Cophyla maharipeo, 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 : 73-76

Cophyla maharipeo sp. nov.

( Figs. 7 a, 11 & 12a)

Remarks. This new species was listed as confirmed candidate species Cophyla sp. 1 by Vieites et al. (2009). Not included in Glaw & Vences (2007) and Wollenberg et al. (2008). Referred to as Cophyla sp. Ca1 by Perl et al. (2014). D’Cruze et al. (2008) did not distinguish the three new species described herein; their Cophyla sp. nov. therefore includes records and altitudinal ranges of C. maharipeo , C. noromalalae , and C. puellarum .

Holotype. 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.

Paratypes. 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.

Diagnosis and comparisons. Assigned to the genus Cophyla 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 Anodonthyla .

Distinguished from all other species of Cophyla by advertisement call characteristics (long call duration 1166–1346 ms). Further distinguished from C. phyllodactyla by smaller body size (adult SVL 22–26 mm vs. 27–29 mm); from C. occultans by larger size (SVL 22–26 mm vs. 16–21 mm); and from C. berara by the fourth finger being equally long as third (vs. fourth finger slightly longer). For the distinction from the other two new Cophyla species described herein, see their respective diagnoses below.

Description of the holotype. 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.

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.

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

4902 d’Ambre

…… continued on the next page Colour of holotype in preservation. 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.

Colour of holotype in life. 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.

Variation. 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 C. maharipeo , has a SVL of 31.4 mm, suggesting that this species might reach a larger size than is suggested by the type specimens.

Etymology. The name is derived from Malagasy mahari-peo (equalling maharitra feo): long voice. This species has the longest call duration among all Cophyla species.

Natural history. 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.

Advertisement call. 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 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), 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.

Figures

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 11. Specimens of Cophyla maharipeo sp. nov. alive: (a) male holotype (ZSM 3251 / 2012); (b) calling male (holotype); (c) paratype specimen (assignment to voucher number not possible); (d) juvenile.

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 13. Audiospectrogram and oscillogram of two advertisement calls (part of a longer call series) of Cophyla maharipeo sp. nov. from paratype UADBA-A 60232, recorded at Joffreville on 17 January 2012, ca 25 ˚ C, number of recording DR 089. The sounds above 3000 Hz represent other frogs or insects, and the initial intensity peaks at each call probably represent an artefact of the recording device or an abnormal sound emission of the recorded specimen.

Tables

TABLE 3. Morphometric measurements (all in mm) of Cophyla form Joffreville and Montagne d’Ambre and of three specimens of C. phyllodactyla from Nosy Be, one specimens from Manongarivo and one specimen of C. occultans from Nosy Be, all from the north of Madagascar. For abbreviations of measured variables, see Materials and Methods. Further abbreviations used: RHL, relative hindlimb length; (1) reaches between forelimb and tympanum, (2) reaches tympanum. MANP, Montagne d'Ambre National Park.

Museum number C. maharipeo ZSM 3251/2012	Field number ZCMV 12195	HT	Sex male	Locality Joffreville	SVL 22.6	HW 6.6	HL 5.8	TD 1.5	ED 2.9	END 1.8	NSD 1.3	NND 1.5	FORL 9.8	HAL 6.0	HIL 28.2	FOTL 12.6	FOL 8.0	TIL 8.9	RHL 1
UADBA–A60232	ZCMV 12196	PT	male	Joffreville	23.2	6.7	5.9	1.1	3.1	1.3	0.9	2.1	10.5	5.3	29.1	14.2	8.9	9.2	1
UADBA-A60233	ZCMV 13501	PT	??	Joffreville	25	6.8	6.6	1.1	3.0	1.9	1.1	2.0	11.9	5.6	30.0	14.0	9.5	9.9	1
UADBA-A60231	ZCMV 12193	PT	female??	Joffreville	24.3	7.9	7.4	1.2	2.9	1.8	1.4	1.9	14.8	6.3	31.2	15.4	9.2	11.1	1
ZSM 3252/2012	ZCMV 12194	PT	female??	Joffreville	26.5	7.5	6.1	1.1	2.9	1.3	1.0	1.7	13.9	5.5	29.3	14.7	9.7	10.2	1
ZSM 3253/2012	ZCMV 12200	PT	??	Joffreville	26.5	7.9	7.0	1.2	2.9	1.5	0.9	1.7	12.6	6.0	33.7	15.3	10.1	10.9	1
ZSM 1659/2008	FGZC 1866	PT	subadult?	Forêt d’Ambre	17.3	6.2	5.6	1.2	2.3	1.7	1.0	1.9	11.7	4.0	11.3	11.3	7.0	8.3	1
C. noromalalae ZSM 3250/2012	ZCMV 13518	HT	male	MANP entrance	21.6	6.2	5.9	1.2	2.8	1.6	1.2	1.9	12.9	6.2	25.2	12.8	8.5	9	1
UADBA-A60235	ZCMV 13519	PT	??	MANP entrance	23.4	7	6.3	1.2	3.3	1.8	1.1	1.8	11.2	5.8	28.5	12.7	8.5	9.7	1
ZSM 2070/2007	FGZC 1013	PT	??	Montagne d’Ambre	28.9	9.8	7.9	1.4	3.5	1.6	1.2	2.2	15.1	7.4	37.4	16.4	11.5	12.2	1
UADBA-A 60236	FGZC 4901	PT	male	Montagne d’Ambre	25.4	7.8	6.7	1.6	3.2	2.2	1.6	1.6	13.1	6.4	31.4	14.6	9.7	10.2	1