Eleutherodactylus grunwaldi, Reyes-Velasco, Jacobo, Ahumada-Carrillo, Ivan, Burkhardt, Timothy R. & Devitt, Thomas J., 2015

Reyes-Velasco, Jacobo, Ahumada-Carrillo, Ivan, Burkhardt, Timothy R. & Devitt, Thomas J., 2015, Two new species of Eleutherodactylus (subgenus Syrrhophus) from western Mexico, Zootaxa 3914 (3), pp. 301-317 : 303-309

Eleutherodactylus grunwaldi , new species

Holotype. MZFC 27472. Adult male, collected by Jacobo Reyes-Velasco and Alexander Hermosillo-Lopez on July 17, 2011, 3.5 km ESE of El Sauz, on road to El Terrero, Municipality of Minatitlán (19.43161 N, - 103.97871 W, 1,329 m; datum = WGS84), Colima, Mexico ( Fig. 1 ).

Paratypes. MZFC 27467–27471, five adult males, collected along the side of a dirt road between El Sauz and El Terrero, Municipality of Minatitlán, Colima, Mexico, collected by Chris I. Grünwald, on July 14, 2008; MZFC 27473–27475, two adult males and a young adult, dirt road between El Sauz and El Terrero, collected on the same day as the holotype by Jacobo Reyes-Velasco and Alexander Hermosillo-Lopez; MZFC 27483, adult male, 6.7 km SW of El Terrero, on road to El Sauz, collected by Jacobo Reyes-Velasco and Gabriela Zamora-Silva on July 15, 2012 ( Fig. 2 ).

Diagnosis. Eleutherodactylus grunwaldi is a member of the E. longipes species series of the subgenus Syrrhophus as defined by Hedges et al. (2008). It is one of the largest members of the species series, with adult males measuring 28.4–32.4 mm SVL. Vocal slits are present in males; digital discs are greatly expanded, usually three times the width of narrowest part of digit on fingers three and four ( Fig. 3 A); shape of digital discs is similar to the condition seen in E. longipes in Lynch (1970; Fig. 1 F), which were described as triangular; first finger shorter than second finger; snout angular in dorsal view (as defined by Savage, 2002 p. 171), and acuminate in profile; the head slightly longer than wide; tympanum width 40% –50% of eye width, never more than 50%; body robust but thinner than head; no compact lumbar glands; dorsal coloration consisting of greenish or yellowish irregular blotches, spots, or reticulations on dark background; no mid-dorsal stripe or light interorbital bar are present; dorsal and ventral skin smooth; ventral coloration white; no dark or pale interorbital bar or bars on thighs; tympanum medium to small, diameter of tympanum maximum 50% of diameter of eye; iris copper-green.

Eleutherodactylus grunwaldi differs from the other members of the subgenus, with the exception of E. dennisi , E. longipes and E. saxatilis by the presence of digital discs on the hands which are three times the width of the narrowest part of the digit ( Fig 3 A). It can readily be distinguished from E. saxatilis by the absence of compact lumbar glands, as well as broader digital discs on the fingers, usually double the narrowest width of the third and fourth digits in E. saxatilis and from two and a half times to three times the width of the narrowest part of the third and fourth digits in E. grunwaldi . This species can be distinguished from E. dennisi and E. longipes by having a smaller tympanum to eye ratio, always 50% or less, as opposed to 50–65% in male E. dennisi and 60–90% in male E. longipes ; E. dennisi also has a conspicuous light interorbital bar, which E. grunwaldi lacks. While E. longipes has a light ground color with darker blotches and/or spots, and a dark brown interorbital bar or triangle, E. grunwaldi shows a dark ground color with yellowish marbling or reticulations and no interorbital marks. The new species is distinguished from most other members of Syrrhophus , except for E. saxatilis and some members of the E. marnockii species group of Hedges et al. (2008) by its dorsal coloration.

