region, Madagascar. A 16S barcode sequence of the holotype was included in the analysis Diagnosis.— Mantidactylus manerana antsanga ssp. nov. is a lineage here considered as subspecies of M. manerana due to its high morphological similarity. It is the sister group of the clade comprising M. m. manerana and M. m. fotaka according to our phylogenomic analysis. See Table 4 and the diagnosis of M. manerana above for a list of diagnostic morphological characters and of differences to other species of Brygoomantis. Morphologically, this poorly known taxon may differ from the other two subspecies by larger body size (SVL of the only examined female 38 mm, vs 28–29 mm female SVL in the other two subspecies). It may also differ from the nominal subspecies by slightly shorter hindlimbs and more developed foot webbing (Table 4). A full list of molecular diagnostic sites in the 16S gene of M. manerana antsanga ssp. nov. in pairwise comparisons to all other Brygoomantis species and subspecies is provided as Supplementary appendix. Description of the holotype.—Adult female in excellent state of preservation (Fig. 73). Fourth toe of right foot removed as tissue sample. Body stout. Head as wide as body. Snout rounded in dorsal and lateral views. Nostrils directed laterally, slightly protuberant, nearer to tip of snout than to eye. Canthus rostralis weakly recognisable, slightly concave; loreal region slightly concave.Tympanum distinct, moderately sized, as wide as high, horizontal diameter of tympanum 69% of horizontal eye diameter. Supratympanic fold recognisable above the tympanum, slightly curved, not recognisable posterior to tympanum. Tongue ovoid, distinctly bifid posteriorly. Maxillary teeth present. Vomerine teeth form two prominent rounded aggregations, positioned posterolateral to choanae and almost as large as these. Choanae rounded. Subarticular tubercles single. Inner and outer metacarpal tubercles present. Fingers without webbing. Relative length of fingers: I Colour in preservative: dorsally brown with some indistinct darker markings, and triangular more distinct dark marking between eyes. More irregularly dark-light marbled on flanks, and some lighter spots on upper lip and underneath the eye. Weakly defined dark crossbands on limbs. Fingers and, less expressed, toes with alternating pattern of light and dark colour. Ventrally beige, without any dark pigmentation except for a light-dark pattern ventrally on lower lip. Colour in life not documented. Natural history.—Largely unknown. Specimens were collected from rainforest. Calls.— The call of this subspecies has not been recorded. Tadpoles.— The tadpole of this subspecies has not been described. Distribution.— Widespread in the Northern Central East (Fig. 7). Currently known from Sahavontsira, Babitanety and Antsahataloka in the Befanjana forest, and Ambatobe. Elevation range: 14–466 m a.s.l. Etymology.—The subspecies name is derived from the Malagasy word antsanga, meaning ‘trash and mud deposited by flood water’, in reference to the riparian habits of this species (and other Brygoomantis species). The name is used as a noun in apposition.

Mini ature sp. nov. urn:lsid:zoobank.org:act: 0C1C4CE2-419A-4030-86D1-B1B89793697D (Figs 1–3 and 9, Table 1) Remark. This species was previously listed as Stumpffia sp. Ca53 MV 2017(MF 867231) by Tu et al. [20], though with incorrect accession number (correct number is MF 768231). Holotype (Figs 2 and 9). ZSM 86 /2004 (FGZC 0 151, GenBank accession numbers MF768231 and MK 459307 for 5’ and 3’ fragments of the 16S rRNA gene, respectively, and MF768147 for cox1 gene), a presumed adult specimen collected in Andohahela National Park between Isaka and Eminiminy above ‘ Camp 1’ (ca. 24.75&ring;S, 46.85&ring;E, ca. 350 m) between 29 and 31 January 2004 by F. Glaw, M. Puente, M. Teschke (née Thomas), and R.D. Randrianiaina. Diagnosis. A highly miniaturised frog assigned to Mini gen. nov. on the basis of its small size, curved clavicles, laterally displaced and reduced nasals, and fusion or loss of carpal 2. This assignment is supported by its genetic affinities (Fig 1; [20]). It is separated by uncorrected pdistances of 10.0–10.6% in the analysed 3’ fragment of the 16S rRNA gene from other members of the genus Mini gen. nov., and 10.8–13.7% from all members of the genus Plethodontohyla. Mini ature sp. nov. is characterised by the unique combination of the following characters (n = 1 specimen): (1) SVL 14.9 mm; (2) ED/HL 0.38; (3) HW/SVL 37.2; (4) FARL /SVL 0.32; (5) TIBL/SVL 0.34; (6) HIL/SVL 1.18; (7) finger 1 strongly reduced, 2 and 4 reduced; (8) toe 1 absent, toes 2 and 5 quite reduced; (9) maxillary and premaxillary teeth present; (10) vomerine teeth absent; (11) lateral colour border absent; (12) black inguinal spots present; (13) postchoanal vomer present, spatulate, medially fused to parasphenoid; (14) nasal cultriform and laterally displaced; (15) quadratojugal-maxilla contact strong and broad; (16) zygomatic ramus of squamosal long, thin, curved, and horizontal; (17) clavicles present, curving with simple lateral articulations, medially not bulbous; (18) prepollex thin and cultriform; (19) carpal 2 absent or fused to post-axial carpal 3–5 element; (20) finger phalangeal formula 1-2-3-3; (21) toe phalangeal formula 1-2-3-4-3; (22–26) calls