Revision of the Subgenus Ochthomantis Frogs from Madagascar (Amphibia: Mantellidae) with the Description of Four Species and Resurrection of Mantidactylus catalai and M. poissoni

Simple Summary The genus Mantidactylus spp. is one of the exceptionally diverse amphibian clades from Madagascar. Currently, 57 species in 6 subgenera are recognized. One subgenus, Ochthomantis, is the focus of the present study. Here, we revise this taxonomic group to recognize the presence of cryptic species through an assessment of morphological and, where available, molecular (16S mitochondrial gene) variation of 637 sexed adult specimens. Our results show that Ochthomantis contains eleven species, including the resurrected Mantidactylus catalai and M. poissoni, and four newly described species. We cannot confirm that M. majori should be considered as part of the subgenus Ochthomantis, so we do not include it in our descriptions. Following our taxonomic revision, we also present a practical simple key to identify all of the species belonging to this subgenus. Abstract The subgenus Ochthomantis is an obligate forest and stream-dwelling group of mantellid frogs, endemic to Madagascar, with six species currently recognized. However, this group suffers from ongoing taxonomic confusion due to low numbers of examined specimens, and failure to consider morphological variation from development and sexual dimorphism. Here, we examined the morphology of 637 sexed adult specimens collected by us in the field and from other museum collections. We also sequenced a DNA fragment of the 16S mtDNA gene for each lineage to determine congruence between morphological and molecular data sets and to help delimit species. Our results demonstrate that the subgenus Ochthomantis includes eleven valid species: five already recognized, M. catalai and M. poissoni that we resurrect from synonymy, and four new species which we describe for the first time here. In some analyses, Mantidactylus majori groups with other Mantidactylus subgenera, so we do not consider it a member of the subgenus Ochthomantis in this study. All species have restricted distributions and elevational ranges in the humid forests of Madagascar. This study demonstrates the utility of assessing cryptic species using both diagnostic morphological characters and molecular data. The discovery of this new cryptic biodiversity, and the taxonomic revision herein, will likely require conservation activities for those species with the most restricted distributions.

