Morphological and ecological convergence at the lower size limit for vertebrates highlighted by five new miniaturised microhylid frog species from three different Madagascan genera

Miniaturised frogs form a fascinating but poorly understood amphibian ecomorph and have been exceptionally prone to taxonomic underestimation. The subfamily Cophylinae (family Microhylidae), endemic to Madagascar, has a particularly large diversity of miniaturised species which have historically been attributed to the single genus Stumpffia largely based on their small size. Recent phylogenetic work has revealed that several independent lineages of cophyline microhylids evolved towards highly miniaturised body sizes, achieving adult snout- vent lengths under 16 mm. Here, we describe five new species belonging to three clades that independently miniaturised and that are all genetically highly divergent from their relatives: (i) a new genus (Mini gen.nov.) with three new species from southern Madagascar, (ii) one species of Rhombophryne, and (iii) one species of Anodonthyla. Mini mum sp. nov. from Manombo in eastern Madagascar is one of the smallest frogs in the world, reaching an adult body size of 9.7 mm in males and 11.3 mm in females. Mini scule sp.nov. from Sainte Luce in southeastern Madagascar is slightly larger and has maxillary teeth. Mini ature sp.nov. from Andohahela in southeast Madagascar is larger than its congeners but is similar in build. Rhombophryne proportionalis sp.nov. from Tsaratanana in northern Madagascar is unique among Madagascar's miniaturised frogs in being a proportional dwarf, exhibiting far less advanced signs of paedomorphism than other species of similar size. Anodonthyla eximia sp.nov. from Ranomafana in eastern Madagascar is distinctly smaller than any of its congeners and is secondarily terrestrial, providing evidence that miniaturisation and terrestriality may be evolutionarily linked. The evolution of body size in Madagascar's microhylids has been more dynamic than previously understood, and future studies will hopefully shed light on the interplay between ecology and evolution of these remarkably diverse frogs

Long-term effect of sheep and goat grazing on plant diversity in a semi-natural dry grassland habitat

Semi-natural dry grassland sites are of great importance for nature conservation because they support high species diversity and the abundance of “Red-List” species. Grazing has proved to be a successful management tool in terms of maintenance and restoration of biodiversity. For a deeper understanding of the effects of different grazers on species biodiversity in dry grasslands, it is necessary to study the long-term effects of major changes in grazing management. In a semi-natural dry grassland habitat, which was formerly grazed by cattle, we investigated the changes in plant species composition due to long term grazing by sheep and goats. Specifically we asked: a) How does long-term grazing by sheep and goats change the composition of all plant species and particularly those that are on the Red-List? Are changes caused mainly by species turnover? b) How does long-term grazing by sheep and goats influence the fertility and acidity of the soil? To address these questions, we compared the composition and diversity of plants as well deriving Ellenberg indicator values of the species. Long-term grazing by sheep and goats subsequent to a year-round cattle grazing changed the plant species composition of the dry grasslands resulting in a high species turnover rate. It did not, however, lead to an increase in plant species diversity even though Red-List species were considerably more abundant in 2013. Overall, the grazing regime studied positively influenced vegetation composition. The effects on local species composition due to species turnover might further be influenced by local factors like soil nitrogen availability

Water quality and biotic interaction of two cavefish species: Typhleotris madagascariensis Petit, 1933 and Typhleotris mararybe Sparks & Chakrabarty, 2012, in the Mahafaly Plateau groundwater system, Madagascar

The karstic subterranean aquatic system of the Mahafaly Plateau in south-western Madagascar is inhabited by two species of cavefish: Typhleotris madagascariensis and Typhleotris mararybe. Knowledge about both cavefish species is scant. In order to learn more about the distribution of the two species, 15 caves and sinkholes spread over the Mahafaly Plateau were inventoried for their presence. Abiotic water quality and interspecific relations of the two species were investigated in six of these caves and five of the sinkholes during the dry and the rainy seasons. Typhleotris madagascariensis was present in all sampled water bodies while T. mararybe was restricted to five sites in the region around the town of Itampolo. The inventories extend the known range of both species of Typhleotris on the Mahafaly Plateau. Abiotic water characteristics did not differ between seasons. The abundances of both species were negatively correlated with iron concentrations. Further correlations between the abundance of either fish species and abiotic water characteristics remained inconclusive as these water characteristics co-varied with geographical latitude that in turn was correlated with fish abundance. For both species neither the abundance nor a condition factor based on body mass showed any significant seasonal variation. Also the presence of T. mararybe had no influence on the abundance and the condition of T. madagascariensis. Thus, no evidence for competition was noticed between the two species

