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Rakotoarison, Andolalao, Scherz, Mark D., Bletz, Molly C., Razafindraibe, Jary H., Glaw, Frank, Vences, Miguel (2019): Description of the lucky Cophyla (Microhylidae, Cophylinae), a new arboreal frog from Marojejy National Park in north-eastern Madagascar. Zootaxa 4651 (2): 271-288, DOI: https://doi.org/10.11646/zootaxa.4651.2.4

Uroplatus finaritra sp. nov. Figs. 1–5, Table 1 Uroplatus sp. G—Raxworthy et al. (2008) Uroplatus sp. Ca9— Ratsoavina et al. (2013) LSID: urn:lsid:zoobank.org:act: E6A1407B-7C9E-46F6-8FEC-D120A111A344 Holotype. UADBA-R 70489 (field number APR 12522), adult female, collected on the eastern slope of Marojejy by a tributary of the Ampanasatongotra River, Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.4559°S, 49.7749°E (780 m a.s.l.) on the night of 15 May 2016 by A. P. Raselimanana. Paratypes. UADBA-R 70493 (field number APR 12591), adult male, UADBA-R 70490 (APR 12590), adult male, and UADBA-R 70491 (APR 12589), adult female, three specimens collected in the same locality as the holotype (750–810 m a.s.l.) on 18 May 2016 by A. P. Raselimanana; UADBA-R 70492 (APR 12691), adult male, and UADBA-R 70494 (APR 12692), juvenile female, two specimens collected on the western slope of Marojejy by Tsiasisa River, 2.2 km east of Antanimbaribe, Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.5074°S, 49.6153°E (845 m a.s.l.) on the night of 28 May 2016 by A. P. Raselimanana; UADBA-R 70501 (MSZC 0253), subadult female, ZSM 458 /2016 (MSZC 0271), subadult female, two specimens collected near the path below Camp 1 (Camp Mantella) of Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.438°S, 49.776°E (450 m a.s.l.) on 21–23 November 2016 by M. D. Scherz, M. Bletz, A. Rakotoarison, J. Razafindraibe, A. Razafimanantsoa, and M. Vences. Etymology. Finaritra is a Malagasy word for special greetings but also means healthy and happy. We refer to our delight in describing this splendid and exceptionally large species from a clade of generally small-sized leaftailed geckos. The name is an invariable noun in apposition. Diagnosis. Uroplatus finaritra sp. nov. is assigned to the Uroplatus ebenaui group based on its relatively small size and its triangular head with supraocular spines, and overall leaf-mimicking aspect. It is characterised by (1) a dark red oral mucosa, (2) a comparatively large body size (82.5–95.3 mm SVL in adults), (3) 7–8 lamellae (exceptionally 6) under the third finger and toe, and (4) a long (length 53–65% of SVL), wide (width 16–18% of SVL), leaf-shaped tail. It differs from all members of the U. fimbriatus group (U. fimbriatus, U. giganteus, U. henkeli, U. sikorae, and U. sameiti) and U. lineatus by its smaller size (SVL 82.5–95.3 mm vs. 85–200 mm) and the lack of lateral dermal fringes on any part of the body (vs. presence except in U. lineatus). Also, Uroplatus finaritra sp. nov. diverges from the species of the U. fimbriatus group by their lack of lateral body compression. The prominent triangular head, supraocular spines, and rather smooth skin distinguish Uroplatus finaritra sp. nov. from U. alluaudi, U. guentheri, U. malahelo and U. pietschmanni. Amongst the U. ebenaui group, the rather long tail (TAL/SVL 0.53–0.68) differentiates Uroplatus finaritra sp. nov. from U. ebenaui, U. fiera, U. fotsivava, and U. kelirambo (TAL/SVL 0.22–0.44), and furthermore it differs from U. fiera and U. fotsivava by a pigmented oral mucosa (vs. unpigmented). The new species differs from U. finiavana by a wider tail (TAW/SVL 0.16–0.18 vs. 0.05–0.14), and by its pigmented oral mucosa (vs. unpigmented). By its large tail and pigmented oral mucosa, the new species is most similar to U. malama and U. phantasticus. It differs from U. phantasticus by larger size (SVL 83–95 mm vs. 52–76 mm), a generally shorter tail (TAL/SVL 0.53–0.65 vs. 0.62–0.76), a typically higher number of lamellae under the third finger and toe (6–8 vs. 5–7), and the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black). It differs from U. malama by a spinier integument in some adult males, by the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black), and probably by a narrower tail (TAW/SVL 0.16–0.18 vs. 0.24). In the mitochondrial phylogeny, the new species occupies an isolated position within the U. ebenaui group, without affinities to the morphologically similar U. phantasticus and U. malama (which are not each other's closest relatives either). The nuclear gene c-mos reveals haplotype sharing of the new species with U. kelirambo and two undescribed candidate species (U. sp. Ca3 and Ca4) occurring in the same region of Madagascar, but morphologically very different by their short tails. TABLE 1. Morphological measurements and scale counts of Uroplatus finaritra sp. nov. For abbreviations used, refer to Methods; additional abbreviations: M, male; F, female; sa, subadult; j, juvenile; HT, holotype; PT, paratype; na, not applicable (due to missing original tail); nm, not measured. All measurements in mm. Description of the holotype. Adult female in fair condition, with a fully intact tail, both of its hind limbs fractured (Fig. 2). SVL 95.3 mm, tail length 50.9 mm, maximum tail width 17.0 mm; for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis indistinct and concave; snout sloping weakly downwards anteriorly; snout weakly depressed, fairly long (1.7 times longer than eye diameter); eyes large (eye diameter 6.5 mm), bulging slightly above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small (horizontal diameter 1.1 mm), its opening facing posterolaterally, but also posteroventrally (ear opening clearly visible in ventral view but not in dorsal view); nostrils laterally oriented; body somewhat laterally compressed, without lateral dermal fringes; limbs without fringes and very few, miniscule spines on the hind limbs (one on the knee, as well as a small flap of skin), and two miniscule spines on the forelimb (on the elbow and near the wrist); forelimb reaches the canthus rostralis when adpressed forward and midbody when adpressed backwards along body (forelimb length/axilla–groin distance 33.4/ 50.2 mm = 67%), hindlimb reaches ¾ up the body when adpressed forward along body (hindlimb length/axilla–groin distance 41.4/ 50.2 mm = 83%); original tail length 53% of snout–vent length, membranous borders of the tail symmetrical, without any emarginations, long and leafshaped, with a thin, borderless spatulate tip. Toes bear small claws, with the distal phalange not much wider than the rest of the digit; the third finger bears 7 lamellae, the third toe 8 lamellae. Nares separated from each other by at least 10 small granular scales, from the first supralabial by 3 scales, and from the rostral scale by 5 scales; first supralabial not taller than the others; rostral entire, much wider than tall; mental scale very small, not differentiated from infralabial scales; 25/26 (right/left) supralabials and 25/25 infralabials; no enlarged postmental scales or chin shields; dorsal and ventral scales of head, neck, body, limbs, and tail small, granular, juxtaposed and largely of uniform size, except on the posterior ventral abdomen, where they are slightly