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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

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

Based on sequences of three mitochondrial and two nuclear-encoded genes, we examine genetic variation in the leaf tailed geckos Uroplatus sameiti and U. sikorae, and morphological and chromatic characteristics of the genetic clusters identified. The mitochondrial phylogeny reveals a puzzling diversity of 16 deep lineages (4 in U. sameiti and 12 in U. sikorae) differing by 2.9–9.9% uncorrected pairwise distance in a fragment of the 16S rRNA gene. Populations from Analalava in the North and Zahamena in the Northern Central East form two mitochondrial lineages clustering with U. sameiti but being deeply divergent (>8% 16S distance); however, no information on their morphology is available. In U. sikorae, the mitochondrial lineages identified form several major geographic clades, two of which (from the northernmost and southernmost populations, respectively) received substantial support in the phylogenetic analysis. No instance of sympatry of two or more mitochondrial lineages was observed, precluding an unambiguous assessment of species status under the biological species criterion without experimental approaches or detailed hybrid zone analyses. In the fragment of the nuclear encoded gene SACS we observed haplotype sharing between species and mitochondrial lineages, while in the fragment of KIAA1239 no haplotype sharing was detected although neither species nor mitochondrial lineages formed coherent phylogroups in the respective network. A screening of colour patterns from live photos, partly of the genotyped individuals, confirmed a large variation within species and populations, with a possible sexual dichromatism where a longitudinally striped phenotype is restricted to males. All individuals from populations of the U. sikorae clade from the Southern Central East and South East had an unpigmented oral mucosa just like U. sameiti, while all other U. sikorae populations are characterized by a black oral mucosa pigmentation. The extremely strong phylogeographic structure in the U. sikorae complex without obvious species-level divergences is unprecedented for large-sized squamates in Madagascar and calls for further taxonomic scrutiny using phylogenomic approaches; and it exemplifies how the loss of any major block of the remaining rainforests in Madagascar will inevitably lead to a substantial loss of genetic diversity – even if often intraspecific – in rainforest-specialized species.

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."]}