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In the era of human-driven climate change, understanding whether behavioural buffering of temperature change is linked with organismal fitness is essential. According to the ‘cost–benefit’ model of thermoregulation, animals that live in environments with high frequencies of favourable thermal microclimates should incur lower thermoregulatory costs, thermoregulate more efficiently and shunt the associated savings in time and energy towards other vital tasks such as feeding, territory defence and mate acquisition, increasing fitness. Here, we explore how thermal landscapes at the scale of individual territories, physiological performance and behaviour interact and shape fitness in the southern rock agama lizard (Agama atra). We integrated laboratory assays of whole organism performance with behavioural observations in the field, fine-scale estimates of environmental temperature, and paternity assignment of offspring to test whether fitness is predicted by territory thermal quality (i.e. the number of hours that operative temperatures in a territory fall within an individual's performance breadth). Male lizards that occupied territories of low thermal quality spent more time behaviourally compensating for sub-optimal temperatures and displayed less. Further, display rate was positively associated with lizard fitness, suggesting that there is an opportunity cost to engaging in thermoregulatory behaviour that will change as climate change progresses., This research was financially and logistically supported by the National Research Foundation (NRF) of South Africa (CPRR no. 98880). K.A. was supported by the NRF Doctoral Scholarship for Full-time Studies. M.L.L. was supported by a United States National Science Foundation Postdoctoral Fellowship in Biology (award number DBI-1402497). S.T. received funding from grant PID2020-117115GA-100 funded by MCIN/AEI/10.13039/501100011033 and by Ramón y Cajal grant RYC2021-03152-I, funded by MCIN/AEI/10.13039/501100011033 and the European Union NextGenerationEU/PRTR.

In the era of human-driven climate change, understanding whether behavioural buffering of temperature change is linked with organismal fitness is essential. According to the ‘cost–benefit’ model of thermoregulation, animals that live in environments with high frequencies of favourable thermal microclimates should incur lower thermoregulatory costs, thermoregulate more efficiently and shunt the associated savings in time and energy towards other vital tasks such as feeding, territory defence and mate acquisition, increasing fitness. Here, we explore how thermal landscapes at the scale of individual territories, physiological performance and behaviour interact and shape fitness in the southern rock agama lizard (Agama atra). We integrated laboratory assays of whole organism performance with behavioural observations in the field, fine-scale estimates of environmental temperature, and paternity assignment of offspring to test whether fitness is predicted by territory thermal quality (i.e. the number of hours that operative temperatures in a territory fall within an individual's performance breadth). Male lizards that occupied territories of low thermal quality spent more time behaviourally compensating for sub-optimal temperatures and displayed less. Further, display rate was positively associated with lizard fitness, suggesting that there is an opportunity cost to engaging in thermoregulatory behaviour that will change as climate change progresses.

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"]}