larger than the rest of the body, and arranged almost uniformly. Two curved lines (rows of slightly enlarged scales) extending from the posterolateral parts of the head (nuchal region) converge on the neck forming a V-shaped pattern (neck triangular line). A similar, moderately distinct, fairly straight line (also formed by a row of slightly enlarged scales) is present between the eyes and connects the supraocular spines. Another, less distinct line is present connecting the anterior angle of the eyes over the snout in a bowed line. The body possesses very few dermal spines—a prominent pointed supraocular spine, a single spine at the posterior angle of the head, and a small number of spines on each limb (described above). No axillary pits present. Colouration. After two years of preservation in 70% ethanol the colour is slightly faded (Fig. 2). All dorsal surfaces of the specimen are a ruddy brown, fading to a salmon brown in the neck region and at the insertion of the hind limbs. A fine network of blackish lines is present across the body, with the following notable features: a somewhat straight line connecting the suprocular spines (described above); a v-shaped marking on the neck, which extends into a vertebral stripe that reaches all the way to the tail tip; a number of posteriorly oriented chevrons from the mid-dorsum to the belly, becoming thinner posteroventrally, three of which are most distinct. The exterior of the shank and outermost two toes are grey-black, whereas the inner foot is brown. The dorsal snout is darker than the rest of the head. The tail is not different in colour or pattern from the dorsal body. A distinct lateral line runs along the ventrolateral edges of the abdomen from the insertion of the leg to the axilla. The ventral colouration is uniformly orange-brown, with a faint, symmetrical pattern of dark lines on the chin. The soles of the hands and feet are grey. The ventral tail is the colour of the trunk mottled with grey splotches. A faint cream tear-like marking is present at the posterior corner of the eye above the supralabial. The oral mucosa is dark red, with a pink tongue and cream under the eyes and along the lower jaw. In life, the iris colour was silver externally and rusty around the pupil (Fig. 3d). Variation. Molecular data are also available from a previous paper (Raxworthy et al. 2008) for one specimen (RAN 42274) from Marojejy as Uroplatus sp. G, included for 12S and cytochrome b as U. sp. Ca 9 in Ratsoavina et al. (2013). The available sequence for 12S agrees with those of our specimens. The paratypes were sexed as adult males by the presence of everted hemipenes (UADBA-R 70493) or presence of distinctly enlarged tail base (UADBA-R 70490 and 70492), and distinctly notched tail edges; or as females based on the absence of hemipenes or hemipenial bulges, presence of visible oviduct or ovary structures after dissection, and generally smooth tail edges. In the smallest specimen (UADBA-R 70494) female inner reproductive organs were observed but were only weakly developed, and we therefore consider this specimen as juvenile. The sexual maturity of two further specimens (UADBA-R 70501 and ZSM 458/2016) is unclear, as they are considerably smaller than the large adult females (UADBA-R 70491 and 70489) but have similar gonad development. We tentatively consider these to be subadults. In general, the paratypes agree well with the holotype in morphology. For measurements, see Table 1. The number of head spines varies from 2–9, and is more in males than in females, as males are more spiny than females in general, and have more heterogeneous scalation. Forelimbs can bear a minimum of two spines, or as many as eight, with a small spine present on the elbow. Hind limbs can bear 2–15 spines, and the dermal knee flap almost always bears at least a small spine, which is smaller in females than males. In males the tail is moderately emarginated, with semicircular emarginations that tend to be asymmetrical. In females the tail is without emarginations and has smooth edges. Internarial scales range from nine to ten, supralabials from 25–29, and infralabials from 21–25. The third finger bears 7–8 lamellae (6 in one juvenile specimen, UADBA-R 70494), the third toe bears 7–8 lamellae. The colouration is varied, as is commonly the case in Uroplatus species (Figs. 3–4). The line on the head connecting the supraocular spines is generally bowed, but can be slightly pointed posteriorly, and in UADBA-R 70492, it is irregular at its midpoint. The dark vertebral stripe is only present in the holotype, UADBA-R 70491 and 70494, and in other specimens is either absent, or is cream and not dark (in UADBA-R 70492 and 70490). Overall dorsal colouration differs dramatically, from almost completely reddish brown (UADBA-R 70501) to highly mottled greys and blacks (UADBA-R 70493). Irregular flecks can be present or absent on any part of the body. The different colouration of the outer shank and foot is always present, and a cream tear-like marking below the eye is almost always present (absent in UADBA-R 70494 and 70501). The ventrolateral stripe is always present but not always distinct (indistinct in UADBA-R 70493). Ventral colouration is as variable as dorsal colouration. UADBA-R 70491 is extremely similar to the holotype, and to a lesser degree 70492, 70494, and 70501 resemble the holotype ventrally. UADBA-R 70490 and 70493 differ dramatically in being darker in colour, having irregular flecks of cream and reddish brown, and lacking the symmetrical markings on the chin. In life, the colouration of the iris is typically silvery but can be red (Figs. 3c, f), and the oral mucosa was consistently dark red (Fig. 5). Distribution. At present this species has only been collected on the Marojejy Massif, at altitudes from 450–845 m a.s.l. Natural History. The holotype was collected active at night on a stem 0.5 m above the forest floor in nearly closed canopy humid forest at the bottom of a slope, in the vicinity of a Uroplatus lineatus specimen. UADBA-R 70490, 70491, and 70493 were collected on small branches and leaves between 1.5 and 2.5 m above the ground in similar forest. UADBA-R 70492 and 70494 were also active on branches 3–4.5 m from the forest floor in nearly closed-canopy humid forest with a thick understory and ferns. UADBA-R 70501 and ZSM 458/2016 were collected active at night on thin branches (up to 4 m above the ground) in closed-canopy, low-altitude rainforest. We consider the various collection localities to represent a single Threat-Defined Location in the sense of the International Union for the Conservation of Nature (IUCN 2012), as we have recently done for several frog species from low- to mid-elevation forests in Marojejy (Scherz et al. 2016, Rakotoarison et al. 2017). In line with these assessments, we recognise that there is on-going habitat degradation due to illegal logging activity, especially at low altitudes in and around Marojejy National Park. The Extent of Occurrence and Area of Occupancy of the new species is certainly greater than 10 km 2, but is likely less than 500 km 2, and as such we recommend a status of Endangered for it under criterion B1ab(iii).

