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Blommersia bara sp. nov. Figures 4, 8 Identity. This species was considered Mantidactylus wittei by Guibé (1974) and Blommers-Schl̂sser & Blanc (1991) for specimens from Ankarafantsika, and called Mantidactylus cf. wittei by Glos (2003), Blommersia sp. aff. wittei “Isalo” by Glaw & Vences (2007), Blommersia sp. aff. wittei by Mercurio et al. (2008), Blommersia sp. (aff. wittei) by Bora et al. (2010), Blommersia sp. 5 by Vieites et al. (2009) and Wollenberg et al. (2011), Blommersia sp. Ca5 by Perl et al. (2014), Penny et al. (2016) and Glaw et al. (2019), Blommersia sp. Ca05 by Penny et al. (2017), and Blommersia sp. aff. wittei Ca 05 “Isalo” by Cocca et al. (2018). Holotype. ZSM 31/2004 (field number FGZC 47), an adult male, collected by F. Glaw, M. Puente, M. Thomas & R. Randrianiaina at a stream near Ranohira, Isalo Massif (22.5856°S, 45.3997°E, 813 a.s.l.), southwestern Madagascar, on 21 January 2004. Paratypes. A total of 30 specimens, all from western Madagascar. ZSM 22/2004 (FGZC 34) and ZSM 24/2004 (FGZC 38) two adult males with the same collecting data as the holotype; ZSM 706/2001 (FGMV 2001.279), adult male, collected by M. Vences, D.R. Vieites, G. Garcia, V. Raherisoa, & A. Rasoamamonjinirina at Ampijoroa, Ankarafantsika National Park (approximately 16.3°S, 46.82°E) on 24 February 2001; ZSM 246/2003 and ZSM 247/2003, two adult males, collected by J. Glos at Kirindy Forest (site “G2”) in January 2001; ZSM 13/2006 (FGZC 691), ZSM 30/2006 (FGZC 727), ZSM 52/2006 (FGZC 779), and ZSM 68/2006 (FGZC 809), two adult females and two adult males, collected by F. Glaw, J. K̂hler, P. Bora & H. Enting at Tsingy de Bemaraha National Park, Andranopasazy, “Camp 1” (18.7086°S, 44.7189°E, 146 m a.s.l.), on 17, 19, and 24 March 2006, respectively; ZSM 2284/2007 (ZCMV 5801) and ZSM 2285/2007 (ZCMV 5804), one adult male and one adult female, collected by M. Vences and collaborators at Isalo National Park, on 17 February 2007; ZSM 2281/2007 (ZCMV 5625), ZSM 2282/2007 (ZCMV 5626), ZSM 2283/2007 (ZCMV 5643), and ZSM 2320/2007 (ZCMV 5621), collected by L. du Preez, C. Weldon & L. Raharivololoniaina at Ankarafantsika, on 9 February 2007; ZSM 3222/2012 (ZCMV 14143), adult male collected by A. Rakotoarison, J. Erens & E. Rajeriarison at Mariarano (15.4978°S, 46.6943°E, 11 m a.s.l.), on 28 December 2012; MRSN A2957 and A2958 (FAZC 11807–11808), two males, collected by F. Andreone, G. Aprea & V. Mercurio in the Isalo Massif, Andohasahenina (22.8333°S, 45.1880°E, 876 m a.s.l.) on 15 January 2004; MRSN A5349 (FAZC 12550), female, and MRSN A5350 (FAZC 12551), male, collected by F. Andreone, F. Mattioli & V. Mercurio in the Isalo Massif, Zahavola (22.6215°S, 45.3587°E, 881 m a.s.l.), on 17 November 2004; MRSN A5353 and A5354 (FAZC 12968 and 12970), two males, collected by F. Andreone, F. Mattioli & V. Mercurio in the Isalo Massif, Sakamalio (22.4348°S, 45.2552°E, 726 m a.s.l.) on 16 December 2004; MRSN A5351 (FAZC 12591), female, collected by F. Andreone, F. Mattioli & V. Mercurio in the Isalo Massif, Andriamanero (22.3672°S, 45.3920°E, 663 m a.s.l.), on 20 November 2004; as well as seven specimens deposited in the UADBA collection (uncatalogued but accessible via field tags): UADBA-ZCMV 5620, 5641, 5642, 5644, four specimens collected by L. du Preez, C. Weldon & L. Raharivololoniaina at Ankarafantsika, on 9 February 2007; and UADBA-ZCMV 5800, 5802, 5803, three specimens collected by collected by M. Vences and collaborators at Isalo National Park, on 17 February 2007. Diagnosis. A species of the genus Blommersia in the subfamily Mantellinae of the family Mantellidae based on presence of intercalary elements between penultimate and ultimate phalanges of fingers and toes (verified by external examination), occurrence in Madagascar, relatively small body size (male SVL From other species of Blommersia, the new species is mainly distinguished as follows: from B. angolafa by the presence of vomerine teeth in many individuals (vs. absence) and a totally different color pattern without whitish spots on the flanks and on the finger- and toe-tips (vs. presence); from B. dejongi by a femoral gland placed centrally on the thigh (vs. distally); from B. blommersae by having lateral metatarsalia separated by webbing (vs. connected by dense tissue); from B. domerguei by having lateral metatarsalia separated by webbing (vs. connected by dense tissue), by the presence of vomerine teeth in many individuals (vs. absence), a larger body size (SVL> 20 mm vs. B. dupreezi by having lateral metatarsalia separated by webbing (vs. connected by dense tissue) and absence of distinct black lateral stripe (vs. presence); from B. galani by presence of vomerine teeth in many individuals (vs. absence); from B. grandisonae by presence of vomerine teeth in many individuals (vs. absence) and a largely different color pattern (but see discussion in Vences et al. 2023 on the identity of B. grandisonae); from B. kely by having lateral metatarsalia separated by webbing (vs. connected by dense tissue), presence of vomerine teeth in many individuals (vs. absence), and larger body size (SVL> 20 mm vs. B. nataliae by ovoid femoral glands in small distance to each other (vs. more rounded, distant glands) and absence of a dark face mask (vs. presence); from B. sarotra by having lateral metatarsalia separated by webbing (vs. connected by dense tissue); from B. transmarina by a narrower head (male HW/SVL ratio 0.29‒0.34 vs. 0.34–0.36) and somewhat shorter hands (male HAL/SVL ratio 0.29–0.33 vs. 0.30–0.35); from B. variabilis by having lateral metatarsalia separated by webbing (vs. connected by dense tissue in most individuals) and probably a smaller distance between femoral gland. Furthermore, differentiated from all these species by differences in advertisement calls and genetic distances>5% in the analyzed mitochondrial 16S rRNA gene fragment of these two species. It differs from B. wittei, however, in a substantial divergence of mitochondrial genes (5.0‒8.2% uncorrected pairwise distance in 16S) and lack of haplotype sharing in RAG1 and SACS, and in advertisement calls lacking distinct pulses vs. being clearly pulsed in B. wittei (except in the Sambava population whose taxonomic status is in need of revision). Description of the holotype. Adult male specimen with distinct femoral glands, in a good state of preservation (Fig. 8). Tongue removed as tissue samples for molecular analysis. SVL 18.5 mm, for further measurements see Table 1. Body slender; head longer than wide, of same width as body; snout rounded in lateral view, obtusely pointed in dorsal and ventral views; nostrils directed laterally, protuberant, nearer to snout tip than to eye; canthus rostralis rounded; loreal region slightly concave; tympanum distinct, round, its diameter 63% of eye diameter; supratympanic fold distinct, curved above tympanum where it follows the tympanum outline; tongue absent, its shape therefore not ascertainable; vomerine teeth not visible; choanae small, round, located toward the front of the palate; maxillary teeth present.Arms slender, subarticular tubercles present, single; fingers without webbing; relative length of fingers 1 After 19 years in preservative (Fig. 8), the dorsum is rather uniformly brownish, with a weakly recognizable dark brownish inverted chevron, a faint small dark transverse bar on the forehead anterior to the eyes, and a very weakly recognizable dark interorbital bar. A dark stripe runs from the nostril to the eye and continues broader posterior to the eye, encompassing the entire tympanic region underneath the supratympanic fold. Underneath these brown elements, a weakly expressed light frenal stripe is visible from the snout tip to the forelimb insertions. Fore- and hindlimbs dorsally with distinct and sharply delimited brown crossbars (three crossbars on thigh and four on shank). Ventral side uniformly beige (unpigmented), throat being a bit lighter than chest, belly and limbs. In life, color was very similar but somewhat more contrasted (Fig. 4A); whitish color was present on the throat; femoral glands granules were somewhat yellowish (Fig. 4B). Variation. ZSM 52/2006 exhibits a broad middorsal light band. Vomerine teeth are clearly visible in paratype ZSM 2281/2007 from Ankarafantsika, also recognizable (although weakly expressed) in the paratypes from Bemaraha. We observed substantial differences in body size among populations, with individuals from the type locality Isalo being distinctly smaller than those from other sites. According to available data (Supplementary