Functional diversity in a fragmented landscape — Habitat alterations affect functional trait composition of frog assemblages in Madagascar

Anthropogenic habitat alterations cause biodiversity loss, which in turn negatively affects ecosystem functioning and services, and thus human well-being. To be able to consider ecosystem functioning in conservation actions, analyzing the effects of habitat alteration on functional diversity is essential. Some altered habitats can maintain a significant part of regional biodiversity, however, functional diversity information in altered habitats is so far mostly lacking. We compared functional richness and functional β-diversity based on resource-use traits of frogs between three land-use categories in a rainforest ecosystem in Madagascar. Land-use categories represent a habitat alteration gradient ranging from continuous forest over forest fragments to matrix habitats including different agricultures. Our study revealed distinct changes in resource-use trait composition and complex patterns in the relationship between species richness and functional richness. Thus, the functional structure of frog assemblages changed due to habitat alterations. However, altered habitats likely provide different, rather than fewer functions compared to intact forest. Streams in all land-use categories were the functionally richest habitats, and thus important for ecosystem functioning. Species richness was one, but not the only driver of functional richness in our system. Functional clustering, potentially due to environmental filters depending on resource availability, was caused by anthropogenic and natural drivers. Our study shows that, even in systems where fragmented landscapes still maintain high species diversity, functional diversity can be altered in human altered habitats, which may affect ecosystem processes like productivity, nutrient cycling, and energy flows

FIGURE 7 in 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

FIGURE 7. Mantidactylus subgenus Brygoomantis species distribution based on verified records, divided by clade (Mantidactylus stelliger sp. nov. is presented alongside the inaudax clade for practicality). Colours for species correspond to those in Figs 2 and 4. Inset map shows the geographic regions of Madagascar identified by Boumans et al. (2007), referred to throughout the text.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 162, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/744102

FIGURE 2 in 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

FIGURE 2. (Continued).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 155, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/744102

