Cophyla noromalalae, Rakotoarison, Andolalao, Crottini, Angelica, Müller, Johannes, Rödel, Mark-Oliver, Glaw, Frank & Vences, Miguel, 2015

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, Zootaxa 3937 (1), pp. 61-89 : 77-78

Cophyla noromalalae sp. nov.

( Figs. 7 b, 12b &14)

Remark. Referred to as Cophyla sp. Ca2 by Perl et al. (2014).

Holotype. ZSM 3250/2012 ( ZCMV 13518), adult male, collected in a bamboo patch near the entrance office of Montagne d’Ambre National Park ( MANP), -12.616625, 49.150129, 900 m a.s.l., on 24 January 2012, by A. Rakotoarison and A. Razafimanantsoa.

Paratypes. UADBA-A 60235 ( ZCMV 13519), same locality, collection date and collector as the holotype; UADBA-A 60236 ( FGZC 4901), calling male, collected on 10 December 2012 around MANP entrance by F. Glaw, F. Ratsoavina, O. Hawlitschek and T. Rajoafiarison; ZSM 3273/2012 ( FGZC 4902), male, same locality, collection date and collector as the calling male, ZSM 2070/2007 ( FGZC 1013), sex unknown, collected at MANP (Voie des milles arbres, -12.516667, 49.176667, 1050 m a.s.l.), on 23 February 2007 by P. Bora, H. Enting, F. Glaw, J. Köhler & A. Knoll.

Diagnosis and comparisons. Assigned to the genus Cophyla in the microhylid subfamily Cophylinae based on enlarged terminal discs on fingers and toes, absence of clavicle, 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 a combination of the following character states: small to moderate body size (adult SVL 22–29 mm; verified adult males 22–25 mm); most specimens uniformly yellowish ventrally; discs of fingers and toes pinkish in life; third toe at least slightly but often distinctly shorter than fifth; posterior vomer without clearly recognizable vomerine teeth (rudiments are probably present); males with prepollical tubercule but lacking a finger-like prepollex as typical for Anodonthyla .

Distinguished from C. maharipeo by distinctly shorter calls, and from C. phyllodactyla by a distinctly slower call repetition rate ( Table 2 ). Further distinguished from C. phyllodactyla by smaller body size (verified adult male SVL 24–25 mm vs. 27–29); from C. occultans by larger size (verified adult male SVL 22–25 mm vs. 16–21 mm). Most similar by morphology and bioacoustics to C. berara which however has strong and concordant differences in mitochondrial and nuclear DNA ( Figs. 2–3 ). For the distinction from C. puellarum , see the respective diagnosis of that species below.

Description of the holotype. Adult male in good state of preservation, some muscle tissue removed from right thigh. Snout-vent length 21.6 mm; body slender; head slightly longer than wide, 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 broadly rounded, loreal region concave; tympanum moderately distinct, 41% of eye diameter; supratympanic fold distinct, almost straight, running from posterior edge of eye to forelimb insertion; tongue broadly rounded, without lingual papilla, not bifid or notched; maxillary teeth present; vomerine teeth not visible. Forelimbs slender; subarticular tubercles on all fingers single, curved; outer metacarpal tubercle relatively small and flat; inner metacarpal tubercle oblong, forming large and distinct protuberance at base of first finger; hand without webbing; fingers distinctly broadly rounded to slightly bilobate, with lateral fringes, smaller than discs of fingers; relative length of fingers 1<2=4<3; finger discs broadly rounded to slightly bilobate; nuptial pads absent. Hindlimbs slender; tibiotarsal articulation reaching between forelimb and tympanum when hindlimb adpressed along body; tibia length 41% of SVL; inner metatarsal tubercle small and flat, outer metatarsal tubercle absent; webbing between toes weakly developed, with traces of webbing between third and fourth toe; subarticular tubercles on toes single; toes flattened and their discs relatively broad, broadly rounded to slightly bilobate; relative length of toes 1<2<3<5<4; third toe slightly (left) to distinctly (right) shorter than fifth. Dorsal skin smooth, without dorsolateral folds. Ventral skin smooth on throat and chest and moderately granular on belly.

