Eleutherodactylus feichtingeri, Díaz, Luis M., Hedges, Blair & Schmid, Michael, 2012

Díaz, Luis M., Hedges, Blair & Schmid, Michael, 2012, A new cryptic species of the genus Eleutherodactylus (Amphibia: Anura: Eleutherodactylidae) from Cuba, Zootaxa 3220, pp. 44-60 : 46-52

Eleutherodactylus feichtingeri , new species

Fig. 1A

Holotype. MNHNCu 1181, adult male from the surroundings of Paso de Lesca (21º35'26" N, 77º47'21" W), on the southern slope of Sierra de Cubitas, municipality of Cubitas, Camagüey province, collected by Luis M. Díaz and Antonio Cádiz on August 5, 2010.

Paratypes (n=67). Males. Camagüey Province: MNHNCu 1182–1189, with the same data as the holotype, collected by L. M. Díaz, A. Cádiz, and Dariel López; MNHNCu 1190–1191, from Hoyo de Bonet (21º36'N, 77º47'W), Sierra de Cubitas, collected by L. M. Díaz on September 9, 1996; MNHNCu 1192–1193, same locality as the holotype, collected by L. M. Díaz on September 16–19, 2002. Sancti Spiritus Province: MNHNCu 1212, surroundings of Hotel Zaza, Río Zaza, collected by L. M. Díaz on July 11, 1999. Granma Province: MNHNCu 1194– 1208, Marea de Limones (19º58'35"N, 77º36'60"W), Niquero, Cabo Cruz, collected by L. M. Díaz and A. Cádiz on October 7, 2010; MNHNCu 1209–1211, same locality, but collected by L. M. Díaz, Freddy Hordies, and André van Hecke, on September 3, 2009; BSC.H 659, 2 km W de Los Muertos, Alegría de Pío, Niquero, collected by Ansel Fong on April 17, 1996. Santiago de Cuba Province: MNHNCu 1213–1217, Gran Piedra (20º00'47"N, 75º37'39"W), Sierra Maestra, 800–900 m a.s.l., Santiago de Cuba, collected by L. M. Díaz and A. Cádiz along the road borders, on October 8, 2010; MNHNCu 1218–1219, surroundings of Santiago de Cuba airport, Santiago de Cuba, collected by L. M. Díaz on May 29, 2000; BSC.H 2476–2480, La Tabla, Tercer Frente, collected by A. Fong and Rolando Viña on April 19, 2000; BSC.H 3176, surroundings of Paso la Mina, 4.5 km S of Cruce de los Baños, Tercer Frente, collected by A. Fong and Rolando Teruel on May 14, 2004; BSC.H 2671–2674, gardens of Hotel Sierramar, Sevilla, Guamá, collected by A. Fong and David Maceira on November 21, 2001; BSC.H 2342–2345, El Olimpo, 4km WNW of Gran Piedra, Santiago de Cuba, collected by A. Fong on February 9, 2000; BSC.H 3156–3157, Providencia, 13.5 km WSW from El Caney, Santiago de Cuba, collected by B. Lauranzón and D. Maceira on April 29, 2004. Guantánamo Province: MNHNCu 1220–1226, City of Guantánamo, crossroad to El Salvador (20º11'17"N, 75º13'40"W), collected by L. M. Díaz and A. Cádiz on October 6, 2010; MNHNCu 330, Arroyón, San Antonio del Sur, collected by Orlando H. Garrido on June 25, 1990; MNHNCu 1227–1229, San Rafael (20º 23'09"N, 74º56'40"W), Yateras, 400 m a.s.l., collected by L. M. Díaz on May 24, 2000. Female. Santiago de Cuba Province: BSC.H 3271, 1.2 km NNW of Nuevo Mundo, main road to La Caoba, San Luis, collected by R. Teruel and A. Fong on May 26, 2005.

