Holotype.—

MNRJ 21356, adult male, collected at Parque Estadual do Ibitipoca (21°43′S, 43°55′W; 1180 m altitude), Conceição do Ibitipoca, Municipality of Lima Duarte, State of Minas Gerais, Brazil, 19–21 November 1987, by U. Caramaschi.

Paratopotypes.—

MNRJ 21347, adult male, collected 4–7 November 1986, by R. N. Feio; MNRJ 21348–21349, adult males, collected 17–18 November 1986, by R. N. Feio; MNRJ 21350–21352, adult males, collected 19–23 January 1987, by R. N. Feio; MNRJ 21353 and 21355, adult males, and MNRJ 21354, adult female, collected with the holotype; MNRJ 21364, adult male, collected 6–9 December 1986, by U. Caramaschi, R. N. Feio, M. C. Britto Pereira, and H. R. Silva; MNRJ 21365–21366, adult males, collected 16–20 March 1987, by R. N. Feio; MNRJ 24826 and 24828, adult males, and MNRJ 24827, adult female, collected 26–28 October 1999, by M. F. Napoli, R. N. Feio, U. Caramaschi, J. P. Pombal, L. B. Nascimento, and C. A. G. Cruz; MNRJ 24842, adult male, collected 01 November 1987, by R. N. Feio; MNRJ 26285–26286, 26288–26297, adult males, and MNRJ 26287, adult female, collected 15–20 January 2001, by J. P. Pombal, C. A. G. Cruz, and R. N. Feio.

Diagnosis and comparison with other species.—

Species characterized by the following combination of traits: small size (SVL 37.0–40.3 mm in adult specimens); head longer than wide, its width 2.8–3.0 times smaller than SVL; tympanum medium size, its diameter 1.6–2.0 times smaller than eye diameter; thigh and tibia lengths 0.48– 0.51 and 0.48–0.52 of SVL, respectively; a dorsolateral dark brown stripe from tip of snout to inguinal region; flanks and anterior/posterior faces of thighs immaculate; dorsum light to medium brown in preservative and reddish brown in life, with distinct transverse dark brown bars; distinct vocal slits in adult males.

A dorsolateral blackish brown stripe from tip of snout to inguinal region distinguishes H. feioi from all other members of the group, which never have such a pattern (some specimens of H. nanuzae have a vestigial stripe). Hyla feioi lacks transverse brown stripes on the flanks and anterior and posterior faces of thighs, while in the other taxa of the group with similar size they often occur (exception within H. nanuzae, in which the stripes may be absent or only present on the superior third of the posterior face of thighs). Hyla feioi is smaller than H. circumdata, H. carvalhoi, H. hylax, H. gouveai, H. luctuosa, and H. ahenea sp. nov. (combined SVL 45.9–71.6 mm in adult specimens); H. feioi is larger than H. sazimai (SVL 30.1–35.1 mm in adult males). The tympanum diameter of H. feioi (ED/TD 1.6–2.0) is smaller than in H. circumdata, H. izecksohni, H. luctuosa (ED/TD 1.0–1.5), H. gouveai, and H. ravida (ED/TD 1.2–1.6). The narrower head distinguishes H. feioi (SVL/HW 2.8– 3.0) from H. ravida (SVL/HW 2.5–2.6), H. luctuosa, and H. circumdata (SVL/HW 2.5–2.8). The shorter thigh and tibia lengths distinguish H. feioi (THL/SVL 0.48–0.51; TL/SVL 0.48– 0.52) from H. izecksohni and H. ravida (THL/ SVL 0.51–0.55; TL/SVL 0.50–0.55). The internarial distance is greater than eye–nostril distance in H. feioi, whereas equal or smaller than in H. sazimai. The presence of vocal slits in adult male H. feioi distinguishes this species from H. izecksohni, which lacks such structures. In life, the dorsum is reddish brown with transverse dark brown bars in H. feioi, whereas in H. ibitipoca and H. sazimai, it is cream color, the latter with reticulated thin brown stripes and random brown dots over it; also, H. gouveai, H. carvalhoi, and H. ahenea sp. nov. have immaculate dorsa, and H. nanuzae lacks distinct transverse brown bars over it. Hyla feioi and H. nanuzae are morphologically very similar but are distinguished from each other by differences in color patterns and number of tooth rows in tadpoles, and by having different advertisement calls.

Description.—

The description is based on adult males as only two adult females were sampled. Descriptive statistics are presented in Appendix 1. Small species (SVL 37.0–40.2 mm). Head length larger (80 %), equal (13.3 %), or smaller (6.6%) than head width, the latter 2.8–3.0 times smaller than snout–vent length; internarial distance smaller than eye–nostril distance and eye diameter, the latter larger than eye–nostril distance; snout rounded or acuminate in dorsal outline, and rounded in lateral outline; canthus rostralis distinct, loreal region oblique; tympanum medium-sized, nearly circular, its width 1.6–2.0 times smaller than eye diameter; supratympanic fold weakly visible, covering partially the tympanum; nostrils directed laterally and slightly forward; internarial region straight, not furrowed; vomerine teeth present in two angular patches between choanae; tongue cordiform; vocal sac single, subgular; vocal slits distinct. Forearm hypertrophied, with a weak, smooth dermal fringe; third finger disk diameter larger than fourth toe disk; subarticular tubercles rounded; supernumerary tubercles present; palmar tubercle absent; prepollex large, curved, single (not bifid), and with protruding prepollical spine; modal webbing formula, I 2 3/4–2 3/4 II 2⁺–3⁺ III 3⁻–2⁺ IV. Thigh and tibia of about the same length, their sum nearly equals the snout–vent length; thigh and tibia lengths 0.48–0.51 of snout–vent length; foot with weak, smooth dermal fringe and calcar; subarticular tubercles rounded; supernumerary tubercles distinct; inner metatarsal tubercle ovoid; modal webbing formula, I 1 1/2–2 II 1⁺– 2⁺ III 1⁺–2 1/2 IV 2⁺–1⁺ V. Belly, chest, and ventral surfaces of thighs glandular; dorsal surfaces smooth.

