Eleutherodactylus cattus, Rodríguez, Ariel, Dugo-Cota, Álvaro, Montero-Mendieta, Santiago, Gonzalez-Voyer, Alejandro, Bosch, Roberto Alonso, Vences, Miguel & Vilà, Carles, 2017

Rodríguez, Ariel, Dugo-Cota, Álvaro, Montero-Mendieta, Santiago, Gonzalez-Voyer, Alejandro, Bosch, Roberto Alonso, Vences, Miguel & Vilà, Carles, 2017, Cryptic within cryptic: genetics, morphometrics, and bioacoustics delimitate a new species of Eleutherodactylus (Anura: Eleutherodactylidae) from Eastern Cuba, Zootaxa 4221 (5), pp. 501-522 : 509-512

Eleutherodactylus cattus sp. n.

( Fig. 5 A,B )

Holotype. CZACC14.14152, adult male collected while vocalizing in the trail to Pico El Gato, Sierra del Cobre, 20.01364 N, 76.04809 W, 844 m a.s.l, by A. Rodríguez and R. Alonso in May 2010.

Paratypes. CZACC 14.14150–51 , 14.14153–60, adult males with the same data as the holotype .

Etymology. The species name is an invariable noun in apposition to the genus name, derived from Latin cattus = cat. It refers to the type locality Loma del Gato (Cat Mountain Ridge) in the Sierra Maestra Mountains, a locality that was neglected for long time in herpetological explorations of Cuba and that surely deserves further attention.

Diagnosis. A small species of Eleutherodactylus that can be assigned to the subgenus Eleutherodactylus based on its genetic, acoustic, and morphological similarities with members of the E. auriculatus species group ( Hedges, et al., 2008; Rodríguez, et al., 2010b). It is most closely related to E. glamyrus with which it shares several morphological, ecological, and behavioral features. However, males of Eleutherodactylus cattus can be readily differentiated from E. glamyrus by their larger size (SVL = 23.1–24.7 mm in E. cattus vs 17.4–21.3 mm in E. glamyrus ) and the following combination of advertisement call features: longer duration (call duration = 128.4– 243.1 ms in E. cattus vs 47.2–112 ms in E. glamyrus ), longer rise time (call rise time = 16–173.7 ms in E. cattus vs 1.9–32.2 ms in E. glamyrus ), lower dominant frequency (dominant frequency = 3036.2–3337.6 Hz in E. cattus vs 3006–3799.6 Hz in E. glamyrus ), and greater frequency modulation (frequency modulation = 140.9–332.8 Hz in E. cattus vs 73.2–262.3 Hz in E. glamyrus ). Additionally, E. cattus differs from E. glamyrus in the studied mitochondrial DNA sequences by 8.2% (cob) and 3.2% (16S) and by one substitution in the nuclear Rag-1 gene (see results).

Description. Head as wide as body, width smaller than length; snout subacute in dorsal and lateral views overhanging the jaw; narines laterally oriented and moderately protuberant; rostral canthus rounded and straight, loreal region smooth and abruptly tilted, lips not enlarged. Superior eyelids with conical and small tubercles also present but less evident in the interorbital area. Tympanum present without supratympanic fold, postrictal tubercles present. Coanae oval, partially concealed by the maxillary arch in ventral view. Vomerine odontophores short, straight, separated and nearly perpendicular to the longitudinal body axis. Tongue rounded, longer than wide with notched posterior edge, fixed by the anterior edge and free in its 3/4 of length. Vocal slits and enlarged vocal sac present in males.

Dorsal skin slightly tuberculated, without dorsolateral folds, and less tuberculated in the lower half of flanks. Ventral skin slightly areolated but without folds. Cloacae not expanded. Glandular regions not evident. Hands with ulnar tubercle subconical, palmar tubercle simple and of similar size as the thenar; thenar tubercle oval and pronounced. Supernumerary tubercles absent; fingers with rounded and subconical subarticular tubercles and without lateral expansions. Finger tips rounded and expanded in all cases with the ventral surface forming a circular digital pad bordered by a circumferential groove in 2/3 of its distal edge. Width of the largest toe pad (finger III) roughly similar to tympanum diameter. Finger order III> II> IV> I. One moderate tubercle present on heel, no tubercles on the outer edge of the tarsal, metatarsal tubercles subconical and smaller than subarticular tubercles; internal oval, same size as external; plantar supernumerary tubercles absent; subarticular tubercles rounded and subconical. Toes not webbed, toe tips rounded and expanded in all cases with the ventral surface forming a circular digital pad bordered by a circumferential groove in 2/3 of its distal edge. Heels overlap when thighs are placed perpendicularly to the longitudinal body axis. Toe size order IV> III> V> II> I.

Color in alcohol. Color pattern was similar among the individuals of the type series. Dorsum varies from brownish to beige with darker mottled and two mostly well-defined dark brownish or black anterior dorsolateral marks; the flanks show a similar pattern, lighter from dorsum to belly, a relatively wide dark brown strip from the insertion of the forelimb to the eye; belly pale and throat (vocal sac) from yellowish to light brown, underside of the limbs slightly yellowish (see Figure 5 A–B for dorsal and ventral views of the holotype).