Table 2; available at DOI 10.5281/zenodo.8049142) male SVL across all populations ranges between 18.2–25.7 mm and female SVL between 22.0– 26.4 mm. Etymology. Named after the Bara people, the ethnic group living in the area of Madagascar that includes the type locality of the new species, the Isalo Massif. The name is used as a noun in apposition. Natural history. A relatively common species in Madagascar, living in areas of a certain humidity such as streams or swamps, including areas of secondary vegetation. During the rainy season, males call during day and night from the low vegetation of these water bodies. In Isalo this species is commonly found on the ground of gallery forests (e.g., Andriamanero, Sakamalio), along temporary rivers and in Pandanus swamps in the open savannah (e.g., Ilakaka, Zahavola). Tadpoles develop in temporary ponds and individuals of this species have been observed in egg-guarding behaviour both in Ankarafantsika (Fig. 4C) and Isalo (Mercurio et al. (2008). In the Sahamalaza Peninsula, this species seems to be abundant and it is found along streams and ponds in intact forested areas as well as in paddy fields in cleared areas. In Bemaraha, specimens were found in the leaf litter. In Kirindy, Glos (2003) found the species in the largest out of 200 ponds studied, and observed males calling at night from a bush at the water edge between 1–4 m above ground. Vocalizations: Advertisement calls recorded on 29 January 1994 at Isalo (air temperature 21.5°C) consist of a single short pulsatile note repeated in call series at rather regular intervals within series (Fig. 6). Each call exhibits some irregular amplitude modulation, sometimes with 2–4 peaks recognizable within each call, with the second peak usually having the highest energy. Numerical parameters of 23 analyzed calls of one male are as follows: call duration (= note duration) 13–18 ms (15.2 ± 1.6 ms); inter-call intervals within regular call series 69–78 ms (74.8 ± 3.0 ms); duration of regular call series 2034 ms (n = 1); call rate within call series approximately 670 calls/minute; dominant frequency 5813–6115 Hz (5968 ± 119 Hz); second dominant frequency peak around 3000 Hz; prevalent bandwidth 2600–9500 Hz. Advertisement calls recorded on 24 February 2001 at Ankarafantsika (air temperature 28°C) consist of a single short pulsatile note repeated in call series at regular intervals within series (Fig. 6). In some calls (= notes) two barely separated peaks of amplitude are recognizable, but call energy is distributed rather equally along the first two thirds of the call’s duration, then dropping rapidly towards its end. Numerical parameters of 18 analyzed calls of one male are as follows: call duration (= note duration) 19–28 ms (23.2 ± 2.5 ms); inter-call intervals within regular call series 94–107 ms (97.3 ± 4.8 ms); duration of regular call series 2078 ms (n = 1); call rate within call series approximately 500 calls/minute; dominant frequency 4694–4758 Hz (4735 ± 27 Hz); second frequency peak around 2370 Hz; prevalent bandwidth 1900–9700 Hz. Distribution. Known from Isalo in the South-West up to Sahamalaza Peninsula in the North-West (see Fig. 2 for genetically confirmed sites). The precise contact zone with B. wittei has not yet been identified but can be expected to be located between Sahamalaza and Ambanja.

Blommersia wittei (Guibé, 1974) Figures 3, 8 Identity and type material. Clarifying the taxonomy of the B. wittei complex requires first ascertaining the identity of B. wittei sensu stricto. The species was described by Guibé (1974) as Mantidactylus wittei, and type specimens are deposited in the Paris museum (acronym MNHN, in earlier times written MNHNP), with the following verbatim information in the original description: “ Holotype: no 1953-60 MNHN Paris. Environs d’Ambanja. J. Guibé (11- 1951). Paratypes: nos 1953-60 A à 1953-60 L MNHN Paris. Même provenance. Nos A.682 à A.684: Ampijora , station forestière à 45 km de Marovaoy (Ouest). Nos 1973-944 à 1973-951 MNHN Paris: forêt d’Ankarafantsika. Ch. P. Blanc (11-1973). ” Due to some imprecise information in the handwritten MNHN catalogue and extensive re-labelling of several of these specimens, there has been confusion on the type series in subsequent publications and in the current digital MNHN catalogue. Frost (2023) correctly states that the holotype is MNHNP 1953.60, by original designation, but merges two separate sites in the type locality account: “ ‘ Environs d’Ambanja’, Forest Ankarafantsika, Madagascar. ” In contrast, the digital type catalogue of the MNHN (downloaded from gbif.org in 2022) as well as the MNHN online catalogue (https://science.mnhn.fr/institution/mnhn/collection/ra/; accessed 11 February 2023) state that the holotype originated from “ Vallée de la Tsiribihina ” and furthermore provide the following list of paratypes: MNHN 1973.944–1973.951 (8 specimens) from Ankarafantsika, and 1993.682–1993.684 (3 specimens) from Ampijoroa. Vences et al. (2010) provided measurements of part of the type series of B. wittei, purportedly all from Ambanja: the holotype MNHN 1953.60, and the following paratypes: MNHN 1991.2529–1991.2533 (5 specimens; previously labelled MNHN 1953.60 A to E) and MNHN 1991.2536–1991.2539 (4 specimens, previously labelled MNHN 1953.60 H to K). After inspecting the original (handwritten) MNHN catalogues (partly available from https://science.mnhn.fr/ catalogues; accessed 11 February 2023) we conclude that: (1) The holotype of Mantidactylus wittei Guibé, 1974 (Fig. 8) unambiguously is the specimen MNHN 1953.60 for which morphometric measurements were provided by Vences et al. (2010) and whose locality, according to the original description and original catalogue entry is “ Environs d’Ambanja”, with the addition in the catalogue: “Cacaoyéres” (= cacao trees, thus indicating the specimen was collected in a cacao plantation). (2) A series of 11 paratypes (originally labelled MNHN 1953.60 A to 1953 K, currently re-labelled as MNHN 1991.2529–1991.2539) originates from the same locality as the holotype. (3) A series of 8 paratypes (MNHN 1973.944–1973.951) originate from Ankarafantsika, (4) A series of 3 paratypes (MNHN 1993.682–1993.684) originate from Ampijoroa, which represents a forestry station (currently local headquarters of Madagascar National Parks) within the Ankarafantsika forest. As explained in the Taxonomic conclusion account above, based on the provenance of the holotype from Ambanja, we continue assigning the name B. wittei to the genetic lineage occurring across much of northern Madagascar and collected by us at Ambanja and nearby localities. The paratype series is however mixed: specimens from Ankarafantsika (MNHN 1973.944–1973.951) and Ampijoroa (MNHN 1993.682–1993.684) are here assigned to B. sp. Ca 5. Only the 11 paratypes from the type locality are likely conspecific with the holotype. Morphology. Measurements of the type series of B. wittei have been published by Vences et al. (2010) and measurements of additional specimens by Pabijan et al. (2011). Measurements of further specimens are included in Table 1 herein.A full set of measurements including those from the previous publications is given as Supplementary Table 2 (available from the Zenodo repository under DOI 10.5281/zenodo.8049142). Based on these data, male SVL is 21.0–27.0 mm and female SVL is 20.7–25.0 mm. In the specimens examined for the present study (listed in Table 1), vomerine teeth are usually recognizable (clearly visible in specimens from the type locality Ambanja) but weakly expressed in several specimens from Montagne d’Ambre. In ZSM 563/2000 from Sambava, vomerine teeth are weakly recognizable, and this specimen apparently has more developed webbing than many other individuals examined. Vocalizations.Advertisement calls recorded in February 1991 at the type locality Ambanja (recording temperature not taken) consist of a single short pulsed note repeated in call series at regular intervals and very fast succession (Fig. 5). Recording quality is poor and the detailed call structure is probably partly masked by background noises. However, each call (= note) seems to contain two pulse groups which are separated from each other, with the first pulse group being of lower amplitude. Pulse structure in not very obvious in the recording and most pulses appear basally fused. Maximum call energy is distributed towards the middle of the call’s duration. Numerical parameters of 16 analyzed calls of one male are as follows: call duration (= note duration) 62–77 ms (66.8 ± 5.3 ms); inter-call intervals within regular