Coloration in life: The iris has a golden ring on its outer area. A dark brown transversal bar is present between the eyes. A dorsal surface of the body may have or not have a pale yellow vertebral line. The inguinal region has a distinct oblique and large yellow patch, sometimes, it extends onto the ventral side. The ventral surface of the body is often pale brown on the anterior parts and brown yellowish color on the belly. The throat sometimes has a pair of dark brown parallel stripes. The inner surface of the thigh has a brown color and its outer part has yellow spots. Hindlimbs have alternate transversal bands, brown and black.
Habits: semi-aquatic rainforest species living close to rivers or small streams with slow-flowing water and rarely, either observed outside the rainforest or along the streams close to relict forests (e.g., Tsarafidy). This species was found on rocks, leaves, and branches or on the ground along river banks between 9.00 am and 23.30 pm. However, it is more active at night. During the day, it can be found hidden in holes and rock crevices along rivers. At night, the females are generally found sitting on shrubs living along streams and more rarely observed on the rocks far from the riverbank. Males can be found in all areas along rivers, except on branches and plant stems. The vertical distribution of individuals on shrubs might be different between the sexes: we observed males up to 1 m above the ground and females up to 2 m. Individuals prefer streams with stream beds between 1-10 m in width and water depth between 10-150 cm. Eggs are laid in masses outside the water, either on a leaf or a branch overhanging of the stream or the river. The tadpoles live in the calm water far from torrents. The period of reproduction is during the cold winter season (e.g., June and July in Mantadia). The calls of males are low and hardly audible. In the Moramanga Region, metamorphosis appears to occur from September to October. At Andranomanamponga in August 2002, the tadpoles of different development stages were observed in calm and transparent water, and these tadpoles' habitats were protected by rocks. These pools have surface water between 1-3 m 2 and a water depth between 20-100 cm. When disturbed, the tadpoles swim quickly and obliquely to hide in mud under dead leaves or retreat into rock crevices.
Distribution: Eastern and northern rainforests, including the highland of Madagascar with an elevation range between 230 to 1600 m above sea level. The species occurs as far north as the Sambirano Region (14 • S) and as far south as the Anosy Mountains (25 • S) ( Figure 3, Table S13).
M. ambreensis was synonymized with M. femoralis by [22], and later recognized again as a good species by [10,31].
Holotype: MNHN 1893.241: Montagne d'Ambre, Madagascar. fold running posteriorly to one point of three large granules above the upper arm and shoulder girdle articulation. Dark tympanum with a small notch in its median superior area.  5 (0.50). The total sum of free phalanges 7.75. The relative toe length is 1 < 2 < 3 < 5 < 4. Lateral surfaces of the body, belly, and sacral areas with very small granules. Oblong femoral glands are well-developed and its medio-proximal area with a pore surrounded by many granules, giving it a crater-like pattern. Internally, femoral gland type 3 [2,28]. In the preservative, the dorsal surface of the head, body, and limbs is dark in color. The lateral surface of the body is less dark than the dorsal surface of the body. Lips with white band colors continue along the lateral surface of the body to above hindlimb articulation. Dark iris surrounded by a white ring. The ventral surface of the body is whitish with dark marbling except in the belly region. Throat with dark-brown parallel bands. Ventral surface of the forelimb with some dark spots. Thigh with large dark spots except on the femoral gland region. Hind and forelimbs with alternate and transverse dorsal bands, dark and light gray colors. Variation: Morphometric variation is summarized in Table S3. Sexual dimorphism is evident: males have smaller SVL (33.3-39 mm versus 38.2-42 mm), relatively longer tibia and feet, and shorter toe 3; ratio td/ed larger in males than females (0.80-1.16 vs. 0.52-0.79); femoral glands more swollen in males and smaller and more circular in females. The free phalanges on toes vary: toe 1 (0-1), internal edge of toe 2 (1), external edge of toe 2 (0-0.50), internal edge of toe 3 (1-1.50), external edge of toe 3 (0-1), internal edge of toe 4 (1-2), external edge of toe 4 (1)(2), and in toe 5 (0-1). Small skin granules may be present or not on the dorsal surface of the body and above the eyes. Some individuals with granules on the sacral area.
Coloration in life: The iris has a gold ring on its outer area. A dark crossbar may be present between the eyes. There is no vertebral line on the dorsum. A white or yellow band is running along the flank. The dorsal surfaces of the head, body, and limbs may be dark green, brown, or grayish brown in color. The ventral surfaces of the body are usually mottled brown, but for some individuals, there is almost no mottling. Some individuals have a diverged pair of short longitudinal dark bands on the throat and this fades on the thorax. Large and round dark spots may also be present on the throat and thorax.
Habits: semi-aquatic forest species living next to flowing streams or rivers generally and close to rocks. This species was found between 9.00-23.00 h, but it is rather nocturnal than diurnal frogs. At night, females rested on leaves rather than on branches, and during the day, we observed them sometimes resting on the banks of the river and very rarely on the ground. For males, during the day they rested on banks and at night on leaves, and sometimes we can find them on rocks. The vertical distribution of individuals on shrubs is different between both sexes: males between 10-200 cm, whereas females between 100-200 cm. Distinguished from M. femoralis, this taxon prefers rivers to small streams. At Montagne d'Ambre (Station les Roussettes), it is observed near the irrigation canal. Calling males were heard in March in the afternoon from the ground along the forest brook [7]. A clutch consisting of about 100-120 eggs was found deposited on a rock edge close to a calm and shallow stream in April 2006 at Vohibola Tsaratanana and those eggs were hatched after a couple of days in a plastic bag. The egg diameter is about 2-3 mm.
Distribution: species of low and mid-altitude forests of Northwestern and Northern Madagascar, with elevational ranges between 200 to 1150 m. This species is found as far north as Montagne d'Ambre and across humid forests in the northern highlands such as Manongarivo, Tsaratanana, Andramanalana, and Sorata; and extending to the northern limit of the High Plateau at Irony River and Ambohibola Forest ( Figure 4, Table S13).
sometimes we can find them on rocks. The vertical distribution of individuals on shrubs is different between both sexes: males between 10-200 cm, whereas females between 100-200 cm. Distinguished from M. femoralis, this taxon prefers rivers to small streams. At Montagne d'Ambre (Station les Roussettes), it is observed near the irrigation canal. Calling males were heard in March in the afternoon from the ground along the forest brook [7]. A clutch consisting of about 100-120 eggs was found deposited on a rock edge close to a calm and shallow stream in April 2006 at Vohibola Tsaratanana and those eggs were hatched after a couple of days in a plastic bag. The egg diameter is about 2-3 mm.
Distribution: species of low and mid-altitude forests of Northwestern and Northern Madagascar, with elevational ranges between 200 to 1150 m. This species is found as far north as Montagne d'Ambre and across humid forests in the northern highlands such as Manongarivo, Tsaratanana, Andramanalana, and Sorata; and extending to the northern limit of the High Plateau at Irony River and Ambohibola Forest ( Figure 4, Table S13). Comments: Our molecular analyses group all our M. ambreensis samples with the M. ambreensis samples reported by [10]: AY324822 (ZSM 492/2000); [11]: HQ610870 (FGMV 2002(FGMV .1950; and by the holotype MNHN 1893.241 (sequence accession MT982173) [5]. Comments: Our molecular analyses group all our M. ambreensis samples with the M. ambreensis samples reported by [10]: AY324822 (ZSM 492/2000); [11]: HQ610870 (FGMV 2002(FGMV .1950; and by the holotype MNHN 1893.241 (sequence accession MT982173) [5]. Razafimanantsoa Diagnosis: A small sized Ochthomantis (adult male SVL 30.0-31.8 mm; adult female 34.0-37.9 mm) with sharply white to yellow stripe, well-defined along the lateral head and lateral surface of body, running from snout tip to groin area. Distinguished from M. ambreensis by its smaller size and from the remaining species by the presence of a sharply white to yellow well-defined lateral stripe. Character diagnostics in Tables 1 and 2.
Description of reference specimen: UADBA 8406 (RAX 2871): Adult male (SVL = 31.8 mm) in excellent state of preservation. Measurements are presented in Table S4. In the dorsal view and lateral view, the snout tip relatively pointed. Snout tip with 1.7 mm ventral extension beyond mouth. In the dorsal view, the head is clearly longer than large (i.e., 1.35 times longer than wide). Head length 0.47 times SVL. Canthus rostralis indistinct. Loreal area concave. Tympanum diameter 1.19 times eye. The round tympanum is in contact with the supratympanic fold along their borders and running posteriorly above towards the upper arm and shoulder girdle articulation. Large and dark tympanum with a small notch in its median superior area. Internarial distance 0.30 times head width. Tongue ovoid anteriorly and bifid in posterior part. Round nostrils with lateral aperture. Eye to nostril distance is 1.40 times nostril to snout distance. Forearm length 0.47 times SVL. Hand length (including discs) 0.19 times SVL. The inner metacarpal is not obvious and the outer metacarpal has obvious granule. Fingers are without webbing. The relative finger length is 1 < 2 < 4 < 3. Digits with large terminal discs (widest part twice of basal disc width). Tibiotarsal articulation between eye and nostril. Lateral metatarsal separated. Hindlimb 1.69 times SVL. Thigh length 0.90 times the tibia length. Foot including tarsus 0.73 times the SVL. Inner metatarsal tubercle not obvious (length 1.35 mm) at base of toe 1. Outer metatarsal tubercle absent. Toe with less extensive webbing and webbing formula: 1 (1) 2i (1) 2e (0.50), 3i (1.25) 3e (1) 4i (2) 4e (1.50) 5 (0.50), and total sum of free phalanges 8.75. Relative toe length 1 < 2 < 3 < 5 < 4. The surfaces of the body: flank, belly, and sacral areas with very small granules and dorsal with obvious granules. Oblong femoral glands are well-developed and pores in its medio-proximal area are surrounded by many granules, giving it a crater-like pattern. Internally, femoral gland type 3 [2,28]. In preservative, the dorsal surface of the head, body, and limbs has a dark color. The lateral surface of the body is less dark than the dorsal surface of the body. Lips with white band colors run along the lateral surface of the body to the hindlimb insertion. Dark iris surrounded by a white ring. The ventral surface of the body is whitish with dark marbling except in the belly region. Throat with dark brown parallel bands. Forelimb with some dark spots on its ventral surface. Thigh with large dark spots except in the femoral gland region. Fore and hindlimbs with dorsal transverse bands, alternating dark and light gray colors.
Coloration in life: The iris has a gold ring on its outer area. A dark crossbar may be present between the eyes. No vertebral line on dorsum. A white or yellow band runs along the flank. The dorsal surfaces of the head, body, and limbs may be dark green or brown in color. The ventral surfaces are usually mottled brown. All individuals have a diverged pair of short longitudinal dark bands on the throat and fade on the thorax. Large and round dark spots were absent on the throat and thorax.
Habits: semi-aquatic forest species living next to quiet rivers with rocks. It was observed between 11.00-20.00 h. Like M. ambreensis, this taxon prefers rivers over small streams. At Montagne d'Ambre (Station les Roussettes), it was observed near the irrigation canals. According to [5], this is a rheophilous species and frequently terrestrial by day, sitting on the ground, on rocks, wood, lichen, or hiding under rocks. At night, often observed on perches above the water, once at 2 m height, and sitting on substrates like leaves, rocks, dead wood, and plant stems.
Distribution: Species of low and mid-altitude forests of Northwestern and Northern Madagascar, with an elevational range between 300 to 1150 m above sea level. It was observed from a relict deciduous forest, south of Maevatanana, to a humid forest at Montagne d'Ambre, extreme Northern Madagascar. All of the sites for their presence are presented in Figure 4 and Table S13.
Comments: This species occurs only at Montagne d'Ambre [5]. Even though our single molecular state this analysis, the specimens that we have recorded outside Montagne d'Ambre based on morphological characters only should be considered too, according to character diagnosis from [5]. This is why we have grouped all our M. ambony samples (molecular and morphological characters) with the M. ambony samples reported by [5] supratympanic fold. This supratympanic fold has an umbrella-like pattern and runs to above one point between the shoulder girdle and upper arm articulation but behind one large strong granule. The anterior half part of the tympanum with light background color and the posterior part dark color. Internarial distance 0.27 times head width. Tongue ovoid anteriorly and bifid posteriorly. Non-protruding nostril with relatively close lateral aperture. Eye to nostril distance 1.59 times nostril to snout distance. Forearm length 0.47 times SVL. Hand length (including discs) 0.29 times SVL. Fingers without webbing. Outer and inner metacarpals are poorly developed. Finger relative length size 1 <2 <4 <3. Digits with a large terminal disc (the widest part twice the width of the basal disc). Tibiotarsal articulation reaching nostril. Lateral metatarsal separated. Hindlimb 1.71 times SVL. The thigh is the same length as the tibia. Foot including tarsus 0.72 times SVL. Inner metatarsal tubercle in bell-like pattern (length 1.6 mm) at base of toe 1. Outer metatarsal tubercle absent. Webbing formula 1 (0), 2i (1), 2e (0), 3i (1), 3e (0), 4i (1.5), 4e (1), 5 (0) and total sum of free phalanges 4. Relative toe length 1 <2 <3 <5 <4. Importance and repartition of body granules differently in shape and color: side and edge of the dorsal surface and above tympanum highly granulated; inguinal area, basal of flanks, and posterior of upper mouth with white evident granules; dorsum with little granules and belly finely granular. Oblong femoral glands are relatively developed with centro-distal pores surrounded by many granules, giving a crater-like pattern form. Internally, femoral glands type 3 [2,28]. In preservative, dorsum with black color. Upper lip and flanks with clean spots. The throat and the thorax with white and some silver reticulated dark brown pigments, and the belly with light yellow color. Thorax with parallel dark bands: X-like pattern on its left side and divided into two forms, spot and "+/−like" pattern on its right side. The ventral surface of the thigh is partially mottled with brown color and weak in the femoral gland. The lower part of the hindlimb is completely pigmented. The ventral side of the forelimb with clean shape and yellowish color. Evident alternate and transverse dorsal bands, shiny and dark, on the hindlimb and indistinct bands on the forelimb, but with some pinkish reticles indifferently distributed in their dorsal surfaces.
Coloration in life: The iris has a golden ring on its outer area. Just the dorsum can have a black crossband in V or Y-like pattern. The dorsum is either dark brown or thoroughly black in color. The upper mouth is either dark brown or blackish. Whitish gray spots may be present or not in the inguinal region. The ventral surface shape is very heterogeneous: throat and thorax with black color and usually with numerous small white spots; belly without pigments; and throat either with two dark spots or not. The lateral surface of the body is unicolor with no evident white spots and few granules. The ventral surface of the thigh has at least some clean pattern surfaces. Hind and forelimbs have alternate transversal bands, black and brown.
Habits: Semi-aquatic rainforest species can live in open and degraded forests, especially in Northern Madagascar. This species was observed resting along the riverbanks and streams at different stages of water speeds, between 9.00 a.m to 23.00 p.m. However, it appears rather diurnal than nocturnal and prefers rocky areas to trees. On trees, it prefers resting on leaves and branches.