unknown. Within the genus Mini gen. nov., the new species can be distinguished by its distinctly larger body size (14.9 mm vs 8.2–11.3 mm), shorter relative hindlimb length (HIL /SVL 1.18 vs 1.41–1.72) and phalangeal formula of fingers (1-2-3-3 vs 1-2-3- 2 in M. mum sp. nov. and 0-2- 3- 2 in M. scule sp. nov.). Additionally, it can be distinguished from M. mum sp. nov. by the presence of maxillary and premaxillary teeth (vs absence), and less distinct lateral colour border, and M. scule sp. nov. by proportionally smaller nasals and braincase, broader quadratojugal-maxillary contact, and vertical dorsal process of premaxilla (vs anterior). This species is particularly similar to some highly miniaturised Stumpffia species, but it can be distinguished from all Stumpffia based on the condition of the carpals and the presence of curved clavicles, and most Stumpffia by the presence of maxillary and premaxillary teeth. Holotype description. Specimen in a moderately good state of preservation, the left arm removed as a tissue sample, the whole body somewhat dorsoventrally flattened in preservative. Body oblong; head wider than long, narrower than body width; snout slightly pointed in dorsal view, pointed in lateral view; nostrils directed laterally, not protuberant, equidistant between tip of snout and eye; canthus rostralis rounded, indistinct, slightly concave; loreal region flat, vertical; tympanum indistinct, round, ~48% of eye diameter; supratympanic fold absent; tongue long, broadening posteriorly, attached anteriorly, not notched; maxillary teeth present; vomerine teeth present; choanae small and round. Forelimb slender; subarticular tubercles single, elongated; outer/palmar metacarpal tubercle small and round; inner metacarpal tubercle slightly smaller than outer/palmar; hand without webbing; first finger strongly reduced, second and fourth fingers reduced, third finger basally broadened; relative lengths of fingers 1 < 2 = 4 < 3, fourth and second finger equal in length; finger tips not expanded into discs. Hindlimbs stocky; TIBL 34% of SVL; lateral metatarsalia strongly connected; inner metatarsal tubercle indistinguishable from first toe; outer metatarsal tubercle absent; no webbing between toes, second and fifth toes reduced; relative lengths of toes 2 < 5 < 3 < 4, fifth toe distinctly shorter than third; toe tips slightly pointed distally. Skin on dorsum smooth without a distinct dorsolateral fold, but with a distinct colour border, see below. Ventral skin smooth. After 14 years in 70% ethanol, the dorsum is light brown, paler—almost beige—laterally, and slightly translucent, with a thin beige vertebral stripe and a darkened area on the posterior head. The skin above the eyes is translucent and dark in colour through the presence of the eyes beneath. The dorsal forelimb is beige flecked with brown, the hand is lighter medially, and the fingers have faint cream annuli. The dorsal hindlimb is beige in base colour with several brown crossbands on the thigh and shank that line up when the leg is folded together. A trapezoid of brown is present around the vent. The foot is dorsally as the forelimb with a light annulus before each distal phalanx. A distinct colour border that is not straight is present laterally, running along the canthus rostralis from the nostril through the eye, through the supratympanic region along the torso to the inguinal region. Small oblong dark brown spots are present in the inguinal region. The side of the head is dark brown. Ventral to this colour bor- der the frog is mocha speckled with beige, lightening ventrally to beige with loose cream speckles. The ventral skin is translucent, and some of the organs can be seen through it. The chin is not differently coloured than the rest of the ventral body. This pattern continues onto the ventral limbs. No data on life colouration are available. Variation. This species is currently known from a single specimen only. Osteology (Fig 9). Based on ZSM 86/2004 (figured). Cranium (Fig 9 D–9G). Shape and proportions. Skull almost equilateral, roughly as wide as long, widest at quadratojugal-squamosal junction, lateral to the otic region. Braincase moderately broad, rostrum not shortened. Neurocranium. Well ossified, including the otic capsules. The anterior cone of the sphenethmoid is ossified and contacts the frontoparietal dorsally but is not in contact with any other bones; small isolated lateral mineralisations of this bone are also present anterodorsal to the anterior tip of the cultriform process of the parasphenoid. Prootic in dorsal contact with lateral flange of frontoparietal, ventral contact with parasphenoid alae, not approaching contralateral ventrally. Septomaxilla miniscule, relatively poorly mineralised, and therefore not further discussed here. Columella (stapes) well ossified, pars media plectra (stylus) long and nearly straight, weakly posteriorly and dorsally oriented toward the reniform, dorsally elongated pars interna plectra (baseplate). Nasal narrow and cultriform, laterally displaced, curved downward laterally, acuminate maxillary process not closely approaching maxillary pars facialis, broadly separated from contralateral. Frontoparietal with rounded anterior edge, laterally bulging, with short lateral flange covering prootic, posteriorly