call series 15–31 ms (18.6 ± 5.6 ms); pulses/note 9–18 (13.8 ± 3.3); duration of regular call series 1275 ms (n = 1); call rate within call series approximately 740 calls/minute; dominant frequency 4694–4886 Hz (4797 ± 82 Hz); second frequency peak around 2500 Hz; prevalent bandwidth 1800–6000 Hz. Advertisement calls recorded on 7 February 1992 from north of Andoany, Nosy Be (air temperature 25°C), consist of a single short pulsed note repeated in short call series at regular intervals and very fast succession (Fig. 5). Each call (= note) exhibits two pulse groups which are clearly separated from each other, with the first pulse group being of lower amplitude. Pulses are partly fused, but countable. Maximum call energy is distributed in the middle of the call, namely the beginning of the second pulse group. Numerical parameters of 10 analyzed calls of one male are as follows: call duration (= note duration) 65–83 ms (74.8 ± 6.3 ms); inter-call intervals within regular call series 16–32 ms (21.5 ± 5.6 ms); pulses/note 14–19 (16.3 ± 1.9); duration of regular call series 490 and 500 ms (n = 2); call rate within call series approximately 640 calls/minute; dominant frequency 4823–5240 Hz (5033 ± 163 Hz); second frequency peak around 2500 Hz; prevalent bandwidth 2000–8200 Hz. Advertisement calls recorded on 15 March 1994 on Montagne d’Ambre (air temperature 22°C), consist of a single short, distinctly pulsed note repeated in call series at regular intervals and very fast succession (Fig. 5). Each call (= note) exhibits two pulse groups which are clearly separated from each other, with the first pulse group being of lower amplitude. Pulses are barely fused and rather distinctly separated. Maximum call energy is distributed in the middle of the call, namely the beginning of the second pulse group. Numerical parameters of 19 analyzed calls of one individual are as follows: call duration (= note duration) 59–73 ms (65.4 ± 4.2 ms); inter-call intervals within regular call series 22–39 ms (29.6 ± 7.1 ms); pulses/note 7–11 (8.9 ± 1.3); duration of regular call series 1821 ms (n = 1); call rate within call series approximately 600 calls/minute; dominant frequency 4489–4597 Hz (4542 ± 41 Hz); prevalent bandwidth 1800–7500 Hz. Advertisement calls recorded from specimen ZSM 50/2018 (MSZC 521) on 25 December 2017 on Montagne d’Ambre (air temperature ca 19°C), consist of a single short, distinctly pulsed note repeated in call series at regular intervals and very fast succession (Fig. 5). Each call (= note) exhibits clearly separated pulses. Intervals between pulses are somewhat irregular, in some calls resulting in two pulse groups separated by a larger interval. Maximum call energy is evident in the middle of the call. Numerical parameters of 16 analyzed calls of one male are as follows: call duration (= note duration) 68–88 ms (78.2 ± 5.7 ms); inter-call intervals within regular call series 28–50 ms (41.4 ± 4.2 ms); pulses/note 7–11 (9.0 ± 1.3); duration of regular call series 1590 ms (n = 1); call rate within call series approximately 500 calls/minute; dominant frequency 5066–5232 Hz (5137 ± 80 Hz); prevalent bandwidth 2000–10000 Hz. Advertisement calls recorded on 20 March 2000 in Sambava (air temperature 24.7°C), here tentatively allocated to B. wittei, consist of a single, short, pulsed note repeated in call series at regular intervals within series. Calls (= notes) exhibit a pulsed structure, but most pulses are largely fused, resulting in an irregular pulse pattern and varying number of countable pulses (Fig. 7). Maximum call energy is distributed among the first two thirds of the call’s duration. Numerical parameters of 22 analyzed calls of one male are as follows: call duration (= note duration) 27– 59 ms (39.4 ± 11.8 ms); inter-call intervals within regular call series 54–76 ms (67.9 ± 7.7 ms); pulses/note 2–6 (4.1 ± 1.0); duration of regular call series 2209 ms (n = 1); call rate within call series approximately 600 calls/minute; dominant frequency 5316–5598 Hz (5440 ± 145 Hz); second frequency peak around 2600 Hz; prevalent bandwidth 2100–8600 Hz.

Blommersia sp. Ca12 Identity. This candidate species name refers to a new lineage discovered in the course of this study and found only at two sites along the Sambirano river (Maevatanana and Antsirasira). Morphology. Unknown. One voucher specimen from Antsirasira (ZSM 589/2001; see Table 1) could not be genotyped as no tissue sample for molecular analysis was taken in the field from this specimen. This specimen is an adult male with rather indistinct femoral glands and weakly expressed vomerine teeth. Photos in life are not available. Vocalizations: Advertisement calls were recorded on 31 January 2001 at Antsirasira (air temperature 26°C), but call vouchers were not collected and calls are therefore not assignable to B. sp. Ca12 with full reliability. These calls consist of a single, short, pulsed note repeated in call series at slightly irregular intervals within series (Fig. 7). Calls (= notes) generally exhibit a pulse structure, although some pulses are not clearly separated but rather largely fused, resulting in an irregular pulse pattern and varying number of countable pulses. Numerical parameters of 10 analyzed calls of one male are as follows: call duration (= note duration) 34–66 ms (47.8 ± 10.8 ms); inter-call intervals within regular call series 48–72 ms (61.4 ± 10.2 ms); pulses/note 3–10 (6.9 ± 2.3); duration of regular call series 1434 and 995 ms (n = 2); call rate within call series varied between 500–580 calls/minute; dominant frequency 5316–5598 Hz (5440 ± 145 Hz); second frequency peak around 2850 Hz; prevalent bandwidth 2200–9000 Hz.

Vences, Miguel, Multzsch, Malte, Köhler, Jörn, Crottini, Angelica, Andreone, Franco, Rakotoarison, Andolalao, Scherz, Mark D., Glaw, Frank (2023): Integrative revision of the Blommersia wittei complex, with description of a new species of frog from western and north-western Madagascar. Zootaxa 5319 (2): 178-198, DOI: 10.11646/zootaxa.5319.2.2, URL: http://dx.doi.org/10.11646/zootaxa.5319.2.2

4.3. Lygodactylus verticillatus Lygodactylus verticillatus from Madagascar and from Europa Island do not differ in scalation (Boettger 1913; Pasteur 1965). The slight difference in the number of precloacal pores mentioned by Pasteur (1965) was not supported by this study. Both populations show a very similar variation in coloration and in patterning. Thus, L. verticillatus from Madagascar and Europa cannot be distinguished by their external appearance or by the number of precloacal pores. The most basal node within L. verticillatus separates specimens from the southern Malagasy localities Sakaraha and Isalo from all other samples. Sakaraha lies approximately 135 km from Toliara on the main road to the Isalo National Park. The remaining sister subgroup consists of specimens exclusively from the coastal regions of Madagascar, approximately from Morondava southwards to Toliara, and the insular population. We cannot exclude an inland origin of the L. verticillatus from Europa Island. However, based on our morphological and genetic dataset, it seems more likely that the nearest relatives of the insular L. verticillatus are conspecifics from populations in the coastal regions of south-western Madagascar. We found no relevant morphological differences, and only weak genetic differences between the Malagasy and the insular population, and only weak genetic variation within the monophyletic lineage of Europa Island. This agrees with the hypothesis of a relatively recent colonization as suggested by Pasteur (1965). Europa Island is 355 km west-northwest from Toliara or 300 km southwest from Cap Saint-Vincent close to Morombe, 529 km east-northeast from Inhambane (Mozambique) and approximately 600 km south of Juan de Nova (Caceres 2003; Fricke et al. 2013). It is nearly circular (6 × 7 km) with a total area of 30 km 2 (Caceres 2003) and represents the largest island of the Îles Éparses. Europa Island belongs to the group of ‘modern’ isolated seamounts in the Channel, possibly developed during Oligocene to Miocene times (Courgeon et al. 2016). Lygodactylus verticillatus could have reached Europa Island recently by natural transoceanic dispersal or by human-mediated dispersal. While a natural dispersal would be supported by flows westward from Madagascar to mainland Africa (Ali & Huber 2010), a much simpler and thus more parsimonious alternative explanation is ship-borne dispersal. Lygodactylus verticillatus could have arrived at Europa Island e.g., with ships of Malagasy fishermen seasonally searching for