Diagnosis. The new species is a member of the genus Eleutherodactylus , subgenus Euhyas ( Eleutherodactylus gundlachi species group, of the Eleutherodactylus planirostris species series), based on molecular phylogenetic relationships and morphological similarity to other species, especially E. varleyi , following classification of Hedges et al. (2008). Males reach 17.4 mm SVL and females 18.5 mm. The new species is similar to E. varleyi (Fig. 1B) in the following combination of characters: (1) very small size; (2) partially areolate venter; (3) ventral disc present; (4) dorsolateral rows of enlarged tubercles; (5) a black stripe crossing the supratympanic fold, surrounding a lower highlighted glandular area; (6) external vocal sac; (7) small digital discs; and (8) accentuated polychromatism. Morphologically, the new species differs from E. varleyi by having: (1) smaller tympanum [8– 15% (x=12%) of head length, vs. 18–26% (x=21%) in E. varleyi ; Fig. 2 ]; (2) tympanum is partially or totally pigmented and embedded in the supratympanic stripe which looks like an extended mask, having a sharp contrast with the underlying light stripe (in E. varleyi , the tympanum tends to be paler and conspicuous, and mostly interrupts the stripe because of its larger size) ( Fig. 3 ); and (3) different advertisement calls (basically one-note call vs. two-notes call in E. varleyi ). Both species differ genetically as shown below.

Description. Head as wide as long, its length 32.4−37.7% (x=34.7%) of SVL; snout subacuminate in dorsal view and in profile, slightly overlapping the lower jaw; snout length 36.7−59.2% (x=41.3%) of head length; nostrils rounded, not protuberant, directed laterally, and separated by a distance equivalent to 19.6−29.6% (x=24.8%) of head width; canthus rostralis almost straight in dorsal view; rounded in profile; loreal region gradually sloping to the labial border; lips not flared; interorbital distance 1.2−2.5 (x=1.7) times the upper eyelid width, without enlarged tubercles; eyelid skin smooth or with very small granules; loreal area smooth; tympanum superficial, rounded to oval, with poorly distinct annulus, 21.9−42.4% (32.6%) of eye diameter, separated from eye by a distance equivalent to 0.4−1.4 (x=0.6) times its own diameter; supratympanic fold distinct; 2−4 postrictal tubercles with glandular features, and very often one of them is very enlarged; choanae 75−100% of third finger disc diameter, oval, partially concealed by palatal shelf of maxillary arch; vomerine odontophores moderate in size, arched, separated from each other by 1/2 times their length, barely reaching the external margins of choanae; tongue suboval, its posterior 1/2 not adherent to floor of mouth; external vocal sac simple, hemispherical, extended onto chest.

Dorsal skin with granules and tubercles; two dorsolateral rows of tubercles extend from sacrum to the level of forelimb insertion; many of these tubercles are elongated. Flanks areolate. Supraaxillary, postfemoral, and inguinal glands present, variably evident; inguinal glands commonly highlighted in yellow, yellowish-green, or orange. Venter partially areolate, except on chest and throat; anal opening not extended in sheath; inner part of thigh areolate. Palmar tubercle oval, smooth, 1.5 times longer than thenar tubercle; supernumerary palmar tubercle scarce; subarticular tubercles of fingers oval, not very prominent, and rounded in profile. Finger length order: III> IV> II> I; digital discs small, oval to conical, diameter 33−77% (x=49.9%) of tympanum width. Heels without enlarged tubercles; inner metatarsal tubercle narrow and smooth, 1.5 to 2.5 times longer than the conical outer metatarsal tubercle; supernumerary tubercles flat, scarce, and inconspicuous; subarticular tubercles oval to slightly conical, moderately projected in profile. Toes without defined lateral ridges or basal webbing; circumferential groove bordering the distal half of toe pad; heels touching or barely overlapping each other when flexed legs are held at right angles to sagittal plane; toe length order: IV> III> V> II> I. Hand length 17.2−22.4% (x=19.3%) of SVL; foot length 35.2−45.5% (x=40.3%) of SVL; thigh length 32.8−43.4% (x=38.2%) of SVL; shank length 32.6−44.7% (x=40.1%) of SVL; tarsal length 20.6−28.1% (x=26.3%) of SVL. Measurements of E. feichtingeri n. sp. are summarized in Table 1 and compared with those of E. varleyi . Only males were included for comparative purposes, considering the still very low number of females available for both species in collections. Holotype measurements are shown independently in Table 1 .

