Leaping towards a saltatorial lifestyle? An unusually long-legged new species of Rhombophryne (Anura, Microhylidae) from the Sorata massif in northern Madagascar

The Madagascar-endemic microhylid genus Rhombophryne consists of a range of partly or completely fossorial frog species. They lead a poorly known, secretive lifestyle, and may be more diverse than previously thought. We describe a new species from the high altitude forests of the Sorata massif in north Madagascar with unusual characteristics for this genus; R. longicrus sp. n. has long, slender legs, unlike most of its fossorial or semi-fossorial congeners. The new species is closely related to R. minuta, a much smaller frog from the Marojejy massif to the southeast of Sorata with similarly long legs. We discuss the morphology of these species relative to the rest of the genus, and argue that it suggests adaptation away from burrowing and toward a more saltatorial locomotion and an accordingly more terrestrial lifestyle. If this is the case, then these frogs represent yet more ecological diversity within the already diverse Cophylinae. We recommend an IUCN Red List status of Endangered B1ab(iii) for R. longicrus sp. n., because it is known only from a single site in a forested area of roughly 250 km2, which is not yet incorporated into any protected area.Field research was conducted under permit No. 265/12/MEF/SG/DGF/DCB.SAP/SCB (dated 18 Oct. 2012), exportation of specimens under permit No. 163N-EA12/MG12 (dated 17 Dec. 2012), (both issued by the Direction Générale des Forêts de Madagascar), and funded by the Mohamed bin Zayed Species Conservation Fund (project 11253064) and BIOPAT.Peer reviewe

Flavonoïde d’Abrahamia Suarezensis Randrian. & Lowry (Anacardiaceae)

RésuméDans le cadre de la valorisation de la flore malagasy, ce travail vise à rechercher par bioguidage des composés biologiquement actifs présents dans les rameaux feuillés d'Abrahamia suarezensis (Anacardiaceae) endémique de Madagascar. Le screening phytochimique a révélé la présence dans la plante des stéroïdes lactoniques, stérols insaturés, flavonoïdes du type flavonol, et de tanins condensés ce qui a conduit à un partage liquide-liquide de la solution aqueuse de l’extrait éthanolique successivement avec hexane, acétate d’éthyle et butanol. Le criblage biologique des extraits acétate d’éthyle (ESAc) et butanolique (ESBu) ont permis de mettre en évidence des activités antihistaminique et antioxydante intéressantes de la plante. Les extraits ESAc (CI50= 6,05 µg.ml-1) et ESBu (CI50 = 7,19 µg.ml-1) ont montré une activité antioxydante plus forte que celle du témoin positif, la vitamine E (CI50 = 11,39 µg.ml-1). L’évaluation de la toxicité aigüe de l’ESAc sur les souris Swiss n’a provoqué aucun décès jusqu’à la dose de 5000 mg.kg-1. Une diminution de l’activité motrice a été constatée à partir de 500 mg.kg-1. Le processus de fractionnement bioguidé de l’extrait ESAc, le plus actif, a permis l’obtention de la fraction E9 (100% d’inhibition de la contraction en test antihistaminique, CI50 = 3,32 µg.ml-1 en test antioxydant) dont sa purification a conduit à l’isolement du composé F4.La détermination de structure des composés utilisant une combinaison des expériences RMN 1D et 2D incluant COSY, HSQC, HMBC et spectrométrie de masse, a permis d’identifier le kaempférol-3-O-α-rhamnopyranoside et de déceler par ESI+ la présence d’un kaempéferol-O-pentoside.

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

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

A distinctive new frog species (Anura, Mantellidae) supports the biogeographic linkage of two montane rainforest massifs in northern Madagascar

We describe a new species of the genus Gephyromantis, subgenus Vatomantis (Mantellidae, Mantellinae), from moderately high elevation (1164–1394 m a.s.l.) on the Marojejy, Sorata, and Andravory Massifs in northern Madagascar. The new species, Gephyromantis (Vatomantis) lomorina sp. n. is highly distinct from all other species, and was immediately recognisable as an undescribed taxon upon its discovery. It is characterised by a granular, mottled black and green skin, reddish eyes, paired subgular vocal sacs of partly white colour, bulbous femoral glands present only in males and consisting of three large granules, white ventral spotting, and a unique, amplitude-modulated advertisement call consisting of a series of 24–29 rapid, quiet notes at a dominant frequency of 5124–5512 Hz. Genetically the species is also strongly distinct from its congeners, with uncorrected pairwise distances ≥10 % in a fragment of the mitochondrial 16S rRNA gene to all other nominal Gephyromantis species. A molecular phylogeny based on 16S sequences places it in a clade with species of the subgenera Laurentomantis and Vatomantis, and we assign it to the latter subgenus based on its morphological resemblance to members of Vatomantis. We discuss the biogeography of reptiles and amphibians across the massifs of northern Madagascar, the evidence for a strong link between Marojejy and Sorata, and the role of elevation in determining community sharing across this landscape