nesting chelonians, or with ships of European (French) settlers who started their settlement from Toliara and lived on the island from approximately 1860 to the 1920s (Fricke et al. 2013). The amount of genetic variation found within Europa (three haplotypes differing by up to two mutations in 16S) would be in agreement with either (i) ship-borne dispersal or (ii) natural dispersal of multiple individuals, or (iii) natural dispersal of a single individual at a somewhat deeper point in time, with subsequent in-situ genetic diversification. Support for either of these hypotheses could come from a denser sampling of Malagasy populations: the presence of haplotypes identical to those on Europa in Malagasy populations would allow to reject the third hypothesis, and given that simultaneous or repeated natural dispersal to such a small islet is rather unlikely, ship-borne introduction would in this case remain as the most probable hypothesis. Only if the presence of the Europa haplotypes in Madagascar could be reasonably excluded, the third hypothesis would be supported. Lygodactylus verticillatus is adapted to dry climatic conditions. In south-western Madagascar, the species inhabits shrubs and trees in semi-arid areas as well as fences and walls in human settlements (Puente et al. 2009). On Europa Island, it is widely distributed and occurs in most natural habitats on rocks, shrubs, trees and in a dry euphorbia forest as well as in human settlements (Brygoo 1966; Sanchez et al. 2015, 2019). The conditions for L. verticillatus in the coastal regions of south-western Madagascar and on Europa are very similar, facilitating the survival on the island., Published as part of Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver & Vences, Miguel, 2023, Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean, pp. 232-250 in Zootaxa 5311 (2) on pages 245-246, DOI: 10.11646/zootaxa.5311.2.4, http://zenodo.org/record/8094276, {"references":["Boettger, O. (1913) Reptilien und Amphibien von Madagascar, den Inseln und dem Festland Ostfarikas. In: Voeltzkow, A. (Ed.), Reise in Ostafrika in den Jahren 1903 - 1905. Wissenschaftliche Ergebnisse. Vol. 3. Systematische Arbeiten. Schweizerbartsche Verlagsbuchhandlung, Nagele und Sprosser, Stuttgart, pp. 269 - 375.","Pasteur, G. (1965) Recherches sur l'evolution des lygodactyles, lezards afromalgaches actuels. Travaux de l'Institut Scientifique Cherifien, Serie Zoologie, 29, 1 - 132.","Caceres, S. (2003) Etude prealable pour le classement en Reserve Naturelle des Iles Eparses. Memoire de DESS Sciences et Gestion de l'Environnement Tropical. Univ. Reunion (Laboratoire ECOMAR) & DIREN Reunion, 136 pp. Available from: http: // poupin. joseph. free. fr / pdf / caceres- 2003 - iles-eparses-classement. pdf (accessed 13 June 2023)","Fricke, R., Durville, P., Bernardi, G., Borsa, P., Mou-Tham, G. & Chabanet, P. (2013) Checklist of the shore fishes of Europa Island, Mozambique Channel, southwestern Indian Ocean, including 302 new records. Stuttgarter Beitrage zur Naturkunde, A, Neue Serie, 6, 247 - 276.","Courgeon, S., Jorry, S. J., Camoin, G. F., BouDagher-Fadel, M. K., Jouet, G., Revillon, S., Bachelery, P., Pelleter, E., Borgomano, J., Poli, E. & Droxler, A. W., (2016) Growth and demise of Cenozoic isolated carbonate platforms: New insights from the Mozambique Channel seamounts (SW Indian Ocean). Marine Geology, 380, 90 - 105. https: // doi. org / 10.1016 / j. margeo. 2016.07.006","Ali, J. R. & Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 463, 653 - 656. https: // doi. org / 10.1038 / nature 08706","Puente, M., Glaw, F., Vieites, D. R. & Vences, M. (2009) Review of the systematics, morphology and distribution of Malagasy dwarf geckos, genera Lygodactylus and Microscalabotes (Squamata: Gekkonidae). Zootaxa, 2103, 1 - 76. https: // doi. org / 10.11646 / zootaxa. 2103.1.1","Brygoo, E. R. (1966) Note sur les reptiles terrestres recoltes a Europa en avril 1964. Memoires du Museum national d'histoire naturelle, Serie A, 41, 29 - 32.","Sanchez, M. & Probst, J. - M. (2015) L'herpetofaune terrestre de l'ile d'Europa (Ocean Indien, Canal du Mozambique): synthese des connaissances et nouvelles donnees sur la repartition de l'ecologie des especes en vue de leur conservation. Bulletin de la Societe Herpetologique de France, 145, 63 - 76."]}

4.1. Lygodactylus pakenhami At present, the taxon ‘ pakenhami’ is considered a subspecies of L. grotei in most publications. In our study, L. grotei and L. g. pakenhami are genetically distinct lineages, confirming previous results (Gippner et al. 2021). Lygodactylus g. pakenhami from Pemba Island differs greatly in 16S and ND2 pairwise distances from mainland individuals considered as L. grotei. While L. grotei and L. g. pakenhami as far as known cannot be distinguished in scalation, they clearly differ in coloration. Furthermore, hatchlings of L. grotei and L. g. pakenhami are easily identifiable and distinguishable by the coloration of their trunks and especially of their tails. The reciprocal monophyly suggested by mitochondrial DNA, conspicuously different coloration—both in adults and hatchlings—and the high genetic distance suggest that these taxa most likely represent two distinct species. This is also in agreement with differences in the RAG1 sequences (although these data are not fully conclusive due to low sample sizes: one sequence available each for grotei and pakenhami). Therefore, we elevate L. g. pakenhami to full species status, as Lygodactylus pakenhami Loveridge, 1941, being aware that this taxonomic hypothesis requires further testing, especially from nuclear-encoded DNA data sets, ideally at the phylogenomic level. Lygodactylus pakenhami is endemic on Pemba Island, the northernmost and second largest island of the Zanzibar Archipelago. Pemba lies off the continental shelf and is surrounded by water 500 to 850 m deep (Moreau & Pakenham 1941). Geological evidence indicates that Pemba Island was separated from the African mainland by faulting that produced the Pemba Channel, possibly during the late Miocene or early Pliocene, 6 million years ago (Stockley 1942; Clarke & Burgess 2000). Corresponding to its long period of isolation, Pemba is characterized by a remarkable number of endemic species, including plants, mammals and reptiles (R̂dder et al. 2010). The simplest explanation for the existence of L. pakenhami on Pemba is vicariance. Its nearest extant relative, L. grotei, is widely distributed in south-eastern Tanzania and northern Mozambique, including the coastal regions. Presumably, a population of an ancestor of L. grotei and L. pakenhami already existed on Pemba Island before its separation from the mainland. Thereafter, there was no further genetic exchange between the continental and the insular population, so that the latter evolved isolated from its continental congener. While a natural dispersal from Tanzania after the separation of Pemba Island cannot be excluded, it is considered as unlikely because the geographic position of Pemba suggests that the East African Coastal Current or a similar paleocurrent conveyed any drifting material into the open ocean very rapidly (Moreau & Pakenham 1940; Hawlitschek et al. 2016a). A recent, human-mediated transportation is unlikely, as after such a short period of isolation we would expect the sharing of mitochondrial haplotypes between mainland and island populations, which we did not detect. Although only two samples yielding DNA sequences of L. grotei have precise georeferenced information, one of these was collected on the mainland directly opposite Pemba Island. However, as is always the case with inferences of allopatric occurrence, we cannot fully exclude that undiscovered populations of L. pakenhami may occur on the African mainland, which would invalidate our biogeographic hypotheses but not our taxonomic conclusions. In contrast to Pemba Island, Zanzibar and Mafia islands lie on the continental shelf and are separated from the mainland only by shallow waters with an average depth of 30 to 35 m, rarely 50 m (Moreau & Pakenham 1941). Geological data of coastal eastern Africa point to a land connection of Zanzibar and Mafia islands with the African mainland up to the end of the Pleistocene, probably only 10,000 –18,000 years ago (Stockley 1942; Clarke & Burgess 2000). Zanzibar and Mafia are inhabited by L. grotei, L. picturatus and L. viscatus, all conspecific with continental populations. Presumably, the distribution of these three species on the islands is probably best explained by recent vicariant events. In contrast to L. pakenhami on Pemba Island, the species on Zanzibar and Mafia have been isolated on the islands for a relatively short time, about 10,000 years. Additionally, both natural dispersal over water after the separation of Zanzibar and Mafia from the mainland and/or a human-mediated transportation due to long-standing trading in the recent past cannot be excluded., Published as part of Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver & Vences, Miguel, 2023, Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean, pp. 232-250 in Zootaxa 5311 (2) on page 244, DOI: 10.11646/zootaxa.5311.2.4, http://zenodo.org/record/8094276, {"references":["Gippner, S., Travers, S. L., Scherz, M. D., Colston, T. J., Lyra, M. L., Ashwini, V. M., Multzsch, M., Nielson, S. V., Rancilhac, L., Glaw, F., Bauer, A. M. & Vences, M. (2021) A comprehensive phylogeny of dwarf geckos of the genus Lygodactylus, with insights into their systematics and morphological variation. Molecular Phylogenetics and Evolution, 165, 107311. https: // doi. org / 10.1016 / j. ympev. 