Color in life: Coloration is variable; the species is polymorphic and exhibits several patterns. Representative patterns ( Fig. 4 ) are: Pattern A: almost uniformly colored frogs or with faint evidence of suprascapular chevron or middorsal diamond shaped figure. Pattern B: presence of dark chevrons, triangles, or rhomboidal dark blotches, sometimes interconnected to each other; these blotches vary in pigment intensity; lighter areas could be present on head and middorsum. Pattern C: wide dorsal longitudinal light areas; this pattern could be evident as two wide paravertebral stripes (gray, greenish, light brown); but it could give the impression of a single dorsal light area depending on extension and contrast of the dark pigment along the vertebral line. Pattern D: two dorsolateral light stripes, which are usually light orange, gray, greenish tan, or yellowish tan.

The frequencies of the different color patterns are variable among populations. For example, of 18 specimens collected at Marea de Limones (Cabo Cruz), 16 exhibited pattern D and only two displayed pattern C. In the Sierra de Cubitas, of 13 individuals, 7 showed pattern A, 1 had pattern B (including the holotype), 2 had pattern C, and 3 had pattern D. Of 5 individuals from Gran Piedra, only 1 had pattern A, and the others pattern C. Finally, in a sample of seven frogs from the city of Guantánamo (crossroad to El Salvador), 1 had pattern A, 2 had pattern B, 2 had pattern C, and 2 had pattern D.

E. feichtingeri n. sp. Holotype E. varleyi

The overall background coloration varies from gray, green, purplish, dark brown, light brown, reddish brown to yellowish brown. Lower flanks are slightly to intensely darker than the dorsum, very often with scattered light areoles. Brown bands or bars on fore- and hindlimbs are well defined, fragmented, or even absent. The supratympanic fold and the tympanic membrane are crossed by a black stripe. The postrictal area is conspicuously white or yellowish green. A half-moon shaped area or a more or less continuous narrow line (usually of the same color as postrictal area) is evident under the eyes. Sides of snout with black pigment, more concentrated along the lower margin of the canthus rostralis. Tip of snout with a vertical pale bar. Venter white or greenish white. Vocal sac usually of same color as belly.

Color in alcohol: Patterns remain after preservation, but most specimens turn gray to greyish brown, with no evidence of green or yellow tones.

Advertisement call description and comparisons. Typical calls of Eleutherodactylus feichtingeri n. sp. consist of one short metallic note ( Fig. 5 A) with a duration of 8–41 milliseconds. In the sonagram, each call usually rises sharply in the first 30–47% of signal duration and then remains constant or lowers slightly in the last part. Maximum amplitude is reached in the first 7–50% (x=16.6%) of call length, after which signal intensity decreases. Some calls show two peaks, the first always more intense than the second. Call duration showed 25.8% of variation within the entire sample. The dominant frequency only varied 4.6%. During a one-hour survey (8 October 2010) along the road from Gran Piedra to the town of Siboney (Santiago de Cuba Province), at an altitudinal gradient from 900 m to sea level (19–26ºC), call pattern was the same. Table 2 summarizes call variables for three populations of E. feichtingeri from which DNA information is available to support species recognition. In all three cases, the accoustic properties were similar.

ranges (in parentheses). Sample sizes are given as N1= number of recorded males, and N2= total number of analyzed calls. Tem-

perature data were taken during field recordings.