Uroplatus finaritra sp. nov. Figs. 1���5, Table 1 Uroplatus sp. G���Raxworthy et al. (2008) Uroplatus sp. Ca9��� Ratsoavina et al. (2013) LSID: urn:lsid:zoobank.org:act: E6A1407B-7C9E-46F6-8FEC-D120A111A344 Holotype. UADBA-R 70489 (field number APR 12522), adult female, collected on the eastern slope of Marojejy by a tributary of the Ampanasatongotra River, Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.4559��S, 49.7749��E (780 m a.s.l.) on the night of 15 May 2016 by A. P. Raselimanana. Paratypes. UADBA-R 70493 (field number APR 12591), adult male, UADBA-R 70490 (APR 12590), adult male, and UADBA-R 70491 (APR 12589), adult female, three specimens collected in the same locality as the holotype (750���810 m a.s.l.) on 18 May 2016 by A. P. Raselimanana; UADBA-R 70492 (APR 12691), adult male, and UADBA-R 70494 (APR 12692), juvenile female, two specimens collected on the western slope of Marojejy by Tsiasisa River, 2.2 km east of Antanimbaribe, Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.5074��S, 49.6153��E (845 m a.s.l.) on the night of 28 May 2016 by A. P. Raselimanana; UADBA-R 70501 (MSZC 0253), subadult female, ZSM 458 /2016 (MSZC 0271), subadult female, two specimens collected near the path below Camp 1 (Camp Mantella) of Marojejy National Park, Sava Region, northern Madagascar, at ca. 14.438��S, 49.776��E (450 m a.s.l.) on 21���23 November 2016 by M. D. Scherz, M. Bletz, A. Rakotoarison, J. Razafindraibe, A. Razafimanantsoa, and M. Vences. Etymology. Finaritra is a Malagasy word for special greetings but also means healthy and happy. We refer to our delight in describing this splendid and exceptionally large species from a clade of generally small-sized leaftailed geckos. The name is an invariable noun in apposition. Diagnosis. Uroplatus finaritra sp. nov. is assigned to the Uroplatus ebenaui group based on its relatively small size and its triangular head with supraocular spines, and overall leaf-mimicking aspect. It is characterised by (1) a dark red oral mucosa, (2) a comparatively large body size (82.5���95.3 mm SVL in adults), (3) 7���8 lamellae (exceptionally 6) under the third finger and toe, and (4) a long (length 53���65% of SVL), wide (width 16���18% of SVL), leaf-shaped tail. It differs from all members of the U. fimbriatus group (U. fimbriatus, U. giganteus, U. henkeli, U. sikorae, and U. sameiti) and U. lineatus by its smaller size (SVL 82.5���95.3 mm vs. 85���200 mm) and the lack of lateral dermal fringes on any part of the body (vs. presence except in U. lineatus). Also, Uroplatus finaritra sp. nov. diverges from the species of the U. fimbriatus group by their lack of lateral body compression. The prominent triangular head, supraocular spines, and rather smooth skin distinguish Uroplatus finaritra sp. nov. from U. alluaudi, U. guentheri, U. malahelo and U. pietschmanni. Amongst the U. ebenaui group, the rather long tail (TAL/SVL 0.53���0.68) differentiates Uroplatus finaritra sp. nov. from U. ebenaui, U. fiera, U. fotsivava, and U. kelirambo (TAL/SVL 0.22���0.44), and furthermore it differs from U. fiera and U. fotsivava by a pigmented oral mucosa (vs. unpigmented). The new species differs from U. finiavana by a wider tail (TAW/SVL 0.16���0.18 vs. 0.05���0.14), and by its pigmented oral mucosa (vs. unpigmented). By its large tail and pigmented oral mucosa, the new species is most similar to U. malama and U. phantasticus. It differs from U. phantasticus by larger size (SVL 83���95 mm vs. 52���76 mm), a generally shorter tail (TAL/SVL 0.53���0.65 vs. 0.62���0.76), a typically higher number of lamellae under the third finger and toe (6���8 vs. 5���7), and the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black). It differs from U. malama by a spinier integument in some adult males, by the absence of black pigment of the oral mucosa (oral mucosa dark red vs. black), and probably by a narrower tail (TAW/SVL 0.16���0.18 vs. 0.24). In the mitochondrial phylogeny, the new species occupies an isolated position within the U. ebenaui group, without affinities to the morphologically similar U. phantasticus and U. malama (which are not each other's closest relatives either). The nuclear gene c-mos reveals haplotype sharing of the new species with U. kelirambo and two undescribed candidate species (U. sp. Ca3 and Ca4) occurring in the same region of Madagascar, but morphologically very different by their short tails. TABLE 1. Morphological measurements and scale counts of Uroplatus finaritra sp. nov. For abbreviations used, refer to Methods; additional abbreviations: M, male; F, female; sa, subadult; j, juvenile; HT, holotype; PT, paratype; na, not applicable (due to missing original tail); nm, not measured. All measurements in mm. Description of the holotype. Adult female in fair condition, with a fully intact tail, both of its hind limbs fractured (Fig. 2). SVL 95.3 mm, tail length 50.9 mm, maximum tail width 17.0 mm; for further measurements see Table 1. Head triangular in dorsal view; canthus rostralis indistinct and concave; snout sloping weakly downwards anteriorly; snout weakly depressed, fairly long (1.7 times longer than eye diameter); eyes large (eye diameter 6.5 mm), bulging slightly above dorsal surface of cranium, directed laterally, pupil vertical with crenate borders; ear opening very small (horizontal diameter 1.1 mm), its opening facing posterolaterally, but also posteroventrally (ear opening clearly visible in ventral view but not in dorsal view); nostrils laterally oriented; body somewhat laterally compressed, without lateral dermal fringes; limbs without fringes and very few, miniscule spines on the hind limbs (one on the knee, as well as a small flap of skin), and two miniscule spines on the forelimb (on the elbow and near the wrist); forelimb reaches the canthus rostralis when adpressed forward and midbody when adpressed backwards along body (forelimb length/axilla���groin distance 33.4/ 50.2 mm = 67%), hindlimb reaches �� up the body when adpressed forward along body (hindlimb length/axilla���groin distance 41.4/ 50.2 mm = 83%); original tail length 53% of snout���vent length, membranous borders of the tail symmetrical, without any emarginations, long and leafshaped, with a thin, borderless spatulate tip. Toes bear small claws, with the distal phalange not much wider than the rest of the digit; the third finger bears 7 lamellae, the third toe 8 lamellae. Nares separated from each other by at least 10 small granular scales, from the first supralabial by 3 scales, and from the rostral scale by 5 scales; first supralabial not taller than the others; rostral entire, much wider than tall; mental scale very small, not differentiated from infralabial scales; 25/26 (right/left) supralabials and 25/25 infralabials; no enlarged postmental scales or chin shields; dorsal and ventral scales of head, neck, body, limbs, and tail small, granular, juxtaposed and largely of uniform size, except on the posterior ventral abdomen, where they are slightly larger than the rest of the body, and arranged almost uniformly. Two curved lines (rows of slightly enlarged scales) extending from the posterolateral parts of the head (nuchal region) converge on the neck forming a V-shaped pattern (neck triangular line). A similar, moderately distinct, fairly straight line (also formed by a row of slightly enlarged scales) is present between the eyes and connects the supraocular spines. Another, less distinct line is present connecting the anterior angle of the eyes over the snout in a bowed line. The body possesses very few dermal spines���a prominent pointed supraocular spine, a single spine at the posterior angle of the head, and a small number of spines on each limb (described above). No axillary pits