A new limestone-dwelling species of Micryletta (Amphibia: Anura: Microhylidae) from northern Vietnam

We report on a new species of the genus Micryletta from limestone karst areas in northern Vietnam, which is described on the basis of molecular and morphological evidence. Micryletta nigromaculata sp. nov. is restricted to narrow areas of subtropical forests covering karst massifs in Cat Ba National Park (Hai Phong Province) and Cuc Phuong National Park (Ninh Binh Province) at elevations of 90–150 m a.s.l. In the phylogenetic analyses, the new species is unambiguously positioned as a sister lineage to all remaining species of Micryletta. We also discuss genealogical relationships and taxonomic problems within the genus Micryletta, provide molecular evidence for the validity of M. erythropoda and discuss the taxonomic status of M. steinegeri. We suggest the new species should be considered as Endangered (B1ab(iii), EN) following the IUCN’s Red List categories. A discussion on herpetofaunal diversity and conservation in threatened limestone karst massifs in Southeast Asia is provided

Rhombophryne diadema Scherz, Hawlitschek, Andreone, Rakotoarison, Vences & Glaw, 2017, sp. nov.

<i>Rhombophryne diadema</i> sp. nov. <p>Suggested common name: Diadem saw-browed diamond frog (Figs 4, 5, 11, 12, S8)</p> <p> <b>Holotype.</b> ZSM 1629 /2012 (FGZC 3604), adult female, collected between the 26th and 30th of November by F. Glaw, O. Hawlitschek, T. Rajoafiarison, A. Rakotoarison, F.M. Ratsoavina, and A. Razafimanantsoa on the Sorata Massif at 13.6817°S, 49.4411°E, at 1339 m a.s.l., in the Sava Region, northeastern Madagascar (Figs 4, 5, S8).</p> <p> <b>Paratypes.</b> ZSM 1628 /2012 (FGZC 3731), adult male, and UADBA-A 60289 (FGZC 3611), adult female with large eggs, same data as holotype except collected at a creek above the campsite in Sorata (13.6780°S, 49.4404°E) at 1407 m a.s.l.</p> <p> <b>Diagnosis and comparisons.</b> A species assigned to the genus <i>Rhombophryne</i> on the basis of molecular phylogenetic affinities (Fig. 3), and the possession of a clavicle combined with the absence of T- or Y-shaped terminal phalanges (vs. either absence of a clavicle or possession of a clavicle combined with T- or Y-shaped terminal phalanges in the morphologically similar <i>Plethodontohyla</i>). Within the genus <i>Rhombophryne</i>, it is assigned to the <i>R. serratopalpebrosa</i> group on the basis of possessing superciliary spines and molecular phylogenetic data (Fig. 3).</p> <p> <i>Rhombophryne diadema</i> <b>sp. nov.</b> is distinguished from all congeners by the following unique suite of characters: SVL 22.7–23.4 mm; tympanum indistinct, TDH/ED = 0.59–0.64); weak supratympanic fold extending from the rear corner of the eye over and behind tympanum toward the axilla; three superciliary spines, the posterior-most considerably smaller than the anterior two; unreduced fingers, second finger distinctly shorter than fourth; tibiotarsal articulation reaching eye; TIBL/SVL = 0.44–0.46; and fifth toe distinctly shorter than third. Osteologically, it is characterised by an anteriorly broadening parasphenoid cultriform process, prechoanal portion of vomer non-radiate, postchoanal portion straight, broad quadratojugal-squamosal contact, stepped anterior edge of ventral ramus of squamosal, short humeral crista ventralis, weak dorsal prominence on iliac shafts, and partially ossified pubis. Additionally, <i>R. diadema</i> is separated from all other <i>Rhombophryne</i> species for which molecular data are available by uncorrected pairwise distances of at least 5.1% in a segment of the <i>16S</i> rRNA mitochondrial gene (Table 1).</p> <p> Within the genus <i>Rhombophryne</i>, <i>R. diadema</i> <b>sp. nov.</b> differs from all species except members of the <i>R. serratopalpebrosa</i> group by the possession of superciliary spines. Within the <i>R. serratopalpebrosa</i> group, it differs from <i>R. serratopalpebrosa</i> by smaller size (SVL 22.7–23.4 vs. 28.5 mm), smaller relative tympanum size (TDH/ED = 0.59–0.64 vs. 0.78), a weak (vs. strong) supratympanic fold, three superciliary spines (vs. four), shorter relative forelimb length (FORL/SVL 0.59 vs. 0.71), and shorter relative hindlimb length (HIL/SVL = 1.66 vs. 1.77); from <i>R. vaventy</i> by much smaller size (SVL 22.7–23.4 vs. 51.9 mm), larger relative tympanum size (TDH/ED = 0.59–0.64 vs. 0.46), narrower head (HW/HL = 1.46–1.59 vs. 1.70), a weak (vs. distinct) supratympanic fold (see Fig. 5), three (vs. four) superciliary spines, tibiotarsal articulation reaching eye (vs. beyond snout tip), shorter relative forelimb length (FORL/SVL = 0.59 vs. 0.76), smaller relative tibia size (TIBL/SVL = 