Mantidactylus ambohimitombi subsp. ambohimitombi Boulenger 1919

Mantidactylus ambohimitombi ambohimitombi Boulenger, 1919 Type material.— Mantidactylus ambohimitombi Boulenger, 1919 is based on a series of syntypes that include BMNH 1947.2.26.25–32 from ‘ Ambohimitombo Forest’. Blommers-Schl̂sser and Blanc (1991) noted that syntypes BMNH 1947.2.26.31– 32 may be referable to Mantidactylus curtus, without justification. We here designate the DNA-barcoded specimen BMNH 1947.2.26.25, a large-sized individual of 65.0 mm SVL, as lectotype. Lectotype designation is justified by the need to stabilize this and other nomina in Brygoomantis, given the uncertain identity and morphological similarity of many taxa in the subgenus. Identity.— Mantidactylus ambohimitombi Boulenger, 1919 is typically considered a valid species (BlommersSchl̂sser & Blanc 1991; Frost 2021; Glaw & Vences 1992a, 1994), although it was considered of uncertain status by Glaw and Vences (2007). It is morphologically close to M. curtus but distinguished by its distinctly larger body size. It was not assigned to a genetic lineage by Vieites et al. (2009). A 16S sequence of the lectotype surprisingly clusters with a lineage predominantly known from the Ankaratra Massif. We therefore redefine this lineage provisionally as corresponding to M. ambohimitombi. In previous studies the populations from the Ankaratra Massif have been considered as confirmed candidate species M. sp. 19 by Vieites et al. (2009), and M. sp. Ca19 by Perl et al. (2014). They were referred to as ‘ M. sp. aff. curtus “Ankaratra”’ by Schmidt et al. (2009). We emphasize that this attribution is preliminary; since only mtDNA data (no genomic information) are available from the lectotype, and no fresh samples are available from Ambohimitombo forest where the original syntype series was collected, we cannot exclude that mitochondrial introgression has taken place, potentially blurring a hypothetical differentiation between the Ankaratra and Ambohimitombo populations. Our Phylonetworks analysis provided evidence for gene flow of syntopic M. curtus into M. ambohimitombi marefo ssp. nov. from Itremo (Fig. 5), providing a first hint that reticulated evolution may have played a role in the origin of the various morphologically divergent frogs that we here subsume in the species M. ambohimitombi. A more in-depth analysis of ranges, and of gene flow among various lineages of the M. curtus clade (M. curtus, M. bourgati, M. ambohimitombi) is necessary to understand their evolutionary history and verify their taxonomy. In our phylogenomic tree, two other lineages form a monophyletic group with specimens of M. ambohimitombi from Ankaratra, and these are in subsequent accounts described as subspecies of M. ambohimitombi; see the rationale in the respective accounts below. Diagnosis.— Mantidactylus ambohimitombi is a member of the M. curtus clade; it is here defined as containing three deep genetic lineages considered as subspecies, and is sister to the morphologically very distinct M. madecassus. See Table 4 for a list of diagnostic morphological characters. The following account only diagnoses the nominal subspecies M. a. ambohimitombi from other species in Brygoomantis; see below for diagnoses and comparisons of the two other subspecies. The combination of relatively large body size of up to 51 mm at Ankaratra (up to 65 mm in the type series from Ambohimitombo forest), slightly granular skin without dorsolateral ridges in most specimens, strongly developed foot webbing with fully webbed fifth toe, small tympanum diameter with a maximum of 11% of SVL in males, distinguishes M. a. ambohimitombi from species of the other clades. Within the M. curtus clade, M. alutus, M. madecassus and M. pauliani have smaller body sizes and are distinguished by either a usually shorter snout (M. madecassus, M. pauliani), or advertisement calls emitted in regular series (in M. alutus, vs single notes) (Table 4); M. curtus usually has a smoother skin and a somewhat shorter snout; M. bourgati is morphologically very similar but appears to occur only on the Andringitra Massif. For detailed distinction from new species described herein, see the respective species accounts.A full list of molecular diagnostic sites in the 16S gene of M. a. ambohimitombi in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Variation.