Mantidactylus zolitschka [10]
Holotype: ZFMK 60110: close to An'Ala Forest (18 • 6-37.7 mm); tibiotarsal articulation at least up to the nostril. Width of terminal disc 1.78 times basal disc. Distinguished from all other species, except M. ambony, by its smaller size (adult male SVL < 31 mm, female < 38 mm) and from all species by foot with less well-developed webbing (total sum of free phalanges, WS > 9). Distinguished from M. ambony by the absence of lateral stripe running from snout tip to inguinal region. Character diagnostics are summarized in Tables 1 and 2.
Description of reference specimen UADBA 6966: Adult male (SVL = 27.65 mm) in good state. Measurements are presented in Table S6. In the dorsal view, the body is clearly slender. In lateral and dorsal view, the snout tip is pointed. Snout tip with 1.20 mm ventral extension beyond the mouth. Head 1.40 times longer than wide. Head length 0.47 times SVL. Canthus rostralis are distinct and straight. Loreal is weakly concave. Tympanum diameter 0.83 times eye diameter. Round tympanum distinctly from the supratympanic fold. This tympanic fold runs straight first and rather curves in midway before reaching the forepart of the shoulder girdle and upper arm articulation. Internarial distance 0.24 times head width. Tongue ovoid anteriorly and distinctly bifid posteriorly. Small and round nostril without protuberant lateral aperture. Eye to nostril distance 1.44 times nostril to snout distance. Forearm length 0.52 times SVL. Hand length (including discs) 0.33 times SVL. Fingers without webbing. Relative finger length 1 < 2 < 4 < 3. Inner and outer metacarpal with tubercles. Digits with slightly enlarged terminal discs (widest part 1.78 times of basal disc width). Legs slender. Tibiotarsal articulation reaching nostril. Lateral metatarsal separated. Hindlimb 1.85 times SVL. Thigh the same length as the tibia. Foot including tarsus 0.78 times SVL. The inner metatarsal tubercle is rather small at the base of toe 1 (0.85 mm). Metatarsal with small tubercle. Webbing formula: 1(1), 2i (1.25), 2e (1), 3i (1.5), 3e (1), 4i (2), 4e (2), 5 (1) and total sum of free phalanges 9.75. Relative toe length 1 < 2 < 3 < 5 < 4. Skin is rather smooth on its upper surface and slightly granular on its flanks. Ventral side smooth one. Obvious femoral glands are in contact with the cloacal area and sharply delimited by granules with irregular tubercle-like patterns, and the presence of a central porus gives it a crater-like pattern. Internally, femoral gland type 3 [2,28]. In preservative, gray-brownish dorsal color with irregular dark and light marble shape. Upper lip and loreal area whitish color. Tympanic region dark brown color. Lower lip with alternate spots, light and dark. The lateral surface of the body is dark in color and the ventral surface of the body is light in color. The ventral surface of the body with different color pattern: the throat is whitish and becoming more yellowish on the belly. Throat with two longitudinal brown markings from the lip to the thorax and both merge at the shoulder girdle as a "Y-like pattern". One light longitudinal stripe runs from the inguinal area and fades towards forelimb insertion. Light brown forelimb and hindlimb with different numerous dark crossbands (six on hand including third finger, four on thigh, three on tibia, and five on tarsus and foot). Hindlimbs with irregular dark mottling.
Coloration in life: The iris has a golden ring on its outer area. The dorsal surface of the body has a strong gray-brownish color with a small light stripe running along it. There is a shiny yellow blotch on the inguinal region.
Habits: semi-aquatic rainforest species living close to stream, around An'Ala forest. The female ZFMK 30116 contains 49 eggs with yellow and dark brown center markings and diameter is 2 mm [10].

Resurrected Species
Based on our molecular and morphological results, we find strong evidence to recognize two species of Mantidactylus (Ochthomantis) that correspond to taxa that currently are considered junior synonyms of M. femoralis. After examining their type specimens and our new materials, we here recognize Mantidactylus catalai [35] (33 specimens) and Mantidactylus poissoni [36] (12 specimens) as valid species and provide new descriptions for both species below ( Figure 6).

Resurrected Species
Based on our molecular and morphological results, we find strong evidence to recognize two species of Mantidactylus (Ochthomantis) that correspond to taxa that currently are considered junior synonyms of M. femoralis. After examining their type specimens and our new materials, we here recognize Mantidactylus catalai [35] (33 specimens) and Mantidactylus poissoni [36] (12 specimens) as valid species and provide new descriptions for both species below ( Figure 6).

Mantidactylus catalai [35]
Mantidactylus catalai [35] (p. 203) (Holotype MNHN 1935.153, according to the original publication and [6] (p. 220); Mantidactylus femoralis: [22] (p. 26); Mantidactylus (Hylobatrachus) femoralis: [27] (p. 312); Mantidactylus (Ochthomantis) femoralis: [7] (p. 400), [3] M. catalai has previously been considered by [22] as a synonym of M. femoralis but [10]    Diagnosis: A medium to large-sized Ochthomantis (adult male SVL 41-45 mm, female SVL 51-62 mm); tibiotarsal articulation between eyes and nostril (or very rarely at snout); toes fully webbed, except on toe 4 where 1-1.5 phalanges are free; no stripe line along superior lip; inguinal region with clean pattern area; snout tip very pointed in lateral view with large extension beyond mouth (1.75-3.45 mm); head wider and flattened but very sharp as "fish-like" pattern. Distinguished from other species by the following characters: M. ambreensis and M. ambony by the absence of white stripes along the lateral body; M. femoralis, M. zolitschka, and M. danieli n. sp. by the number of free phalanges in the internal edge of toe 4 (1-1.5) and absence of prominent pale yellow or white stripes in the inguinal region (horizontal or oblique); M. mocquardi by the number of free phalanges on the internal edge of toe 4 (1-1.5) and flanks without whitish spots; M. olgae n. sp. by the absence of obvious black granules on flanks and absence of crossbars in V-or Y-like pattern on dorsum in preservative; M. tavaratra n. sp. by digits with large terminal discs (widest part > 1.80 times of basal disc width), lack of prominent and pale inguinal streak, and absence of white strip on superior lip; M. poissoni by the absence of white spots below the eye and tibiotarsal articulation between eyes and nostrils; M. macrotympanum n. sp. by smaller adult male SVL (<60 mm). Character diagnostics are summarized in Tables 3 and 4.  Table S7. In the dorsal and lateral view, the snout tip is pointed. Snout tip with 2.60 mm ventral extension beyond the mouth. Head 1.40 times longer than wide. Head length 0.50 times SVL. Canthus rostralis well distinct. Loreal region with evident indentation. Tympanum diameter 1.03 times eye. Slightly round tympanum in contact with supratympanic fold, except in its posterior part, and this tympanic fold runs behind tympanum towards the small granule above shoulder girdle and forearm articulation. Tympanum with a small notch in its median superior area. Internarial distance 0.27 times head width. Tongue ovoid anteriorly and bifid posteriorly. Nostrils with distinct cutaneous fold and lateral oblique aperture. Eye to nostril distance 1.58 times nostril to snout distance. Forearm length 0.61 times SVL. Hand length (including discs) 0.42 times SVL. Fingers without webbing. Inner and outer metacarpal tubercles are evident. Relative fingers length 1 < 2 < 4 < 3. Digits with large terminal discs (widest part 2.07 times of basal disc width). Tibiotarsal articulation between the eye-nostril. Lateral metatarsal separated. Hindlimb 1.84 times SVL. The thigh is the same length as the tibia. Foot including tarsus 0.78 times SVL. The inner metatarsal tubercle is obvious along toe 1 (2.15 mm). The outer metatarsal tubercle is small and granule-shaped. Webbing formula: 1 (0.75) 2i (1) (1) 5 (0) and total sum of free phalanges 6. Relative toe length 1 < 2 < 3 < 5 < 4. Body granules vary: flank with obvious granules, dorsum with granules irregularly distributed, granules of tympanum more concentrated above its superior part, and sacral area and belly finely granulated. Oblong femoral glands are relatively developed and its medio-proximal part with pores surrounded by some granules, giving it a crater-like pattern. Internally, femoral gland type 3 [2,28]. In preservative, the dorsal surface of the body is brown in color and the white longitudinal vertebral band runs from the snout tip to the anal pore. Lips with two vertical light stripes on the loreal region. Shiny line behind the eyes. The ventral surface of the body is orange-brown in color but has a clean pattern. The throat has a couple of dark brown spots (in front of the shoulder girdle). The inguinal region with white spots or not. The ventral surface of the forearm is bordered by a brown-sided but clean pattern. Fore and hindlimbs with obvious alternate crossbands, dark and shiny.
Variation: Morphometric variation is summarized in Table S7. Sexual dimorphism is evident: males have smaller SVL than females (41.10-45.40 mm versus 51.85-61.50 mm), wider head, bigger tympanum, longer fore and hindlimbs, and metatarsal tubercle not evident. The ratio of td/ed in males is larger than in females (0.89-1.08 vs. 0.50-0.69).
The femoral glands are more swollen in males and smaller and more circular in females. A majority of all specimens are brown darker color, except for the holotype and UADBAs Habits: semi-aquatic rainforest species living in bamboo forests. This species is diurnal and/or nocturnal, observed between 9.00 a.m to 0.15 a.m, which is adapted to a "burrowing life", inside of holes and interstice of rocks, but close to the waters (stagnant water to the river but very rarely in a fast stream). It is mainly observed on rocks and sometimes on the ground, especially during the day. No individuals were seen on leaves or branches, but one individual UADBA 4521 was collected on roots.

