Corresponding author: Franco Andreone (
Academic editor: Natalia Ananjeva
We describe a new mantellid frog of the subfamily
The intense herpetological activity carried out in Madagascar during the last decades, together with the wider use of integrative taxonomic tools has led to the description of an astonishingly high number of new amphibians species (
Although, the highest species richness of amphibians is typically found along the eastern rainforest belt (
While the species already described from the arid West mostly belong to radiations of explosive breeders reproducing in ephemeral ponds, a special attention has been given to species ascribed to clades that are more typical of humid habitats and rainforest biomes. This was the case, for example, with the recent discovery of two new mantellines at the Isalo Massif (
During recent herpetofaunal inventories we discovered a further new species of a rather inconspicuous
Thirty-six described species are currently ascribed to
Due to morphological and external similarities, the new frog was formerly believed to be related to
Unfortunately, the secretive life of this new species prevented us from obtaining much biological informations and we still lack information about its acoustic repertoire, breeding behaviour and larval morphology. Notwithstanding these challenges, the ongoing collaborative effort generated by the ACSAM (A Conservation Strategy for the Amphibians of Madagascar,
We present here the formal description of this new
Schematic map of Madagascar with images and distribution of the four described species of the genus
The Tsingy de Bemaraha is a karstic plateau in the Melaky Region, five to 15 km wide and about 100 km long, located in western Madagascar. Numerous long, sharp pinnacles of rock, that may reach 45 meters in height, outcrop along the plateau and form the characteristic landscape (the so-called “tsingy” or “atsingy” in Malagasy language). Dry, deciduous forest is the most common vegetation type but humid areas occur within some of the larger canyons. An extensive area of forest and rock outcrop is included within two adjacent protected areas (Parc National Tsingy de Bemaraha and Réserve Naturelle Intégrale du Tsingy de Bemaraha). Savanna grasslands surround the plateau and there are numerous marshy depressions, caves and gorges associated with the main outcrop. This area has been object of some herpetological surveys that led to the discovery and description of several new species of amphibians (
We searched for frogs at night with the aid of hand torches and headlamps. Geographic coordinates were taken using a GPS device. Toponyms often follow the indications by local people, and must be therefore seen as largely unofficial names. Frogs were collected by hand and euthanised by immersion in chlorobutanol solution, fixed in 5% formalin or in 90% ethanol and finally stored in 75% ethanol solution. Voucher specimens (
Morphological measurements were made with a digital calliper to the nearest 0.1 mm. The following biometric measurements were taken (according to
A fingertip, or part of the muscle of the tongue, was cut from each collected individual and stored in 99% ethanol. Total genomic DNA was extracted from the tissue samples using proteinase K digestion (10 mg/ml concentration) following
The specific noun “
This species has been referred to as
MRSN A5487 (NFN), subadult male, collected at Tsingy de Bemahara National Park, western Madagascar, Andamozavaky (Bekopaka commune, Antsalova district, Melaky region, Mahajanga province), 19°01.86'S, 44°46.80'E; 122 m a.s.l., collected by J. E. Randrianirina on 23 May 2003.