strongly connected to exoccipital, anteroventrally contacting sphenethmoid, lacking any dorsal process. Parasphenoid with narrow, rather straight-edged cultriform process and broad perpendicular alae, considerably shorter than frontoparietals, in contact with exoccipitals posterodorsally, prootics dorsally along the edges of the alae, anteroventrally in contact with postchoanal vomer and not in contact with neopalatine; posteromedial process excluded from participating in foramen magnum by exoccipitals. Vomer divided into pre- and postchoanal portions; prechoanal portion narrow, arcuate, triradiate, with a short lateral and anterior ramus and long, curving posterior ramus; postchoanal portion spatulate and edentate, contacting its contralateral on the midline, in dorsal contact with the parasphenoid proximally and the neopalatine distally, lacking an anterior projection. Neopalatine simple, straight, almost indistinguishable from lateral postchoanal portion of vomer, laterally broadly separated from the maxilla, not exceeding the lateral-most point of the postchoanal vomer. Maxillary arcade fairly slight, maxilla and premaxilla bearing numerous diminutive teeth, anterior extension of maxilla exceeding lateral extent of premaxilla but not in contact with it. Premaxilla with a broad acuminate dorsal alary process rising laterally, 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 running parallel to but not touching the anterior ramus of the pterygoid. Pterygoid with a short medial ramus, long anterior ramus, and broad posterior ramus and posterolaterally calcified to the quadratojugal complex. Quadratojugal weakly bowed laterally, broadly connected to the ventral ramus of the squamosal, bearing a small posteroventral knob; the articulation of the mandible is apparently fortified by the mineralisation of the posterior ramus of the pterygoid+squamosal+quadratojugal posteroventral knob. Squamosal with a slender ventral ramus, broadened otic ramus, and long, thin zygomatic ramus, the otic ramus oriented slightly dorsally and posteriorly, the zygomatic ramus horizontal and strongly curved. Mandible slim and edentate, largely unremarkable, with a low coronoid process on the angulosplenial. Mentomeckelians separated from the dentary, with slightly bulbous, almost hooked ventrolateral projections. Posteromedial processes of hyoid proximally rounded with a broad medial crista. Postcranial skeleton (Fig 9 A–9C, 9H and 9I). Eight procoelous unfused presacrals, much broader than long, lacking neural spines, with round posterior articular processes, presacral I with a complete neural arch, presacrals II–IV with thicker and longer transverse processes than V–VIII. Sacrum with expanded diapophyses, the leading and trailing edges roughly equally angled, the articulation type IIB sensu Emerson [42]. Urostyle bicondylar, long, not broadening posteriorly, without lateral processes and with a low dorsal ridge. Pectoral girdle without ossified prezonal or postzonal elements, with ossified clavicles. Clavicle thin and curved, with a simple lateral junction, shorter than the coracoid. Coracoid fairly narrow, not flared laterally, strongly flared medially with a large medial articular surface with the contralateral. Scapula slender, with a thin pars acromialis, the cleithral border concave. Cleithrum ossified for two thirds the width of the scapular border, thickened anteriorly. Suprascapula unossified. Humerus with a well-developed crista ventralis and no medial or lateral cristae. Radioulna slender with a distinct sulcus intermedius. Carpals poorly ossified, composed of radiale, ulnare, prepollical element, element Y, and large post-axial element formed by carpals 3–5. Carpal 2 has either been lost or fused to the latter element. Finger phalangeal formula is reduced (1-2-3- 3), and the terminal phalanx of the first finger is a small, round element. Small distal knobs on terminal phalanx of finger 3. Prepollex thin and cultriform and extending only to the base of the first metacarpal. Pubis 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, almost lacking a posterior crest. Tibiofibula shorter than femur, with a sulcus intermedius. Tibiale and fibulare fused proximally and distally. T1 and T2+3 tarsals present, T1 considerably smaller than T2+3, plus a small additional ossification (possibly a sesamoid) between the bases of metatarsals 2 and 3. Centrale present, roughly the size of T2+3. Prehallux subtriangular. Phalangeal formula reduced (1-2-3-4-3). Terminal phalanges of toes 2–4 with almost T-shaped knobs. Distribution, natural history, and conservation status. This species is known only from a single specimen from Andohahela National Park in southeast Madagascar (Fig 6). The species is larger than the other members of the genus Mini, and its ecology may differ accordingly. Advertisement calls were not recorded. At present it is not possible to estimate its distribution or population status, and we prefer to suggest this species be considered Data Deficient until more information is available. Etymology. We use the specific epithet ‘ature’ as an arbitrary combination of letters, in order to form a pun on ‘miniature’ from the name in apposition, in reference to the small size of this species. It is to be regarded as an invariable noun.