0.44–0.46 vs. 0.53), and smaller inner metacarpal tubercle size (IMCL/HAL 0.13–0.14vs. 0.19); from <i>R. guentherpetersi</i> by smaller size (SVL 22.7–23.4 vs. 27.5–35.1 mm), three superciliary spines, the anterior two of which are of medium size, the posterior-most of which is considerably smaller (vs. two to three small superciliary spines), broader head (HW/HL = 1.46–1.59 vs. 1.34–1.41), tibiotarsal articulation reaching the eye (vs. reaching the insertion of the arms), longer relative tibia length (TIBL/SVL 0.44–0.46 vs. 0.32–0.36), partially ossified pubis (vs. unossified), and broader pectoral girdle (compare Fig. 7 b with Fig. 12 b); from <i>R. ornata</i> by smaller size (SVL 22.4–23.4 mm vs. 33.0 mm), a weak (vs. distinct) supratympanic fold, three superciliary spines (vs. two), longer relative hindlimb length (HIL/SVL = 1.66 vs. 1.45–1.63), smaller relative inner metacarpal tubercle length (IMCL/HAL = 0.13–0.14 vs. 0.15–0.19), absence of reddish colour on the hidden portions of the legs (vs. presence), and ossified carpals and limb bone epiphyses (vs. unossified); from <i>R. tany</i>, which it most strongly resembles, by its weaker supratympanic fold (see Fig. 5 and compare Figs 1 d and 11), three superciliary spines (vs. two), slightly shorter forelimbs (FORL/SVL 0.59 vs. 0.63), slightly longer relative tibia length (TIBL/SVL 0.44–0.46 vs. 0.43), prootics in contact with parasphenoid alae (vs. not in contact), parasphenoid cultriform process broadening anteriorly (vs. having parallel edges), nasals not anterolaterally displaced (vs. displaced), quadratojugal-squamosal contact broad (vs. narrow), anterior edge of ventral ramus of squamosal distinctly stepped (vs. weakly stepped), dorsal prominence of iliac shafts weak (vs.</p> <p> strong), and partially ossified pubis (vs. unossified); from <i>R. regalis</i> by anterior-most superciliary spine sitting atop eye (vs. anterior to eye; see Fig. 5 and compare Figs 9 and 11), slightly shorter relative forelimb length (FORL/SVL = 0.59 vs. 0.59–0.70), shorter relative tibia length (TIBL/SVL 0.44–0.46 vs. 0.47–0.56), tibiotarsal articulation reaching the eye (vs. reaching the snout tip or beyond), exoccipitals ventromedially not in contact (vs. in contact), anterior edge of ventral ramus of squamosal stepped (vs. smoothly sigmoidal), dorsal prominence of iliac shafts weak (vs. strong), and partially ossified pubis (vs. ossified); and from <i>R. coronata,</i> which it also closely resembles, by much smaller size of the posterior-most superciliary spine (vs. three roughly equal-sized superciliary spines), larger relative tympanum size (TDH/ED 0.59–0.64 vs. 0.37–0.59), longer relative tibia length (TIBL/SVL 0.44–0.46 vs. 0.35–0.39), first toe unreduced (vs. sometimes reduced to a short nub; see Fig. 5), anterior edge of squamosal ventral ramus stepped (vs. straight), and sphenethmoids not exceeding postchoanal vomers (vs. exceeding postchoanal vomers).</p> <p> <b>Description of the holotype.</b> (Figs 4, 5) An adult female specimen in a very good state of preservation. Tissue taken from the left thigh for genetic sequencing. A transverse incision present in the posterior abdomen.</p> <p>Body robust. Head wider than long (HW/HL = 1.59). Pupils round. Snout rounded in dorsal and lateral views. Canthus rostralis concave. Loreal region slightly concave. Nostril nearer to snout tip of than to eye (END/NSD = 0.47), directed laterally, slightly protuberant. Internarial distance greater than distance from eye to nostril. Tympanum indistinct, TDH/ED = 0.64. Supratympanic fold weak, extending from the middle back of the eye over and behind the tympanum toward the axilla. A small granular bump is present posteroventral to both tympana. Three superciliary spines above each eye, the anterior two roughly equal in size, the posterior-most almost imperceptible without magnification. Vomerine teeth present, in straight rows either side of the palate, separated medially by a small gap. Tongue broad and unlobed, attached anteriorly.</p> <p>Arms fairly slender. Fingers not reduced. Fingers without webbing; relative lengths 1<2<4<3; fourth finger distinctly longer than second. Finger tips not enlarged. Nuptial pads absent; inner metacarpal tubercle present, outer metacarpal tubercle absent, subarticular tubercles weak. Hindlimbs strongly built. Tibiotarsal articulation reaches the eye; TIBL/SVL = 0.46. Inner metatarsal tubercle present; outer metatarsal tubercle absent. Toes unwebbed; relative lengths 1<2<5<3<4; fifth toe distinctly shorter than third. Dorsal skin granular, rugose in life. Dorsolateral folds absent.