MRSN A5486 (BMR 001), subadult male without evident femoral glands, MRSN A5484 (NFN), adult female, MRSN A5482 (BMR 008), MRSN A5483 (BMR 031), MRSN A5485 (BMR 002), three juveniles (sex unknown) sampled from the same locality, collector and date of the holotype (tissue sample taken for genetical analysis for all individuals); ZSM 23/2006 (FGZC 0715), adult female, from Grotte Crystal, close to Andranopasazy, Tsingy de Bemaraha National Park (Antsalova commune, Antsalova district, Melaky region, Mahajanga province), 18°42'31"S, 44°43'08"E, 146 m a.s.l., collected by F. Glaw, J. Köhler, P. Bora and H. Enting on 19 March 2006, fixed in ethanol (tissue sample taken for genetical analysis), individual found at night on limestone cliffs, close to the entrance of the cave; ZSM 37/2006 (FGZC 0746), juvenile (unknown sex) from Grotte Anjohimbazimba, Tsingy de Bemaraha National Park (Antsalova commune, Antsalova district, Melaky region, Mahajanga province), 18°41'34"S, 44°42'36"E, 160 m a.s.l., collected by F. Glaw, J. Köhler, P. Bora and H. Enting on 20 March 2006 (tissue sample taken for genetical analysis), individual found in the cave; ZSM 107/2006 (FGZC 0886), juvenile (sex unknown) from Bendrao Forest (“Camp 3”), Tsingy de Bemaraha National Park (Antsalova commune, Antsalova district, Melaky region, Mahajanga province), 18°47'04"S, 44°51'37"E, 427 m a.s.l., collected by F. Glaw, J. Köhler, P. Bora and H. Enting on 26–27 March 2006; (tissue sample taken for genetical analysis). All these specimens were fixed in 90% ethanol and preserved in 70% ethanol. UADBA 28112 (RBJ 708), female from Ranotsara (Bekopaka commune Antsalova district, Melaky region, Mahajanga province), 19°02'08"S, 44°46'29"E, 65 m a.s.l., collected by R. Andriantsimanarilafy on 18 November 2006; UADBA 28116 (RBJ 792), female from Ankilogoa (Bekopaka commune, Antsalova district, Melaky region, Mahajanga province), 19°07'52"S, 44°48'32"E, 57 m a.s.l., collected by R. Randrianavelona on 13 December 2006; UADBA 28120 (RBJ 791), female from Ankilogoa (Bekopaka commune, Antsalova district, Melaky region, Mahajanga province), 19°07'52"S, 44°48'32"E, 57 m a.s.l., collected by R. Randrianavelona on 13 December 2006; UADBA 28127 (RBJ 718), female from Ranotsara (Bekopaka commune, Antsalova district, Melaky region, Mahajanga province), 19°02'08"S, 44°46'29"E, 65 m a.s.l., collected by R. Randrianavelona on 19 November 2006; UADBA 39057 (RBJ 660), female from Anjaha (Antsalova commune, Antsalova district, Melaky region, Mahajanga province), 18°39'43"S, 44°49'33"E, 403 m a.s.l., collected by J.C. Randrianantoandro, R. Randrianavelona, R.K.B. Jenkins, R.R. Andriantsimanarilafy and E.F. Hantalalaina and Madagascar National Parks personnel on 15–24 February 2006; UADBA 39081 (RBJ 609), female from Andranopasazy (Melaky region, Mahajanga province), 18°42'31"S, 44°43'02"E, 146 m a.s.l. collected by J.C. Randrianantoandro, R. Randrianavelona, R.K.B. Jenkins, R.R. Andriantsimanarilafy and E.F. Hantalalaina and Madagascar National Parks personnel on 13–30 January 2006; UADBA 39082 (RBJ 630), female from Andranopasazy (Antsalova commune, Antsalova district, Melaky region, Mahajanga province), 18°42'31"S, 44°43'02"E, 146 m a.s.l. collected by J.C. Randrianantoandro, R. Randrianavelona, R.K.B. Jenkins, R.R. Andriantsimanarilafy and E.F. Hantalalaina and Madagascar National Parks personnel on 13–30 January 2006; UADBA 39099 (RBJ 627), adult male (with developed glands) from Andranopasazy (Antsalova commune, Antsalova district, Melaky region, Mahajanga province), 18°42'31"S, 44°43'02"E, 146 m a.s.l. collected by J.C. Randrianantoandro, R. Randrianavelona, R.K.B. Jenkins, R.R. Andriantsimanarilafy and E.F. Hantalalaina and Madagascar National Parks personnel on 13–30 January 2006; UADBA 39100 (RBJ 658), female from Anjaha (Antsalova commune, Antsalova district, Melaky region, Mahajanga province), 18°39'43"S, 44°49'33"E, 403 m a.s.l., collected by J.C. Randrianantoandro, R. Randrianavelona, R.K.B. Jenkins, R.R. Andriantsimanarilafy and E.F. Hantalalaina and Madagascar National Parks personnel on 15–24 February 2006.