E. feichtingeri Call duration Call rise time Call interval Call rate Dominant fre-

n. sp. (seconds) (seconds) (seconds) (calls/minute) quency (kHz) In contrast to Eleutherodactylus feichtingeri n. sp., calls of E. varleyi have two notes and are less intense (not metallic calls). The first note has a duration of 9–32 milliseconds and the second note 8–25 milliseconds, with a note interval of 12–25 milliseconds. The second note tends to be higher pitched than the first note. Sporadically, single notes are given at the begining of long sequences of typical two-note calls, which are uttered more or less uniformly spaced or in a faster sequence of two or more calls separated from another group by a longer period of time. Eleutherodactylus varleyi shows a high variation in the duration, intensity, and pattern of the call. The only individual recorded at Sierra de Cubitas showed a tendency to utter notes with a descending pattern of frequency ( Fig. 5 B). In a total of 20 calls from this individual, the first note rose in frequency in one call, was sinuous three calls, descended in 12, and had an inverted “U”-shaped pattern in four. The second note of that individual’s calls descended in frequency in 16 calls and rose in four ( Fig. 5 C-D). In the first note, maximum amplitude was reached at 27–60% (x=40%) of the call length, and in the second note at 12–44% (27%). These differences in the call rise time correspond to signal shape in the oscillogram ( Fig. 3 C-D). Sympatric individuals of both species were recorded while calling at the same temperature (27.4ºC). Table 2 shows call variables for three populations of E. varleyi .

Molecular phylogeny. The tree obtained from the Cyt-b sequences ( Fig. 6 ) shows that Eleutherodactylus feichtingeri n. sp. and E. varleyi form two well-supported clades. DNA analyses also confirm the sympatry of both species in the vicinity of Sierra de Cubitas, Camagüey Province. Eleutherodactylus varleyi from Cubitas belongs to a clade of central populations (typical E. varleyi ) as well as sampled specimens from the Mayabeque Province. However, populations from Artemisa Province (western Cuba) reveal some complex relationships that may indicate the presence of yet another cryptic species, requiring further study. Eleutherodactylus feichtingeri shows little genetic variation among populations.

Cytogenetic comparisons. With conventional Giemsa staining, all specimens present a diploid number of 2n = 32 with two small metacentric chromosomes 1 and 2 and 14 pairs of telo- or subtelocentric chromosomes 3–16 of continuously decreasing sizes ( Fig. 7 ). No differences can be found in the conventionally stained karyotypes of both species. C-banding reveals distinct differences in the amount and chromosomal location of the constitutive heterochromatin in the karyotypes of the specimens from La Chorrera and Soroa populations ( Eleutherodactylus varleyi ) in northwestern Cuba and the Santo Domingo population ( E. feichtingeri n. sp.) in southeastern Cuba. In the karyotypes of E. feichtingeri from Santo Domingo the constitutive heterochromatin is located in the centromeric and telomeric regions of all chromosomes. Additionally, distinct interstitial C-bands are present in the larger telocentric chromosome pairs 3–8 ( Fig. 8 A). In contrast, the constitutive heterochromatin is confined to the centromeric and telomeric regions in the chromosomes of E. varleyi from Soroa ( Fig. 8 B).

Furthermore, the sizes of centromeric C-bands in the chromosomes from Eleutherodactylus feichtingeri n. sp. of Santo Domingo are generally larger than those of E. varleyi from Soroa. C-banding did not reveal the existence of heteromorphic XY3/XXƤ or ZZ3/ZWƤ sex chromosomes in the populations examined.

Silver staining of the nucleolus organizer regions (NORs) reveals that there are two NOR cytotypes in Eleutherodactylus feichtingeri n. sp. from Santo Domingo. The more frequent one has the standard NORs in the long-arm paracentromeric region of both telocentric homologues 11 ( Fig. 9 A). In the other, rarer one, the NOR in one of the homologues 11 is shifted to the short arm by a class II pericentric inversion ( Fig. 9 B). Individuals homozygous for the inverted NOR have not yet been observed among the low number of examined individuals, but are suspected to occur in this population. Since the NORs in E. varleyi from Soroa are also located in the long-arm paracentromeric regions of telocentric homologues 11 ( Fig. 9 C), the inverted NOR in E. feichtingeri is considered to be the derived condition. Several individuals in the Soroa population show a deletion of one of the homologous NORs ( Fig. 9 C).

In both species, quinacrine staining does not reveal brightly fluorescing heterochromatin in the karyotypes. The chromosomes fluoresce with a uniform intensity over their entire lengths. This observation indicates that the constitutive heterochromatin detected by C-banding is not enriched in AT base pairs ( Schmid et al., 2010). Male meiotic chromosomes in the stage of diakinesis show either a ring-like or cross-like pairing arrangement, typical for male meiosis in the highly evolved Anura ( Morescalchi, 1971, 1973; Schmid et al., 2010), or else, after terminalization of the chiasmata at the ends of their paired arms, a rod-like or v-shaped end-to-end association.