</p> <p>Colouration of the holotype: After two and a half years in preservative, dorsal body colour is dark brown flecked with black—in life, the body was a more reddish brown (Fig. 11 a). The posterior half is lighter in colour than the anterior, particularly around the midline. Two light-brown lines run posteromedially from the eyes, approaching one another toward the midline but ending in the suprascapular region. These lines merge anterolaterally at the posterior of the eye with the lighter flank colouration, extending onto the lateral portions of the head. A whitish line runs from the posterior edge of the eye to the corner of the mouth. The arms are dorsally light brown flecked with black. The legs are dorsally light brown, with one dark crossband on the thigh, two on the shank—the proximal of which is less distinct than the distal—one on the tarsus, one on the metatarsals, and some black spots on the toes.</p> <p>The ventral surface is cream, with a little brown speckling on the chin and below the insertions of the arms. The legs are ventrally flecked with light brown and cream. The iris is gold with black reticulations and a black periphery.</p> <p> <b>Variation.</b> Morphologically, ZSM 1628/2012 agrees strongly with the holotype. Its tympanum is slightly smaller (TDH/ED = 0.59), and its tibia shorter (TIBL/SVL = 0.44). Its colouration is starkly different, however. Dorsally, it is uniformly an earthy brown. Crossbands on the shanks are faint but present, as is the light line running between the corner of the mouth and the posterior of the eye. Based on colour photographs (Fig. 11), UADBA-A 60289, which is female, also closely agrees in morphology with the rest of the type material. Its colouration is roughly intermediate between the other specimens, its dorsum being muddy brown flecked with white and black spots; a pair of small white spots being present between the anterior edges of the eyes, and again between the posterior edges, as well as over the suprascapular region. Its inguinal region is a yellowish cream. The crossbands on its legs are as in the holotype.</p> <p> <b>Etymology.</b> The specific epithet <i>diadema</i> is the latinized Greek word for diadem, a small crown typically worn by female royalty. It refers to the superciliary spines borne by this species. It is a feminine nominative singular noun in apposition.</p> <p> <b>Natural history.</b> Individuals of this species were captured when active during the day jumping among the leaflitter or near pitfall traps, suggesting a terrestrial, possibly partially fossorial lifestyle. The holotype contained at least 13 well-developed, yellow eggs (diameter 2.45 ± 0.25 mm), suggesting that the species was reproductively active at the end of November, around the start of the rainy season.</p> <p>The montane rainforest of Sorata is under high human-disturbance pressure, especially due to the high number of zebu cattle, which are responsible for widespread forest disturbance in the area. The area where the specimens of this species were discovered was exceptionally intact, with dense leaf litter.</p> <p> <b>Distribution and conservation status.</b> This species is known only from high altitudes (1339–1407 m a.s.l.) in the forest of the Sorata Massif in northern Madagascar (Fig. 8). This forest is unprotected and therefore threatened by deforestation and degradation without restriction. Additionally, species at high altitude may be threatened by climate change (Raxworthy 2008; Raxworthy <i>et al.</i> 2008), although this threat is most likely less imminent than that of deforestation. If this species is restricted to the Sorata Massif, then its extent of occurrence and optimistically estimated area of occupancy constitutes an area of only ~ 250 km 2 (calculated in Google Earth® Pro 6.1.0.500, Google Inc., Mountain View, CA). Due to its likely restriction to a small area of unprotected forest that is under threat from deforestation and possible long-term threat of climate change, <i>R. diadema</i> <b>sp. nov.</b> qualifies as Endangered under the IUCN Red List Criteria (2012) B1ab(iii) as defined for <i>R. guentherpetersi</i> above, similar to <i>R. longicrus</i>, which was described from the same area (Scherz <i>et al.</i> 2015b).</p>Published as part of <i>Scherz, Mark D., Hawlitschek, Oliver, Andreone, Franco, Rakotoarison, Andolalao, Vences, Miguel & Glaw, Frank, 2017, A review of the taxonomy and osteology of the Rhombophryne serratopalpebrosa species group (Anura: Microhylidae) from Madagascar, with comments on the value of volume rendering of micro-CT data to taxonomists, pp. 301-340 in Zootaxa 4273 (3)</i> on pages 318-321, DOI: 10.11646/zootaxa.4273.3.1, <a href="http://zenodo.org/record/803049">http://zenodo.org/record/803049</a&gt