2021.107311","Loveridge, A. (1941) New geckos (Phelsuma and Lygodactylus), snake (Lepotyphlops) and frog (Phrynobarachits) from Pemba Island, East Africa. Proceedings of the Biological Society, Washington, 54 (8), 175 - 178.","Stockley, G. M. (1942) The geology of the Zanzibar Protectorate and its relation to the East African mainland. Geological Magazine, 79 (4), 233 - 240. https: // doi. org / 10.1017 / S 0016756800073921","Clarke, G. P. & Burgess, N. D. (2000) Geology and geomorphology. In: Burgess, N. D. & Clarke, G. P. (Eds.), Coastal Forests of Eastern Africa. IUCN, Gland, pp. 29 - 40.","Moreau, R. E. & Pakenham, R. H. W. (1940) The land vertebrates of Pemba, Zanzibar, and Mafia: a zoogeographical study. Proceedings of the Zoological Society, London, 110 (A), 97 - 128. https: // doi. org / 10.1111 / j. 1469 - 7998.1941. tb 08463. x","Hawlitschek, O., Garrido, S. R. & Glaw, G. (2016 a). How marine currents influenced the widespread natural overseas dispersal of reptiles in the Western Indian Ocean. Journal of Biogeography, 44, 1426 - 1440. https: // doi. org / 10.1111 / jbi. 12940"]}

4.2. Lygodactylus insularis Lygodactylus insularis is an endemic species on Juan de Nova, a very small island with a total area of approximately 5 km 2, 280 km from mainland Africa (East Mozambique) and 120 km to the nearest point on the Malagasy coast (Caceres 2003; Lambs et al. 2016). Pasteur (1965) already included L. insularis in the L. capensis group and assumed that L. insularis originated from either L. c. capensis or L. c. grotei. Lygodactylus capensis and L. insularis share the same number of precloacal pores and a superficially similar coloration of the adults. However, they strongly differ in the status of the subcaudals and in the appearance of their hatchlings. It is thus impossible to discern the relationships of L. insularis, L. capensis and L. grotei by means of scalation or coloration alone. Our molecular phylogenetic analyses reveal that L. insularis is embedded within the L. capensis group, clearly indicating its African origin and, in so far, supporting Pasteur (1965). While the single gene as well as the multigene analyses fully support closer relationships between L. insularis and L. capensis than between L. insularis and L. grotei, the position of L. insularis within the clade formed by L. insularis, L. nyaneka, L. capensis sensu stricto and six L. aff. capensis lineages (provisionally named as “groups”; here, the last two of these are referred to as L. capensis sensu lato) is not clearly resolved. For a taxonomic revision of the L. capensis complex—one is stated to be in progress by Travers et al. unpublished (in Marques et al. 2020)—we strongly recommend a broader incorporation of genetic markers to resolve the conflicting topologies since our multigene dataset consists of only few markers especially for L. nyaneka and for the numerous groups forming L. capensis sensu lato. The clade including L. grotei and L. pakenhami is defined as the sister clade of L. insularis, L. nyaneka and L. capensis sensu lato. Lygodactylus insularis is recovered as a monophyletic group based on mtDNA, probably resulting from a single dispersal event. Furthermore, the species based on the sampling herein exhibits little intraspecific genetic variability, possibly due to the tiny size of the island resulting in a panmictic population. Members of the genus Lygodactylus successfully crossed the Mozambique Channel from mainland Africa back to Madagascar one or perhaps two times (R̂ll et al. 2010; Mezzasalma et al. 2017; Gippner et al. 2021). The ancestor of L. insularis probably began at least one further traverse in eastward direction ending on Juan de Nova, an island in the middle of the channel. Juan de Nova developed as the uplifted top of a seamount that probably was formed during the Cretaceous along the Davie Fracture (Riaux-Gobin & Witkowski 2012). Ali & Hedges (2022) suggest that it may have remained emerged as a low elevation atoll since the Early Palaeocene. In such a scenario, the dispersal of the ancestors of L. insularis may have taken place as early as 22.3 (confidence interval 15.6–30.2) mya, the stem age of the L. capensis group estimated in the Lygodactylus timetree of Gippner et al. (2021). Results of palaeogeographic reconstructions and palaeo-oceanographic modelling suggest strong surface currents in the Mozambique Channel flowing from northeast Mozambique and Tanzania eastward towards Madagascar during the Palaeogene (66 to 23 mya), which would support this scenario (Ali & Huber 2010). While the main contemporary direction of oceanic surface currents is westward, the situation in the Mozambique Channel is very complex with numerous eddies and possible countercurrents (Hawlitschek et al. 2016a). This also suggests that dispersal could have occurred during a more recent period of mainly westward marine currents. An alternative hypothesis, dispersal via short-lived Cenozoic land-bridges between mainland Africa and Madagascar (e.g., Mazza et al. 2019; Masters et al. 2020), is disputed and has been ruled out based on current geological reconstructions (e.g., Ali & Huber 2010; Ali & Hedges 2022). It also must be emphasized that despite a reasonable sampling of Eastern Africa, especially Mozambique (with L. capensis group sequences from 14 localities available so far), it is possible that a still undiscovered species representing the sister lineage of L. insularis may exist in this region. A somewhat similar situation was uncovered in the frogs of the Ptychadena mascariensis complex in which the sister lineage of the Malagasy species has only been recorded from a small area of Malawi and could have easily been overlooked without intensive sampling efforts (Zimkus et al. 2017). If a closer relative of L. insularis was found on the African mainland, it would lead to younger age estimates of the colonization event of Juan de Nova by these geckos., Published as part of Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver & Vences, Miguel, 2023, Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean, pp. 232-250 in Zootaxa 5311 (2) on pages 244-245, DOI: 10.11646/zootaxa.5311.2.4, http://zenodo.org/record/8094276, {"references":["Caceres, S. (2003) Etude prealable pour le classement en Reserve Naturelle des Iles Eparses. Memoire de DESS Sciences et Gestion de l'Environnement Tropical. Univ. Reunion (Laboratoire ECOMAR) & DIREN Reunion, 136 pp. Available from: http: // poupin. joseph. free. fr / pdf / caceres- 2003 - iles-eparses-classement. pdf (accessed 13 June 2023)","Lambs, L., Mangion, P., Mougin, E. & Fromard, F. (2016) Water cycle and salinity dynamics in the mangrove forests of Europa and Juan de Nova Islands, southwest Indian Ocean. Rapid Communications in Mass Spectrometry, 30, 311 - 320. https: // doi. org / 10.1002 / rcm. 7435","Pasteur, G. (1965) Recherches sur l'evolution des lygodactyles, lezards afromalgaches actuels. Travaux de l'Institut Scientifique Cherifien, Serie Zoologie, 29, 1 - 132.","Marques, M. P., Ceriaco, L. M. P., Buehler, M. D., Bandeira, S. A., Janota, J. M. & Bauer, A. M. (2020) A revision of the dwarf geckos, genus Lygodactylus (Squamata: Gekkonidae), from Angola, with description of three new species. Zootaxa, 4853 (3), 301 - 352. https: // doi. org / 10.11646 / zootaxa. 