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
treatment provided by
Cophyla fortuna sp. nov.
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 View FIGURE 5 ) 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 View Materials /2016 ( MSZC 0260) ( Fig. 4b View FIGURE 4 ), 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. GoogleMaps
Paratypes. One female, ZSM 468 View Materials /2016 ( ZCMV 15082), and one male, ZSM 469 View Materials /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 GoogleMaps ; three specimens of unverified sex, UADBA-A ( ZCMV 15139), ZSM 470 View Materials /2016 ( ZCMV 15142) ( Fig. 4 View FIGURE 4 c–d), UADBA-A ( ZCMV 15146), and two males, ZSM 471 View Materials /2016 ( ZCMV 15147) ( Fig. 4 View FIGURE 4 e–f) and ZSM 542 View Materials /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 GoogleMaps ; 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 GoogleMaps ; ZFMK 59940–59941 View Materials , 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 GoogleMaps
Referred specimens. ZSM 1617/2012 ( FGZC 3951), adult male (call voucher, Fig. 6 View FIGURE 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.
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.View Figure View Figure
After two years in preservative ( Fig. 5 View FIGURE 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 View FIGURE 4 ) 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 View FIGURE 3 , 6 View FIGURE 6 ) to this species, based on molecular evidence ( Fig. 1 View FIGURE 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 View FIGURE 4 ). 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 View FIGURE 4 ), 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 View FIGURE 4 ), 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 View FIGURE 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.
|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|
|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|
No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.