Cophyla puellarum Rakotoarison, Crottini, Müller, Rödel, Glaw & Vences, 2015, sp. nov.

<i>Cophyla puellarum</i> sp. nov. <p>(Figs. 7 c, 12c & 16)</p> <p> <b>Remark.</b> This species (ZFMK 57465) has been figured in Glaw & Vences (1994: Fig. 361) as <i>Platypelis</i> sp. and in Glaw & Vences (2007: 135, Fig. 5 d) as unidentified <i>Platypelis</i> from Montagne d’Ambre. Referred to as <i>Cophyla</i> sp. Ca3 by Perl <i>et al</i>. (2014).</p> <p> <b>Holotype.</b> ZSM 3249/2012 (ZCMV 13521), adult male, collected at Point de Vue du Grand Moulin in Montagne d’Ambre National Park, -12.54077, 49.15424, 1246 m a.s.l., on 25 January 2012, by A. Rakotoarison and A. Razafimanantsoa.</p> <p> <b>Paratypes.</b> ZSM 0782/2009 (ZCMV 13005), male, and ZSM 0781/2009 (ZCMV 13004) juvenile, both collected at Lac Maudit, Montagne d’Ambre National Park, -12.5856, 49.1457, 1306 m, on 19 and 20 November 2009 by A. Crottini, S. J. Hauswaldt, A. Lima, E. Rajeriarison, F. M. Ratsoavina and A. Razafimanantsoa; UADBA-A 60237 (FGZC 4903), calling male, ZSM 3274/2012 (FGZC 4904), calling male, ZSM 3275/2012 (FGZC 4905), gravid female, UADBA-A 60238 (FGZC 4906) male, all four collected at Point de Vue du Grand Moulin in Montagne d’Ambre National Park, 12.54077, 49.15424, 1246 m, on 10 December 2012, by A. Rakotoarison and A. Razafimanantsoa; ZFMK 57465, adult female (SVL 33.3 mm), collected at high altitude of Montagne d’Ambre National Park (no coordinates available) on 18–21 March 1994 by F. Glaw, N. Rabibisoa & O. Ramilison.</p> <p> <b>Diagnosis and comparisons.</b> Assigned to the genus <i>Cophyla</i> in the microhylid subfamily Cophylinae based on enlarged terminal discs on fingers and toes, posterior vomer undivided and not overlapping with neopalatines, absence of nuptial pads and molecular phylogenetic relationships. Among other arboreal species of cophylines the species can be identified by combination of the following character states: body size comparatively large for <i>Cophyla</i> (SVL 27.3–33.6 mm); most of the specimens possess a uniformly yellow-greenish ventral colour; fifth toe slightly longer than third; males with prepollical tubercule but lacking a finger-like prepollex as typical for <i>Anodonthyla</i>; posterior vomer with well developed and clearly recognizable vomerine teeth; clavicle present.</p> <p> Distinguished from all other species of <i>Cophyla</i> by a unique advertisement call, characterized by a combination of short calls with long inter-call intervals; (Table 2). Further distinguished from <i>C. phyllodactyla</i>, <i>C. maharipeo</i> and <i>C. noromalalae</i> by the presence of a clavicle (no osteological data available for <i>C. berara</i> and <i>C. occultans</i>). Further distinguished from <i>C</i>. <i>occultans</i> and <i>C</i>. <i>berara</i> by larger body size (adult male SVL 27–29 mm <i>vs</i>. respectively 16–21 mm and 23–26 mm).</p> <p> <b>Description of the holotype.</b> Adult male in good state of preservation, some muscle tissue removed from right thigh. Snout-vent length 27.3 mm; body slender; head wider than long, not wider than body; snout rounded dorsally and laterally; nostrils not protuberant, nearer to tip of snout than to eye; canthus rostralis almost indistinct, loreal region very slightly concave; tympanum slightly distinct, 46% of eye diameter; supratympanic fold running from posterior edge of eye to forearm; tongue broadly rounded, without papilla, not bifid or notched; weakly developed maxillary teeth present; vomerine teeth not visible, but palpable. Forelimbs slender; subarticular tubercles on all fingers recognizable (single and rounded); outer metacarpal tubercle relatively large and flat; inner metacarpal tubercle oblong and large, forming distinct protuberance at base of first finger; hand without webbing; fingers distinctly flattened and broad along entire length; relative length of fingers 1<2=4<3, fourth finger slightly longer than second; finger discs broadly rounded to slightly bilobate, with lateral fringes; nuptial pads absent. Hindlimbs slender; tibiotarsal articulation reaching between forelimb and tympanum when hindlimb adpressed along body; tibia length, 42% of SVL; inner metatarsal tubercle oblong; outer metatarsal tubercle absent; webbing between toes very weakly developed, with traces of webbing between toes 3–4 and 4–5; subarticular tubercles on toes hardly recognizable; toes flattened and their discs relatively broad, broadly rounded to slightly bilobate, with lateral fringes, smaller than discs of fingers; relative length of toes 1<2<3<5<4 left toes and 1<2<3=5<4 right toes; fifth toe slightly (left) to distinctly (right) longer than third. Dorsal skin smooth, without dorsolateral folds. Ventral skin smooth on throat and chest and heavily granular on belly.