present. Colouration. After two years of preservation in 70% ethanol the colour is slightly faded (Fig. 2). All dorsal surfaces of the specimen are a ruddy brown, fading to a salmon brown in the neck region and at the insertion of the hind limbs. A fine network of blackish lines is present across the body, with the following notable features: a somewhat straight line connecting the suprocular spines (described above); a v-shaped marking on the neck, which extends into a vertebral stripe that reaches all the way to the tail tip; a number of posteriorly oriented chevrons from the mid-dorsum to the belly, becoming thinner posteroventrally, three of which are most distinct. The exterior of the shank and outermost two toes are grey-black, whereas the inner foot is brown. The dorsal snout is darker than the rest of the head. The tail is not different in colour or pattern from the dorsal body. A distinct lateral line runs along the ventrolateral edges of the abdomen from the insertion of the leg to the axilla. The ventral colouration is uniformly orange-brown, with a faint, symmetrical pattern of dark lines on the chin. The soles of the hands and feet are grey. The ventral tail is the colour of the trunk mottled with grey splotches. A faint cream tear-like marking is present at the posterior corner of the eye above the supralabial. The oral mucosa is dark red, with a pink tongue and cream under the eyes and along the lower jaw. In life, the iris colour was silver externally and rusty around the pupil (Fig. 3d). Variation. Molecular data are also available from a previous paper (Raxworthy et al. 2008) for one specimen (RAN 42274) from Marojejy as Uroplatus sp. G, included for 12S and cytochrome b as U. sp. Ca 9 in Ratsoavina et al. (2013). The available sequence for 12S agrees with those of our specimens. The paratypes were sexed as adult males by the presence of everted hemipenes (UADBA-R 70493) or presence of distinctly enlarged tail base (UADBA-R 70490 and 70492), and distinctly notched tail edges; or as females based on the absence of hemipenes or hemipenial bulges, presence of visible oviduct or ovary structures after dissection, and generally smooth tail edges. In the smallest specimen (UADBA-R 70494) female inner reproductive organs were observed but were only weakly developed, and we therefore consider this specimen as juvenile. The sexual maturity of two further specimens (UADBA-R 70501 and ZSM 458/2016) is unclear, as they are considerably smaller than the large adult females (UADBA-R 70491 and 70489) but have similar gonad development. We tentatively consider these to be subadults. In general, the paratypes agree well with the holotype in morphology. For measurements, see Table 1. The number of head spines varies from 2���9, and is more in males than in females, as males are more spiny than females in general, and have more heterogeneous scalation. Forelimbs can bear a minimum of two spines, or as many as eight, with a small spine present on the elbow. Hind limbs can bear 2���15 spines, and the dermal knee flap almost always bears at least a small spine, which is smaller in females than males. In males the tail is moderately emarginated, with semicircular emarginations that tend to be asymmetrical. In females the tail is without emarginations and has smooth edges. Internarial scales range from nine to ten, supralabials from 25���29, and infralabials from 21���25. The third finger bears 7���8 lamellae (6 in one juvenile specimen, UADBA-R 70494), the third toe bears 7���8 lamellae. The colouration is varied, as is commonly the case in Uroplatus species (Figs. 3���4). The line on the head connecting the supraocular spines is generally bowed, but can be slightly pointed posteriorly, and in UADBA-R 70492, it is irregular at its midpoint. The dark vertebral stripe is only present in the holotype, UADBA-R 70491 and 70494, and in other specimens is either absent, or is cream and not dark (in UADBA-R 70492 and 70490). Overall dorsal colouration differs dramatically, from almost completely reddish brown (UADBA-R 70501) to highly mottled greys and blacks (UADBA-R 70493). Irregular flecks can be present or absent on any part of the body. The different colouration of the outer shank and foot is always present, and a cream tear-like marking below the eye is almost always present (absent in UADBA-R 70494 and 70501). The ventrolateral stripe is always present but not always distinct (indistinct in UADBA-R 70493). Ventral colouration is as variable as dorsal colouration. UADBA-R 70491 is extremely similar to the holotype, and to a lesser degree 70492, 70494, and 70501 resemble the holotype ventrally. UADBA-R 70490 and 70493 differ dramatically in being darker in colour, having irregular flecks of cream and reddish brown, and lacking the symmetrical markings on the chin. In life, the colouration of the iris is typically silvery but can be red (Figs. 3c, f), and the oral mucosa was consistently dark red (Fig. 5). Distribution. At present this species has only been collected on the Marojejy Massif, at altitudes from 450���845 m a.s.l. Natural History. The holotype was collected active at night on a stem 0.5 m above the forest floor in nearly closed canopy humid forest at the bottom of a slope, in the vicinity of a Uroplatus lineatus specimen. UADBA-R 70490, 70491, and 70493 were collected on small branches and leaves between 1.5 and 2.5 m above the ground in similar forest. UADBA-R 70492 and 70494 were also active on branches 3���4.5 m from the forest floor in nearly closed-canopy humid forest with a thick understory and ferns. UADBA-R 70501 and ZSM 458/2016 were collected active at night on thin branches (up to 4 m above the ground) in closed-canopy, low-altitude rainforest. We consider the various collection localities to represent a single Threat-Defined Location in the sense of the International Union for the Conservation of Nature (IUCN 2012), as we have recently done for several frog species from low- to mid-elevation forests in Marojejy (Scherz et al. 2016, Rakotoarison et al. 2017). In line with these assessments, we recognise that there is on-going habitat degradation due to illegal logging activity, especially at low altitudes in and around Marojejy National Park. The Extent of Occurrence and Area of Occupancy of the new species is certainly greater than 10 km 2, but is likely less than 500 km 2, and as such we recommend a status of Endangered for it under criterion B1ab(iii)., Published as part of Ratsoavina, Fanomezana Mihaja, Raselimanana, Achille P., Scherz, Mark D., Rakotoarison, Andolalao, Razafindraibe, Jary H., Glaw, Frank & Vences, Miguel, 2019, Finaritra! A splendid new leaf-tailed gecko (Uroplatus) species from Marojejy National Park in north-eastern Madagascar, pp. 563-577 in Zootaxa 4545 (4) on pages 566-572, DOI: 10.11646/zootaxa.4545.4.7, http://zenodo.org/record/2618341, {"references":["Ratsoavina, F. M., Raminosoa, N. R., Louis, E. E. Jr., Raselimanana, A. P., Glaw, F. & Vences, M. (2013) An overview of Madagascar's leaf tailed geckos (genus Uroplatus): species boundaries, candidate species and review of geographical distribution based on molecular data. Salamandra, 49, 115 - 148.","Raxworthy, C. J., Pearson, R. G., Zimkus, B. M., Reddy, S., Deo, A. J., Nussbaum, R. A. & Ingram, C. M. (2008) Continental speciation in the tropics: contrasting biogeographic patterns of divergence in the Uroplatus leaf-tailed gecko radiation of Madagascar. Journal of Zoology, 275, 423 - 440. https: // doi. org / 10.1111 / j. 1469 - 7998.2008.00460. x","IUCN. (2012) IUCN Red List Categories and Criteria: Fersion 3.1. IUCN, Gland and Cambridge.","Scherz, M. D., Glaw, F., Vences, M., Andreone, F. & Crottini, A. (2016) Two new species of terrestrial microhylid frogs (Microhylidae: Cophylinae: Rhombophryne) from northeastern Madagascar. Salamandra, 52, 91 - 106.","Rakotoarison, A., Scherz, M. D., Glaw, F., Kohler, J., Andreone, F., Franzen, M., Glos, J., Hawlitschek, O., Jono, T., Mori, A., Ndriantsoa, S. H., Raminosoa Rasoamampionona, N., Riemann, J. C., Rodel, M. - O., Rosa, G. M., Vieites, D. R., Crottini, A. & Vences, M. (2017) Describing the smaller majority: integrative taxonomy reveals twenty-six new species of tiny microhylid frogs (genus Stumpffia) from Madagascar. Fertebrate Zoology, 67, 271 - 398."]}