Distribution. Eleutherodactylus feichtingeri n. sp. is widely distributed in part of the central and most of the eastern regions of Cuba ( Fig. 10 ), from sea level to 900 m a.s.l.

Etymology. We take pleasure in naming this species after Dr. Wolfgang Feichtinger from the Biocenter of the University of Würzburg ( Germany), for his enthusiastic contributions during field work and improvements in the herpetological work in Cuba.

Natural history. In the Sierra de Cubitas and surrounding plains, Eleutherodactylus feichtingeri n. sp. and E. varleyi are primarily associated with herbaceus vegetation. Most specimens were collected in an open savanna on serpentine rock, a vegetation complex known as cuabal ( Fig. 11 A). In forested areas of Cubitas, the new species also inhabits the leaflitter and grass stratum of a semideciduous forest on limestone. Through its distributional range, this frog is very common around disturbed habitats, occupying pastures, sugar cane plantations, other agricultural lands, and gardens. In higher mountains, like the Gran Piedra (Sierra Maestra, Santiago de Cuba; ~ 900 m a.s.l.), E. feichtingeri is found in the grass-covered road borders and open herbaceus situations within tropical rain forests.

Males call mainly from grass ( Fig. 11 B) and vocal activity is more intense after rains. The species has been heard throughout the year at many localities in the mountains of the eastern part of the island, or seasonally (April to December) in the driest lowlands. Sometimes dense choruses occur late in the afternoon (~16:00–19:00 hours). Vocal activity gradually decreases after midnight, and a second peak occurs shortly before dawn. Males call either at a very close distance from each other (0.5–1.0 m) or several meters appart. At Marea de Limones, Cabo Cruz, 16 calling individuals were collected in less than 100 m 2 at around 20:00 hours (7 October 2010). Vocalizing frogs were found in the following situations: (1) directly on the ground, (2) hidden in the base of grass clumps, (3) exposed at different levels on the grass, (4) on broad leaves of vines (exposed or hidden between two leaves), (5) on small bush leaves 0.2–1m high, or (6) inside a rolled leaf on the ground of the forest. Typically, males call in a horizontal position. Specimen MNHNCu 1226 contains a large encysted nematode between the arm and the chest cavity, measuring 17.3 mm when distended (frog SVL = 16.4 mm).

Figures

FIGURE 2. Plotting graph of Tympanum width vs. head length in samples (by population) of two closely related species of Cuban Eleutherodactylus.

FIGURE 3. Variation in the size of the tympanum and supratympanic stripe in some specimens of Eleutherodactylus varleyi (A-I) and E. feichtingeri n. sp. (J-R). See Diagnosis to understand the figure. (A) Holotype of E. varleyi (MCZ 10601), from Soledad, Cienfuegos Province (only tympanum size is informative, the specimen is discolored); (B) MNHNCu 1231 and (C) MNHNCu 1232, both from Sierra de Cubitas, Camagüey Province; (D) LMD 279, Sabanas Llanas, Alturas de Pizarras del Sur, Pinar del Río Province; (E) MNHNCu 613, Cueva Cheta, Majagua-Canteras, Sierra de San Carlos, Pinar del Río Province; (F) MNHNCu 1235, Campismo La Chorrera, Artemisa Province; (G) Boca de Canasí, Mayabeque Province; (H) LMD 251 and (I) LMD 252, Jardín Botánico de Cienfuegos (= Soledad), Cienfuegos Province; (J) Holotype of E. feichtingeri n. sp. (Sierra de Cubitas, Camagüey Province); (K) MNHNCu 1193 and (L) MNHNCu 1192, paratopotypes; (M) MNHNCu 1195 and (N) MNHNCu 1210, Marea de Limones, Cabo Cruz, Granma Province; (O) MNHNCu 1224, Guantánamo City (cross-road to El Salvador), Guantánamo Province; (P) MNHNCu 1217, Gran Piedra, Sierra Maestra, Santiago de Cuba Province; (Q) MNHNCu 1219, Santiago de Cuba City (airport), Santiago de Cuba Province; (R) MNHNCu 1227, San Rafael, Yateras, Guantánamo Province.