—Variation in measurements is given in Table 5. See Fig. 13 for colouration in life and its variation. A light vertebral stripe occurs in few individuals. There is pronounced sexual size dimorphism (at Ankaratra, largest confirmed male SVL 37.7 vs female SVL 50.6 mm). As discussed above, the type series is comprised of particularly large-sized animals reaching 65.0 mm in SVL. Femoral glands are only distinct in some male specimens, possibly due to seasonal effects; in two specimens, ZSM 190/2021 (ACZCV 334) and ZSM 195/2021 (ACZCV 342), FGL and FGW are 4.7 mm x 3.8 mm and 4.0 mm x 3.0 mm, respectively. Natural history. — A common species on theAnkaratra Massif, especially above the tree line along streams and swamp in montane savanna and heatland, but also in rainforest (reported by Vences et al. 2002 under the name M. curtus). Males were found calling during the day underwater. Calls.— The advertisement call of M. a. ambohimitombi, recorded on 16 February 2006, 14:50 h, at Ankaratra, consists of a short, pulsed note (Fig. 14), emitted in series at slow succession and somewhat irregular intervals. The calls were emitted from several shy animals at the border of a mountain stream, partly underwater, and identification of the calling specimen was therefore impossible. Notes exhibit slight amplitude modulation, with maximum call energy occurring either at first third of the note’s length or at the centre of the note, and the terminal pulse of the note always being of lowest energy. The initial pulse is sometimes separated from the second pulse by a slightly longer inter-pulse interval. Numerical parameters of 20 analysed calls are as follows: call duration (= note duration) 136– 218 ms (180.2 ± 18.7 ms); 9–14 pulses per note (10.9 ± 1.5); pulse duration 5–8 ms (6.3 ± 1.0 ms); pulse repetition rate within notes 56.1–69.3 pulses/s (62.3 ± 5.1); dominant frequency 925–1012 Hz (959 ± 30 Hz); prevalent bandwidth 500–3100 Hz; call repetition rate (= note repetition rate) in regular series ca 11–20 calls/ min. Tadpoles. —A tadpole of M. a. ambohimitombi was described under the name ‘ M. sp. aff. curtus “Ankaratra”’ by Schmidt et al. (2009). Distribution.— Apparently endemic to a small area of the central highlands of Madagascar (Fig. 7). The nominal form is known from Ankaratra. Mitochondrial sequences assignable to this lineage have also been recorded from Analafohy and Antoetra; however, confirmation is needed, especially at Antoetra, as to whether this is evidence of true co-occurrence, or is a result of introgression with M. curtus, which is common in this locality. The type locality Ambohimitombo forest is close to Antoetra. Elevation range: 1150–2380 m a.s.l. Etymology.—Formulated from the type locality, ‘Ambohimitombo Forest’.Published as part of Scherz, Mark D., Crottini, Angelica, Hutter, Carl R., Hildenbrand, Andrea, Andreone, Franco, Fulgence, Thio Rosin, Köhler, Gunther, Ndriantsoa, Serge Herilala, Ohler, Annemarie, Preick, Michaela, Rakotoarison, Andolalao, Rancilhac, Loïs, Raselimanana, Achille P., Riemann, Jana C., Rödel, Mark-Oliver, Rosa, Gonçalo M., Streicher, Jeffrey W., Vieites, David R., Köhler, Jörn, Hofreiter, Michael, Glaw, Frank & Vences, Miguel, 2022, An inordinate fondness for inconspicuous brown frogs: integration of phylogenomics, archival DNA analysis, morphology, and bioacoustics yields 24 new taxa in the subgenus Brygoomantis (genus Mantidactylus) from Madagascar, pp. 113-311 in Megataxa 7 (2) on pages 178-182, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/744102