Hutter, Carl Richard, Andriampenomanana, Zo F., Razafindraibe, Jary H., Rakotoarison, Dr. Andolalao, Scherz, Mark D. (2018): New dietary data from Compsophis and Alluaudina species (Squamata: Lamprophiidae: Pseudoxyrhophiinae), and implications for their dietary complexity and evolution. Journal of Natural History 52 (39-40): 2497-2510, DOI: 10.1080/00222933.2018.1543732, URL: http://dx.doi.org/10.1080/00222933.2018.1543732

Gephyromantis tohatra sp. nov. (Figs. 2–3) Holotype. ZSM 422/2016 (ZCMV 15245), adult male, collected at a site along the main trail leading from Camp Simpona to the summit of the Marojejy Massif, Sava Region, northeastern Madagascar, at geographical coordinates 14.44400°S, 49.73791°E and an elevation of 1758 m above sea level (Fig. 2, 3a–b) by M. D. Scherz, M. Vences, J. H. Razafindraibe, and M. Bletz, in the evening of the 19th of November 2016. Paratype. UADBA uncatalogued (ZCMV 15236), adult male, with the same collection data as the holotype. Diagnosis. The new species is assigned to the genus Gephyromantis and subgenus Duboimantis on the basis of its mostly connected lateral metatarsalia, moderately enlarged finger tips, type 2 femoral glands (Glaw et al. 2000), inner and outer dorsolateral ridges sensu Vences & Glaw (2001), presence of webbing between the toes, absence of a distinct white spot in the centre of the tympanum, and presence of small supraocular spines. It is characterised by the following unique suite of characters: (1) SVL at least up to 33 mm, (2) orange to yellowish ventral colouration in males, (3) type 2 femoral glands with 10–21 small granules, (4) strongly distinct inner and outer dorsolateral ridges sensu Vences & Glaw (2001), (5) a pale loreal region and broad pale marking covering most of the anterior edge of the tympanum, (6) second finger much shorter than fourth, (7) third toe much shorter than fifth, and (8) 7– 10 distinct pulsed notes per call. Gephyromantis tohatra sp. nov. differs from the majority of Duboimantis species by smaller body size: adult male SVL 33 vs.> 35 mm in G. granulatus, G. cornutus, G. redimitus, G. tschenki, G. salegy, G. tandroka, G. luteus, G. sculpturatus, G. plicifer, and G. zavona. It can also be distinguished from all other species of Duboimantis by its orange to yellowish venter (typically whitish in all other species). Additionally, the presence of distinct inner and outer dorsolateral ridges distinguishes the new species in particular from G. granulatus, G. redimitus, and G. zavona in which these folds (in particular the inner dorsolateral ridges) are absent or very weakly expressed. Gephyromantis luteus, G. plicifer, and G. sculpturatus have dark concave markings in the suprascapular region (absent in the new species) and much more extensive webbing on the feet. Gephyromantis cornutus, G. tschenki, and G. tandroka have strongly expressed interocular tubercles (absent in the new species). Three species of Duboimantis overlap with the new species in body size. Gephyromantis leucomaculatus is typically larger (adult male SVL 32–41 mm) and differs by absence of inner dorsolateral ridges and straight outer dorsolateral ridges (vs. anteriorly curved). Gephyromantis moseri (adult male SVL 27–40 mm) has a granular dorsum without or with interrupted dorsolateral ridges (vs. smooth or reticulated dorsum and continuous dorsolateral ridges in the new species) and interocular tubercles (vs. absence). Gephyromantis schilfi (adult male SVL 27–30 mm) occurs in close syntopy with the new species but differs from it by the absence of distinct inner and outer dorsolateral ridges (vs. presence in the new species). Bioacoustically, the new species differs from all species in Duboimantis by advertisement call structure. Most species in this subgenus emit regularly repeated calls, each of which consists of a single, typically pulsed or pulsatile note (G. granulatus, G. cornutus, G. leucomaculatus, G. moseri, G. redimitus, G. schilfi, G. tschenki, G. tandroka, G. plicifer, G. zavona). In many of these species, notes can also be more irregularly arranged in short series, but calls do not consist of a regular number of 7–10 short and rapidly repeated notes as in G. tohatra sp. nov. This characteristic is particularly helpful in its distinction from the sympatrically occurring and morphologically similar G. schilfi, which emits a single, densely pulsed note. Gephyromantis sculpturatus and G. luteus emit a fast series of squeaking, unpulsed notes that are very distinct from the call of G. tohatra. The only other Duboimantis with a surprisingly similar call structure to the new species is G. salegy, which however differs by a substantially larger body size of 45–48 mm in adult males and by a larger number of granules in femoral glands (36–38 vs. 10– 21). G. tohatra sp. nov. differs from its probable sister species G. schilfi (Fig. 1) by a substantial genetic differentiation of 4.3% uncorrected pairwise distance (p-distance) in the 16S gene, and from all other species of Duboimantis by a 16S p-distance>6%. The bioacoustically similar G. salegy differs by 6.8% 16S p-distance and is not the sister taxon of the new species. Holotype description. A specimen in a good state of preservation, the right thigh muscle taken for DNA tissue samples. SVL 32.7 mm. For other measurements see below. Body gracile; head longer than wide, wider than body; snout pointed in dorsal and lateral view; nostrils directed laterally, protruding slightly, much nearer to tip of snout than to eye; canthus rostralis distinct, straight; loreal region concave and weakly oblique; tympanum indistinct, oval, its horizontal diameter 42% of eye diameter; supratympanic fold distinct, straight, following the line of the eyelid to above the insertion of the arm; tongue fairly narrow, posteriorly bifid; vomerine teeth clearly distinct, arranged in two small aggregations on either side of the midline of the palate at the level of the anterior edge of the eye, posteromedial to choanae; choanae small and rounded and laterally displaced. Dark, translucent dermal fold below each jaw starting at the level of the mid-eye. Arms slender, subarticular tubercles single, highly distinct; outer metacarpal tubercle small and indistinct and inner metacarpal tubercle relatively well developed; fingers without webbing; relative length of fingers 1 et al. (2000), consisting of roughly 16 granules on the right thigh and 21 on the left thigh. Measurements, all in mm: SVL, 32.7; HW, 9.5; HL, 12.0; TD, 1.9; ED, 4.5; END, 3.3; NSD, 2.7; NND, 3.8; UAL, 6.7; LAL, 8.0; HAL, 11.0; THIL, 17.7; TIBL, 19.4; TARL, 9.5; FOL, 18.2. In life (Fig. 3a,b), the dorsum was a medium brown with the external dorsolateral ridges distinctly tan, the internal ridges less distinct but also light in colour. The dorsal surface of head was not distinct from the dorsum, with a slightly darker brown band between the eyes. The flank fades to a more grey-brown ventrally. The dorsal surfaces of hindlimbs were as the flank in colour, with three dark brown crossbands on the thigh, three on the shank, and no crossbands on the foot. The dorsal surface of forelimb was similar to the lower leg in colour. No