Images of
A medium sized frog species (adult SVL 35–43 mm), assigned to the genus Morphometric measurements (in mm) of specimens of Catalogue number Field number Species Locality GenBank Rank SEX SVL HW HL ED END NSD NND TD HAL HIL FORL FOTL FOL IMTL IMTH FGL FGW NG GD MRSN A5487 NFN G. atsingy Andamozavaky HQ640419 HT M 34.8 14.1 14.5 5.7 3.8 2.6 3.6 3.3 11.6 37.5 18.6 24.8 17.7 1.1 1.1 - - 5 0.7 MRSN A5486 BMR 001 G. atsingy Andamozavaky HQ640421 PT M 31.3 12.4 13.5 5.3 3.9 1.9 3.1 2.8 10.6 33.1 16.2 23.6 15.5 1.7 1.2 - - - - MRSN A5484 NFN G. atsingy Andamozavaky HQ640418 PT F 43.4 15.9 17.3 6.4 4.5 3.1 3.9 3.6 13.3 43.2 20.1 30.9 20.4 2.2 1.0 - - - - ZSM 23/2006 FGZC 0715 G. atsingy Grotte Crystal, close to Andranopasazy HQ640414 PT F 38.5 14.3 15.2 6.1 4.3 2.2 3.6 2.7 11.7 39.9 18.0 26.0 16.7 1.6 0.9 - - - - MRSN A5482 BMR 008 G. atsingy Andamozavaky HQ640420 PT J 22.0 8.4 9.6 3.8 2.6 1.6 2.5 2.2 7.4 23.7 11.1 17.1 9.9 1.1 0.6 - - - - MRSN A5483 BMR 031 G. atsingy Andamozavaky HQ640417 PT J 19.6 7.3 8.3 3.5 2.2 1.1 2.1 1.7 6.3 17.9 9.4 13.3 8.1 0.4 0.2 - - - - MRSN A5485 BMR 002 G. atsingy Andamozavaky HQ640415 PT J 23.6 8.9 10.5 4.1 2.9 1.4 2.6 2.3 8.1 25.5 13.2 18.4 11.6 0.6 0.2 - - - - ZSM 107/2006 FGZC 0886 G. atsingy Bendrao Forest HQ640416 PT J 22.5 7.8 9.5 3.1 2.4 0.9 2.1 1.8 6.5 22.4 10.9 16.5 10.0 1.0 0.3 - - - - ZSM 37/2006 FGZC 0746 G. atsingy Grotte Anjohimbazimba - PT J 24.6 8.2 10.1 3.2 2.7 1.5 2.3 1.9 6.9 24.2 11.1 17.1 10.8 1.1 0.5 - - - - UADBA 39099 RBJ 627 G. atsingy Andranopasazy HQ640413 PT M 36.6 11.2 18.6 4.2 3.6 1.9 2.9 3.4 11.5 62.1 16.4 25.5 17.5 1.1 0.8 7.5 3.1 70 0.5 UADBA 39081 RBJ 609 G. atsingy Andranopasazy - PT F 38.4 14.8 16.8 4.3 3.6 1.9 3.4 3.3 11.9 63.5 17.4 27.3 19.3 1.4 0.8 - - - - UADBA 28120 RBJ 791 G. atsingy Ankilogoa - PT F 35.9 11.1 17.1 4.5 3.3 1.6 2.6 3.2 11.1 64.0 16.9 27.1 18.1 1.1 0.5 - - - - UADBA 39100 RBJ 658 G. atsingy Anjaha - PT F 35.1 12.1 17.8 4.4 3.9 1.7 2.7 3.3 11.0 62.2 17.3 26.8 17.3 1.3 0.7 - - - - UADBA 28127 RBJ 718 G. atsingy Ranotsara - PT F 39.1 12.7 17.9 4.6 3.7 1.6 3.2 3.3 12.1 64.0 18.4 28.4 18.7 1.4 0.6 - - - - UADBA 39082 RBJ 630 G. atsingy Andranopasazy - PT F 38.1 12.6 16.9 3.9 4.1 1.6 2.9 2.9 11.6 63.9 17.8 27.4 17.6 1.5 0.5 - - - - UADBA 28112 RBJ 708 G. atsingy Ranotsara - PT F 38.4 11.8 16.8 4.2 4.0 1.1 2.5 3.0 11.4 61.9 18 26.3 16.4 1.2 0.5 - - - - UADBA 39057 RBJ 660 G. atsingy Anjaha - PT F 35.3 11.4 17.4 4.3 3.6 1.4 2.9 2.8 11.6 63.9 16.6 27.5 18.4 0.9 0.5 - - - - UADBA 28116 RBJ 792 G. atsingy Ankilogoa - PT F 33.9 11.0 16.3 3.4 3.7 1.2 2.2 2.5 11.0 58.6 16.4 25.0 15.9 1.0 0.5 - - - - MRSN A5373 FAZC 12859 G. corvus Isalo, Tsiombivositra HQ640423 - M 39.8 16.1 15.3 5.8 4.8 2.9 4.3 3.7 11.8 38.7 16.3 28.6 19.6 2.4 1.4 9.1 4.1 96 0.6 MRSN A5325 FAZC 13000 G. corvus Isalo, Ambovo - - F 