Genus-level taxonomy of the Cophylinae Mini adds a ninth genus to the Cophylinae for a unique clade of miniaturised frogs that falls sister to the large-bodied Plethodontohyla (Fig 1). Although body size is the most obvious character that differentiates these two sister genera, they are also distinguished by a number of osteological features, and can be identified without skeletal analysis by their digital reduction (present in Mini, absent in Plethodontohyla) and vomerine teeth (absent in Mini, present in Plethodontohyla). Despite these differences and consistent recovery of the two genera as being reciprocally monophyletic in genetic analyses, the uncorrected p-distances between these genera in the 3��� fragment of the 16S rRNA mitochondrial gene analysed here are at first glance surprisingly small at 8.3���13.3% (these distances would be distinctly higher if insertions and deletions would be considered in their calculation). Nevertheless, we consider the differences between these clades sufficiently great and robust that we regard them as constituting separate genera. Aside from their morphological and osteological differentiation, a further argument for their classification in distinct genera comes from the strength of support for their sistergroup relationship; while the two clades here seen as genera Plethodontohyla and Mini have been placed sister to each other in most molecular analyses so far, support values for this grouping often were low, and typically lower than the respective support for each of the two clades. The clade stability criterion [36] is therefore better served considering both clades as separate genera. The relationship of Mini to Plethodontohyla is analogous to the relationship of Stumpffia to Rhombophryne: a genus-level sister clade of miniaturised frogs (although Stumpffia also contains several non-miniaturised species), recovered in robust genetic phylogenies as reciprocally monophyletic, and distinguished by several diagnostic characters [14, 15, 23]. Peloso et al. [19] argued for the lumping of Stumpffia, Rhombophryne, and later also Anilany [22] into a single genus, Rhombophryne. In response, we showed that the initial argument for lumping was based on misidentified specimens [15], and subsequently incorrectly coded morphology, ignoring various unique diagnostic features of Anilany, and the relationships of particularly unstable taxa (most notably Stumpffia tridactyla) [23]. We revised the taxonomy of the genus Stumpffia, describing 26 new species, and providing a more robust phylogeny that resolved the phylogenetic position of Stumpffia tridactyla [14]. Despite this progress, the Amphibian Species of the World database (ASW) currently continues to use the lumped taxonomy, in contrast to AmphibiaWeb and other researchers that have adopted our proposed taxonomy (e.g. [20, 53]). The newly described Rhombophryne proportionalis, which is the only miniaturised Rhombophryne so far known, is highly distinct from Stumpffia, lacking, for example, the externally obvious digital reduction that is present in all miniaturised species of Stumpffia, and differing in body shape and proportions [14]. This demonstrates that even miniaturised Rhombophryne species can be distinguished by external morphology from Stumpffia species, providing still further support for the recognition of Rhombophryne, Stumpffia, and Anilany as separate genera., Published as part of Scherz, Mark D., Hutter, Carl R., Rakotoarison, Andolalao, Riemann, Jana C., R��del, Mark-Oliver, Ndriantsoa, Serge H., Glos, Julian, Roberts, Sam Hyde, Crottini, Angelica, Vences, Miguel & Glaw, Frank, 2019, Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera, pp. 1-45 in PLoS ONE 213314 on pages 37-38, DOI: 10.1371/journal.pone.0213314, http://zenodo.org/record/2614083, {"references":["36. Vences M, Guayasamin JM, Miralles A, de la Riva I. To name or not to name: Criteria to promote economy of change in Linnaean classification schemes. Zootaxa. 2013; 3636 (2): 201 - 44. https: // doi. org / 10. 11646 / zootaxa. 363 6.2. 1","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.","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","23. Scherz MD, Vences M, Rakotoarison A, Andreone F, Kohler J, Glaw F, et al. Lumping or splitting in the Cophylinae (Anura: Microhylidae) and the need for a parsimony of taxonomic changes: a response to Peloso et al. (2017). Salamandra. 2017; 53 (3): 479 - 83.","19. Peloso PLV, Frost DR, Richards SJ, Rodrigues MT, Donnellan S, Matsui M, et al. The impact of anchored phylogenomics and taxon sampling on phylogenetic inference in narrow-mouthed frogs (Anura, Microhylidae). Cladistics. 2016; 32 (2): 113 - 40. https: // doi. org / 10.1111 / cla. 12118","22. Peloso PLV, Raxworthy CJ, Wheeler WC, Frost DR. Nomenclatural stability does not justify recognition of paraphyletic taxa: A response to Scherz et al. (2016). Molecular Phylogenetics and Evolution. 2017; 111 (2017): 56 - 64. https: // doi. org / 10.1016 / j. ympev. 2017.03. 0 16 PMID: 28336422","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","53. Feng Y-J, Blackburn DC, Liang D, Hillis DM, Wake DB, Cannatella DC, et al. Phylogenomics reveals rapid, simultaneous diversification of three major clades of Gondwanan frogs at the Cretaceous-Paleogene boundary. Proceedings of the National Academy of Sciences of the USA. 2017; 114 (29): E 5864 - E 70. https: // doi. org / 10.1073 / pnas. 17046 32114 PMID: 286739 70"]}

