Cophyla fortuna, Rakotoarison & Scherz & Bletz & Razafindraibe & Glaw & Vences, 2019

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), pp. 271-288 : 277-281

Cophyla fortuna sp. nov.

( Figs. 4–5 )

LSID: urn:lsid:zoobank.org:act:3C7B6499-D985-4A06-BC6D-55859384F919

Remark. The calls of this species were described as those of Platypelis occultans (Marojejy) in Vences et al. (2005) . This species has been figured as an unidentified Platypelis from Marojejy National Park in Glaw & Vences (2007: 135, Fig. 5c ) and listed as Cophyla sp. Ca4 and Ca04 in Rakotoarison et al. (2015), Scherz et al. (2016b, 2017b) and Peloso et al. (2017), and Cophyla sp. MRSN A 2660 in Tu et al. (2018).

Holotype. ZSM 467 /2016 ( MSZC 0260 ) ( Fig. 4b ), adult male, collected on 22 November 2016, from low elevation of Marojejy National Park (S14.4393, E49.7766, 444 m a.s.l.), north-eastern Madagascar, by M.D. Scherz, J. Razafindraibe, and A. Razafimanantsoa.

Paratypes. One female, ZSM 468 /2016 ( ZCMV 15082 ), and one male, ZSM 469 /2016 ( ZCMV 15115 ) both collected on 15 November 2016 from a campsite called Camp 0 in Marojejy National Park (S14.44633, E49.78523, 310 m a.s.l.), by A. Rakotoarison, M.D. Scherz, M.C. Bletz, J.H. Razafindraibe, A. Razafimanantsoa, and M. Vences ; three specimens of unverified sex, UADBA-A ( ZCMV 15139 ), ZSM 470 /2016 ( ZCMV 15142 ) ( Fig. 4 c–d), UADBA-A ( ZCMV 15146 ), and two males, ZSM 471 /2016 ( ZCMV 15147 ) ( Fig. 4 e–f) and ZSM 542 /2016 ( ZCMV 15151 ), all collected on 16 November 2016 from the vicinity of Camp Mantella in Marojejy National Park (S14.437666, E49.77557, 456 m a.s.l.), by A. Rakotoarison, M.D. Scherz, M.C. Bletz, J.H. Razafindraibe, A. Razafimanantsoa, and M. Vences ; five specimens of unverified sex, UADBA-A ( ZCMV 15309 ), UADBA-A ( ZCMV 15311 ) and UADBA-A ( ZCMV 15313 – ZCMV 15315 ), all collected on 21 November 2016, from the vicinity of Camp Mantella in Marojejy National Park with the same coordinates and collectors as the previous specimens ; ZFMK 59940–59941 , two adult males (call vouchers), collected on 22 February 1995 from low elevation of Marojejy National Park (Camp 1, ca. 300 m a.s.l., no coordinates available) by F. Glaw & O. Ramilison

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Referred specimens. ZSM 1617/2012 (FGZC 3951), adult male (call voucher, Fig. 6 ), collected on 7 December 2012, in Fanambana forest (approximately at 13.61–13.62°S, 49.99–50.00°E, 53–67 m a.s.l.), by F. Glaw, O. Hawlitschek, F.M. Ratsoavina, A. Rakotoarison, T. Rajoafiarison, and A. Razafimanantsoa. MRSN A2660, a specimen that was not examined by us, but that was used for genetic analyses in Andreone et al. (2005 b) and Rakotoarison et al. (2012) and is genetically non-divergent. ZFMK 59959–59960, two adult males (call vouchers), collected on 18 February 1995 ca. 15 km south of Sambava (no coordinates available) by F. Glaw & O. Ramilison.

Diagnosis. Assigned to the genus Cophyla in the microhylid subfamily Cophylinae based on enlarged terminal discs on fingers and toes, elongated body, absence of nuptial pads, and molecular phylogenetic relationships. The species can be distinguished from all other cophylines by the combination of the following character states: (1) Small size; adult male snout–vent length (SVL) 17.5–21.6 mm, female SVL up to 23.7 mm; (2) manus with four fingers (second finger slightly shorter than fourth) and pes with five toes (third toe slightly shorter than fifth); (3) discs of fingers and toes orangish in life; (4) males with prepollical tubercle but lacking a finger-like prepollex as typical for Anodonthyla .