Mantidactylus mahery Scherz & Crottini & Hutter & Hildenbrand & Andreone & Fulgence & Köhler & Ndriantsoa & Ohler & Preick & Rakotoarison & Rancilhac & Raselimanana & Riemann & Rödel & Rosa & Streicher & Vieites & Köhler & Hofreiter & Glaw & Vences 2022, sp. nov.

Mantidactylus mahery sp. nov. Identity and justification.—This lineage has been considered as confirmed candidate species M. sp. 14 by Vieites et al. (2009) and M. sp. Ca14 by Perl et al. (2014). This is a relatively large-sized species of Brygoomantis fromwesternMadagascarwithamorphologysuperficially similar to M. ulcerosus which, however, is not its closest relative and concordantly differs in mitochondrial and nuclear genes and in advertisement call structure. Both species occur syntopically at least at one site (Makira). Although M. mahery sp. nov. mainly occurs in western Madagascar unlike the other species of the M. curtus clade which live in the central highlands, the phylogenomic data unambiguously support its inclusion in the group. Even more surprising, the phylogenomic tree places the species sister to M. pauliani, which is a montane endemic from the Ankaratra Massif that differs in numerous morphological characters. The status of M. mahery sp. nov. as a separate species is well supported by multiple lines of evidence. Holotype. — ZSM 23/2004 (field number FGZC 37), adult male collected by F. Glaw, M. Puente, R. Randrianiaina, and M. Teschke (née Thomas) on 21 January 2004 in Isalo at a creek near Ranohira (22.5856°S, 045.3997°E, 813 m a.s.l.), Ihorombe Region, Madagascar. A 16S barcode sequence of the holotype is available from GenBank (accession AY848286). Paratypes.—A total of 12 paratypes: ZSM 25/2004 (FGZC 39), adult male, and ZSM 26/2004 (FGZC 42), adult female, with the same collection data as the holotype; ZSM 567/2009 (ZCMV 11457) and ZSM 569/2009 (ZCMV 11484), two adult females, collected by M. Vences, D.R. Vieites, F.M. Ratsoavina, R.D. Randrianiaina, E. Rajeriarison, T. Rajofiarison, and J. Patton on 20 June 2009 in Sahaovy (‘Camp 0’), Makira (15.4889°S, 049.0785°E, 607 m a.s.l.); ZSM 9/2006 (FGZC 682), adult male, and ZSM 61/2006 (FGZC 793), adult female, collected by F. Glaw, J. K̂hler, P. Bora, and H. Enting on 18 and 23 March 2006, respectively, at Antranopasazy (‘Camp 1’), Tsingy de Bemaraha National Park (18.7086°S, 044.7189°E, 146 m a.s.l.); ZSM 134/2006 (FGZC 938), ZSM 135/2006 (FGZC 939), two adult males, and ZSM 136/2006 (FGZC 940), adult female, collected by F. Glaw, J. K̂hler, P. Bora, and H. Enting on 31 March 2006 at Andafiabe on the Beboka River (‘Camp 2’), Tsingy de Bemaraha National Park (18.7842°S, 044.7794°E, 177 m a.s.l.); ZSM 927/2003 (FG / MV 2002.1421), putative male, ZSM 941/2003 (FG / MV 2002-1485), adult male, and ZSM 942/2003 (FG / MV 2002-1486), adult female, collected by G. Aprea, M. Puente, L. Raharivololoniaina, M. Teschke (née Thomas), and D.R. Vieites between 29 January and 1 February 2003 at Hotel Isalo Ranch (22.5929°S, 045.3928°E, ca 800 m a.s.l.). Diagnosis.— Mantidactylus mahery sp. nov. is a member of the M. curtus clade as revealed by the phylogenomic analysis, and sister to the morphologically strongly different M. pauliani. While all other species in the M. curtus clade occur on the central plateau of Madagascar, M. mahery is distributed in western Madagascar, including some rather arid areas where it appears to be the only Brygoomantis present, and it is also present in one locality in the North West (western slope of Makira). See Table 4 for a list of diagnostic morphological characters. The combination of a large body size of up to 49 mm, slightly granular skin with (weakly expressed) dorsolateral ridges, absence of white spots on flanks and of white marking on snout tip, and short pulsed advertisement calls emitted in regular series distinguishes the new species from species of the other clades. In the North West it can occur syntopically with the similarly sized M. ulcerosus which however has a distinctly more tubercular dorsal skin, and more pulses per note in advertisement calls. Within the M. curtus clade, the new species differs by its larger tympanum diameter in males (11–13% of SVL) from all other species (Table 4). Mantidactylus alutus, M. madecassus and M. pauliani furthermore are smaller and have a shorter snout (Table 4). For detailed distinction from other new species described herein, see the respective species accounts. A full list of molecular diagnostic sites in the 16S gene of M. alutus in pairwise comparisons to all other Brygoomantis species is provided as Supplementary appendix. Description of the holotype. —Adult male in good state of preservation (Fig. 9). Tongue excised as tissue sample (tongue no longer present); femoral glands partly detached for examination in internal view. Body rather slender (in this, differing from many other specimens of this species which are stouter). Head as wide as body. Snout rounded in dorsal and lateral views. Nostrils directed laterally, slightly protuberant. Nostrils nearer to tip of the snout than to eye. Canthus rostralis weak, slightly concave. Loreal region weakly concave. Tympanum distinct, large, elliptical, diameter about 93% of eye diameter. Supratympanic fold present, beginning straight,with a rather distinct bend midway towards forelimb insertion. Maxillary teeth present. Vomerine teeth present in two rounded aggregations, positioned posterolateral to choanae. Choanae rounded. Subarticalur tubercles single. Outer metacarpal tubercle present, inner metacarpal tubercle present. Fingers without webbing. Relative length of fingers: I=II<IV<III. Finger discs slightly enlarged. Nuptial pads absent. Foot longer than tibia (115%). Lateral metatarsalia separated. Inner metatarsal tubercle present. Outer metatarsal tubercle not