4853.3.1","Mezzasalma, M., Andreone, F., Aprea, G., Glaw, F., Odierna, G. & Guarino, F. M. (2017) Molecular phylogeny, biogeography and chromosome evolution of Malagasy geckos of the genus Lygodactylus (Squamata, Gekkonidae). Zoological Scripta, 46, 42 - 54. https: // doi. org / 10.1111 / zsc. 12188","Gippner, S., Travers, S. L., Scherz, M. D., Colston, T. J., Lyra, M. L., Ashwini, V. M., Multzsch, M., Nielson, S. V., Rancilhac, L., Glaw, F., Bauer, A. M. & Vences, M. (2021) A comprehensive phylogeny of dwarf geckos of the genus Lygodactylus, with insights into their systematics and morphological variation. Molecular Phylogenetics and Evolution, 165, 107311. https: // doi. org / 10.1016 / j. ympev. 2021.107311","Riaux-Gobin, C. & Witkowski, A. (2012) Small-sized and discoid species of the genus Cocconeiopsis (Bacillariophyta) on Holothuria atra (Juan de Nova, Mozambique Channel). Phytotaxa, 54, 43 - 58. https: // doi. org / 10.11646 / phytotaxa. 54.1.5","Ali, J. R. & Hedges, S. B. (2022) A review of geological evidence bearing on proposed Cenozoic land connections between Madagascar and Africa and its relevance to biogeography. Earth-Science Reviews, 232, 104103. https: // doi. org / 10.1016 / j. earscirev. 2022.104103","Ali, J. R. & Huber, M. (2010) Mammalian biodiversity on Madagascar controlled by ocean currents. Nature, 463, 653 - 656. https: // doi. org / 10.1038 / nature 08706","Hawlitschek, O., Garrido, S. R. & Glaw, G. (2016 a). How marine currents influenced the widespread natural overseas dispersal of reptiles in the Western Indian Ocean. Journal of Biogeography, 44, 1426 - 1440. https: // doi. org / 10.1111 / jbi. 12940","Mazza, P. P. A., Buccianti, A. & Savorelli, A. (2019) Grasping at straws: a re-evaluation of sweepstakes colonisation of islands by mammals. Biological Reviews, 94, 1364 - 1380. https: // doi. org / 10.1111 / brv. 12506","Masters, J. C., Genin, F., Zhang, Y., Pellen, R., Huck, T., Mazza, P. P. A., Rabineau, M., Doucoure & M., Aslanian, D. (2020) Biogeographic mechanisms involved in the colonization of Madagascar by African vertebrates: Rifting, rafting and runways. Journal of Biogeography, 48, 492 - 510. https: // doi. org / 10.1111 / jbi. 14032","Zimkus, B. M., Lawson, L. P., Barej, M. F., Barratt, C. D., Channing, A., Dash, K. M., Dehling, J. M., Du Preez, L., Gehring, P. - S., Greenbaum, E., Gvozdik, V., Harvey, J., Kielgast, J., Kusamba, C., Nagy, Z. T., Pabijan, M., Penner, J., R ˆ del, M. - O., Vences, M. & L ˆ tters, S. (2017) Leapfrogging into new territory: How Mascarene ridged frogs diversified across Africa and Madagascar to maintain their ecological niche. Molecular Phylogenetics and Evolution, 106, 254 - 269. https: // doi. org / 10.1016 / j. ympev. 2016.09.018"]}

Röll, Beate, Sanchez, Mickaël, Gippner, Sven, Bauer, Aaron M., Travers, Scott L., Glaw, Frank, Hawlitschek, Oliver, Vences, Miguel (2023): Supplementary Materials for Phylogeny of dwarf geckos of the genus Lygodactylus (Gekkonidae) in the Western Indian Ocean. Zootaxa (supp.) 5311 (2): 1-18, DOI: http://doi.org/10.5281/zenodo.8102622

Guibemantis (Pandanusicola) pulcherrimus sp. nov. Figs. 7‒9 Holotype. ZSM 518/2009 (field number ZCMV 11217), adult female, collected by M. Vences, D.R. Vieites, F.M. Ratsoavina, R.D. Randrianiaina, E. Rajeriarison, T. Rajoafiarison and J.L. Patton on 23 June 2009 near a site locally named Hevirina, western slope of the Makira Massif, Analanjirofo Region, Madagascar (GPS coordinates not taken, but coordinates of Hevirina are 15.4490°S, 49.1119°E, 1093 m a.s.l.). Paratypes. Eight specimens. ZSM 519/2009 (ZCMV 11249) and ZSM 520/2009 (ZCMV 11264), two adult females, collected by M. Vences, D.R. Vieites, F.M. Ratsoavina, R.D. Randrianiaina, E. Rajeriarison, T. Rajoafiarison and J.L. Patton on 23–24 June 2009 at a campsite near the source of Fotsialanana river, western slope of the Makira Massif, Analanjirofo Region, Madagascar (coordinates 15.46675°S, 49.128889°E, 1067 m a.s.l.); ZSM 1754/2010 (ZCMV 12505), adult female, collected by M. Vences, D.R. Vieites, R.D. Randrianiaina, F.M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison and T. Rajoafiarison on 27 June 2010 at Bemanevika, campsite Atsirakala, Sofia Region, Madagascar (coordinates 14.43061°S, 48.60179°E, 1466 m a.s.l.); as well as five specimens (field numbers ZCMV 11225, ZCMV 11226, ZCMV 11228, ZCMV 11229), all collected in June 2009 by by M. Vences, D.R. Vieites, R.D. Randrianiaina, F.M. Ratsoavina, S. Rasamison, A. Rakotoarison, E. Rajeriarison and T. Rajoafiarison on the western slope the Makira Massif, not examined morphologically but included in the 16S tree (Fig. 1), integrated but still uncatalogued at the UADBA (MZBA) collection (individually identifiable via their field number tags). Diagnosis. Assigned to the subgenus Pandanusicola of the genus Guibemantis based on its small body size, phytotelm-dwelling habitats (in Pandanus plants), near-absent webbing between toes, connected lateral metatarsalia, the presence of both inner and outer metatarsal tubercles, intercalary elements between ultimate and penultimate phalanges of fingers and toes (verified by external examination), and molecular phylogeny. The new species is characterized by the unique combination of the following characters: (1) medium size with SVL 25‒29 mm in females, (2) bright green dorsal color with translucent shade, with dense fine purplish-brown spots, (3) distinct brown rostral stripe, (4) absence of white or yellow spots in the inguinal region, (5) absence of webbing on hand and only traces of web at foot, (6) vomerine teeth rudimentary. The new species is most similar to its sister species G. pulcher, which differs (despite some overlap) by fewer (12–107, mean 42; vs. 47–223, mean 103) and larger (diameter often up to 0.8 mm vs. 0.4 mm) purplish-brown dorsal spots, and a larger and denser purplish-brown band on the flanks which often is adorned centrally by silvery whitish reticulations, as well as by numerous diagnostic nucleotide positions in the mitochondrial 16S rRNA gene. Description of the holotype. Adult female in good state of preservation. Tissue removed from ventral side of right thigh for DNA extraction, one transverse and two longitudinal cuts on venter for gonad examination. Head longer than wide and slightly wider than body; snout rounded in dorsal, ventral and lateral views; canthus rostralis straight, loreal region flat; nostrils much nearer to tip of snout than to eye, tympanum distinct, 47% of horizontal eye diameter; supratympanic fold poorly expressed, rather straight; vomerine odontophores very poorly expressed and almost not visible, located medially between eye and choanae on either side of head; tongue ovoid, and distinctly bifid at its tip. Arms moderately thin; relative finger length 1 After 14 years in preservative, dorsally beige with many small spots; these spots are light brown with a weakly recognizable reddish/purple shade. From the snout tip over the nostrils to the eye, and then again from the eye over the tympanum to an area above the forelimb insertions, a distinctly delimited dark brown rostral stripe is present. After the forelimb insertion, this stripe becomes broader, lighter and less distinctly delimited, and forms a brown band on the anterior flanks, dissolving into dense brown spotting on the posterior flanks. The ventral side is uniformly beige without any pattern. In life (Fig. 7A) the dorsal side was green with a translucent shade, and the spots and bands were purplish-brown. Variation. The three morphologically examined paratypes are similar to the holotype. See Table 1 for morphological measurements. Sexual dimorphism cannot be reliably assessed as all genotyped specimens are females. Photos from specimens morphologically assigned to this species from Marojejy and Anjanaharibe-Sud agree with the overall trend of rather small dorsal spots and poorly marked purplish-brown flank patches. Etymology. The species epithet pulcherrimus is a Latin adjective meaning “most beautiful”, referring to the appealing color pattern of the new species and alluding to the name of its sister species (G. pulcher). Natural history. All known specimens were collected from the leaf axils of Pandanus screw pines in rainforest. Calls, reproductive behavior and larval stages are unknown, but it is likely that the species reproduces in these phytotelms, as does its sister species G. pulcher. Distribution. Based on genotyped specimens, G. pulcherrimus is known from (1) Hevirina and Fotsialanana source, two nearby sites on the western slope of the Makira Massif, and (2) Bemanevika. Additional specimens morphologically consistent with this taxon are known from three additional localities in northern Madagascar: (3) Anjanaharibe-Sud, (4) Marojejy, and (5) Masoala; these localities need to be genetically verified in the future.