Distinguished from C. maharipeo by distinctly shorter calls and more contrasted orange color of toes; from C. phyllodactyla by smaller male body size (SVL: 17.5–21.6 mm vs. 27–29 mm) and by a distinctly longer duration of silent intervals between calls (inter-call interval 1324–2021 ms vs. 555–605 ms); from C. occultans by slightly shorter call duration (411–531 ms vs. 500–550 ms) and from C. berara by shorter call duration (call duration 411–531 ms vs. 612–820 ms); from C. puellarum by smaller body size (SVL 17.5–21.6 mm vs. 27–29 mm) and by a combination of longer calls with shorter inter-call intervals (411–531 ms vs. 320–390 ms and 1324–2021 ms vs. 1961–3996 ms). The new species is morphologically most similar to (and genetically the sister species of) C. noromalalae but differs from this species by having slightly smaller body size (verified adult male SVL 17.5–21.6 mm vs. 21.6–25.4 mm; Table 1 ), by having the second finger slightly shorter than the fourth, by marginally shorter relative FOTL (FOTL/SVL 0.54–0.55 vs. 0.56–0.58; Table 1 ), by slightly shorter call duration (411–531 ms vs. 555–821 ms), and by higher spectral frequency in advertisement calls (approximate prevalent bandwidth 3450– 3950 Hz vs. 2513–3000 Hz; a variable however possibly influenced by its smaller body size).

Description of the holotype. Adult male in good state of preservation, some muscle tissue removed from right thigh. SVL 17.5 mm; body slender; head slightly longer than wide, not wider than body; snout slightly pointed in dorsal view, rounded in lateral view; nostrils not protuberant, nearer to tip of snout than to eye; canthus rostralis distinct, concave; loreal region slightly concave; tympanum moderately distinct, 54% of eye diameter; supratympanic fold distinct, supratympanic fold starting at the posterior border of the eye and ending at the base of the forearm; tongue long, broadening posteriorly, attached anteriorly, not notched; maxillary teeth present, weakly developed; vomerine teeth not visible or palpable (presence of rudimentary vomerine teeth cannot be excluded and would require osteological examination); choanae rounded. Forelimbs slender; subarticular tubercles single, distinct; outer metacarpal tubercle not visible; inner metacarpal tubercle distinct, forming large and distinct protuberance at base of first finger; hand without webbing; fingers with lateral fringes, smaller than discs of fingers; relative length of fingers 1 <2<4<3; finger discs broadly rounded to slightly bilobed; nuptial pads absent. Hindlimbs slender; tibiotarsal articu- lation reaching between forelimb and tympanum when hindlimb adpressed along body; tibia length 40% of SVL; inner metatarsal tubercle small, oval; 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 with rela- tively broad discs, broadly rounded to slightly bilobed; relative length of toes 1<2<3<5<4; third toe slightly shorter than fifth. Dorsal skin smooth, without dorsolateral folds. Ventral skin smooth on throat and chest and moderately granular on belly.

After two years in preservative ( Fig. 5 ), the specimen has an inconspicuous coloration with little or no pattern. The dorsal side is a muddy beige with diffusely delimited darker brown on the head, and irregular dark brown dots scattered over the dorsum, and darker brown color also irregularly spread over the dorsal sides of the limbs. Ventrally, the specimen is cream with some dark speckles on the abdomen and light beige on throat, chest and limbs. Color in life not recorded under daylight; at night while calling ( Fig. 4b ) pale and translucent, with some scattered yellowish dots on the dorsal surfaces and yellowish tips of fingers and toes.

Distribution. The species is known from lowlands in and close to Marojejy National Park, encompassing an elevational range from 310–456 m a.s.l. and probably including Manantenina village at ca. 65 m a.s.l. (S14.496489, E49.821839) where it was recorded as Cophyla occultans by Glaw & Vences (2007). We furthermore tentatively assign a specimen from Fanambana forest ( Figs. 3 , 6 ) to this species, based on molecular evidence ( Fig. 1 ) (but note its bioacoustic differences as discussed below). We also assign specimens from near Sambava ( ZFMK 59959–59960) to this species; they had similar calls to those from Marojejy ( Vences et al. 2005) and identified call differences might result from higher recording temperatures, but their genetic identity is unknown, and thus their attribution to C. fortuna remains tentative as well.

Natural history. The species occurs in rainforest at low elevations. Active at night, specimens were mostly found in bamboo forest. Calling occurred in the evening at dusk. Males were calling with the head directed upwards from leaves in the vegetation ca. 2 m above the ground. A large expanded vocal sac remains visible in the silent period between two calls. Each call is most probably produced during one expiration. Calls were recorded in November and February, suggesting prolonged reproductive activity in the rainy season. Numerous specimens were found infested with trombiculid mites, probably of the genus Endotrombicula ( Wohltmann et al. 2007) , especially on fingers, toes and limbs but also on the belly and vocal sac (visible in Figs. 4a,b,e,h ). The holotype was observed exhibiting jerky hand and foot movements whilst calling, as though it were fending off small biting insects, but none were observed on or near it during the recording (video deposited at https://youtu.be/Kjw10jofNSY); possibly this behavior is directed at mites.