</p> <p> <b>Colour of holotype in preservation.</b> After several years in 70% ethanol (Fig. 7 c), dorsum brownish with irregular markings, including a darker W-shaped patch extending over the posterior dorsum. Forelimbs and hindlimbs dorsally lighter with dark crossbands. Ventrally, uniformly beige, with some darker colour on venter caused by inner organs shining through the ventral skin. Some dark colour along lower jaws.</p> <p> <b>Colour in life.</b> Dorsum brownish with yellowish patches that have a greenish shade (Fig. 16 b). Ventrally uniformly whitish (Fig. 12 c).</p> <p> <b>Variation.</b> Most of the paratypes are morphologically similar to the holotype (Table 3) except the females ZSM 3275/2012 (FGZC 4905) and ZFMK 57465 which are larger than the males. The third toe is slightly shorter than the fifth toe in all specimens. The female is larger than the males (Table 3). There is large variation of coloration between the specimens. In life, dorsum is beige with yellow patches (Fig. 16 a), or beige and greenish with yellow patches (Fig.16 b), or brown with yellow patches (Fig. 16 c), or yellow with brown patches (Fig. 16 e) for adult and subadult specimens. The paler color pattern probably reflects coloration at night whereas the more contrasted pattern is probably typical for specimens during the day, but our fieldnotes do not allow to confirm unambiguously this hypothesized color change. Dorsum is dark brown with yellow patches and yellow vertebral line in a juvenile specimen (Fig. 16 d).</p> <p> <b>Etymology.</b> The name is derived from Latin puellarum: girls, genitive plural of puella. This species is dedicated to the "girls team" of a joint TU Braunschweig and Université d'Antananarivo expedition who discovered the first specimen in 2009 at Lac Maudit.</p> <p> <b>Natural history.</b> The holotype was found during the day with two metamorphosing tadpoles in a tree hole in highland forest. An ovigerous female (ZSM 3275/2012) contained 21 eggs. The recorded male (UADBA-A 60237) was found on a tree trunk in December 2012 in the same area as the holotype. A second calling male (ZSM 3274/ 2012) was found on a <i>Pandanus</i> leaf. More specimens were regularly observed calling. They were typically heard from low perches of ca. 100–150 cm on leaves, but in a few cases also from higher perches of above 2 m from the ground.</p> <p> <b>Advertisement call.</b> Recorded from paratype UADBA-A 60237 (FGZC 4903), at Point de Vue du Grand Moulin in MANP, on 10 December 2012, at an air temperature of 16.6˚C (Fig. 17). The following call description refers to the calls of this single specimen. As with all other <i>Cophyla</i> and most cophylines, single tonal calls were repeated in long series (continued at least for several minutes but probably much longer in undisturbed specimens). The dominant spectral call frequency ranged from 2300–2400 Hz (N= 47). Temporal call parameters were as follows: call duration 326–390 ms (363±13 ms; N= 47); duration of inter-call intervals 1961–3996 ms (2640±594 ms; N= 46). As in <i>C. maharipeo</i>, each call in the recorded sequence starts with a very short intensity peak, and we hypothesize this represents an artefact of the recording equipment or an abnormal call feature (see call description of <i>C. maharipeo</i>).</p>Published as part of <i>Rakotoarison, Andolalao, Crottini, Angelica, Müller, Johannes, Rödel, Mark-Oliver, Glaw, Frank & Vences, Miguel, 2015, Revision and phylogeny of narrow-mouthed treefrogs (Cophyla) from northern Madagascar: integration of molecular, osteological, and bioacoustic data reveals three new species, pp. 61-89 in Zootaxa 3937 (1)</i> on pages 78-83, DOI: 10.11646/zootaxa.3937.1.3, <a href="http://zenodo.org/record/288186">http://zenodo.org/record/288186</a&gt