Colour of holotype in preservation. After several years in 70% ethanol ( Fig. 7 b), dorsum almost uniformly greyish with a few fine and small dark dots, and faint dark crossbands on toes and lower arm. Forelimbs greyish with moderate brown band; hindlimbs greyish with yellow dots. Finger and toe discs pinkish. Ventrally, brown and white with dark dots on throat, dark brown on chest, dark brown and whitish with dark dots on belly, and brown and yellowish on limbs.

Colour of holotype in life. Dorsum almost uniformly beige with some light yellow patches ( Fig. 14 a). Finger and toe discs light orange. Ventrally almost uniformly whitish with a pinkish shade ( Fig. 12 b).

Variation. The paratypes are of larger body size than the holotype ( Table 3 ). In all paratypes, the third toe is slightly shorter than the fifth toe. Sexual dimorphism uncertain due to lack of data on females. Coloration varies among specimens. Dorsum can be beige with yellowish patch ( Fig. 14 a-b), or almost uniformly greyish with some yellow patches ( Fig. 14 c), or dark grey with pinkish and whitish patches on the flank ( Fig.14 d).

Etymology. This species is dedicated to Noromalala R. Raminosoa, the Malagasy supervisor of the first author, to whom we are immensely grateful for her support over the past years.

Natural history. The holotype was found during the day in a bamboo patch near the entrance office of Montagne d’Ambre National Park. The calling male (paratype 3273/2012) was found on a bamboo stand at night. No further information is available on the natural history and reproduction of this species.

Advertisement call. Recorded from paratype ZSM 3273/2012 (FGZC 4902), around the park entrance of MANP, on 10 December 2012, at an air temperature of ca. 20˚C ( Fig. 15 ). The following call description refers to the calls of this single specimen. As with all other Cophyla and most cophylines, single tonal calls were repeated in long series (continued at least for several minutes but probably much longer in undisturbed specimens). Dominant frequency ranged from 2300–2500 Hz (N= 64). Temporal call parameters were as follows: call duration 662–821 ms (716±38 ms; N= 64); duration of inter-call intervals 874–1882 ms (1217±239 ms; N= 63).

Figures

FIGURE 2. Phylogenetic tree of species and candidate species in the genus Cophyla. Majority-rule consensus tree derived from a partitioned Bayesian inference analysis of concatenated DNA sequences of the 12 S, 16 S, COX 1, COB, and RAG 1 genes (2627 bp), showing relationships among species of Cophyla. Numbers at nodes are posterior probabilities (only values> 0.9 shown). For the three new species described herein the tree also shows their candidate species names according to Perl et al. (2014).

FIGURE 3. Majority-rule consensus tree derived from a partitioned Bayesian inference analysis of DNA sequences of the nuclear RAG 1 gene (503 bp), showing relationships among species of Cophyla. Numbers at nodes are posterior probabilities (only values> 0.9 shown). The tree was rooted with the same outgroups as in Fig. 2 (removed for better graphical representation).

FIGURE 7. Comparative photographs of the three preserved holotypes (scale bar 5 mm): (a) C. maharipeo sp. nov., ZSM 3251 / 2012, (b) Cophyla noromalalae sp. nov., ZSM 3250 / 2012; (c) C. puellarum sp. nov., ZSM 3249 / 2012.

FIGURE 12. Ventral sides of the holotypes of the three newly described Cophyla species from Montagne d’Ambre and Joffreville, photographed from living specimens. Inset photos show the same specimens in dorsolateral view. (a) C. maharipeo sp. nov., male holotype ZSM 3251 / 2012, Joffreville; (b) C. noromalalae sp. nov., male holotype ZSM 3250 / 2012, park entrance, male; (c) C. puellarum sp. nov., male holotype ZSM 3249 / 2012, Point de Vue du Grand Moulin.

FIGURE 14. Specimens of Cophyla noromalalae sp. nov. in life: (a) male holotype, ZSM 3250 / 2012; (b) male observed calling, ZSM 3273 / 2012; (c) male paratype, UADBA-A 60236; (d) paratype specimen, ZSM 2070 / 2007.

FIGURE 15. Audiospectrogram and oscillogram of three advertisement calls (part of a longer call series) of Cophyla noromalalae sp. nov. from paratype ZSM 3273 / 2012, recorded around the entrance of Montagne d'Ambre National Park on 10 December 2012, ca. 20 ˚ C, number of recording DR 90. The sounds above 2500 Hz represent other frogs or insects.