Etymology. The species name is derived from the Latin noun fortuna , meaning luck, and is used as a noun in apposition to the generic name. The new species is fortunate to be found largely within a protected area and apparently may be ecologically lucky in being able to survive in relict bamboo stands and thus less heavily impacted by deforestation than other frogs.

Variation. All morphologically studied paratypes have larger body size than the holotype ( Table 1 ). In all paratypes, the second finger is slightly shorter than fourth and the third toe is slightly shorter than the fifth toe. Based on the single female available, ZSM 468/2016 ( ZCMV 15082), females are larger than males ( Table 1 ). The smallest specimen ZSM 467/2016 ( SVL 17.5 mm) was observed calling, confirming this small body size refers to a mature male. During the day, all specimens have a beige ground coloration with a dark brown pattern of irregular speckles and markings of different intensity. In some specimens ( Fig. 4c ), only small irregularly scattered dark spots are spread over the dorsal surface, with a dark tympanic fold, a dark crossline between the eyes, some yellowish color in the inguinal region, and faint yellowish-orange discs of fingers and toes. In other individuals ( Fig. 4g ), the dark pattern is much denser, forming a dark teddy-bear shaped marking on the head and irregular crossbands on the limbs. The iris is light metallic brown. The ventral side is largely unpigmented and translucent, which causes a pinkish tone on the throat and especially on the belly and ventral side of hindlimbs; some small and indistinct darker spots are often present in the chest region.

Call. The following call description refers to the calls of seven different specimens. The calls were recorded in three different localities at Marojejy National Park, at air temperatures of approximately 18–20°C and 25°C, respectively. The advertisement call consists of a single-note tonal call repeated in a long series ( Fig. 7 ). Calls were recorded (1) by M.D. Scherz on 22 November 2016 at Marojejy National Park low elevation from the holotype, specimen ZSM 467/2016 ( MSZC 260), (2) by M.D. Scherz on 16 November 2016 at Camp Mantella, Marojejy National Park from specimen ZSM 470/2016 ( ZCMV 15142), (3) by M. Vences on 16 November 2016 at Camp Mantella, Marojejy National Park from specimen ZSM 471/2016 ( ZCMV 15147), (4) by M. Vences on 16 November 2016 at Camp Mantella, Marojejy National Park from specimen ZSM 472/2016 ( ZCMV 15151), and (5) by F. Glaw on 22 February 1995 at 25°C air temperature at Marojejy National Park, low elevation (ca. 300 m a.s.l.) from three males (among them ZFMK 59940–59941, pooled in Table 2 as Rakotoarison et al. 2015). Detailed call parameters for the new species and new comparative temporal and spectral call data for C. noromalalae and the specimens referred to C. cf. fortuna from Sambava and Fanambana are provided in Table 2 . Summarizing the data for all these specimens (but not considering the tentatively assigned recordings from south of Sambava and Fanambana; Table 2 ), the call of C. fortuna sp. nov. has a call duration of 411–531 ms, a duration of inter-call intervals of 1126–2999 ms, and an approximate prevalent bandwidth of 3450–3950 Hz.

Figures

FIGURE 1. Maximum likelihood tree of Cophyla species from an unpartitioned analysis based on 2123 bp of four mitochondrial gene fragments (12S, 16S, cob, cox1). Values at nodes represent bootstrap support from 500 replicates (not shown if <50%) and Bayesian posterior probabilities from a partitioned BI analysis of the same data set (not shown if BI consensus tree did not include the respective node). Note that the position of the candidate species, C. sp. Ca5 and Ca6, is based on the 16S gene only as the other genes have not yet been sequenced for these lineages. The tree was rooted with Platypelis tuberifera (removed graphically from the tree to improve visibility of intra-generic relationships and branch lengths). In some cases of branch length differences between individuals from the same site, these are due to missing data for some genes and do not reflect strong genetic differences. The new species, C. fortuna sp. nov., corresponds to the lineage previously named C. sp. Ca4.

FIGURE 3. Map of northern Madagascar, showing the reliably known occurrence records of species of Cophyla.

FIGURE 4. Cophyla fortuna sp. nov. in life: a. Calling male, specimen not collected; b. Holotype ZSM 467/2016 (MSZC 260) calling (see video at https://youtu.be/Kjw10jofNSY); c–d. ZSM 470/2016 (ZCMV 15142); e–f. ZSM 471/2016 (ZCMV 15147); g–h. ZSM 472/2016 (ZCMV 15151).

FIGURE 5. Cophyla fortuna sp. nov., preserved holotype ZSM 467/2016 (MSZC 260) in dorsal and ventral views.

FIGURE 6. Cophyla cf. fortuna sp. nov. in life from Fanambana in dorsolateral (a) and ventral (b) views; call voucher, ZSM 1617/2012 (FGZC 3951).