FIGURE 4. Color patterns of Eleutherodactylus feichtingeri n. sp. and E. varleyi (drawings from Díaz and Cádiz, 2008). See text for pattern descriptions.

FIGURE 5. Oscillograms and sonagrams (A – D) of advertisement calls of two cryptic species of Cuban Eleutherodactylus: E. feichtingeri n. sp. (A), from the type locality, voucher male MNHNCu 1181 (holotype), air temperature: 27.4 ºC, 01: 00 h, 6 Aug. 2010; call variation of E. varleyi (B – D), same locality, voucher male MNHNCu 1230, 27.4 ºC, 01: 23 h, 6 Aug. 2010. (E) Occurrence of one, two, and three note calls in sound recordings (two minutes long) of male Eleutherodactylus varleyi and E. feichtingeri n. sp. from the same localities listed in Table 2. Males with shorter periods of recording were not included in the graph. Voucher specimens: Eleutherodactylus varleyi (excepting Jardín Botánico Nacional, La Habana): Sierra de Cubitas: MNHNCu 1230; Jardín Botánico de Cienfuegos: LMD 266 – 269, 22 – 23 ° C. Eleutherodactylus feichtingeri n. sp.: Sierra de Cubitas: MNHNCu 1181, 1184, and 1189, 27.4 – 27.9 ° C; Marea de Limones, Cabo Cruz: MNHNCu 1208 and 1209, 26 ° C; Gran Piedra: MNHNCu 1213, 1214, and 1216, 20.6 ° C

FIGURE 6. Phylogenetic tree of Cuban grass frogs, Eleutherodactylus varleyi and its close relative E. feichtingeri n. sp. The tree was constructed from DNA sequences of the mitochondrial Cyt-b gene, using maximum likelihood. Bootstrap support values are indicated on nodes. In the tree, the holotype of E. feichtingeri is GenBank JN 015155.

FIGURE 7. Conventionally (Giemsa) stained karotype of a male specimen of Eleutherodactylus varleyi collected in Soroa, Sierra del Rosario (Artemisa Province). Note the two small metacentric chromosomes 1 and 2 and the 14 telo- or subtelocentric chromosomes 3 – 16 of continuously decreasing sizes.

FIGURE 8. C-banded karyotypes of (A) a male of Eleutherodactylus feichtingeri n. sp. specimen from Santo Domingo, Sierra Maestra (Granma Province) and (B) a female specimen of E. varleyi from Soroa, Sierra del Rosario (Artemisa Province). Note the distinct interstitial C-bands in the larger telocentric chromosome pairs 3 – 8 in the karyotype of the individual from the Santo Domingo population.

FIGURE 9. Silver-stained nucleolus organizer regions (NORs) in the karyotypes of (A) a male and (B) a female specimen of Eleutherodactylus feichtingeri n. sp. collected in Santo Domingo, Sierra Maestra (Granma Province), as well as (C) a male specimen of E. varleyi from Soroa, Sierra del Rosario (Artemisa Province). Note the two NOR cytotypes in the Santo Domingo population and the deletion of one of the homologous NORs in the karyotype of the specimen from the Soroa population.

FIGURE 10. Distribution of E. varleyi (triangles) and E. feichtingeri n. sp. (circles). Numbered localities are those where type specimens and tissue samples were collected; those not numbered are accoustic records: (1) surroundings of Hotel Zaza, Sancti Spiritus Province; (2) Paso de Lesca, Sierra de Cubitas, Camagüey Province; (3) Marea de Limones, Cabo Cruz, Granma Province; (4) Los Muertos, Alegría de Pío, Niquero, Granma Province; (5) Minas del Frío, Sierra Maestra, Granma Province; (6) Santo Domingo, Sierra Maestra Granma Province; (7) Hotel Sierramar, Sevilla, Guamá, Santiago de Cuba Province; (8) La Tabla, Sierra Maestra, Santiago de Cuba Province; (9) 4.5 km S Cruce de los Baños, Tercer Frente, Santiago de Cuba Province; (10) Ciudad de Santiago de Cuba, Santiago de Cuba Province; (11) Providencia, 13 km WNW of El Caney, Santiago de Cuba Province; (12) 1.2 km NNW of Nuevo Mundo, mainroad to La Caoba, San Luis, Santiago de Cuba Province; (13) Gran Piedra, Sierra Maestra, Santiago de Cuba Province; (14) Santa Rita, Guantánamo Province; (15) El Salvador, Guantánamo Province; (16) City of Guantánamo, Guantánamo Province; (17) Arroyón, San Antonio del Sur, Guantánamo Province; (18) El Palmar, Guantánamo Province; (19) San Rafael, Yateras, Guantánamo Province; (20) La Munición, Guantánamo Province; (21) Loma de La Mensura, Holguín Province.