Mantidactylus poissoni [36]
Mantidactylus poissoni [36] Table S8. In the dorsal view and lateral view, the snout tip is relatively obtuse. Snout tip with 1.35 mm ventral extension beyond the mouth. Head 1.25 times longer than wide. Head length 0.45 times SVL. Canthus rostralis distinct. Loreal with groove. Tympanum diameter 0.98 times eye. Slightly round tympanum with a vivid small notch in the middle of the superior area and in contact with the supratympanic fold in its anterior part and separate in its posterior part, and this supratympanic fold runs towards before reaching the shoulder girdle and upper arm articulation. Internarial distance 0.25 times head width. Tongue ovoid anteriorly and bifid posteriorly. Round nostrils with distinct cutaneous fold and lateral aperture. Eye to nostril distance 1.81 times nostril to snout distance. Forearm length 0.49 times SVL. Hand length (including discs) 0.28 times SVL. Fingers without webbing. Inner and outer metacarpals exist. Fingers without webbing. Relative finger length 1 < 2 < 4 < 3. Digits with large terminal discs (widest part > 1.70 times of basal disc width). Tibiotarsal articulation between nostril and snout tip. Lateral metatarsal separated. Hindlimb 1.76 times SVL. The thigh is the same length as the tibia. Foot including tarsus 0.72 times SVL. The inner metatarsal tubercle is shield-shaped (length 2 mm) at the base of the toe 1. Outer metatarsal tubercle in the small granule. Webbing formula: 1 (0), 2i (1), 2e (0), 3i (1.25), 3e (0.50), 4i (1.75), 4e (1.75), 5 (0.25) and total sum of free phalanges 6.50. Relative toe length is 1 < 2 < 3 < 5 < 4. Lateral surfaces of the body and dorsum with numerous granules. Little and swollen femoral glands elongated, and its pore on the media-distal area was surrounded by many granules, giving it a crater-like pattern. Internally, femoral glands type 3 [2,28]. In preservative, the dorsal surface of the body with blackish brown color. Upper lip with shiny transverse and interrupted band oriented to eyes. The belly and ventral surface of the thigh with yellowish-white spots. Obvious white pigments on boundary surfaces of the thorax and abdomen and making them together an 8-like pattern. Thorax and throat were almost pigmented by whites with some scattered brown spots and the thorax with two brownish parallel dark bands. The inguinal region with whitish L-shaped bed pattern. Dorsal transverse bands are rather indistinct on the hind and forelimbs.
Variation: Morphometric variation is summarized in Table S8. Sexual dimorphism is evident: males have a smaller SVL (39.7-46.8 mm), thicker snout, shorter hand, larger terminal disc, and loreal less elongated. The ratio of td/ed is larger in males than in females (0.62-0.82 vs. 0.51-0.68). Tibiotarsal articulation between nostril and snout tip except UADBAs (19786,26411,26412)  Coloration in life: The iris has a golden ring on its outer area with some black spots. The dorsal surface of the body with a dark brown color has more small granules. There is a more or less round yellow spots in the inguinal region for females and a stick-like pattern for males. The ventral surface of the body is pigmented by a white color with yellowish border bands in an 8-like pattern. Hind and forelimbs with alternate dark and shiny brown transverse bands.
Habits: Semi-aquatic rainforest species living close to small streams with rock. Stream depth does not reach 1 m. All specimens were observed during the day and night between 16.00 pm to 21.30 pm. In the daytime, they were observed between 5-10 m far from the bank on the ground and at night 3-5 m far from the river, and roosting on leaves between 50-100 cm in height. UADBA 7125 was seen on rocks in the middle of the river where water is very speedy, maybe ending up there by misfortune. It is rather a ground and tree-dwelling than an aquatic frog.
Distribution: Mid and high elevations of the eastern slope forest of Madagascar ( Figure 6, Table S13).
Comments: The only genetic data known for this species are the sequence from the reference specimen AMNH A174653 (RAX 9367) ( Table S1). The LSID number is 57C53062-B2DF-41E5-88A3-EFE7B083DA84.

New Species Descriptions
Based on our molecular and morphological results, we find strong evidence to recognize four species of Mantidactylus (Ochthomantis) that correspond to taxa that cannot be assigned to any nominal species (or to names considered as junior synonyms). After examining our new materials, and developing diagnoses for each taxon, we here provide descriptions for each of these new species, such as Mantidactylus danieli n. sp. (54 specimens), M. macrotympanum n. sp. (5 specimens   Diagnosis: A small to medium-sized Mantidactylus (Ochthomantis) species (adult SVL male 33-42 mm, female 42-59 mm), with dark brown color on the dorsal surface of the body and often with black spotted granules. The inguinal region with a soft pale yellow streak and may be partly broken up or narrow; no continuous pale stripe along the lower flank; lack of obvious white spot below the eye; moderately developed foot webbing with 1.5-2 free phalanges at the internal edge of toe 4 and WS > 7; tibiotarsal articulation extends beyond nostril; and maximum width of a terminal disc on fingers < 1.70 disc width base.    Table S9. In the dorsal view, the head is longer than the width. Head length 1.18 times width length. Head length 0.41 times SVL. In dorsal view, the snout tip is pointed and rounded lateral, with a 3 mm ventral extension beyond the lower lip. Canthus rostralis evident. Loreal concave. The tympanum and supratympanic fold are distinct from each other, and this supratympanic fold runs towards the front of the upper arm and shoulder girdle articulation. Round tympanum diameter 0.74 times eye diameter. Tongue ovoid anteriorly and bifid posteriorly. Internarial distance 0.21 head width. Round nostril with lateral aperture. Eye-nostril distance 1.27 times nostril-snout distance. Forearm length 0.51 times SVL. Hand length (including discs) 0.26 times SVL. Fingers without webbing. Outer and inner metacarpal with developed tubercles. Relative finger length 1 < 2 <4 < 3.
The terminal is disc relatively large (the widest part is 1.45 width of its basal disc). Tibiotarsal articulation between nostril and snout tip. Hindlimb 2.01 times SVL. Thigh length 0.98 times tibia length. Foot including tarsus 0.92 times SVL. Lateral metatarsal separated. Inner metatarsal tubercle as a skin-like pattern with three-sided (1.35 mm) on base of toe 1. Outer metatarsal tubercle present. Webbing formula: 1 (1), 2i (1), 2 (0.25), 3i (1.75), 3e (0.50), 4i (2), 4e (2), 5 (0.50), and total sum of free phalanges is 9. Relative toe length 1 < 2 < 3 < 5 < 4. The dorsal surface of the thigh, the posterior area above the eye, above the tympanum, and around the femoral glands with granule skin-like patterns. Round femoral glands poorly developed with central pores. Internally, femoral gland type 3 [2], but without small granules in the proximal area, characteristics of male femoral glands [28]. In the preservative, the dorsal surface of the body is dark brown in color and has small darker spots. Upper lip dark brown color except under tympanic region clean pattern. The throat is pale brown in color with two dark parallel bands in an L-shape. The belly is pale brown in color and is almost entirely covered with fine dark brown spots, which decrease in density posteriorly. Brown forelimbs without dark bands. Brown hindlimb with darker transversal bands on its dorsal surface and pale brown with dark-brown spots on its ventral surface. The inguinal region with pale and discontinuous oblique streak.
Coloration in life: The iris color is golden in the superior area and fades to the lower area of the eye. The dorsum and dorsal surfaces of the head and limbs have a dark brown color and sometimes black spots for females. The upper lip has a white band that is most developed in the posterior area. Between the eyes, there is a darker brown cross-bars in a V-like pattern. The ventral surfaces with dark brown spot pigments have a different background color: the belly is creamy or yellowish but the paler pattern is almost dark reddish on its anterior part, two dark reddish parallel stripes may be present on the throat and become narrower in its posterior part and decline on the thorax or is absent (UADBA 8382, 26361, 26366, 26367, 26371, 26373). The inner area of the thigh is completely covered with dark reddish-brown spots. A pale yellow oblique streak in the inguinal region is typically discontinuous with all of the specimens, except for UADBA 19595, 26359, and 26367. Transversal bands, black and darker brown were observed on the dorsal surface of the hindlimb, but indistinctly clear with some dark specimens (e.g., UADBA 26406). One adult female (UADBA 3770) has a pale vertebral line running from the snout to the anal pore.
Habits: Living in rainforests between 350-1580 m elevation and semi-aquatic species; occupying the banks of small streams and rivers. Observed between 10.30-23.00 h, although more active at night and appears to be mostly nocturnal. Unlike other Ochthomantis species, males rest mostly on river banks, more rarely on leaves of trees and bushes, and very rarely on rocks. Females are more tree-dwelling than rupicolous. The vertical distribution appears to be different between sexes: males between 20-100 cm, whereas females are between 50-300 cm above the ground. This species appears to prefer rivers to small fast-flowing streams and females were observed mostly in areas of slow-moving water.
Etymology: The specific name danieli is a patronym, i.e., a noun in the genitive case honoring Daniel Rakontondravony for his substantial contributions to the knowledge of the Malagasy fauna.