40.0 15.2 16.2 6.4 4.3 2.6 4.1 3.2 10.7 40.3 18.2 30.1 20.0 1.6 0.6 - - - - MRSN A5323 FAZC 12661 G. corvus Isalo, Malaso HQ640422 - F 39.0 15.0 15.7 6.1 4.4 2.8 4.2 3.6 11.9 39.3 17.6 29.7 20.3 1.6 0.8 - - - - MRSN A5324 FAZC 12758 G. corvus Isalo, Zahavola HQ640424 - F 40.1 15.0 16.3 6.1 4.4 2.8 4.2 3.3 10.8 37.2 16.9 28.0 19.8 1.3 0.7 - - - - MRSN A2786 FAZC 11964 G. corvus Isalo, Andranomena HQ640425 - F 40.8 15.0 15.8 6.3 4.7 2.7 4.3 3.6 11.1 40.5 17.7 30.2 20.4 1.9 0.7 - - - - MRSN A5310 FAZC 12568 G. azzurrae Isalo, Andriamanero EF222301 HT M 41.1 16.9 13.4 6.1 3.9 2.5 4.0 4.0 12.1 41.1 20.0 30.0 18.8 2.0 1.1 6.3 2.0 45 0.5 MRSN A5309 FAZC 12567 G. azzurrae Isalo, Andriamanero EF222300 PT M 38.5 15.3 12.8 5.2 4.3 2.2 3.7 3.7 11.1 41.1 19.9 26.7 18.9 1.9 1.3 6.5 2.7 38 0.5 MRSN A5311 FAZC 12569 G. azzurrae Isalo, Andriamanero EF222302 PT M 40.2 15.8 14.1 6.0 4.0 2.7 4.0 4.1 11.2 41.0 19.9 27.7 19.9 2.1 1.1 6.7 2.7 40 0.6 MRSN A5312 FAZC 12910 G. azzurrae Isalo, Iambahatsy EF222304 PT M 23.3 8.8 8.8 4.1 2.8 1.4 2.2 2.5 8.8 24.5 11.1 17.7 12.1 1.1 0.5 _ _ _ _ SMF 85859 NFN G. azzurrae Isalo, Sakamalio EF222305 PT M 42.7 16.4 14.3 5.4 3.7 2.6 3.9 3.5 13.4 41.1 21.0 29.9 20.0 1.9 1.1 7.0 3.0 42 0.6 SMF 85860 NFN G. azzurrae Isalo, Sakamalio EF222303 PT M 43.7 16.4 13.5 5.7 4.0 2.5 3.8 4.0 12.2 42.3 21.1 27.7 20.0 1.9 1.0 7.5 2.7 42 0.5 MRSN A3415 NFN G. pseudoasper Nosy Be - - M 33.3 12.1 13 4.9 3.8 2.6 2.8 3.1 10.4 52.4 15.7 23 17.5 2.1 1.1 6.1 2.5 43 0.3 MRSN A3416 FN 6696 G. pseudoasper Nosy Be - - M 37.4 12.9 15 4.9 4.4 3 3.8 4.4 10.3 54.2 16.4 25 15.8 2.4 1.1 7.2 2.9 39 0.4 MRSN A3417 FN 6423 G. pseudoasper Nosy Be - - F 33.1 11.9 14 5.3 3.7 2.7 2.9 3 9.9 56.5 16.5 24 18.2 1.9 1 - - - -
Subadult male in mediocre state of preservation, with the belly opened for gonadal inspection and part of the ventral surface of thighs cut and opened to check the glands. SVL 34.8 mm; for other measurements see
We based the current description of variability upon some specimens (paratypes and complementary individuals), part of which (ZSM 23/2006, 37/2006, 107/2006, MRSN A5486 and MRSN A5483) were also photographed in nature, and thus provided more diagnostic characters. The female ZSM 23/2006 (
According to our observations, the species lives in habitats that retain some humidity, such as rock cavities and along the walls of the canyon-like formations. One important notation comes from the fact that several of the collectors, independently (JER, FG, JCR) found this species within the caves which are typical of the area. We suspect that the species uses caves because these sites presumably have a higher humidity than the surrounding areas. In such a sense it behaves similarly to
Only known from the localities of the type specimens within the Tsingy de Bemaraha National Park.