Anodonthyla eximia sp. nov. urn:lsid:zoobank.org:act: 2C419E74-C13D-447D-BDCF-312324515EAE (Figs 1, 2, 3, 5, 12 and 13, Tables 1 and 2) Remark. This species was previously listed as A. sp. Ranomafana (Maharira) [24], A. sp. Ca04 Ranomafana (ZCMV 204) [15] and Anodonthyla sp. 4 Ranomafana [20]. Holotype. ZMA 20246 (ZCMV 204, GenBank accession number GU177052 and FJ559111 for the 5��� and 3��� fragments of the 16S rRNA gene, respectively, and GU177063 for the cox1 gene), an adult male specimen (vocal sac inflated when collected) collected from a campsite at the base of Maharira mountain (21.3258&ring;S, 47.4025&ring;E, approx. 1248 m a.s.l.) in Ranomafana National Park, Vatovavy-Fitovinany Region, former Fianarantsoa province, southeastern Madagascar on 25 January 2004 by M. Vences, I. de la Riva, and E. Rajeriarison. Diagnosis. An extremely miniaturised frog assigned to the genus Anodonthyla on the basis of the possession of a large, cultriform prepollex, T-shaped terminal phalanges, absence of postchoanal vomer, and by its genetic affinities. It is separated by uncorrected p-distances of 9.3���17.0% in the analysed 3��� fragment of the 16S rRNA gene from all other members of the genus Anodonthyla. Anodonthyla eximia sp. nov. is characterised by the unique combination of the following characters (n = 1 male specimen): (1) male SVL 11.3 mm; (2) ED/HL 0.45; (3) HW/SVL 0.31; (4) FARL /SVL 0.30; (5) TIBL/SVL 0.39; (6) HIL/SVL 1.34; (7) finger 1 highly reduced, other fingers small; (8) toe 1 absent, toes 2 and 5 quite reduced; (9) maxillary and premaxillary teeth absent; (10) vomerine teeth absent; (11) lateral colour border absent; (12) black inguinal spots absent; (13) postchoanal vomer absent; (14) nasal subrectangular with an acuminate maxillary process, not displaced; (15) quadratojugal-maxilla contact absent; (16) zygomatic ramus of squamosal short, thin, and anterodorsally oriented; (17) clavicles present, curving with simple lateral articulations, medially bulbous; (18) prepollex cultriform, longer than first metatarsal; (19) carpal 2 present; (20) finger phalangeal formula 1-2-3-3; (21) toe phalangeal formula 2-2- 3-4-3; (22) single-note, unpulsed calls, not emitted in series; (23) weakly frequency modulated calls; (24) call dominant frequency 8406 �� 78 Hz (n = 5); (25) call duration 59.6 �� 6.5 ms (n = 5); (26) inter-call interval 3749.0 �� 1149.9 ms (n = 4). This species is considerably smaller than all other Anodonthyla species (11.3 mm vs 15���34 mm), and as such is only possible to confuse with juveniles of its congeners. In addition to its small size, it can be distinguished from all other Anodonthyla species by the absence of flared crests on the humerus in adult males. Among similarly sized adult frogs in Madagascar, it can be distinguished by the possession of a large inner metacarpal tubercle with a large, cultriform prepollex (present only in male Anodonthyla, much smaller in all other species except Anilany, where it is broad and triangular instead of cultriform), and a laterally displaced neopalatine not in contact with the sphenethmoid or vomer (neopalatine either in contact with sphenethmoid or in contact with vomer in all other species; in some species of Stumpffia, the neopalatine is lost, unpublished data). Holotype description. Specimen in a good state of preservation, a piece of the right thigh removed as a tissue sample. Body oblong; head wider than long, narrower than the body width; snout rounded in dorsal and lateral view; nostrils directed laterally, not protuberant, closer to eye than to tip of snout; canthus rostralis rounded, straight; loreal region concave, vertical; tympanum indistinct, round, about 46% of eye diameter; supratympanic fold weak, not raised, straight from posterior corner of eye to axilla; tongue long, thin, attached anteriorly, not notched; maxillary teeth absent, vomerine teeth absent; choanae oblong, very small. Forelimbs relatively broad; subarticular tubercles indistinct, single; outer metacarpal tubercle small, indistinct, single; inner metacarpal tubercle large and distinct, bulging outward strongly, underlain by cultriform prepollex; hand without webbing; all fingers short, first highly reduced and only marginally longer than the prepollex; relative length of fingers 1 < 2 = 4 < 3, fourth finger equal in length to second; tip of third finger marginally expanded, other fingers not expanded into discs. Hind limbs robust; TIBL 39% of SVL; lateral metatarsalia strongly connected; inner metatarsal tubercle rounded, indistinguishable from first toe; outer metatarsal tubercle absent; no webbing between toes; first toe practically absent, second and fifth toes shortened; relative length of toes 2 < 5 < 3 < 4; fifth toe distinctly shorter than third. Skin on dorsum smooth, without distinct dorsolateral folds. Ventral skin smooth. After 13 years in 70% ethanol, the dorsum is pale brown with a faint darker brown chevron over the scapular region (Fig 2). The dorsal legs have faint dark brown crossbands, especially on the shanks. The dorsal colouration fades over the shanks to cream, which is continuous on the venter. The lateral head is dark brown, with a distinct border to the dorsum and flank formed by the supratympanic fold. The chin is a lighter brown than the dorsum but distinctly not the cream of the abdomen. The ventral legs are translucent cream. Colour in life as in preservative but more vibrant; the venter was slate grey with blue-cream flecks. Bioacoustics. Calls recorded from an unknown specimen by M. Vences at 08h45, on 26 January 2004 at Maharira in Ranomafana National Park (21.3258&ring;S, 047.4025&ring;E, 1248 m a.s.l.) in the leaf litter of primary rainforest, referred to this species (Fig 4D, Table 2). The holotype and single known specimen of this species was collected during careful searches of the leaf litter from the exact spot where similar calls were heard, and upon capture, had an apparent partially inflated vocal sac. No other