light annulus was present before the tip of each toe or finger. The lateral head was distinctly differentiated from the dorsal head, being almost black along the canthus rostralis, from the nostril to the tip of the snout, over the dorsal portion of the tympanic region beneath the supratympanic fold, and beneath the eye. An immaculate white spot was present anterior to the eye, with a more faded spot posterior to the eye over part of the tympanum. The iris was whitish above, and dark brown below the pupil. The lower lip was yellowish. Ventrally, the hands, forearms, tarsus and feet were all burnt umber in colour, with pale yellowish subarticular tubercles on the fingers, the subarticular tubercles of the toes being more orange in colour. The ventral chin and throat were burnt umber, especially on the jaw, making a distinct line anterior to the pectoral girdle, posterior to which the abdomen was orange with a few dark brown flecks anteriorly. The ventral thighs were also orange. The femoral glands were a creamy orange. The ventral humerus was semi-translucent. After six months in preservative, the coloration has faded. Dorsally, areas that were brown have become grey to silver, with light grey dorsolateral ridges. The head colouration is mostly retained. The ventral colouration has lost its orange hue, and is now a yellow-cream. All elements of the colour pattern are however retained. Variation. The paratype was not available for measurements but was of similar size as assessed in the field. Its colour in life (Fig. 3c,d) was generally similar to the holotype, but it lacked the bright whitish marking anterior to the eye, had only one dark brown marking below the eye (two in the holotype), had slightly broader crossbands on the legs, and its outer dorsolateral ridges were less distinctly coloured than those of the holotype. Ventrally it was also similar to the holotype but its abdomen was more yellowish, with yellow-cream patches posterior to the pectoral girdle. Etymology. The specific name is a noun in apposition, derived from the Malagasy word ‘ tohatra,’ meaning ‘stairs,’ in reference to the difficulty of hiking up Marojejy on very steep trails with stair-like stretches between Camp Marojejia and Camp Simpona, and especially from Camp Simpona to the summit, necessary to discover this species. Natural history. Calling males were heard at night at a site between Camp 3 (= Camp Simpona) and the summit of the Marojejy Massif, within montane rainforest of rather low canopy height and in an area of very steep slopes. Most specimens were calling in a dry headwater area of a small stream, in between very dense bushy undergrowth, at perch heights of 0.5–1 m above the ground from leaves and branches. In some areas the call could be heard from far outside the fores. Advertisement calls. Call recordings were made in an area where several (>5) males were calling relatively close to one another, and were obtained from the holotype and the paratype. The calls are considered advertisement calls because they were loud, stereotyped calls emitted by males without any sign of conflict with conspecifics. Calls consist of 7–10 pulsed notes (Fig. 4). Calls are separated by long and rather irregular intervals. In the holotype, call duration was 547–640 ms (594 ± 38 ms; n = 9), inter–call interval 1258–2306 ms (1622 ± 398 ms; n = 8), dominant frequency 3014–3273 Hz, and approximate prevalent band width 2000–5000 Hz. Each note had three to four recognizable pulses, with the last note of the call having more pulses (about 8). Characteristics of notes were measured in two calls. Pulses were not separated by distinct silent intervals. Intensity of each note was highest in the first pulse of each note. Note and interval duration are only tentatively presented, as there remains a weak background sound energy during these periods. Call 1: Note duration 34–78 ms (53 ± 14 ms, n = 7), internote interval duration 19–40 ms (30 ± 8 ms, n = 6), pulses per note 3–8 (4 ± 2, n = 7). Call 2: Note duration 45–77 ms (57 ± 11 ms, n = 7), inter-note interval duration 7–36 ms (26 ± 12 ms, n = 6), pulses per note 3–9 (4 ± 2, n = 8). Calls of the paratype were similar in most parameters but differed in that the pulses in notes were neither distinct nor clearly distinguishable; clear silent intervals between notes were not present, possibly due to background noise, and a weak frequency modulation was apparent in each note. Call duration was 502–798 ms (698 ± 117 ms, n = 5), inter-call interval duration 1458–1461 (n = 2), notes per call 7–10 (9 ± 1 ms, n = 5), dominant frequency of first three notes in a call 2454–2713 Hz (2637 ± 124 Hz, n = 4), dominant frequency of last 2–3 notes in a call 3014– 3229 Hz (3122 ± 124 Hz, n = 4), time between start of one note and subsequent note 73–86 ms (82 ± 4, n = 17 notes from 2 calls). Comparative data for syntopic G. schilfi. A few individuals of G. schilfi were heard emitting advertisement calls very close to the site in which G. tohatra sp. nov. individuals were heard and collected, on the same day and ca. 15 minutes after collecting G. tohatra sp. nov. One adult calling male was recorded and collected (voucher ZSM 415/2016, field number ZCMV 15246) from dense vegetation at a perch height of ca. 1 m above the ground, in a rainforest area on a steep mountain slope. Measurements: SVL, 29.8; HW, 9.5; HL, 11.6; TD, 1.9; ED, 4.6; END, 2.9; NSD, 2.5; NND, 2.6; UAL, 4.8; LAL, 7.5; HAL, 9.3; THIL, 15.6; TIBL, 18.3; TARL, 9.2; FOL, 16.5. As previously described (Glaw & Vences 2000), the call is a single pulsed note emitted after long and rather irregular silent intervals (Fig. 5). Call duration is 483–538 ms (518 ± 25 ms; n = 4), inter-call interval duration is 2917–5121 ms (3672 ± 1255 ms, n = 3), one note consists of ca. 73–89 pulses (80 ± 7; n = 5). Pulses were not clearly separated by silent intervals and in some cases not unambiguously distinguishable from each other. The intervals between pulse intensity maxima were somewhat irregular, becoming longer towards the end of the call, with a duration of about 5 ms at the beginning of the call and 7–12 ms at the end of the call. Dominant frequency was 3402–3445 Hz and approximate prevalent bandwidth was between 2000–4000 Hz. Range extension for G. schilfi and G. tandroka. The goal of this study is not a comprehensive reassessment of the phylogeny within the subgenus Duboimantis and molecular analyses were thus limited to a subset of taxa. In exploratory analyses, the new species clustered with G. salegy, G. schilfi, and G. tandroka, three species that formed a highly supported subclade of Duboimantis in the analyses of Wollenberg et al. (2011) and Kaffenberger et al. (2012). For our analysis we thus expanded the sampling for these three species with additional individuals. We found molecular evidence for new populations of G. schilfi and G. tandroka, both of which were so far known only from Marojejy. Both these species were now also found on the Sorata Massif north of Marojejy, and had different degrees of genetic differentiation. While the populations of G. schilfi (Sorata vs. Marojejy) differed by only 0.6– 0.8% uncorrected 16S p-distance, those of G. tandroka had a higher divergence of 5.4% which suggests their taxonomic status requires revision and we therefore here flag the Sorata population as new Unconfirmed Candidate Species, Gephyromantis sp. Ca32, rationalised following Vieites et al. (2009). Another candidate species of Duboimantis (G. sp. Ca30 from the Bealanana district of northern Madagascar) that was also recently identified (Scherz et al. in press) was not included in our analysis here, but we have confirmed that it is not conspecific with G. sp. Ca32 (data not shown).