FIGURE 11. Sierra de Cubitas and surrounding plains (A), Camagüey Province, where Eleutherodactylus feichtingeri n. sp. and E. varleyi are primarily associated with herbaceus vegetation. Calling male (B) of E. feichtingeri from Marea de Limones, Cabo Cruz, Granma Province; note the distended vocal sac. Photos: L. M. Díaz.

Tables

TABLE 1. Morphological measurements (in mm) of Eleutherodactylus feichtingeri n. sp., and E. varleyi. Values are means and ranges (in parentheses). Only measurements from adult males are provided.

	(N=40)		(N=40)
Snout-vent length	14.7 (13.3–17.4)	15.2	14.1 (12.2–18.5)
Head length	5.2 (4.6–5.9)	5.4	5.4 (4.6–7.0)
Head width	4.9 (4.4–5.8)	5.3	5.1 (4.2–6.6)
Snout length	2.1 (1.8–3.0)	2.1	2.1 (1.8–3.0)
Upper eyelid width	0.9 (0.8–1.2)	0.8	1.1 (0.8–1.4)
Interocular distance	1.7 (1.3–2.1)	2.1	1.5 (1.2–2.4)
Internarial distance	1.2 (1.0–1.5)	1.3	1.2 (1.0–1.5)
Tympanum width	0.6 (0.4–0.7)	0.7	1.1 (0.8–1.7)
Tympanum height	0.7 (0.3–1.0)	1.0	1.2 (1.0–1.6)
Thigh length	5.6 (4.7–6.5)	6.4	5.8 (4.7–7.5)
Shank length	5.9 (4.9–7.0)	6.0	6.1 (5.5–8.3)
Tarsal length	3.8 (3.1–4.6)	4.0	4.0 (3.4–5.0)
Foot length	5.9 (4.7–7.3)	5.5	5.9 (5.1–8.3)
Hand length	2.8 (2.3–3.5)	2.9	3.0 (2.3–3.9)
Eye-tympanum distance	0.4 (0.3–0.6)	0.3	0.4 (0.1–0.7)
Eye diameter	1.8 (1.6–2.1)	2.0	1.8 (1.4–2.8)
Eye-naris distance	1.2 (1.0–1.5)	1.3	1.4 (1.1–2.0)
Finger disc (III) width	0.3 (0.2–0.4)	0.3	0.3 (0.2–0.4)
Toe disc (IV) width	0.4 (0.3–0.5)	0.3	0.4 (0.2–0.5)

TABLE 2. Variation of some acoustic variables in Eleutherodactylus feichtingeri n. sp., and E. varleyi. Values are means and

Sierra de Cubitas 0.019 N1=4; N2=40 (0.008–0.028) 27.4–27.9ºC	0.003 (0.002–0.004)	3.0 (1.7–4.7)	27.1 (21.4–37.0)	3.9 (3.7–4.5)
Marea de Limones, 0.032 Cabo Cruz (0.021–0.041) N1=3; N2=30 26ºC	0.004 (0.003–0.006)	2.4 (1.8–3.1)	27.9 (23.0–31.6)	3.7 (3.2–3.9)
Gran Piedra 0.032 N1=4; N2=40 (0.020–0.041) 20.6ºC	0.004 (0.002–0.006)	2.0 (1.5–3.1)	32.8 (26.2–35.7)	3.7 (3.5–3.9)