calls assignable to a miniature frog were heard in the area at this time in the morning. Assignment of the calls to this species therefore is the most likely hypothesis but remains tentative. The call bears remarkable resemblance to that of Stumpffia miery (Table 2), which is also known from lower elevations in Ranomafana National Park, and detailed future field study will be required to confirm its assignment to Anodonthyla eximia sp. nov. Calls were emitted as part of a large chorus in the early hours of the morning following cyclonic rainfall after the retreat of major flooding of the area. Calls consisted of a single note and were emitted at regular intervals without defined call series. Calls were weakly frequency modulated with an increase in pitch, but recording quality is too poor for detailed analysis. For approximate call parameters, see Table 2. Osteology (Fig 13). Based on ZMA 20246 (figured). Cranium (Fig 13 D���13G). Shape and proportions. Skull short and rounded, longer than wide, widest at the bowing of the quadratojugal roughly in line with the anterior face of the prootic. Braincase proportionally broad, with an extremely short rostrum. Neurocranium. Moderately ossified, otic capsules partly ossified. Sphenethmoid unossified. Prootic in dorsal contact with lateral flange of frontoparietal, ventral contact with parasphenoid alae, not approaching contralateral. Septomaxilla miniscule, very tightly curled, not further discussed due to low ossification and insufficient resolution. Columella (stapes) well ossified, pars media plectra (stylus) long and slightly curved, posteriorly and dorsally oriented toward the broadened, somewhat posteriorly oriented pars interna plectra (baseplate). Nasal narrow, retaining the shape of larger cophylines: subrectangular with an elongated, acuminate maxillary process that does not closely approach the maxillary pars facialis; displaced laterally, broadly separated from contralateral. Frontoparietal with rounded anterior edge, laterally rather straight-edged, with short lateral flange covering prootic, posteriorly strongly connected to exoccipital, lacking any dorsal process, separated from contralateral by a narrow gap with a small rhomboid facet at the level of the prootics, which may represent a pineal foramen. Parasphenoid with narrow, rather straight-edged cultriform process and roughly equally broad perpendicular alae, considerably shorter than frontoparietals, in contact with exoccipitals posterodorsally, prootics dorsally along the edges of the alae, anteroventrally free; posteromedial process long but not participating in foramen magnum. Vomer lacking postchoanal portion (typical of Anodonthyla); prechoanal portion strongly curved, bearing a small lateral ramus. Neopalatine simple and very thin, laterally displaced, not contacting any other ossified elements. Maxillary arcade gracile, maxilla and premaxilla edentate, anterior extension of maxilla not exceeding lateral extent of premaxilla. Premaxilla with a narrow dorsal alary process rising laterally, pars palatina shallowly divided into a narrow palatine process and broad lateral process. Maxilla practically lacking a pars facialis and bearing a narrow pars palatina, its posterior tip acuminate and not contacting the quadratojugal, the lingual surface of the pars palatina contacting the anterior ramus of the pterygoid, which has taken over the articulation. Pterygoid with a short medial ramus, long and strongly curved anterior ramus, and broad posterior ramus, posteriorly calcified to the quadratojugal complex. Quadratojugal bowed laterally, rather short, broadly connected to the ventral ramus of the squamosal, bearing a small posteroventral knob, anteriorly dividing and with decreasing ossification, not connected to the maxilla; the articulation of the mandible is apparently somewhat fortified by the mineralisation of the posterior ramus of the pterygoid+squamosal+quadratojugal posteroventral knob. Squamosal with a slender, rather straight ventral ramus, thin, posterodorsally oriented otic ramus, and short, thin, anterodorsally oriented zygomatic ramus. Mandible slim and edentate, largely unremarkable, with a weakly raised coronoid process on the angulosplenial. Mentomeckelians separated from the dentary, with small, poorly ossified hooked ventrolateral projections. Posteromedial processes of hyoid proximally rounded without an obvious medial crista. Postcranial skeleton (Fig 13 A���13C, 13H and 13I). Eight procoelous presacrals, all much broader than long, lacking neural spines, with round posterior articular processes, presacral I with a complete neural arch, presacrals II���IV with thicker and longer transverse processes than V���VIII. Sacrum with flared diapophyses, the leading and trailing edges roughly equally curved, the articulation type IIB sensu Emerson [42]. Urostyle bicondylar, long, broadening gently posteriorly, with a somewhat flared head and a low dorsal ridge. Pectoral girdle without ossified prezonal or postzonal elements, with ossified clavicles. Clavicle thin and curved, with a simple lateral junction, equal in length to the coracoid, medially bulbous. Coracoid broad, flared laterally and strongly flared medially with a curving medial articular surface with the contralateral. Scapula robust, with a broad pars acromialis, the cleithral border straight. Cleithrum ossified for three-quarters the width of the scapular border, thickened anteriorly. Suprascapula unossified. Humerus with a well-developed crista ventralis and no medial or lateral cristae. Radioulna slender with a distinct sulcus intermedius. Carpals composed of radiale, ulnare, element Y, prepollex, carpal 2, and a large post-axial element formed by