Mantidactylus macrotympanum
Description of holotype: Adult male (SVL 61 mm) in excellent state of preservation. Measurements are presented in Table S10. In dorsal view, the head is longer than wide: head length 1.29 times head width. Head length 0.44 times SVL 0.44. In the dorsal view, the snout tip is pointed, and in the lateral view almost acuminate and curved down with 2.20 mm ventral extension beyond the lower lip. Obvious canthus rostralis and concave loreal making lip well evident. Tympanum and supratympanic fold distinct and supratympanic fold running vertically in its posterior part to one evident granule above the upper arm and shoulder girdle articulation. Round tympanum diameter 0.94 times eye diameter. Tongue ovoid anteriorly and bifid posteriorly. Internarial distance 0.23 times head width. Non-protruding nostril with lateral slanting aperture. Eye-nostril distance 1.86 times nostril-snout tip distance. Forearm length 0.50 times SVL. Hand length (including discs) 0.32 times SVL. Fingers without webbing. Outer and inner metacarpal tubercles flattened and widened. Relative finger lengths 1 < 2 <4 <3. Terminal discs are large (the widest part is 2.20 times of basal disc width). Tibiotarsal articulation largely beyond the snout tip. Hindlimb 1.89 times SVL. Thigh and tibia same length. Foot including tarsus 0.76 times SVL. Lateral metatarsal separated. Outer metatarsal tubercle absent. The inner metatarsal tubercle is not evident at the base of toe 1. Webbing formula 1 (0), 2i (0.5), 2e (0), 3i (1), 3e (0), 4i (1.25), 4e (1), 5 (0) and total sum of free phalanges 3.75. Relative toe length 1 < 2 < 3 < 5 < 4. Flanks with few prominent granules in its superior part and a white spot-like pattern posteriorly. Numerous epidermic granules in the sacral area and finely granules above the eyes. Round femoral glands are not swollen. Internally, femoral gland type 4 [2,28]. In preservative, the dorsal surface of the body is blackish and marbled with some shiny spots. Ventral surfaces of the body and superior lip homogenous pattern with a brownish color. The throat is darker with two parallel bands. Fore and hindlimbs with alternate transversal bands, shiny and dark. The inguinal region has some scattered dark spots.
Variation: Morphometric variation is summarized in Table S10. Species are only known from six male specimens. SVL between 57.9 and 62.1 mm. The ventral surface of the body is heterogeneous according character of the belly: individuals with a clean pattern belly have black spots and those can be either present or not with dark belly specimens. Flanks are shiny brownish backgrounds with white plates in the central area like the holotype, but sometimes white spots are observed for specimens that have black flank backgrounds. Three internal phalanges vary between 0.50-1 for toe 2i and 1-1.50 for toe 4i.
Coloration in life: The iris has a constant black color surrounded by white color in its superior and lower parts. The body is dark brownish. The superior lip has a black crossbar anteriorly. Above the eyes, there is a white line. The belly is darker with a black spot. Thorax has two parallel bands. The vivid brown flanks have a white plate-like. A white oblique line can be present in the inguinal area. Two alternate bands, shiny and dark, are rather distinct on the fore and hindlimbs.
Habits: This is a rainforest species living along calm streams and fast rivers with scattered rocks. Individuals were also found outside the forest, e.g., in plantations at Manongarivo. Individuals were observed between 12.00-20.00 h but they are more nocturnal than diurnal. At night, they roost and rest on branches between 10-200 cm in height and during the day, they rest on the rocks. All the known specimens observed are males.
Etymology: The specific name macrotympanum is composed of the Latin words macro (large) and tympanum (tympanum), referring to the very big tympanum of this species. The name is used as a noun in apposition.
Distribution: Known only from relict forest close to Ambalafary village, the Manongarivo Special Reserve at 250 m, and from the Ramena River (Tsaratanana Reserve) between 180-800 m ( Figure 10, Table S13).
Comments: The only genetic data known for this species were obtained from AMNH A167589 (RAX 2715), UMMZ 201416 (RAN 39170), and UMMZ 213447 (RAN 39125) ( Table S1). The LSID number is 957B5BBF-17E1-4175-A499-2953919D7F89. Diagnosis: A small-to medium-sized Mantidactylus (Ochthomantis) species (adult SVL male  with gray color of the body; dorsal and lateral surfaces of the body with black granules; no pale yellow streak in the inguinal region; no broad pale line or white spots on the lateral surface of the body; upper lip with a pale line; the presence of crossbars in V or Y-like pattern on dorsum; snout tip pointed in lateral view; well-developed foot webbing with 1 free phalange on internal edge of toe 4 (WS < 7); tibiotarsal articulation between eye and nostril; and maximum width of terminal disc on fingers > 1.8 times of basal disc width. Distinguished from all others species by presence of obvious black skin granules both on dorsal and lateral surfaces of body and by the following characters: M. femoralis by one free phalange on internal edge of toe 4, no yellow patch on inguinal region, and tibiotarsal articulation between eye and nostril; M. ambreensis and M. ambony by lacking of broad pale line on lateral surface of body; M. mocquardi by gray body color and no white spots on lateral surface of body; M. zolitschka by larger male adult SVL > 33 mm and female adult > 44 mm; M. poissoni by lack of white spots below eye; M. catalai by presence of crossbars in V or Y-like pattern on dorsum; M. danieli n. sp. by following characters:webbing more developed (WS < 7), one free phalange at internal edge of toe 4, no pale yellow streak in inguinal region, and tibiotarsal articulation between eye and nostril; M. macrotympanum n. sp. by smaller male adult SVL < 60 mm and smaller male tympanum eye diameter < 0.94; and M. tavaratra n. sp. by larger terminal disc and maximum width of terminal discs on finger > 1.8 times of basal disc width. Character diagnostics are summarized in Tables 5 and 6.
Description of holotype: Subadult female (SVL = 40.70 mm) in excellent state of preservation. Measurements are presented in Table S11. In the dorsal view, the head is rather longer than the width: the head length is 1.23 times the head width. Head length 0.46 times SVL. In dorsal and lateral view, the snout tip is pointed with evident ventral extension 2.20 mm beyond the lower lip giving it a shell-like pattern. Canthus rostralis not evident. Loreal furrowed. The tympanum was distinct from a well-developed suratympanic fold which ran and reached towards one point before upper arm and shoulder girdle articulation. Tympanum diameter 0.56 times eye diameter. Tongue ovoid anteriorly and bifid posteriorly. Internarial distance 0.21 times head width. Round nostrils with cutaneous fold anterolateral aperture. Eye-nostril distance 1.74 times nostril-snout tip distance. Forearm length 0.55 times SVL. Hand length (including discs) 0.33 times SVL. Fingers without webbing. Outer and inner metacarpal tubercles are poorly developed. Relative fingers length 1 < 2 < 4 < 3. Larger terminal disc, i.e., widest part twice of basal disc width). Tibiotarsal articulation between eye and nostril. Hindlimb 1.98 times SVL. Thigh length 1.07 times tibia length. Foot including tarsus 0.77 times SVL. Lateral metatarsal separated. Inner metatarsal tubercle in half moon-like pattern (1.3 mm length) at base of the toe 1. Outer metatarsal with small granule. Webbing formula: 1 (0), 2i (1), 2e (0), 3i (1), 3e (0), 4i (1.25), 4e (1), 5 (0) and total sum of free phalanges 4.25. Relative toe length 1 < 2 < 3 < 5 < 4. Skin granules on the flank. Round femoral gland poorly developed with a central pore. Internally, femoral gland type 3 [2] but without small granules in its proximal area characteristics of the male femoral gland [28]. In preservatives, coloration is grayish dorsally with a Y-like pattern. Upper lip with white pigments. Throat with two dark parallel bands. Belly, ventral surface of the thigh, and forelimb clean patterns. Hind and forelimbs with gray color and alternate dorsal bands, shiny and dark, crossing them. An inguinal region without a pale inguinal streak. Flank with some black granules.