This species has been collected in sympatry with Eleutherodactylus nivicolimae and E. modestus , and at slightly higher elevation than E. nitidus . Eleutherodactylus grunwaldi differs from E. nivicolimae in size, with males over 28 mm in E. grunwaldi and under 23.5 mm in E. nivicolimae , as well as by having a dark gray ground color with yellow or yellowish-green spots and reticulations, whereas E. nivicolimae has a grayish, reddish or yellowish ground color without spots ( Fig. 4 A). Eleutherodactylus nivicolimae occasionally has a pale middorsal stripe, which is not present in E. grunwaldi . Eleutherodactylus modestus is smaller, with males never reaching over 22 mm, and has a shorter, narrower head with a rounded snout (as defined by Savage, 2002). Eleutherodactylus modestus further differs from E. grunwaldi by having a reddish or orange ground color, which is covered in darker gray or black spots, dashes or reticulations ( Fig. 4 B–C), very distinct from the greenish or yellowish spots present in E. grunwaldi . The new species can be distinguished from E. nitidus by size (males> 28 mm in SVL vs. <27 mm in SVL in E. nitidus ); they also differ by the very expanded toe pads of E. grunwaldi , reaching more than twice the width of the narrowest part of the digit in the new species, but only to less than one and a half the width of the narrowest part of the digits in E. nitidus , and by the presence of compact lumbar glands in the latter species ( Fig. 4 D).

Description of the holotype. Male of moderate size (29.7 mm SVL); head as wide as long, 10.7 mm. in length and width, wider than body; snout truncate, angular, non-rounded from a dorsal view, acuminate from a lateral profile; lip slightly flared; tympanum rounded, upper edge indistinct, no supratympanic fold, greatest tympanum diameter 1.7 mm; greatest eye diameter 3.6 mm; tympanum-to-eye ratio 0.5; eyelid 2 mm wide, about half as wide as interorbital distance; first finger shorter than second; finger lengths from shortest to longest 1-2-4-3; digital pads on fingers three and four greatly expanded, approximately three times the narrowest point of the digit; three palmar tubercles, inner palmar tubercle just over a third as large as middle tubercle, outer palmar tubercle about a fifth as large as middle palmar tubercle; toe lengths from shortest to longest 1-2-5-3-4. Both dorsal and ventral skin texture smooth. FL 13.8 mm, TL 15.3 mm, FeL 13.5 mm, FoL 8.3 mm, IND 2.8 mm, IOD 4 mm, END 3.6 mm. FeL to SVL 45%, TL to SVL 52%, FL to SVL 46%, HL to SVL 36%, HW to SVL 36%. Dorsal, lateral and ventral skin texture smooth. Vocal slits present. See Figure 1 for a photograph of the holotype in life. Figures 5 A–B show the holotype in preservative.

Coloration in life pale gray on the lateral surfaces, with almost no pattern, while dorsal surfaces dark gray to blackish, with network of greenish-yellow botches forming reticulations on the back, head, neck, forelimbs and hind limbs; venter white; hands and feet pale gray without markings, though some faint markings present on hands; uninterrupted gray line running from snout through nares and lower half of eye, continuing past tympanum and axilla where it grades into lateral gray coloration. Coloration in preservative dark, ground coloration of varying shades of brown, with pale blotches sometimes forming uniformly pale grayish tan reticulations; ventral coloration is white.

Variation. Mensural variation is presented in Table 1 . This species shows remarkably little color variation when compared to some other species in the subgenus (e.g., E. modestus ). The color pattern consists of greenish yellow or yellowish blotches, spots or reticulations on a black ground color. The amount and intensity of the pale blotches varies such that some specimens appear to be dark frogs with pale colored blotches, while others have more intense reticulation.