Color in life. Dorsum color varies from brownish to greenish tan, with an interocular bar of darker brown color followed by an X shaped mark both darker than the background color. A distinctive pattern of chevron-like bands is evident in the sacral region extending to the hind legs. Eyelids with a greenish wash, pupils creamy to coopery colored with horizontal slit, iris black. The flanks are lighter colored than the dorsum, tympanum creamy white. Loreal region color varies from tan to greenish with an obvious black stripe that extends from snout to supratympanic fold and becomes progressively diffuse towards the insertion of the forelimb. Venter whitish to translucent, vocal sac yellow. A photograph of a living paratype is presented in figure 5 C.

Measurements of the holotype (mm).: (see Methods for abbreviations) SVL 23.9; HW 9.0; HL 9.1; IN 2.1; EN 2.9; IO 4.2; EL 3.0; TyL 1.2; FaL 5.7; HaL 1.8; F1 2.7; F2 3.0; F3 4.6; F4 2.9; FP1 0. 6; FP2 0.7; FP3 0.9; FP4 1.0; ThL 10.7; TL 11.6; FL 7.1; T1 3.6; T2 4.2; T3 6.1; T4 9.4; T5 7.3; TP1 0.7; TP2 0.6; TP3 0.6; TP4 0.7; TP5 0.6.

Remarks. Although the proposed diagnostic features allow a clear discrimination between adult males of Eleutherodactylus cattus and E. glamyrus , a straightforward classification of non-vocal juveniles and females is so far impossible without genetic analyses. It is likely that adult females of E. cattus oversize those of E. glamyrus but no female specimens are available for E. cattus and a proper test of this hypothesis will require the collection of a sufficient number of female specimens of both taxa. It can also be hypothesized that additional diagnostic features for both taxa will be recovered after a detailed examination of osteological and soft tissue examinations are conducted.

Distribution. This species is only known from the type locality but assuming it has specialized to high elevations like its sister taxon, Eleutherodactylus glamyrus , it could well be found in neighboring areas above 800 m a.s.l..

Natural history. Field data indicate that Eleutherodactylus cattus is a nocturnal species that inhabits the montane rainforests and elfin woodlands above 800 m a.s.l., in areas of Loma del Gato-Monte Líbano Ecological Reserve, in the Sierra del Cobre massif, Santiago de Cuba province, in eastern Cuba. In this region, the mean monthly air temperature is 18.4 °C with minimal and maximum mean values around to 15.7 °C and 22.4 °C, respectively. Mean monthly relative humidity is high year round, and ranges between 87–92%, the mean annual precipitation is 1.220 mm, with May and October being the rainiest months ( Potrony, et al., 1994; Reyes, 1999). Males of Eleutherodactylus cattus were observed calling in the vegetation at 1.16 ± 0.47m (mean ± SD; range: 0.50–2.20 m) above the ground in the understory ( Figure 5 C). This forest stratum is dense and rich in shrubs and herbaceous plants, with ferns, liverworts, mosses and terrestrial orchids ( Figure 5 D). Calling males were heard in three nearby localities in Loma El Gato, between 844–1070 m a.s.l (ascent trail on the northern slope of Pico El Gato, 844 m a.s.l; surroundings of Loma de la Cruz, 1070 m a.s.l.; and Loma de La Juana, 900 m a.s.l). E. cattus showed an apparent acoustic activity peak at dusk (between 19:00–21:00 hrs) and shortly before dawn (5:00– 6:30hrs). Calling males appear to show preference for exposed surfaces of leaves and ferns, but they can call at different orientations, facing down or horizontally.

At least another eight species of Eleutherodactylus are known to occur in sympatry with E. cattus in the surroundings of the type locality: Eleutherodactylus atkinsi Dunn , E. auriculatus , E. cuneatus (Cope) , E. dimidiatus (Cope) , E. gundlachi Schmidt , E. ionthus Schwartz , E. limbatus (Cope) , E. ricordii (Duméril and Bibron) , plus the hylid Osteopilus septentrionalis (Duméril and Bibron) . Two species of the Eleutherodactylus auriculatus species group, E. ionthus and E. auriculatus ( Hedges, et al., 2008; Padial, et al., 2014) were heard and observed vocalizing simultaneously with E. cattus at the type locality. The first species, E. ionthus , vocalizes from high perches (above three meters) on the arboreal stratum often on epiphytic plants ( Bromeliaceae ). This species produces a very different advertisement call in terms of temporal and spectral structure (Hedges et al., 1992; Díaz and Cádiz, 2008). The second species, E. auriculatus , uses similar calling sites (substrates and heights) in the understory as E. cattus , but the acoustic features of its advertisement call are distinctly different (e.g. faster call rate, shorter call duration and higher dominant frequency), as already noted by Estrada and Hedges (1997).

Figures

FIGURE 5. A – B) Holotype of Eleutherodactylus cattus (CZACC 14.14152) in dorsal and ventral views. C) Male (CZACC 14.14153, paratype) calling in the trail to Pico El Gato, Sierra del Cobre, 844 m a. s. l .. D) Habitat in the type locality, dominated by montane rainforest, Pico del Gato in the background.