All the described species of Distribution, habitats and diagnostic characters of the nominal species in the genus Species Distribution Habitat SVL Vocal Sacs Dorsal colouration Ventral colouration Dorsal texture Gephyromantis atsingy Tsingy de Bemaraha Karst pinnacles and caves 35–43 mm Non-evident Light brownish with greenish shadings Whitish Slightly warty Gephyromantis azzurrae Isalo Massif Open canyons and permanent rivers 23–44 mm Double and brownish Brownish, sometimes with wide light band Whitish, with reddish shadings Warty with heterogeneous warts Gephyromantis corvus Isalo Massif Close canyons and cave-like canyons 39–41 mm Double and blackish Greyish with darker spots Whitish Warty Gephyromantis pseudoasper Sambirano, N, NW and NE Rainforests, karstic areas 33–37 mm Double and blackish Brownish Whitish Slightly warty
The molecular data confirm the attribution of
This species appears to be restricted to the Bemaraha Plateau, where it has been found in seven localities within the Tsingy de Bemaraha National Park. It may also occur in the Réserve Naturelle Intégrale, which forms the northerly limit of the Bemaraha Plateau, but survey data are lacking from this site. Within the national park, some areas of forest are damaged by conversion to agriculture and charcoal production, but the humid canyons where G. atsingy occur are generally well protected. We therefore recommend assigning a category of Near Threatened because the species nearly qualifies for listing as Vulnerable under D2: the species is confined to a single site, the Bemaraha Massif (1,577 km2), with a plausible threat that could impact the species in the near future. If the threat became operational, the species would be eligible for listing as Endangered since its extent of occurrence is well within the 5,000km2 threshold under the B criterion and it would occur at a single location (where the threat is habitat loss from agricultural activities and charcoal production) and there would be a continuing decline in the quality and area of habitat, qualifying the species for the criteria B1ab(iii).
The Bemaraha plateau is one of the most peculiar areas of western Madagascar in terms of amphibian richness and endemicity (
One powerful tool is the application of an integrated taxonomy approach, which includes aspects of direct field surveys, behavioural assessment, molecular screening and bioacustic analysis. This is what allowed
As already stressed by
We expect that other forest blocks in western Madagascar may host further undescribed species of
The accelerated species discovery in Malagasy amphibians points to the importance of taxonomic surveys (see www.sahonagasy.org), and we like to consider
Many friends and colleagues helped us providing samples that have been used in this study. Thanks to S. Federici and M. Barbuto for their help in the lab. Special thanks to M. Vences, P. Bora, J. Köhler and H. Enting for the help in the field and for providing information on the herpetofauna of Bemaraha. We thank N. Kaffenberger for sharing unpublished information on molecular relationships, and A. Angulo and M. Hoffmann for their precious help on analysing the conservation status of this newly described species. This work has been carried out in the framework of collaboration agreements of the authors’ institutions with UADBA. The Malagasy authorities kindly granted research and export permits. A. Crottini was supported by a Dote Ricercatore of Regione Lombardia fellowship. Fieldwork by RKBJ and JCR was funded by the Darwin Initiative.