carpals 3���5. Prepollex extremely long and acuminate, longer than first metacarpal. Finger phalangeal formula is reduced (1-2- 3-3), and the terminal phalanx of the first finger small and columnar, others bearing T-shaped knobs. Pubis ossified; iliac shafts passing ventral to and beyond sacrum, oblong in cross-section, with a weak dorsal crest, a distinct dorsal prominence, and a shallow oblique groove. Femur weakly sigmoid, bearing a distinct posterior crest. Tibiofibula slightly shorter than femur in length, with a sulcus intermedius. Tibiale and fibulare fused proximally and distally. T1 and T2 +3 tarsals present, T1 considerably smaller than T2+3. Centrale present, larger than other tarsals. Prehallux elongated, half the length of first metatarsal. Phalangeal formula standard (2-2- 3-4-3). Terminal phalanx of toe 1 a small round element, those of toes 2���5 with T-shaped knobs. Etymology. The species epithet eximia is the feminine form of the Latin adjective eximius meaning ���remarkable��� or ���special���, in reference to the surprisingly small body size and terrestrial habits of this Anodonthyla species. Distribution, natural history, and conservation status. Anodonthyla eximia sp. nov. is known only from Maharira in Ranomafana National Park (Fig 6). It is a terrestrial species. Nothing else is known of its natural history. It is likely that this species should be classified as Vulnerable like other species from Maharira (e.g. Gephyromantis runewsweeki), but as we know almost nothing of its range and ecology, we instead recommend that it be considered Data Deficient until more data are available., Published as part of Scherz, Mark D., Hutter, Carl R., Rakotoarison, Andolalao, Riemann, Jana C., R��del, Mark-Oliver, Ndriantsoa, Serge H., Glos, Julian, Roberts, Sam Hyde, Crottini, Angelica, Vences, Miguel & Glaw, Frank, 2019, Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera, pp. 1-45 in PLoS ONE 213314 on pages 33-37, DOI: 10.1371/journal.pone.0213314, http://zenodo.org/record/2614083, {"references":["24. Vences M, Glaw F, Kohler J, Wollenberg KC. Molecular phylogeny, morphology and bioacoustics reveal five additional species of arboreal microhylid frogs of the genus Anodonthyla from Madagascar. Contributions to Zoology. 2010; 79 (1): 1 - 32.","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","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","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"]}

Mini gen. nov. urn:lsid:zoobank.org:act: 67171236-00A2-428B-8194-584BF52E84E6 (Figs 2–9, Table 1) Type species. Mini mum sp. nov. Contents. Mini mum sp. nov., M. scule sp. nov., and M. ature sp. nov. Etymology. The genus name is derived from English prefix ‘mini-’, denoting a small version of an object. We treat this name as an arbitrary combination of letters in the sense of the International Code of Zoological Nomenclature Articles 30.1.4.1 and 30.2.2, and we assign it the feminine gender. We have searched all available taxonomic databases and could not find any evidence that this name has ever been used to refer to a genus of animals, and we therefore conclude that it is available. Diagnosis. Diminutive terrestrial frogs (adult SVL 8.2–14.9 mm), assigned to the Madagascar-endemic subfamily Cophylinae on the basis of divided vomers, procoelous vertebral column, divided sphenethmoids, and genetic affinities. Skin smooth to slightly granular, occasionally iridescent. A lateral colour border is present but varies in intensity among species. Highly reduced fingers and toes, fusion or loss of carpal 2, and paedomorphic skull morphology: laterally displaced narrow nasals, teeth absent from vomer, in some species present on the maxilla and premaxilla, otic capsule sometimes dorsally ossified, brain case comprising most of the skull’s length and width. All members of the genus Mini gen. nov. resemble miniaturised to extremely miniaturised members of the genus Stumpffia. However, all species can be distinguished from Stumpffia on the basis of their curving clavicles and a fused or lost carpal 2. In the species accounts below, we provide detailed distinctions from Stumpffia relevant to each species. Justification. The erection of the genus Mini is justified by significant genetic differentiation from all other major cophyline lineages (see Fig 1), by the fact that it does not form a monophyletic group with the genus Stumpffia, and furthermore by the strong morphological differences (including but not restricted to the much smaller size) to all species of its sister clade, Plethodontohyla. The following characters distinguish the genus from all Plethodontohyla species, including juveniles: digital reduction of the fingers and toes (vs no reduction), laterally displaced and reduced nasals (vs large nasals situated anterior to frontal), parasphenoid cultriform process shorter than frontoparietals (vs roughly equal in length to the frontoparietals) and considerably narrower than alary processes (vs as wide or wider), vomerine teeth absent (vs present), carpal 2 absent (vs present). Uncorrected p-distances between Mini and Plethodontohyla range from 8.3–13.3% in the 3’ fragment of 16S rRNA analysed here, and they have been found to be sister to Plethodontohyla in all phylogenetic analysis since their first inclusion in genetic datasets [15, 17, 20, 22] (except the DNA barcoding study of Vieites et al. [40], where they were placed at the base of Rhombophryne + Stumpffia + Anilany, but that study lacks any resolution at deep nodes, and was not intended to provide phylogenetic hypotheses at deep levels). Distribution. The genus Mini is apparently endemic to low-elevation habitats (0–350 m a. s. l.) of southeastern Madagascar (Fig 3).