FIGURE 7. Spectrograms (upper panels) and oscillograms (lower panels) of advertisement calls of (a–b) Cophyla fortuna sp. nov. holotype ZSM 467/2016 (MSZC 0260), and (c–d) C. noromalalae, ZSM 68/2018 (MSZC 0652). On the left, single calls in sections of 1 s duration; on the right, parts of a call series in sections of 10 s duration.

Tables

TABLE 1. Morphometric measurements (all in mm) of Cophyla fortuna sp. nov., C. noromalalae, C. sp. Ca5 and C. sp. Ca6. C. noromalalae measurements from Rakotoarison et al. (2015). Abbreviations: SVL, snout–vent length; HW, maximum head width; HL, head length from posterior maxillary commissure to snout tip; TD, horizontal tympanum diameter; ED, horizontal eye diameter; END, distance from eye to nostril; NSD, distance from nostril to snout tip; NND, distance between nostrils; FORL, forelimb length from axilla to tip of longest finger; HAL, hand length; HIL, hindlimb length from cloaca to tip of longest toe; FOTL, foot length including tarsus; TIL, tibia length; FOL, foot length; HT, holotype; PT, paratype. NA, information not available.

Museum number	Field number	Type	Sex	Locality	SVL	HW	HL	TD	ED	END	NSD	NND	FORL	HAL	HIL	FOTL	FOL	TIL
C. fortuna sp. nov.
ZSM 467/2016	MSZC 0260	HT	Male	Marojejy	17.5	5.2	5.6	1.3	2.4	1.9	1.2	1.5	9.2	5.0	20.7	10.0	5.6	7.1
ZSM 468/2016	ZCMV 15082	PT	Fe-	Marojejy	23.7	7.1	6.8	1.5	3.0	2.2	1.0	1.7	13.5	6.0	30.2	14.3	8.8	10.2
ZSM 469/2016	ZCMV 15115	PT	male Male	Marojey	19.4	5.6	5.4	1.3	2.3	1.7	1.0	1.6	10.6	4.8	24.7	11.1	6.6	8.6
ZSM 470/2016	ZCMV 15142	PT	Male	Marojejy	21.1	6.8	5.3	1.2	3.0	1.2	1.4	1.5	11.2	5.3	26	11.9	6.8	8.2
ZSM 471/2016	ZCMV 15147	PT	Male	Marojejy	21.4	6.1	6.5	1.2	2.9	1.8	1.3	2.2	10.8	4.9	26.7	12.0	6.4	8.7
ZSM 472/2016	ZCMV 15151	PT	Male	Marojejy	21.6	6.7	6.6	1.2	2.5	1.8	1.6	1.2	12.4	5.0	26.9	11.7	8.0	8.6
C. cf. fortuna
ZSM 1617/2012	FGZC 3951		Male	Fanambana	21.4	6.7	6.5	1.5	2.9	1.9	1.1	2.2	10.9	5.9	29.4	13.0	8.3	10.1
C. noromalalae
ZSM 3250/2012	ZCMV 13518	HT	Male	Montagne d’Ambre	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	9.0
UADBA-A60235	ZCMV 13519	PT	NA	Montagne d’Ambre	23.4	7.0	6.3	1.2	3.3	1.8	1.1	1.8	11.2	5.8	28.5	12.7	8.5	9.7
ZSM 2070/2007	FGZC 1013	PT	Fe-	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
UADBA-A 60236	FGZC 4901	PT	male 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
ZSM 3273/2012	FGZC 4902	PT	Male	Montagne d’Ambre	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
C. sp. Ca5
UADBA-A 60303	FGZC 3825		NA	Near Andrafainkona	21.8	7.3	7.1	1.4	2.3	2.2	1.6	1.9	12.7	6.2	31.1	14.5	8.8	10.1
UADBA-A 60304	FGZC 3838		Fe-	Near Andrafainkona	26.0	8.2	7.3	1.5	2.8	1.7	1.2	2.4	14.4	6.2	33.6	16.1	9.2	11.7
UADBA-A 60305	FGZC 3856		male Male	Near Andrafainkona	23.8	6.6	7.2	1.2	2.5	1.8	1.2	1.5	11.9	6.2	31.5	14.4	8.8	10.2
C. sp. Ca6
UADBA-A 60302	FGZC 3820		NA	Ambodimandresy	23.9	6.8	7.2	1.2	2.8	1.7	1.3	1.9	12.5	6.0	30.5	14.3	8.7	9.2
NA	FGZC 3819		NA	Ambodimandresy	26.9	7.6	7.4	1.5	3.1	1.8	1.6	2.5	14.0	6.8	32.5	15.6	9.0	11.1