Cophyla maharipeo Rakotoarison, Crottini, Müller, Rödel, Glaw & Vences, 2015, sp. nov.

<i>Cophyla maharipeo</i> sp. nov. <p>(Figs. 7 a, 11 & 12a)</p> <p> <b>Remarks.</b> This new species was listed as confirmed candidate species <i>Cophyla</i> sp. 1 by Vieites <i>et al</i>. (2009). Not included in Glaw & Vences (2007) and Wollenberg <i>et al</i>. (2008). Referred to as <i>Cophyla</i> sp. Ca1 by Perl <i>et al</i>. (2014). D’Cruze <i>et al.</i> (2008) did not distinguish the three new species described herein; their <i>Cophyla</i> sp. nov. therefore includes records and altitudinal ranges of <i>C. maharipeo, C. noromalalae,</i> and <i>C. puellarum.</i></p> <p> <b>Holotype.</b> ZSM 3251/2012 (ZCMV 12195), adult male, collected in a small patch of bamboo in the nun garden of Joffreville, -12.49397, 49.20504, 634 m a.s.l., on 17 January 2012, by A. Rakotoarison and A. Razafimanantsoa.</p> <p> <b>Paratypes.</b> UADBA-A 60231 (ZCMV 12193, female), ZSM 3252/2012 (ZCMV 12194, female), UADBA-A 60232 (ZCMV 12196, male), ZSM 3253/2012 (ZCMV 12200), UADBA 60233 (ZCMV 13501), same locality and collectors as the holotype. ZSM 1659/2008 (FGZC 1866), juvenile, collected in the Fontenay Private Nature Park near Joffreville (-12.495278, 49.200833, 720 m a.s.l.), on 26 February 2008, by J. Köhler & M. Franzen.</p> <p> <b>Diagnosis and comparisons.</b> Assigned to the genus <i>Cophyla</i> in the microhylid subfamily Cophylinae based on enlarged terminal discs on fingers and toes, absence of nuptial pads, absence of clavicle, posterior vomer undivided and not overlapping with neopalatines, and molecular phylogenetic relationships. From other arboreal species of cophylines the species can be distinguished by a combination of the following character states: small to medium body size (adult SVL 22–27 mm); usually with yellowish and white ventral colour; third toe length equals fifth toe; posterior vomer probably with weakly developed vomerine teeth; males with prepollical tubercle but lacking a finger-like prepollex as typical for <i>Anodonthyla</i>.</p> <p> Distinguished from all other species of <i>Cophyla</i> by advertisement call characteristics (long call duration 1166–1346 ms). Further distinguished from <i>C</i>. <i>phyllodactyla</i> by smaller body size (adult SVL 22–26 mm <i>vs</i>. 27–29 mm); from <i>C</i>. <i>occultans</i> by larger size (SVL 22–26 mm <i>vs</i>. 16–21 mm); and from <i>C</i>. <i>berara</i> by the fourth finger being equally long as third (<i>vs</i>. fourth finger slightly longer). For the distinction from the other two new <i>Cophyla</i> species described herein, see their respective diagnoses below.</p> <p> <b>Description of the holotype.</b> Adult male in good state of preservation, some muscle tissue removed from right thigh; snout-vent length 22.6 mm; body slender; head wider than long, not wider than body; snout bluntly rounded in dorsal and lateral views; nostrils not protuberant, nearer to tip of snout than to eye; canthus rostralis distinct and somewhat rounded, loreal region slightly concave to straight; tympanum moderately distinct, 47% of eye diameter; supratympanic fold starting at the posterior border of the eye and ending at the base of the forearm (less distinct on left side); tongue broadly rounded, not bifid or notched, no lingual papilla; maxillary teeth present; vomerine teeth small, rudimentary. Forelimbs slender; subarticular tubercles on all fingers single, protuberant; outer metacarpal tubercle relatively large and flat; inner metacarpal tubercle oblong, forming small protuberance at base of first finger; hand without webbing; fingers distinctly flattened and broad along entire length; relative length of fingers 1<2=4<3; finger discs distinctly enlarged, broadly rounded to slightly bilobate, with lateral fringes; nuptial pads absent. Hindlimbs slender; tibiotarsal articulation reaching between forelimb and tympanum when hindlimb adpressed along body; tibia length 39% of SVL; lateral metatarsalia strongly connected; inner metatarsal tubercle small; outer metatarsal tubercle small and flat, difficult to recognize; webbing between toes weakly developed, restricted to traces of webbing between third and fourth toe; subarticular tubercles on toes present, single, toe discs flattened and relatively broad, broadly rounded to slightly bilobate, smaller than discs of fingers; relative length of toes 1<2<5=3<4.