Scherz, Mark D., Razafindraibe, Jary H., Rakotoarison, Andolalao, Dixit, Nadi M., Bletz, Molly C., Glaw, Frank, Vences, Miguel (2017): Yet another small brown frog from high altitude on the Marojejy Massif, northeastern Madagascar (Anura: Mantellidae). Zootaxa 4347 (3), DOI: https://doi.org/10.11646/zootaxa.4347.3.9

Gephyromantis tohatra sp. nov. (Figs. 2���3) Holotype. ZSM 422/2016 (ZCMV 15245), adult male, collected at a site along the main trail leading from Camp Simpona to the summit of the Marojejy Massif, Sava Region, northeastern Madagascar, at geographical coordinates 14.44400��S, 49.73791��E and an elevation of 1758 m above sea level (Fig. 2, 3a���b) by M. D. Scherz, M. Vences, J. H. Razafindraibe, and M. Bletz, in the evening of the 19th of November 2016. Paratype. UADBA uncatalogued (ZCMV 15236), adult male, with the same collection data as the holotype. Diagnosis. The new species is assigned to the genus Gephyromantis and subgenus Duboimantis on the basis of its mostly connected lateral metatarsalia, moderately enlarged finger tips, type 2 femoral glands (Glaw et al. 2000), inner and outer dorsolateral ridges sensu Vences & Glaw (2001), presence of webbing between the toes, absence of a distinct white spot in the centre of the tympanum, and presence of small supraocular spines. It is characterised by the following unique suite of characters: (1) SVL at least up to 33 mm, (2) orange to yellowish ventral colouration in males, (3) type 2 femoral glands with 10���21 small granules, (4) strongly distinct inner and outer dorsolateral ridges sensu Vences & Glaw (2001), (5) a pale loreal region and broad pale marking covering most of the anterior edge of the tympanum, (6) second finger much shorter than fourth, (7) third toe much shorter than fifth, and (8) 7��� 10 distinct pulsed notes per call. Gephyromantis tohatra sp. nov. differs from the majority of Duboimantis species by smaller body size: adult male SVL 33 vs.> 35 mm in G. granulatus, G. cornutus, G. redimitus, G. tschenki, G. salegy, G. tandroka, G. luteus, G. sculpturatus, G. plicifer, and G. zavona. It can also be distinguished from all other species of Duboimantis by its orange to yellowish venter (typically whitish in all other species). Additionally, the presence of distinct inner and outer dorsolateral ridges distinguishes the new species in particular from G. granulatus, G. redimitus, and G. zavona in which these folds (in particular the inner dorsolateral ridges) are absent or very weakly expressed. Gephyromantis luteus, G. plicifer, and G. sculpturatus have dark concave markings in the suprascapular region (absent in the new species) and much more extensive webbing on the feet. Gephyromantis cornutus, G. tschenki, and G. tandroka have strongly expressed interocular tubercles (absent in the new species). Three species of Duboimantis overlap with the new species in body size. Gephyromantis leucomaculatus is typically larger (adult male SVL 32���41 mm) and differs by absence of inner dorsolateral ridges and straight outer dorsolateral ridges (vs. anteriorly curved). Gephyromantis moseri (adult male SVL 27���40 mm) has a granular dorsum without or with interrupted dorsolateral ridges (vs. smooth or reticulated dorsum and continuous dorsolateral ridges in the new species) and interocular tubercles (vs. absence). Gephyromantis schilfi (adult male SVL 27���30 mm) occurs in close syntopy with the new species but differs from it by the absence of distinct inner and outer dorsolateral ridges (vs. presence in the new species). Bioacoustically, the new species differs from all species in Duboimantis by advertisement call structure. Most species in this subgenus emit regularly repeated calls, each of which consists of a single, typically pulsed or pulsatile note (G. granulatus, G. cornutus, G. leucomaculatus, G. moseri, G. redimitus, G. schilfi, G. tschenki, G. tandroka, G. plicifer, G. zavona). In many of these species, notes can also be more irregularly arranged in short series, but calls do not consist of a regular number of 7���10 short and rapidly repeated notes as in G. tohatra sp. nov. This characteristic is particularly helpful in its distinction from the sympatrically occurring and morphologically similar G. schilfi, which emits a single, densely pulsed note. Gephyromantis sculpturatus and G. luteus emit a fast series of squeaking, unpulsed notes that are very distinct from the call of G. tohatra. The only other Duboimantis with a surprisingly similar call structure to the new species is G. salegy, which however differs by a substantially larger body size of 45���48 mm in adult males and by a larger number of granules in femoral glands (36���38 vs. 10��� 21). G. tohatra sp. nov. differs from its probable sister species G. schilfi (Fig. 1) by a substantial genetic differentiation of 4.3% uncorrected pairwise distance (p-distance) in the 16S gene, and from all other species of Duboimantis by a 16S p-distance>6%. The bioacoustically similar G. salegy differs by 6.8% 16S p-distance and is not the sister taxon of the new species. Holotype description. A specimen in a good state of preservation, the right thigh muscle taken for DNA tissue samples. SVL 32.7 mm. For other measurements see below. Body gracile; head longer than wide, wider than body; snout pointed in dorsal and lateral view; nostrils directed laterally, protruding slightly, much nearer to tip of snout than to eye; canthus rostralis distinct, straight; loreal region concave and weakly oblique; tympanum indistinct, oval, its horizontal diameter 42% of eye diameter; supratympanic fold distinct, straight, following the line of the eyelid to above the insertion of the arm; tongue fairly narrow, posteriorly bifid; vomerine teeth clearly distinct, arranged in two small aggregations on either side of the midline of the palate at the level of the anterior edge of the eye, posteromedial to choanae; choanae small and rounded and laterally displaced. Dark, translucent dermal fold below each jaw starting at the level of the mid-eye. Arms slender, subarticular tubercles single, highly distinct; outer metacarpal tubercle small and indistinct and inner metacarpal tubercle relatively well developed; fingers without webbing; relative length of fingers 1 et al. (2000), consisting of roughly 16 granules on the right thigh and 21 on the left thigh. Measurements, all in mm: SVL, 32.7; HW, 9.5; HL, 12.0; TD, 1.9; ED, 4.5; END, 3.3; NSD, 2.7; NND, 3.8; UAL, 6.7; LAL, 8.0; HAL, 11.0; THIL, 17.7; TIBL, 19.4; TARL, 9.5; FOL, 18.2. In life (Fig. 3a,b), the dorsum was a medium brown with the external dorsolateral ridges distinctly tan, the internal ridges less distinct but also light in colour. The dorsal surface of head was not distinct from the dorsum, with a slightly darker brown band between the eyes. The flank fades to a more grey-brown ventrally. The dorsal surfaces of hindlimbs were as the flank in colour, with three dark brown crossbands on the thigh, three on the shank, and no crossbands on the foot. The dorsal surface of forelimb was similar to the lower leg in colour. No light annulus was present before the tip of each toe or finger. The lateral head was distinctly differentiated from the dorsal head, being almost black along the canthus rostralis, from the nostril to the tip of the snout, over the dorsal portion of the tympanic region beneath the supratympanic fold, and beneath the eye. An immaculate white spot was present anterior to the eye, with a more faded spot posterior to the eye over part of the tympanum. The iris was whitish above, and dark brown below the pupil. The lower lip was yellowish. Ventrally, the hands, forearms, tarsus and feet were all burnt umber in colour, with pale yellowish subarticular tubercles on the fingers, the subarticular tubercles of the toes being more orange in colour. The ventral chin and throat were burnt umber, especially on the jaw, making a distinct line anterior to the pectoral girdle, posterior to which the abdomen was orange with