Distribution and ecology. This species is known from the Sierra de Manantlán in the municipalities of Minatitlán, Colima, and Tolimán, Jalisco ( Fig. 6A ). It is probably distributed throughout the Sierra de Manantlán, and may range north into some of the other coastal sierras of Jalisco (Sierra de Cacoma, Sierra de Talpa). This species has been collected between 1300–2200 m in the Sierra de Manantlán Biosphere Reserve and at lower elevations (~ 800m) in the Grutas de Toxín in Jalisco. It inhabits tropical deciduous forest, Madrean pine-oak woodland and the ecotone between tropical deciduous forest and pine-oak woodland. Like E. saxatilis , this species appears to be strictly saxicolous and found almost exclusively on limestone outcrops. All specimens have been found on limestone, either in outcrops, sink holes or caves. A single specimen was found while calling from a tree, however the ground consisted of soil mixed with broken limestone. Frogs were collected during the months of June, July and August, which correspond to the beginning of the rainy season and when breeding appears to take place. One specimen was collected in the dry season (February) from inside a cave, and many other individuals were seen active on the walls of the cave. A photograph of the type locality is shown in Figure 6 B.

Etymology. The species epithet is a patronym for German-Mexican naturalist Christoph I. Grünwald, who collected the original specimen (now lost) on July 18th, 2005. This original specimen came from near 2200 m at 4.8 km E of El Terrero, Municipality of Minatitlán, Colima.

Figures

FIGURE 1. Holotype of Eleutherodactylus grunwaldi sp. nov. in life. MZFC 27472, UNAM. Field number JRV 139.

FIGURE 2. Paratypes of Eleutherodactylus grunwaldi, sp. nov. A) MZFC 27474, UNAM. B) MZFC 27473, UNAM.

FIGURE 3. Underside of the right hands of the holotypes of (A) Eleutherodactylus grunwaldi sp. nov., MZFC 27472, and (B) E. wixarika sp. nov., MZFC 27477. Hands are not drawn to scale.

FIGURE 4. Species of Eleutherodactylus, subgenus Syrrhophus from western México. A) E. nivicolimae, Volcán de Colima, Jalisco. B) E. modestus, adult male, Ixtlahuacán, Colima. C) E. modestus, subadult. Minatitlán, Colima D) E. nitidus orarius, Ixtlahuacán, Colima.

FIGURE 5. Holotypes in preservative. A) Dorsal photo of Eleutherodactylus grunwaldi sp. nov., MZFC 27472, (B) Ventral photo of Eleutherodactylus grunwaldi sp. nov., MZFC 27472, (C) Dorsal photo of Eleutherodactylus wixarika sp. nov., MZFC 27477, (D) Ventral photo of Eleutherodactylus wixarika sp. nov., MZFC 27477.

FIGURE 6. A) Map showing the type localities of Eleutherodactylus grunwaldi sp. nov. (white star) and E. wixarika sp. nov. (white triangle) in western Mexico. White circle represents an additional locality of E. grunwaldi in the state of Jalisco. B) Type locality of E. grunwaldi, 3.5 km ESE of El Sauz, on road to El Terrero, Municipality of Minatitlán, Colima. C) Type locality of E. wixarika, Bajio de los Amoles, Municipality of Mezquitic, Jalisco.

Tables

TABLE 1. Intraspecific morphological variation in Eleutherodactylus grunwaldi and E. wixarika. This table shows only a subset of all measurements taken, which showed to be the most useful for distinguishing between members of Syrrhophus. SVL = snout-vent length, TH = tympanum height, TW = tympanum width,), F 3 L = finger 3 length, F 3 PW = finger 3 pad width. Asterisk marks the type specimens.

Species	Catalogue #	SVL	HW	HL	SL
E. grunwaldi	MZFC 27467	29.8	11.1	10.2	9.6
E. grunwaldi	MZFC 27468	28.8	10.9	10.4	9.6
E. grunwaldi	MZFC 27470	28.1	10.6	9.8	9.5
E. grunwaldi	MZFC 27472*	29.7	10.8	10.7	10.2
E. grunwaldi	MZFC 27473	32.0	11.0	11.1	10.1
E. grunwaldi	MZFC 27474	32.4	11.4	10.7	10.3
E. grunwaldi	MZFC 27475	21.6	7.8	7.6	7.6
E. wixarika	MZFC 27477*	21.9	7.7	7.3	6.4
E. wixarika	MZFC 27478	21.2	7.8	7.2	6.7
E. wixarika	MZFC 27479	24.5	8.9	8.2	7.6