Vences, Miguel, Hutter, Carl R., Glaw, Frank, Rakotoarison, Andolalao, Raselimanana, Achille P., Scherz, Mark D. (2023): A new species of Pandanus-dwelling frog from northern Madagascar related to Guibemantis pulcher. Zootaxa 5306 (1): 97-115, DOI: 10.11646/zootaxa.5306.1.4, URL: http://dx.doi.org/10.11646/zootaxa.5306.1.4

Sigwart, Julia D., Chen, Chong, Tilic, Ekin, Vences, Miguel, Riehl, Torben (2023): Why is there no service to support taxonomy? BioEssays 2300070: 1-4, DOI: 10.1002/bies.202300070, URL: http://dx.doi.org/10.1002/bies.202300070

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 brevirostris sp. nov. Identity and justification.—A deep genetic lineage of the M. biporus clade known from Betampona, and Sahavontsira. This lineage has been considered as unconfirmed candidate species M. sp. 31 by Vieites et al. (2009) and M. sp. Ca31 by Perl et al. (2014). It was referred to as ‘ M. sp. aff. biporus [Ca FJ559260]’ by Rosa et al. (2012). According to the phylogenomic analysis, it represents the sister taxon of another lineage named below (M. eulenbergeri sp. nov.) but differs from that lineage by a 16S distance of 8.6–9.1%, and possibly by at least one morphological difference in foot webbing, suggesting a status as distinct species. Diagnosis.— Mantidactylus brevirostris sp. nov. is a member of the M. biporus clade, sister to the new species M. eulenbergeri sp. nov. (described below) according to our phylogenomic analysis. See Table 4 for a list of diagnostic morphological characters. The combination of a small body size (male SVL 23 mm, female SVL 28 mm), rather smooth dorsal skin without dorsolateral ridges, large tympanum size in males (12% of SVL), presence of white spots on flanks, and absence of a white marking on the snout tip, distinguishes M. brevirostris sp. nov. from species of the M. betsileanus, M. curtus, M. fergusoni, M. tricinctus, and M. ulcerosus clades. The distantly related M. inaudax (M. inaudax clade) is morphologically very similar but appears to reach larger body sizes, has more developed foot webbing, and has in many individuals a pattern where the colour of flanks differs from that on the dorsum; M. biporus occurs syntopically with M. brevirostris sp. nov. but has a larger body size and a more developed foot webbing; M. augustini has longer hindlimbs and a more developed foot webbing; M. bletzae has a more granular dorsal skin with dorsolateral ridges and a more 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. brevirostris sp. nov. in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Holotype.— MRSN A6257 (FAZC 13581), adult male, collected by G.M. Rosa, and F. Andreone on 9 February 2007 at Sahambendrana, Réserve Naturelle Intégrale de Betampona (17.8984°S, 049.2154°E, 458 m a.s.l.), Antsinanana Region, Madagascar. A 16S barcode sequence of the holotype is available from GenBank (accession HM364736). Paratypes.—A single paratype: ZSM 185/2021 (ACZCV 265, extraction ACP 2211; tissue ACZC 6309), adult female, collected by A. Crottini, D. Salvi, E. Scanarini, George, J. N̂el, and F. Andreone on 22 November 2013 at Betampona (Sahabefoza). Description of the holotype.—Adult male in good state of preservation (Fig. 62). Fourth and fifth finger from right foot missing (taken as tissue sample). Body rather stout. Head as wide as body. Snout rounded in dorsal view, somewhat truncate in lateral 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, as wide as high, horizontal diameter of tympanum 89% of horizontal eye diameter. Supratympanic fold not clearly recognisable, basically corresponding to outer edge of tympanum. Tongue ovoid, bifid posteriorly. Maxillary teeth present. Vomerine teeth form two somewhat elongate 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 10. See Fig. 68 for colouration in life and its variation. Too few specimens have been sexed to assess the degree of sexual size dimorphism. Femoral glands in life are not documented (no photographs of the ventral side available). Natural history.—Species usually observed in slowrunning parts of streams and other small courses. Active both day and night. Quite shy and able to hide under the mud or actively swimming when disturbed. Calls.— The call of this species has not been recorded. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to low-elevation (Etymology. — The species epithet is a Latin thirddeclension two-termination adjective, derived from the adjective ‘brevis’, meaning short, and ‘rostrum’, meaning snout, in the genitive singular, and refers to the short snout observed in several individuals 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 273-275, 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","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"]}

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 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: I Finger discs slightly enlarged. Nuptial pads absent. Foot slightly longer than tibia (104%). Lateral metatarsalia separated. Inner metatarsal tubercle present, small. Outer metatarsal tubercle not clearly recognisable. Webbing formula: 1(1), 2i(1.5), 2e(0.75), 3i(1.75), 3e(1), 4i(1.5), 4e(1), 5(0.5). Relative length of toes: I Skin on the upper surface smooth. Ventral side smooth. Femoral gland distinct. Variation.—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.

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 katae sp. nov. Identity and justification.— This lineage has been considered as confirmed candidate species M. sp. 28 by Vieites et al. (2009) and M. sp. Ca28 by Perl et al. (2014). It was reported as ‘ Mantidactylus sp. aff. betsileanus “slow calls”’ by Glaw and Vences (2007), and has previously been considered as M. multiplicatus (e.g. Poth et al. 2012, 2013); however, DNA barcodes obtained in the present study have shown that this assignment was wrong and that the holotype of M. multiplicatus is conspecific with M. betsileanus, and the nomen therefore a junior synonym of M. betsileanus (see account of that species). Mantidactylus katae is a widespread species with a unique call, characterized by a very low pulse repetition rate (or rather, consisting of a single-pulse note repeated regularly). It also is genetically distinct in mitochondrial DNA and does not appear to share Rag-1 haplotypes with other similar species, including the sympatric (and syntopic) M. betsileanus. Unfortunately, due to failure of other samples, M. katae sp. nov. is only represented by two individuals from the South East of Madagascar in our phylogenomic tree, from where no reliable call recordings are available. The samples (FGZC 163 and ZCMV 14842) cluster closely with M. tripunctatus. However, these samples correspond to those alleles that also in the Rag-1 haplotype network cluster close to M. tripunctatus, and we cannot exclude that they represent individuals of M. tripunctatus with an introgressed M. katae mitochondrial genome. The exact phylogenetic position of M. katae therefore remains in need of confirmation; the phylogenetic tree of Wollenberg et al. (2011), based on multiple mitochondrial genes, placed it (as M. sp. 28) in a clade with M. noralottae and M. tripunctatus (as M. sp. 29), which would agree with the affinities suggested by our phylogenomic tree. Typical M. katae from the Southern Central East and Northern Central East of Madagascar differ from both M. noralottae and M. tripunctatus very strongly in advertisement call structure, by femoral gland morphology, and do not share Rag-1 haplotypes with these two species, thus leaving no doubt about the species status of this lineage. Holotype.— ZSM 79/2002 (FGMV 2001.1179), an adult male, collected by M. Vences on 1 December 2001 at Andasibe (ca 18.9333°S, ca 048.4167°E, 920 m a.s.l.), Alaotra-Mangoro Region, Madagascar. A 16S barcode sequence of the holotype was obtained in this study and was included in the analysis. Paratypes.— A total of 17 paratypes: ZSM 79– 81; 83/2002 (FGMV 2001.1197 = 2002.G24; FGMV 2001.1180 = 2002.G25; FGMV 2001.1275 = 2002.G66; FGMV 2001.1173 = 2002.G18), two adult females and two adult males, collected by M. Vences on 1–5 December 2001 at Andasibe (ca 18.9333°S, 048.4167°E, 920 m a.s.l.); ZSM 637/2003 (FG / MV 2002.132), adult male, collected by F. Glaw, M. Puente, L. Raharivololoniaina, M. Teschke (née Thomas), and D.R. Vieites on 15 January 2003 beside a small brook in Ranomafana National Park (21.250°S, 047.450°E, 932 m a.s.l.); ZSM 668/2003 (FG / MV 2002.255), adult male, collected by F. Glaw, M. Puente, L. Raharivololoniaina, M. Teschke (née Thomas), and D.R. Vieites on 16 January 2003 in Ranomafana National Park (21.250°S, 047.450°E, 932 m a.s.l.); ZSM 708/2003 (FG / MV 2002.348), adult male, collected by F. Glaw, M. Puente, L. Raharivololoniaina, M. Teschke (née Thomas), and D.R. Vieites on 20 January 2003 at Kidonafo Bridge, Ranomafana (21.2262°S, 047.3696°E, 1152 m a.s.l.); ZSM 196/2021 (FAZC 15504, extraction ACP 3659, tissue ACZC 8591), ZSM 197/2021 (FAZC 15507, ACP 3662, ACZC 8594), and MRSN A7043 (FAZC 15523, ACP 3662, ACZC 8594), two males and one female collected at Maromizaha (18.9771°S, 048.4682°E, ca 1000 m a.s.l.), in January 2017 by E. Coppola; ZMB 81918 (JCR 105), adult male collected on 16 March 2010 by J.C. Riemann, and S.H. Ndriantsoa at Andalangina, Ranomafana area (21.29844°S, 047.60343°E, 480 m a.s.l.); ZMB 81920 (field NSH 1069; GenBank JCR 1069), adult female, collected on 12 May 2010 by J.C. Riemann, and S.H. Ndriantsoa at Ambatovory, Ranomafana area (21.23966°S, 047.42581°E, 953 m a.s.l.); ZMB 81922 (field NSH 2577; GenBank JCR 2577), adult male, collected on 26 March 2012 by J.C. Riemann, and S.H. Ndriantsoa at Sahamalaotra, Ranomafana area (21.23688°S, 047.39887°E); UADBA-A 43149 (JCR 106), adult male collected on 16 March 2010 by J.C. Riemann, and S.H. Ndriantsoa at Andalangina, Ranomafana area (21.29844°S, 047.60343°E, 480 m a.s.l.); UADBA-A 62106 (JCR 245), subadult collected on 21 April 2010 by J.C. Riemann, and S.H. Ndriantsoa atAmbolo, Ranomafana area (21.26386°S, 047.50862°E, 643 m a.s.l.); UADBA-A 62104 (JCR 320), adult female collected on 21 May 2010 by J.C. Riemann, and S.H. Ndriantsoa at Ambolo, Ranomafana area (21.26307°S, 047.50696°E, 660 m a.s.l.); UADBA-A 62105 (JCR 323), adult female collected on 21 May 2010 by J.C. Riemann, and S.H. Ndriantsoa at Ambolo, Ranomafana area (21.26307°S, 047.50696°E, 660 m a.s.l.). Additional material.