Coloration in life: The iris has a golden ring on its outer area. Body coloration is gray except for brown color, AMNH A167560 and UADBA (8391, 8477). The upper lip is white and dirty except for the holotype which is white in color. No dark parallel bands on the throat except for UADBA 8407. All of the specimens have V or Y-like patterns with dark spots on the dorsum except UADBA (8384, 8387, 8389) with X-like patterns. The ventral surface of the body is clean pattern except for juveniles which are dirty and some specimens have a throat with white pigments including the holotype and AMNH (A167557, A167564, A167568-167569, A167571, A167573-167574). The inguinal region has no pale yellow streak. The dorsal hindlimbs have darker brown transversal bands. Some specimens have a thick pale brown vertebral line.
Habits: Low-and mid-altitude rainforest species close to the edge of streams or rivers with rocky bottoms. We observed it between 10.00-22.30 h and mainly nocturnal except for juveniles. It was observed in various habitats: rock, grounds, riverbanks, and edges of water. Generally, females rest on rocks at night and some specimens overhanging on leaves between 20-50 cm above the ground. Males are more tree-dwelling than rupicola and rest on leaves of shrubs, between 10-30 cm in height. Specimens observed at all types of water, but they have a particular preference for streams or fast rivers.
Etymology: The specific name olgae honors the late Olga Ramilijaona, professor at UADBA and supervisor of the Ph.D. thesis of N.R., in recognition of her substantial mentorship.
Distribution: Species inhabits along Ramena River (Tsaratanana Strict Natural Reserve, and Analabe), Andramanalana, and Sorata in the North of Madagascar. The elevation ranges between 600-1700 m ( Figure 11, Table S13). Comments: The only genetic data known for this species are presented in Table S1. Although M. olgae has been recorded at Andramanalana, no genetic or morphological samples have yet been confirmed from Marojejy, which is a well-sampled site about 40 km to the west. The LSID number is FB2EEFBB-92C7-460E-892A-9B5BBFF65862.  Table S12. In the dorsal view, the head is larger than the width: the head length is 1.34 times the head width. Head length 0.44 times SVL. In the dorsal view, the snout tip is slightly pointed and almost straight in the lateral view with a 1.55 mm ventral extension beyond the lower lip, which makes the snout tip have an obtuse appearance. Canthus rostralis obvious and loreal concave making the jaw well evident. Tympanum and supratympanic fold distinct running almost vertically towards three evident white granules in front of upper arm and shoulder girdle articulation. Round tympanum diameter 0.62 times eye diameter. Tongue ovoid anteriorly and bifid posteriorly. Internarial distance 0.30 times head width. Round nostril with non-protruding lateral aperture. Eye-nostril distance equals nostril-snout distance. Forearm length 0.46 times SVL. Hand length (including discs) 0.30 times SVL. Fingers without webbing. Outer and inner metacarpal tubercles flattened and widened. Relative finger length 1 < 2 < 4 < 3. Fingers without webbing. The terminal disc is relatively large (the widest part is 1.40 times the basal disc width). Tibiotarsal articulation between eye and nostril. Ventral surface of the forearm with three granules on its internal ridge at the base of metacarpal tubercles. Hindlimb 1.90 times SVL. The thigh length I almost equal to the tibia length. Foot including tarsus 0.77 times SVL. Lateral metatarsal separated. Outer metatarsal tubercle indistinct. Inner metatarsal tubercle flattened at base of toe 1. The webbing formula is: 1 (0.25), 2i (1), 2e (0), 3i (1.25), 3e (0.25), 4i (1.5), 4e (1.5), 5 (0), and the total sum of the free phalanges is 5.75. Relative toe length 1 < 2 < 3 < 5 < 4. The dorsal surface of the body with small granules on its edge and on the sacral region. Oblong femoral glands are relatively developed and in their centro-distal area with pores surrounded by numerous granules, giving a craterlike pattern. Internally, femoral gland type 3 [2,28]. In preservative, brown dorsum with some clean pattern or depigmented portions, especially in its posterior region. Upper lip with finely dark brown pigments posteriorly and covering with obvious round pigments anteriorly. The indistinct band runs behind the eyes and crumbles in the anterior half of the dorsum. Throat with two dark short parallel stripes. Ventral surfaces are homogeneous for different parts of the structure: throat and thorax with silver-white pigments and some reticulated dark brown pigments, belly whitish in color with some brown round spots, thigh and forearm no pigments in its part of the area. The inguinal region has fine streaks. The lower portion of the hindlimb with few scattered brown spots. Hindlimb with evident alternate bands, thick dark, and fine shiny. Forelimb with indistinct alternate bands.
Coloration in life: The iris has a golden ring on its outer area. The body is grayish brown, light, dark brown, or sometimes reddish brown. The upper lip has an evident vivid white band or not. Between the eyes, there is a darker brown cross-bars in a rod-like pattern. The dorsal surface of the body has dark brown cross-bars in a V or Y-like pattern. The throat and thorax are silvery white or white in color. The throat is always with distinct dark brown parallel stripes. The belly has a clean pattern with smaller white spots. The inner side of the thighs has some dark reticulations. The inguinal area has a white or yellowish little line or not, but if it exists the line is narrower in shape. White brownish bands can be present on the outer side of the forelimbs and alternate transversal bands, white and brown on the inner side of the forelimbs. The dorsal surface of the hindlimbs has alternate transversal bands, light and dark browns. The black granules may be present or not on the white flank but concentrated in the posterior part of the dorsum and on the side of the forearm, but they are not very obvious like M. olgae.
Habits: This is a rainforest and savannah species living along the riverbank. It inhabits almost all kinds of aquatic habitats: streams, rivers, ponds, and shallow marshes except waterfalls. We observed it between 9.30-23.00 h, but mainly nocturnal except juveniles. Juveniles observed during the day. At night, they rest both on rocks and leaves, but rarely on branches and accidentally on aerial roots and Pandanus sp. The females prefer roosting on rocks rather than on leaves, whereas the males are indifferent to those. The vertical distribution varies between the sexes: males between 5-100 cm and females between 5-150 cm above the ground.
Etymology: The specific name tavaratra refers to the Malagasy word for "north". This name is used as a nonlatinized noun in apposition and is given in reference to the known distribution of this species in Northern Madagascar Distribution: Northern endemic species of Madagascar, elevation varies from 80 to 2450 m elevation ( Figure 12, Table S13).
Etymology: The specific name tavaratra refers to the Malagasy word for "north". This name is used as a nonlatinized noun in apposition and is given in reference to the known distribution of this species in Northern Madagascar Distribution: Northern endemic species of Madagascar, elevation varies from 80 to 2450 m elevation ( Figure 12, Table S13). Comments: Our morphological description agrees with the M. sp. aff. mocquardi shown by [29] (Figure 2, p. 249) from Marojejy. Our molecular analyses group all our M. tavaratra samples (Table S1) with those Manongarivo AY324819 (FGMV 2002.824) and Tsaratanana AY324820 (ZSM 643/2001), reported as M. cf. mocquardi by [10]. FGMV 2002.824 from Manongarivo has more recently been referred to as "UCS sp. 63 Tsaratanana", and this taxon also includes another specimen (FGMV 2001.114) from Tsaratanana [11]. The LSID number is BD3F2F68-88CF-4629-9A91-0196E2CF2F4A.  -Lateral head and body without a broad pale line running continuously from the snout tip to the groin area. 2.