a few dark brown flecks anteriorly. The ventral thighs were also orange. The femoral glands were a creamy orange. The ventral humerus was semi-translucent. After six months in preservative, the coloration has faded. Dorsally, areas that were brown have become grey to silver, with light grey dorsolateral ridges. The head colouration is mostly retained. The ventral colouration has lost its orange hue, and is now a yellow-cream. All elements of the colour pattern are however retained. Variation. The paratype was not available for measurements but was of similar size as assessed in the field. Its colour in life (Fig. 3c,d) was generally similar to the holotype, but it lacked the bright whitish marking anterior to the eye, had only one dark brown marking below the eye (two in the holotype), had slightly broader crossbands on the legs, and its outer dorsolateral ridges were less distinctly coloured than those of the holotype. Ventrally it was also similar to the holotype but its abdomen was more yellowish, with yellow-cream patches posterior to the pectoral girdle. Etymology. The specific name is a noun in apposition, derived from the Malagasy word ��� tohatra,��� meaning ���stairs,��� in reference to the difficulty of hiking up Marojejy on very steep trails with stair-like stretches between Camp Marojejia and Camp Simpona, and especially from Camp Simpona to the summit, necessary to discover this species. Natural history. Calling males were heard at night at a site between Camp 3 (= Camp Simpona) and the summit of the Marojejy Massif, within montane rainforest of rather low canopy height and in an area of very steep slopes. Most specimens were calling in a dry headwater area of a small stream, in between very dense bushy undergrowth, at perch heights of 0.5���1 m above the ground from leaves and branches. In some areas the call could be heard from far outside the fores. Advertisement calls. Call recordings were made in an area where several (>5) males were calling relatively close to one another, and were obtained from the holotype and the paratype. The calls are considered advertisement calls because they were loud, stereotyped calls emitted by males without any sign of conflict with conspecifics. Calls consist of 7���10 pulsed notes (Fig. 4). Calls are separated by long and rather irregular intervals. In the holotype, call duration was 547���640 ms (594 �� 38 ms; n = 9), inter���call interval 1258���2306 ms (1622 �� 398 ms; n = 8), dominant frequency 3014���3273 Hz, and approximate prevalent band width 2000���5000 Hz. Each note had three to four recognizable pulses, with the last note of the call having more pulses (about 8). Characteristics of notes were measured in two calls. Pulses were not separated by distinct silent intervals. Intensity of each note was highest in the first pulse of each note. Note and interval duration are only tentatively presented, as there remains a weak background sound energy during these periods. Call 1: Note duration 34���78 ms (53 �� 14 ms, n = 7), internote interval duration 19���40 ms (30 �� 8 ms, n = 6), pulses per note 3���8 (4 �� 2, n = 7). Call 2: Note duration 45���77 ms (57 �� 11 ms, n = 7), inter-note interval duration 7���36 ms (26 �� 12 ms, n = 6), pulses per note 3���9 (4 �� 2, n = 8). Calls of the paratype were similar in most parameters but differed in that the pulses in notes were neither distinct nor clearly distinguishable; clear silent intervals between notes were not present, possibly due to background noise, and a weak frequency modulation was apparent in each note. Call duration was 502���798 ms (698 �� 117 ms, n = 5), inter-call interval duration 1458���1461 (n = 2), notes per call 7���10 (9 �� 1 ms, n = 5), dominant frequency of first three notes in a call 2454���2713 Hz (2637 �� 124 Hz, n = 4), dominant frequency of last 2���3 notes in a call 3014��� 3229 Hz (3122 �� 124 Hz, n = 4), time between start of one note and subsequent note 73���86 ms (82 �� 4, n = 17 notes from 2 calls). Comparative data for syntopic G. schilfi. A few individuals of G. schilfi were heard emitting advertisement calls very close to the site in which G. tohatra sp. nov. individuals were heard and collected, on the same day and ca. 15 minutes after collecting G. tohatra sp. nov. One adult calling male was recorded and collected (voucher ZSM 415/2016, field number ZCMV 15246) from dense vegetation at a perch height of ca. 1 m above the ground, in a rainforest area on a steep mountain slope. Measurements: SVL, 29.8; HW, 9.5; HL, 11.6; TD, 1.9; ED, 4.6; END, 2.9; NSD, 2.5; NND, 2.6; UAL, 4.8; LAL, 7.5; HAL, 9.3; THIL, 15.6; TIBL, 18.3; TARL, 9.2; FOL, 16.5. As previously described (Glaw & Vences 2000), the call is a single pulsed note emitted after long and rather irregular silent intervals (Fig. 5). Call duration is 483���538 ms (518 �� 25 ms; n = 4), inter-call interval duration is 2917���5121 ms (3672 �� 1255 ms, n = 3), one note consists of ca. 73���89 pulses (80 �� 7; n = 5). Pulses were not clearly separated by silent intervals and in some cases not unambiguously distinguishable from each other. The intervals between pulse intensity maxima were somewhat irregular, becoming longer towards the end of the call, with a duration of about 5 ms at the beginning of the call and 7���12 ms at the end of the call. Dominant frequency was 3402���3445 Hz and approximate prevalent bandwidth was between 2000���4000 Hz. Range extension for G. schilfi and G. tandroka. The goal of this study is not a comprehensive reassessment of the phylogeny within the subgenus Duboimantis and molecular analyses were thus limited to a subset of taxa. In exploratory analyses, the new species clustered with G. salegy, G. schilfi, and G. tandroka, three species that formed a highly supported subclade of Duboimantis in the analyses of Wollenberg et al. (2011) and Kaffenberger et al. (2012). For our analysis we thus expanded the sampling for these three species with additional individuals. We found molecular evidence for new populations of G. schilfi and G. tandroka, both of which were so far known only from Marojejy. Both these species were now also found on the Sorata Massif north of Marojejy, and had different degrees of genetic differentiation. While the populations of G. schilfi (Sorata vs. Marojejy) differed by only 0.6��� 0.8% uncorrected 16S p-distance, those of G. tandroka had a higher divergence of 5.4% which suggests their taxonomic status requires revision and we therefore here flag the Sorata population as new Unconfirmed Candidate Species, Gephyromantis sp. Ca32, rationalised following Vieites et al. (2009). Another candidate species of Duboimantis (G. sp. Ca30 from the Bealanana district of northern Madagascar) that was also recently identified (Scherz et al. in press) was not included in our analysis here, but we have confirmed that it is not conspecific with G. sp. Ca32 (data not shown)., Published as part of Scherz, Mark D., Razafindraibe, Jary H., Rakotoarison, Andolalao, Dixit, Nadi M., Bletz, Molly C., Glaw, Frank & Vences, Miguel, 2017, Yet another small brown frog from high altitude on the Marojejy Massif, northeastern Madagascar (Anura: Mantellidae) in Zootaxa 4347 (3), DOI: 10.11646/zootaxa.4347.3.9, http://zenodo.org/record/1048679, {"references":["Blommers-Schlosser, R. M. A. (1979) Biosystematics of the Malagasy frogs. I. Mantellinae (Ranidae). Beaufortia, 352, 1 - 77.","Kaffenberger, N., Wollenberg, K. C., Kohler, J., Glaw, F., Vieites, D. R. & Vences, M. (2012) Molecular phylogeny and biogeography of Malagasy frogs of the genus Gephyromantis. Molecular Phylogenetics and Evolution, 62, 555 - 560. https: // doi. org / 10.1016 / j. ympev. 2011.09.023","Vieites, D. R., Wollenberg, K. C., Andreone, F., Kohler, J., Glaw, F. & Vences, M. (2009) Vast underestimation of Madagascar's biodiversity evidenced by an integrative amphibian inventory. Proceedings of the National Academy of Sciences of the USA, 106, 8267 - 8272. https: // doi. org / 10.1073 / pnas. 0810821106"]}