present. Webbing formula: 1(0.25), 2i(1), 2e(0), 3i(1), 3e(0.5), 4i(2), 4e(1), 5(0.5). Relative length of toes: I<II<V<III<IV. Skin on the upper surface quite smooth with very few scattered granules on flanks. Ventral side smooth. Femoral glands present, in external view with a distal ulcerous macrogland and a large proximal granular gland field. Colour in preservative: dorsum dark brown. Forelimbs brown with poorly defined darker markings. Hindlimbs brown with poorly defined darker crossbands. Inguinal region without whitish spots. Snout tip without a light dot. Venter beige with brown mottling, throat darker than belly. Lower lip with alternating light and brown spots. Toe discs dark. Colour in life of holotype unknown. Variation. —Variation in measurements is given in Table 5. See Fig. 21 for colouration in life and its variation. In life, dorsum is brown with distinct darker markings. Dark band between eyes is present. Forelimbs brown with very indistinct darker markings; hindlimbs with indistinct darker crossbands. Belly beige; throat with distinct white and brown mottling. A longitudinal white line on abdomen and throat is present. Femoral glands distinctly orange. There is moderate sexual size dimorphism(confirmed male SVL 29.2–37.2mm [n =7] vs confirmed female SVL 34.2–48.6 mm [n = 6]). Horizontal tympanum diameter is 80–100% of eye diameter in males and 69–86% of eye diameter in females. Skin on the back with very few indistinct tubercles on the flanks. Colour on the back varies from light brown with distinct darker markings (e.g. ZSM 927/2003) to uniformly dark brown. Two dark spots on the back at the level of forelimb insertion always more or less distinctly present, except in the holotype (ZSM 23/2004), whose colour is too dark to see any markings.A dark brown more or less triangular band between eyes is always present. A light vertebral band or line is not present. An indistinct light dot on the snout tip is never present except in ZSM 25/2004. Lower lip with more (e.g. ZSM 23/2004) or less (e.g. ZSM 25/2004) distinct alternating light and brown spots. Venter and throat from uniformly beige with faint markings (e.g. ZSM 26/2004) to dark brown mottled (e.g. ZSM 23/2004). A longitudinal light median line on abdomen and throat is present in ZSM 25/2004. Hindlimbs always distinctly striped (e.g. ZSM 941/2003) except in ZSM 23/2004 where hindlimbs are striped indistinctly dark-brown. Forelimbs brown with irregular darker markings and stripes. Femoral glands of males large and prominent with a clear proximal granular gland field in ZSM 25/2004 and ZSM 941/2003, in ZSM 23/2004 and ZSM 927/2003 less prominent with indistinct proximal granular gland field. In external view a central depression in the middle of the femoral gland can be seen, thus indicating a distal ulcerous macrogland. In females femoral glands are always small but distinctly present (e.g. ZSM 942/2003), but a proximal granular gland field is never present. In life, males in reproductive state have femoral glands orange coloured (Fig. 21b, h), and sometimes (Fig. 21b) the proximal granular gland field is larger and more prominent than the distal ulcerous macrogland, which is uncommon among mantellines; the granular gland fields on the two opposite thighs contact each other medially. Natural history.—The species is known from various sites in the West and North West of Madagascar, reaching into the South at Isalo. It has been found along running water (including very slowly running streams) in and outside of forest. At the western slope of the Makira Reserve it was found with M. ulcerosus and M. jonasi sp. nov. (see below) along a relatively large stream in degraded remnants of rainforest. Calls.— The advertisement call of M. mahery, recorded on 28 January 1994 at Isalo National Park, near Ranohira, 23.4°C air temperature (Vences et al. 2006: CD 2, track 76), consists of a short, pulsed note, emitted in regular series at fast succession (Fig. 22). Pulse repetition rate is distinctly higher at the beginning of calls and significantly reduces after approximately one quarter of the call’s duration. Amplitude modulation is present, with highest call energy occurring at the beginning of the call and continuously decreasing towards its end. Numerical parameters of eight analysed calls are as follows: call duration (= note duration) 120– 144 ms (131.8 ± 8.7 ms); 21–26 pulses per note (24.1 ± 2.3); pulse duration 1–2 ms (1.6 ± 0.5); pulse repetition rate within notes 107.8–667.0 pulses/s (331.4 ± 205.3); dominant frequency 1004–1270 Hz (1151 ± 95 Hz); prevalent bandwidth 800–4500 Hz; call repetition rate (= note repetition rate) within regular series ca 211–218 calls/min. Tadpoles.— The tadpole of this species has not been described. Distribution.— Endemic to an eclectic collection of disparate localities, mostly in the West of Madagascar, but reaching the eastern rainforest escarpment at Makira in the North West, and Tsaranoro in the Central region (Fig. 7). This species is known from Isalo (various localities), Forêt de Beanka, Makay, Makira West (Sahaovy, Camp 0), Tsaranoro, and Tsingy de Bemaraha (various localities). Elevation range: 120–960 m a.s.l. Etymology. —The species name is derived from the Malagasy adjective mahery, meaning ‘big’ or ‘strong’, and refers to the rather stout body shape of this species. The name is used as a noun in apposition.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 193-195, DOI: 10.11646/megataxa.7.2.1, http://zenodo.org/record/744102

FIGURE 62 in 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

FIGURE 62. Preserved holotype specimens of newly named species in the Mantidactylus biporus clade and of the M. stelliger clade. Scale bars equal 5 mm.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/744102