4.
Upper lip with a large pale spot under the eye (female) or multiple white spots (male), groin area lacks a short pale line or spots. M. poissoni.
-Upper lip lacks a large pale spot under the eye (female) or multiple white spots (male), groin area with a short pale line or spots. 5.

5.
Large finger pads with disc width > 1.7 × disc base, upper lip with a thin pale continuous line, groin area with a short bold pale line that may be continuous or broken (in life, yellow). M. femoralis.
-Smaller finger pads with disc width < 1.7 × disc base, upper lip without a thin pale continuous line, groin area with a short weak pale line that is often broken or even absent. M. danieli.

6.
Upper lip with a thin pale continuous line. 7.
-Upper lip without a thin pale continuous line but may have small pale spots. 8.

7.
Snout tip sharply angular in lateral view, flanks with an irregular border from the dark to pale venter coloration, dorsal surface of body gray in life (and often in preservation) with black granules, larger finger pads, with disc width > 1.8 × disc base. M. olgae.
-Snout tip not sharply angular in lateral view, flanks typically with a straight border from the dark to pale venter coloration, dorsal surface of body brown without black granules, smaller finger pads, with disc width ≤ 1.8 × disc base. M. tavaratra.

8.
Snout tip angular in lateral view, body dark brown (or black in preservation) with distinct white spots on flanks, groin area with a few or lacking pale spots, foot webbing with 1.5 free phalanges on the internal edge of toe 4 and WS 4. M. mocquardi.
-Snout tip not angular shape in lateral view, body medium brown (never black in preservation) and lacks white spots on flanks, groin area with a prominent short pale line or spots, foot webbing with 1-1.5 free phalanges on the internal edge of toe 4 and WS 6. M. catalai.
The increased species number is in agreement with the previous study focusing on tadpoles [11] which identified numerous candidates for species in the subgenus, several of which were validated by our study with adult specimens through morphological and cladistic analysis [28].
One species currently included in the subgenus Ochthomantis, M. majori (Figure 13), should not be included in this subgenus based on unpublished Principal Component Analysis, which places the species completely separate from the other species of the subgenus Ochthomantis, and on our cladistic analysis on 96 morphological patterns, habitats, and behavioral characters [28].
The divergence of M. majori is evidenced by its different reproductive behavior (parental care, time, and mating behavior). Based on our field observation, we confirm that males of M. majori guard eggs and practice parental care (Figure 13a) as [38,39] observed too; probably to protect eggs against fungi [40], predators, and desiccation. Egg deposition takes place on leaves overhanging the water (Figure 13b) in contrast to other Ochthomantis species where it takes place at edges of riverbanks either on twigs or on rocks, as we observed in M. femoralis and M. ambreensis (Figures 7d and 13c). Furthermore, morphological differences in M. majori to other Ochthomantis are very evident, i.e., it has very developed webbing (toes almost fully webbed, WS = 0-2.5), hindlimbs very short, internal space more developed than interorbital space, presence of the thin line vertebral, toe 3 slightly shorter than toe 5, presence of the thin white line in the side of hindlimbs and especially inguinal region free of obvious pigments (white or yellow spots; line or band patterns). However, our molecular analyses recover M. majori as a sister to M. macrotympanum, and this new subclade forms a paraphyletic group with the remnant of the subgenus Ochthomantis species with moderate support (posterior probability = 0.91); thus, while we do not include M. majori in this revision, we recommend that the placement of M. majori is further investigated in a study with broader taxonomic and genetic sampling. cies with moderate support (posterior probability = 0.91); thus, while we do not include M. majori in this revision, we recommend that the placement of M. majori is further investigated in a study with broader taxonomic and genetic sampling.
We confirm M. flavicrus as a junior synonym of M. femoralis. The analysis of the type of M. flavicrus⎯from the description of [8], the measurement from [10], and an examination made by CJR of the type specimen of M. flavicrus based on our identification key⎯supported that they both represent the same species.  We confirm M. flavicrus as a junior synonym of M. femoralis. The analysis of the type of M. flavicrus-from the description of [8], the measurement from [10], and an examination made by CJR of the type specimen of M. flavicrus based on our identification key-supported that they both represent the same species.
We noticed that two species in our analysis are composed of genetic subclades, Mantidactylus mocquardi and M. ambreensis ( Figure 1); however, the genetic distances between the subclades are small (<3%) but may be further explored in future studies. The first of these species, M. mocquardi, has two subclades representative of populations from the north and south of the Sambirano River, respectively. The north of the Sambirano specimens are characterized by their black color, while those from the south of the Sambirano have a more brownish color. These subgroups also differ by additional characters: for the totally blackish specimens, (1) granules almost missing, body smoothing and homogeneous, (2) white spots or silvery in flanks and missing on superior lips, and (3) ventral face marbled with brown reticulate and white pigment except on belly that is a clean pattern in preservative; for the brownish specimens (1) granules evident on flanks, (2) white pigment very conspicuous on lower flanks continuing to lower lips, (3) some dark spots observed on back with different shape: UADBA 12313 (NR 1372), UADBA 26290 (RAX 8036), UADBA 26238 (RAX 8021), UADBA 26240 (RAN 45476), (4) ventral face completely white in living specimen, while thorax and throat with brown spots and marbled with white or silvery pigments in contrasting of clean pattern belly-shaped in preservative. Additional specimens and observation of M. ambreensis especially from southern populations, Ramena, and Irony, are needed to better discern morphological differences including body size.

Sexual Dimorphism and Size
Within the subgenus Ochthomantis, sexual dimorphism is evident in SVL and tympanum size. Males have a larger tympanum than females (0.62-1.33 vs. 0.39-0.79). The tympanum senses sound waves and detects the position and direction of animal movement [41]. Here, sexual dimorphism in tympanum size may be related to reproductive communication. The tympanum in males can play a role not only for receiving but also for emitting sound by vibration, which especially makes sense given their small vocal sacs, where their call has low intensity [6,35]. The tympanum in females only functions as a receiver, thus its smaller size. This may suggest that attraction by calls over long distances is less significant for Ochthomantis compared to other anuran groups that breed during the warm and rainy season in Madagascar.
Ochthomantis females are characterized by a larger body size than males. This characteristic is a mechanism to reduce the effort spent during egg laying [42] and allows for an increase in the number and size of eggs which in some dissected individuals occupied almost the whole of the ventral cavity [28]. The smaller size of males can be also explained by: 1) a higher energy spending due to their higher activity during reproduction and competition with other males.
Some other morphometric characters may be related to the ecology of each species: (1) longer hindlimbs, and thus longer tibias may be related to improved jumping capabilities and are found in species with more terrestrial behavior (M. femoralis, and M. danieli) than in species more specialized to living on rocks and in aquatic environments (M. olgae, M. mocquardi, M. tavaratra, and M. catalai), (2) M. catalai has pointed snout and flattened head which may be an adaptation to move within in rock fissures, (3) Terminal discs of phalanges are probably used to cling and hang on leaves or rocks. Accordingly, the more tree-dwelling and rock-dwelling species (e.g., M. olgae, M. tavaratra) have a larger terminal disc than those most ground-dwelling species such as M. danieli, (4) well-developed webbing is observed especially in the more aquatic and rock-dwelling species (M. olgae, M. mocquardi, M. tavaratra) compared to the terrestrial and arboreal-dwelling species (M. femoralis, M. danieli). Webbing is used for propulsion in the water and increases the surface to grip the rocks, 5). Generally, species with stocky bodies often have long hindlimbs. In addition, according to our observations, they appear to be more dynamic jumpers (e.g., M. femoralis, M. danieli).

Period of Reproduction and Reproduction
The observations made in the north and the region of Moramanga indicate that the reproduction at least of some species of Ochthomantis is spread throughout the year. Thus, the statement from [6] that reproduction happens during the cool season is partially true for the species of low altitude and mid-altitude in the central eastern region. The beginning of reproduction probably takes place in April during which the temperature falls to 10 • C and metamorphosis of tadpoles occurs between October and November. While for species from the high altitude in the North (>1400 m asl) as M. tavaratra, the reproductive season starts very early when the temperature falls to 8 • C, e.g., at Tsaratanana the tadpoles in advanced stages were observed at the end of February (stage 36). It could be hypothesized that in mid and high altitudes, the activating factor may be the decline of temperature (around 10 • C in Moramanga, and 8 • C in Tsaratanana at high altitude, 2300 m elev. asl). We also noticed that the metamorphosis from the larval stage to the juvenile stages appears to occur fast at high altitudes. This speed may be necessary to avoid predation or other factors [43], e.g., the drying up of streams at high altitudes for instance on the Tsaratanana massif.

Biogeography, Climate, and Conservation
We noticed that Ochthomantis are absent in the West and South of the Island. We propose three hypotheses related to their reproduction habits. The first deals with temperatures, as the factor activating mating in species of the subgenus, appears to be a decline of temperature (between 8-15 • C), and it is clear that these regions of the island have a high temperature all year long. The second is the lack of water during the dry winter season when most of the streams are dried up. Finally, the third possible factor could be the absence of consistently calm water for breeding given that during the summer season, the water velocity can be very quick after torrential rains.
Global climate change has the potential to influence the fate of Ochthomantis as high mountain species have difficulty rapidly adapting to new conditions or migrating to areas with suitable conditions [44]. For Ochthomantis, climate change may also disturb the period of reproduction, particularly in those species where a decrease in temperature is critical to the induction of reproduction. In addition, changes in water regimes in the high mountains may cause the disappearance of breeding sites due to desiccation, or to a reduction in calm stretches of water with an increased frequency of torrential rains.
Data presented during a dedicated workshop held in Madagascar on 28 January 2009 [45] projected warming across the island, with Southern Madagascar being most affected and the coast and the north showing lower projected temperature increases. Precipitation increase was projected to be centered in the northwest while drying was projected in the East.
Integrating climate change into conservation strategies for amphibians, and specifically for the subgenus Ochthomantis, requires more precise distribution model forecasts under alternative climate scenarios, and we anticipate that amphibians occurring at high elevations will show higher sensitivities than those from low elevations [46]. For ground-truthing these scenarios, long-term monitoring efforts are also needed [47].