— The following specimens are assigned to M. katae sp. nov. based on morphology, but have not been DNA barcoded: ZMA 20232 (ZCMV 236), adult male, collected by M. Vences and I. de la Riva on 24 January 2004 at Ranomafana National Park, Maharira base camp (21.3258°S, 047.4024°E, 1248 m a.s.l.); ZFMK 62212, adult female, collected by F. Glaw, D. Rakotomalala, and F. Ranaivojaona on 10 March 1996 at Andasibe; ZMA 6828-863; 6828-864, adult male and female, collected on 23 September 1972, and ZMA 6886-641–643, three adult males, collected by R.M.A. Blommers-Schl̂ sser on 4 April 1972 at Andasibe; ZMA 6833-149 and 6833-156–158, two adult females and two adult males, collected by R.M.A. Blommers-Schl ̂sser on 1 July 1971 at Ranomafana. Furthermore, the following specimen (included in our phylogenomic analysis) from the South of Madagascar agrees with M. katae sp. nov. in mitochondrial DNA but due to the absence of bioacoustic data from this population, its identity is in need of confirmation: ZSM 91/2004 (FGZC 163), adult male, collected by F. Glaw, M. Puente, M. Thomas and R. Randrianiaina on 31 January 2004 at ‘Camp 1’ between Isaka and Eminiminy, Andohahela (24.7586°S, 046.8542°E, 247 m a.s.l.). Diagnosis.— Mantidactylus katae sp. nov. is a member of the M. betsileanus clade, related to M. noralottae and M. tripunctatus. See Table 4 for a list of diagnostic morphological characters. The combination of a relatively small body size (male SVL 22–27 mm), slightly tubercular dorsal skin with distinct continuous dorsolateral ridges, absence of white spots on flanks, presence of a white marking on snout tip, large femoral glands (FGW up to 13% of SVL) that contact each other medially, and advertisement call consisting of a single pulse repeated at a slow rate of 10–16 calls per second distinguishes M. katae sp. nov. from species of all other clades.The only other species with a similar advertisement call structure is M. fergusoni sp. nov. which has a larger size and more tubercular dorsal skin (see account of that species below). Within the M. betsileanus clade, the new species can be distinguished from all other species by its unique call structure and larger femoral glands (Table 4); furthermore from M. noralottae by smaller size of males and presence of a distinct white marking on snout. For a field diagnosis from the syntopic species M. betsileanus, according to our measurements, M. katae differs by a smaller dot on the snout tip (dot on the snout tip is 9–14% of head width vs 13–24%), a smaller distance between the femoral glands (distance between the femoral glands is 0– 14% of SVL vs 13–22%), larger femoral glands (femoral gland length is 19–31% of SVL vs 13–22%), and a higher number of granules per femoral gland (5–8 vs 1–5). 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. katae 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). Some muscle tissue removed from right thigh. Femoral glands partly detached for examination in internal view. Body relatively slender. Head slightly wider than body. Snout rounded in dorsal view, slightly 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, slightly wider than high, horizontal diameter of tympanum73% of horizontal eye diameter.Supratympanic fold rather distinct, running rather straight from behind eye and bending about 45° about midway towards forelimb insertion. Tongue ovoid, distinctly bifid. 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 dark brown with some poorly contrasting, slightly darker markings and a slightly lighter band between eyes. Dark crossbands on limbs. Ventrally light beige with dark pattern on chest and throat, also extending on anterior belly. Throat with light colour medially, forming a very irregular and discontinuous broad medial stripe. Upper lip ventrally dark brown with white spots. Colouration of holotype in life not documented. Variation.—Variation in measurements is given in Table 7. See Fig. 42 for colouration in life and its variation. There is moderate sexual size dimorphism (confirmed male SVL 22.1–27.2 mm [n = 7] vs confirmed female SVL 25.6–35.9 mm [n = 3]). Males have a larger tympanum than females (HTD/ED ratio is 61–68% in females, 73– 100% in males). Femoral glands in males are large and very distinct, of a quite characteristic shape. They consist of a large-sized distal ulcerous macrogland located rather close to the insertion of the thighs and thus almost in contact with each other. Consequently, the proximal granular gland field (by definition located proximally from the ulcerous macrogland) is basically absent in these frogs. Glands are typically coloured yellowish in life. Natural history.—Occurs along running water bodies in rainforest, where specimens can often be seen calling at night from muddy banks of somewhat larger, slow-moving streams. It often occurs in syntopy with M. betsileanus in rainforest, including forest fragments (Riemann et al. 2015), and along streams with at least a narrow gallery forest band surrounded by degraded or unforested areas. However, in contrast to M. betsileanus it is not found in rice fields or other plantations (Ndriantsoa et al. 2017). Usually sitting in shallow water or along the shore, hiding in the leaf litter or perched on low vegetation up to 0.5 m hight in the vicinity of a stream. Found at an elevational range between 450–1142 m a.s.l. in Ranomafana and surrounds. Females with visible eggs (transparent abdominal wall) were observed in May 2010 and from January to June in 2011 in the Ranomafana area. Calls.—The advertisement call of M. katae, recorded on 17 December 1994 at Andasibe, ca 20°C air temperature (Vences et al. 2006: CD2, track 64, cut 2), consists of a simple, very short, single pulse ‘click’ note, emitted in long series at regular intervals and fast succession (Fig. 43). The duration of the call series analysed from Andasibe was 7568 ms. Numerical parameters of 47 analysed calls were as follows: call duration (= note duration) 2–5 ms (3.6 ± 0.8 ms); 1 pulse per note (1.0 ± 0.0); pulse duration = note duration = call duration; dominant frequency 3036–3165 Hz (3108 ± 46 Hz); prevalent bandwidth 1250–3540 Hz; call repetition rate (= note repetition rate = pulse repetition rate) within regular series 696–971 calls/min (840 ± 106 calls/min). Calls recorded on 24 January 2004 at Maharira forest, Ranomafana National Park, 18.4°C air temperature (Vences et al. 2006: CD2, track 64, cut 1) are in agreement with the calls from Andasibe in all spectral and temporal parameters, except for slightly lower dominant frequency ranging from 2713–2993 Hz (2833 ± 123 Hz), and slightly lower call repetition rate, ranging from 515–731 calls/min (626 ± 109 calls/min). Both slight differences can be explained by potentially larger body size of the calling male and lower temperature during recording and leave no doubt about the conspecificity of the calls from the two localities. Another call recording from Andasibe, obtained on 20 March 1995, 23.4°C air temperature (Vences et al. 2006: CD2, track 63) is similar to the calls described above, but differs by shorter call series (2226–2520 ms), slightly longer call duration (= note duration = pulse duration) of 8–10 ms, and lower dominant frequency of 1323–1433 Hz (1365 ± 52 Hz). However, the latter might be due to recording equipment and/or recording conditions, as the calls described above have a second frequency peak that roughly corresponds to the lower dominant frequency range described here. Call repetition rate (= note repetition rate = pulse repetition rate) is in the same range compared to the calls described above, ranging from 630–857 calls/ min (741 ± 90 calls/min). Tadpoles.—The tadpole of M. katae was described by Knoll et al. (2007) under the name ‘ M. sp. aff. betsileanus “very slow calls”’. Distribution.— Widespread in eastern Madagascar (Fig. 7). This species is known from Ambatomandondona, Ambohitsara,An’Ala,Andasibe,Andohahela,Andringitra, Bibiango, Ambatofotsy, Ifandiana, Mahatsara-Mantadia, Mariavatra, Mantady, Marolambo, Maromizaha, Pic d’Ivohibe,Ranomafana(varioussites,includingValohoaka, Ambatolahidimy, Ambatolahy, Ambolo, Ampitavanana, Andalangina, Beremby, Bibiango, Kidonavo, Maharira, Ampangadiamesa, Andranovorimainty, Ambodiriana, Imaloka, Sahamalaotra, Talatakely, and Vohiparara), Sahamalotra, Sahateza, Torotorofotsy, and Vohidrazana. Elevation range: 247–1248 m a.s.l. Etymology.—We dedicate this species to Katharina (‘Kat’) Wollenberg Valero, in recognition of her numerous contributions to the research on little brown frogs of Madagascar, and specifically on the behaviour of this species (under the name M. multiplicatus) in the framework of studies of their femoral gland compounds (Poth et al. 2012).

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."]}