</p> <p>Dorsal skin smooth, without dorsolateral folds. Ventral skin slightly granular on throat and chest and granular on belly. Skin on throat extended, marking the presence of vocal sac.</p> <p>ZSM 3273/2012 FGZC PT male Montagne 24.3 6.8 6.6 1.6 2.4 2.3 1.2 1.7 11.7 5.3 27.5 13.6 8.5 9.7 1</p> <p>4902 d’Ambre</p> <p> <b>……</b> <i>continued on the next page</i> <b>Colour of holotype in preservation.</b> After three years in 70% ethanol (Fig. 7 a), anterior part of dorsum brown with a darker X-shaped pattern. Forelimbs and hindlimbs dorsally brown and beige. Ventrally white on throat with dark dots, brown and yellowish with dark dots on chest, dark brown and whitish with dark dots on belly.</p> <p> <b>Colour of holotype in life.</b> Rather similar to colour in preservative (Fig. 11 a). The beige part of the body was more yellowish. Ventrally (Fig. 12 a) yellow on throat and chest, whitish on belly.</p> <p> <b>Variation.</b> Most of the paratypes are morphologically similar to the holotype (Table 3) except ZSM 1659/2008 (FGZC 1866) which is smaller than the other specimens. In five of them the third and fifth toes are of equal length and in three of them the third is slightly shorter than the fifth. The tibiotarsal articulation in all paratypes reaches between forelimb and tympanum. Males are smaller than females (Table 3). The dorsum is yellow with a brown patch and line (Fig. 11 a), or greyish with a yellowish patch (Fig. 11 c) for adult and subadult specimens. In one juvenile specimen the dorsum is dark beige with a dark brown line and patches (Fig. 11 d). One unsexed specimen (ZSM 233/2004) from low elevations (650 m) of Montagne d’Ambre with uncertain identity, but possibly referable to <i>C. maharipeo</i>, has a SVL of 31.4 mm, suggesting that this species might reach a larger size than is suggested by the type specimens.</p> <p> <b>Etymology.</b> The name is derived from Malagasy mahari-peo (equalling maharitra feo): long voice. This species has the longest call duration among all <i>Cophyla</i> species.</p> <p> <b>Natural history.</b> The holotype was found at night, calling on a bamboo tree. About eight calling males (including the holotype) were heard from the same small patch of bamboo in the garden of an abbey in Joffreville. During the day, at the same site, one female with oocytes and one male were found in a bamboo trunk, together with five juveniles. No further information is available on the natural history and reproduction of this species.</p> <p> <b>Advertisement call.</b> Calls were recorded from paratype UADBA-A 60232 (ZCMV 12196) on 17 January 2012, at an air temperature of ca. 25˚C (Fig. 13). The following call description refers to the calls of this single specimen. As with all other <i>Cophyla</i> and most cophylines, single tonal calls were repeated in long series (continued at least for several minutes but probably much longer in undisturbed specimens), with regular inter-call intervals. Dominant spectral call frequency ranged from 2600–2800 Hz (N= 28). Temporal call parameters were as follows: call duration 1166–1346 ms (1257±40 ms; N= 28); duration of inter-call intervals 2154–3881 ms (2552±387 ms; N= 27). In the recorded call sequence, each call starts with a very short intensity peak visible in the oscillograms and of slightly higher frequency than the rest of the call. While we cannot exclude that this peak is a genuine part of the vocalization of this species, we find it more likely that it represents either an artefact of the recording equipment, or an abnormal feature in sound emission.</p>Published as part of <i>Rakotoarison, Andolalao, Crottini, Angelica, Müller, Johannes, Rödel, Mark-Oliver, Glaw, Frank & Vences, Miguel, 2015, Revision and phylogeny of narrow-mouthed treefrogs (Cophyla) from northern Madagascar: integration of molecular, osteological, and bioacoustic data reveals three new species, pp. 61-89 in Zootaxa 3937 (1)</i> on pages 73-76, DOI: 10.11646/zootaxa.3937.1.3, <a href="http://zenodo.org/record/288186">http://zenodo.org/record/288186</a&gt