Conclusions
This study resolves the taxonomy of various cryptic species in the subgenus Ochthomantis using both morphological characters and molecular data. Our analysis revealed four new species that are formally named and described herein: M. danieli n. sp, M. macrotympanum n. sp, M. olgae n. sp, M. tavaratra n. sp. Furthermore, Mantidactylus catalai and M. poissoni are revalidated from junior synonyms to good species. The subgenus Ochthomantis now contains 11 species, but we suspect that some specimens included here may represent additional distinct species. It is hoped that the discovery and taxonomic revision of this new cryptic biodiversity will initiate conservation activities for those species with the most restricted distributions.

Institutional Review Board Statement:
The research was conducted in Protected Areas (MNP and NAP) and outside of PAs of Madagascar, with research permits from Ministère de l'Environnement et de Développement Durable, MEDD (previous name Ministère de l'Environnement, de l'Écologie et des Forêts, MEEF). During our fieldwork, we followed data collection procedures according to the national legislation regarding permits and international regulations regarding animal welfare. The type specimens used are from AMNH, BMNH, MNHN, and UADBA. The information reported here is the results from our data, and the findings are not reported anywhere else. Data Availability Statement: All holotypes are housed at AMNH and paratypes are shared between AMNH and UADBA according to research permits. Specimens used for molecular analyses are also stored in both institutions. Molecular sequences reported here are available in GENBANK.
Acknowledgments: Field studies in Madagascar were made possible due to the assistance of the Ministère des Eaux et Forêts, Madagascar National Parks, and in particular by the Université d'Antananarivo, Département de Biologie Animale for administrative assistance, and granting access to their zoological collection. MNHN and BM for access to holotype specimens, and UMMZ for sending us specimens. Research and fieldwork support were provided by AMNH. We thank the many people who have contributed to this research program, especially those who participated in the fieldwork, collecting data and specimens, including: N. Harilanto, S. D. Mahaviasy, R. A. Nussbaum, F. Rabemananjara, J. Rafanomezantsoa, D. Rakotomalala, A. Rakotondrazafy, J.B. Ramanamanjato, O. Ramilison, M. Randriambahiniarime, J. Randrianirina, F. Ranjanaharisoa, A.P. Raselimanana, P. Razafimahatratra, A. Razafimanantsoa, A. Razafimanantsoa, local guides, and the Conservation Agents of the Reserves we visited. Maps were drawn with help from B. Randriamahatantsoa.

Conflicts of Interest:
We declare no conflicts of interest. The funders had no role in the design of the study; in the collection, analyses, or interpretation of data; in the writing of the manuscript; or in the decision to publish the results. Table A1. Explanation of the range size abbreviations.

DB
Terminal disc width, measured on dorsal view, ratio between broadest part to its basal part ED Eye diameter, measured horizontally between peripheral orbit EHEAD Head height, measured in lateral view thickness of head in tympanum region EM Snout height, measured in lateral view thickness of snout in nostril region EN Eye-nostril distance; measured as shortest distance between the center of nostril and anterior border of eye EE Prefrontal width, measured in dorsal view area in front of marginal orbits EST Snout tip extension, measured as ventrally extent of snout tip, from the shortest distance between margin of upper lip and to snout tip FE Thigh length, measured in dorsal view from center of knee to margin of cloaca FT Foot length without tarsus, measured in ventral view from base of metatarsal tubercles to tip of toe 4 HD Hand length, measured between point of carpe and radius-ulna articulation to tip of finger n • 3 HDW Head width, measured in dorsal view, distance between tympanums HL Head length, measured in lateral view, from snout tip to mouth corner NN Internostril distance, measuring dorsally between center of each nostril NS Nostril-snout tip distance, measured as shortest distance between center of nostril to snout tip RC Radius-ulna length, measured in latero-external view as distance between base of inner metatarsal granule and center of elbow SVL Snout-vent length, measured in dorsal view from snout tip to cloaca TA Tarsus length, measured from center of tibiotarsal articulation to base of inner metatarsal tubercle TD Tympanum diameter, measured horizontally lateral surface of tympanum TI Tibia length, measured in latero-external view from center of knee to middle of tibiotarsal articulation T1 Outer metatarsal tubercle length along toe 1, measured in lateral view from its base to its top TO3 Toe 3 length, measured in dorsal view from base of proximal phalange to tip of toe 3

TO5
Toe 5 length, measured in dorsal view from base of proximal phalange to tip of toe 5 SVL Snout-vent length, measured in dorsal view from snout tip to cloaca TA Tarsus length, measured from center of tibiotarsal articulation to base of inner metatarsal tubercle TD Tympanum diameter, measured horizontally lateral surface of tympanum TI Tibia length, measured in latero-external view from center of knee to middle of tibiotarsal articulation T1 Outer metatarsal tubercle length along toe 1, measured in lateral view from its base to its top TO3 Toe 3 length, measured in dorsal view from base of proximal phalange to tip of toe 3 TO5 Toe 5 length, measured in dorsal view from base of proximal phalange to tip of toe 5 Figure A1. Showing the acronyms of the measurements of Ochthomantis species. The numbers 1−5 indicate the serial number from the toe 1 to toe 5 and from the finger 1 to finger 4; ED: Eye diameter, measured horizontally between peripheral orbit; EE: Prefrontal width, measured in dorsal view area in front of marginal orbits; EHEAD: Head height, measured in lateral view thickness of head in tympanum region; EM: Snout height, measured in lateral view thickness of snout in nostril region; EN: Eye-nostril distance; measured as shortest distance between the center of nostril and anterior border of eye; EST: Snout tip extension, measured as ventrally extent of snout tip, from the shortest distance between margin of upper lip and to snout tip; FE: Thigh length, measured in dorsal view from center of knee to margin of cloaca; FT: Foot length without tarsus, measured in ventral view from base of metatarsal tubercles to tip of toe 4; HD: Hand length, measured between point of carpe and radius-ulna articulation to tip of finger n°3 ; HDW: Head width, measured in dorsal view, distance between tympanums; HL: Head length, measured in lateral view, from snout tip to mouth corner; NS: Nostril-snout tip distance, measured as shortest distance between center of nostril to snout tip; RC: Radius-ulna length, measured in latero-external view as distance between base of inner metatarsal granule and center of elbow; TA: Tarsus length, measured from center of tibiotarsal articulation to base of inner metatarsal tubercle; TI: Tibia length, measured in latero-external view from center of knee to middle of tibiotarsal articulation ; T1: Outer metatarsal tubercle length along toe 1, measured in lateral view from its base to its top ; TO3: Toe 3 length, measured in dorsal view from base of proximal phalange to tip of toe 3; TO5: Toe 5 length, measured in dorsal view from base of proximal phalange to tip of toe 5. Figure A1. Showing the acronyms of the measurements of Ochthomantis species. The numbers 1-5 indicate the serial number from the toe 1 to toe 5 and from the finger 1 to finger 4; ED: Eye diameter, measured horizontally between peripheral orbit; EE: Prefrontal width, measured in dorsal view area in front of marginal orbits; EHEAD: Head height, measured in lateral view thickness of head in tympanum region; EM: Snout height, measured in lateral view thickness of snout in nostril region; EN: Eye-nostril distance; measured as shortest distance between the center of nostril and anterior border of eye; EST: Snout tip extension, measured as ventrally extent of snout tip, from the shortest distance between margin of upper lip and to snout tip; FE: Thigh length, measured in dorsal view from center of knee to margin of cloaca; FT: Foot length without tarsus, measured in ventral view from base of metatarsal tubercles to tip of toe 4; HD: Hand length, measured between point of carpe and radius-ulna articulation to tip of finger n • 3; HDW: Head width, measured in dorsal view, distance between tympanums; HL: Head length, measured in lateral view, from snout tip to mouth corner; NS: Nostril-snout tip distance, measured as shortest distance between center of nostril to snout tip; RC: Radius-ulna length, measured in latero-external view as distance between base of inner metatarsal granule and center of elbow; TA: Tarsus length, measured from center of tibiotarsal articulation to base of inner metatarsal tubercle; TI: Tibia length, measured in latero-external view from center of knee to middle of tibiotarsal articulation; T1: Outer metatarsal tubercle length along toe 1, measured in lateral view from its base to its top; TO3: Toe 3 length, measured in dorsal view from base of proximal phalange to tip of toe 3; TO5: Toe 5 length, measured in dorsal view from base of proximal phalange to tip of toe 5.