Rhombophryne proportionalis sp. nov. urn:lsid:zoobank.org:act: 1 A79607 B-E9D4-4214-BF47-9BC0B4320E5F (Figs 1, 2, 3, 5, 10 and 11, Tables 1 and 2) Remark. This species was previously referred to as Stumpffia sp. Ca34 [15, 46] and as Stumpffia sp. 39 MV-2012 (KC 351481) [20]. Holotype. ZSM 1826 /2010 (ZCMV 12404, GenBank accession number KC351380 and KU 937808 for 5��� and 3��� fragments of the 16S rRNA gene, respectively, and KF611640 for the cox1 gene), an adult male specimen (seen calling, not recorded) collected at Bepia Campsite on the Tsaratanana massif (Camp 3; 14.1182&ring;S, 48.9782&ring;E, 2294 m a.s.l.), Diana Region, former Antsiranana province, northern Madagascar on 16 June 2010 by M. Vences, D.R. Vieites, R.D. Randrianiaina, S. Rasamison, and E. Rajeriarison. Paratypes. ZSM 1840 /2010 (ZCMV 12405, GenBank accession number KC351481 and MK 459317 for 5��� and 3��� fragments of the 16S rRNA gene, respectively), adult male specimen with the same collection data as the holotype; and ZSM 636 /2014 (DRV 6224), an adult presumed male specimen collected at Andranomadio Campsite in Tsaratanana (Camp 4; 14.0801&ring;S, 48.9854&ring;E, 2503 m a.s.l.) on 16 June 2010 by the same collectors. Diagnosis. A diminutive frog assigned to the genus Rhombophryne 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 7.0���12.9% in the analysed 3��� fragment of the 16S rRNA gene from other members of the genus Rhombophryne. Rhombophryne proportionalis sp. nov. is characterised by the unique combination of the following characters (n = 3 male specimens): (1) male SVL 11.0��� 12.3 mm; (2) ED/HL 0.40���0.48; (3) HW/SVL 0.33���0.37; (4) FARL /SVL 0.33���0.35; (5) TIBL/SVL 0.34���0.36; (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 9���17 calls at irregular intervals; (23) non-frequency modulated calls; (24) call dominant frequency 5460 �� 117 Hz (n = 79); (25) call duration 45.4 �� 8.2 ms (n = 79); (26) inter-call interval 63.0 �� 9.0 ms (n = 73). 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 S. contumelia, S. obscoena, S. davidattenboroughi, S. makira, S. achillei, and S. analanjirofo, and some specimens of S. tridactyla, 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 Rhombophryne species is still possible, but these have much larger teeth proportional to their skull size, and most lack the distinct lateral head colouration and possess clavicles. The call is unique among the frogs of Madagascar and is instantly distinctive in being emitted as a rapid, high-pitched series of tonal notes. Holotype description. 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 < 2 = 4 < 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 < 2 < 5 < 3 < 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 (Fig 10). Ventral skin smooth in preservative, granular in life. 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. Variation. For measurements, see Table 1. The paratypes are in general very similar to the holotype in morphology. ZSM 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. ZSM 1840/ 2010 has a more massive body than the others. The colouration of the specimens is relatively consistent, with the whole venter of ZSM 1840/2010 being darker than those of the other two specimens. The dorsolateral lines of the holotype are present in ZSM 1840/2010, but not in ZSM 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 ZSM 1840/2010 than the other two specimens. Bioacoustics. 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 (Fig 5C, Table 2) at around 11h40 on 15 June 2010 in Camp Bepia (14.11822&ring;S, 048.97822&ring;E, 2294 m a.s.l.), and further calls were heard but not recorded at Camp Andranomadio (14.0801&ring;S, 048.9854&ring;E, 2503 m 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 Stumpffia psologlossa, where the units have a very short duration, as a single call composed of pulses [14], in Rhombophryne mangabensis as a series of notes [52], and in R. minuta two closely spaced units were seen as components of a single note [52]. To allow comparison within Rhombophryne 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 5���50 ms range for which the pulse category was recommended by K��hler et al. [33]. Calls are rapid series of high-pitched slightly frequency modulated notes. Each call consists of a series of 9���17 notes, and calls are repeated at long and irregular intervals. For detailed call parameters, see Table 2. Osteology (Fig 11). Based on ZSM 1826/2010 (figured) and ZSM 636/2014 (not figured). Cranium (Fig 11 C���11F). Shape and proportions. Skull short and stout, relatively wide, widest at quadratojugal anterior to the otic region. Short rostrum. Braincase broad. Neurocranium. Well ossified except the weakly calcified otic capsules. Sphenethmoid well ossified, in contact with the frontoparietal in ZSM 1826/2010 but not in ZSM 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 Rhombophryne (see [25]). 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 ZSM 1826/ 2010). 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. 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. 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. 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. Posteromedial processes of hyoid proximally pointed with a broad medial crista. Postcranial skeleton (Fig 11A, 11B, 11G and 11H). 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 II���IV with thicker and longer transverse processes than V���VIII. Sacrum with expanded diapophyses, the leading and trailing edges roughly equally angled, the articulation type IIB sensu Emerson [42]. Urostyle bicondylar, long, broadening posteriorly, without lateral processes and with a low dorsal ridge. 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. Humerus with a well-developed crista ventralis and no medial or lateral cristae. Radioulna broad with a distinct sulcus intermedius. Carpals well ossified in ZSM 1826/2010 and poorly ossified in ZSM 636/2014, composed of radiale, ulnare, prepollical element, element Y, carpal 2, and large post-axial element formed by carpals 3���5. 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 ZSM 1826/2010 but is not visible in ZSM 636/2014. 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). Distribution, natural history, and conservation status. Rhombophryne proportionalis sp. nov. is known from two localities (Bepia and Andranomadio) on the Tsaratanana massif in northern Madagascar. 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, Platypelis alticola and P. tsaratananaensis, are also known from these locations, and both are currently listed as Endangered, due to their small distribution < 2000 km 2, presence in a single threat-defined location, and potentially on-going decline in habitat quality. We therefore suggest R. proportionalis sp. nov. also to be Endangered, following the same rationale. Etymology. The species epithet is the Latin adjective proportionalis 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., Published as part of Scherz, Mark D., Hutter, Carl R., Rakotoarison, Andolalao, Riemann, Jana C., R��del, Mark-Oliver, Ndriantsoa, Serge H., Glos, Julian, Roberts, Sam Hyde, Crottini, Angelica, Vences, Miguel & Glaw, Frank, 2019, Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera, pp. 1-45 in PLoS ONE 213314 on pages 28-33, DOI: 10.1371/journal.pone.0213314, http://zenodo.org/record/2614083, {"references":["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","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","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","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.","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","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","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","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"]}