Appendix 1. Descriptive Statistics of Adult Males and Females of Hyla feioi and Hyla ahenea

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Color in life.—

Dorsum reddish-brown, with transverse dark brown bars; a blackish brown dorsolateral stripe from the tip of snout over eyes extending to inguinal region; superior half of flanks with dark brown blotches, while inferior half immaculate; proximal and distal faces of flanks purple and pale yellow, respectively; throat and chest pale yellow; belly vivid yellow; anal region cream; superior faces of forearms, upper arms, hands, thighs, shanks, and feet, reddish brown with transverse dark brown bars; hidden areas of forearms and upper arms purple; anterior and posterior faces of thighs reddish brown and immaculate; hidden areas between thighs and flanks purple; ventral surfaces of hands and feet reddish brown; iris orange, surrounded with a black halo; pupil horizontal and black.

Color in preservative.—

Dorsal surfaces medium to light brown; dorsum with dark brown transversal bars; a weak brown vertebral stripe may be present from the tip of snout to the middle of dorsum; some specimens with distinct white blotches on dorsum, elbow, knee, and/or above anus; canthus rostralis delimited by a subcanthal dark brown stripe; a slender blackish brown dorsolateral stripe from the tip of snout to inguinal region; flanks immaculate; anterior and posterior faces of thighs pale cream and immaculate; superior faces of thighs, tibia, and feet with transverse brown bars; ventral surfaces pale cream.

Measurements of holotype.—

SVL 37.7, HL 13.6, HW 13.3, ED 4.4, END 3.5, TD 2.4, UEW 3.2, IOD 4.6, IND 2.9, NSD 2.0, THL 19.4, TL 18.7, FL 27.4, 3FD 2.0, 4TD 1.8.

Sexual dimorphism.—

Adult males of H. feioi have hypertrophied forearms, enlarged prepollices, and vocal slits, while adult females do not have enlargement of forearms and prepollices, and lack vocal slits. The small number of adult females in samples (n = 2) did not allow morphometric analyses.

Natural history.—

Hyla feioi occurs in gallery forests along creeks surrounded by rocky mountain fields called “campo rupestre” (for description of “campo rupestre,” see Rizzini, 1979). One specimen captured by the first author was calling alone on a branch that was standing upright 1 m from the water. R. N. Feio (unpubl. data) observed specimens of H. feioi often on bushes along the brooks, between 0.2 and 2.0 m from the ground, some of them calling from hollow surfaces such as bark and epiphytic bromeliads, and often far from each other; reproductive activity was observed during the wet season (October to March). Males of H. feioi when handled by the first author jabbed their prepollical spine into his thumb and then showed tanathosis.

Advertisement call.—

Call (Fig. 3A) with multiple pulsed notes, with unequal duration and intervals; note duration is 0.32–0.35 sec. First pulse separated from the second pulse by nearly 0.05 sec; second pulse to fourth separated from each other by 0.005–0.01 sec; the last pulse group comprises several pulses very close to each other (about 30 overlaid pulses), and has a duration of nearly 0.15 sec. Dominant frequency between 1.9–2.4 kHz. Apparent frequency modulation in the last overlaid pulses. Distinct sidebands in the last pulse group.

The call note structure of H. feioi is distinguished from that published for H. sazimai, where the latter has the frequency amplitude of the first pulse greater than the others, the last part of the note with about fifteen overlaid pulses, absence of intermediate pulses between the first and the last overlaid ones, presence of isolated series of pulses within unequal intervals, and dominant frequency between 2.3–2.8 kHz (Cardoso and Andrade, 1982). The call note structure of H. feioi differs from that published for H. astartea mainly by the greater number of intermediate pulses between the first (isolated pulse) and the last pulses, and by having a great number of overlaid pulses in the end of the note (Heyer et al., 1990). The call structure of H. nanuzae differs from H. feioi mainly by the greater number of note groups (2–4), and duration of each note between 0.17–0.2 sec (Bokermann and Sazima, 1973). The absence of new recordings of H. nanuzae did not make possible a more detailed comparison between these calls. The call structure of H. feioi also differs markedly from the syntopic H. ibitipoca (see below).

Description and habits of tadpoles.—

Description based on two tadpoles in stage 36 (Gosner, 1960) collected at the type-locality (MNRJ 25167). Body (Fig. 4A–C) oval in dorsal and ventral views and widest posteriorly; eyes dorsolateral, interorbital distance 3.3–3.4 times the eye diameter; nostrils rounded and dorsal, 1.1 times nearer eye than tip of snout; tube of the sinistral spiracle short, directed posterodorsally, its opening 1.1–1.3 times nearer tip of snout than the anal opening; distance from spiracle to snout 33.5–35.0% of body length; anal tube short, dextral, attached to ventral fin; tail length 61.8–63.3% of total length, its height 1.3 times the body height; dorsal fin slightly arched and ventral fin relatively straight; tail musculature tapering gradually to the rounded tail tip. Oral disc (Fig. 4D) anteroventral, its width 32.9– 35.7% of body width, bordered by two rows of rounded papillae, without agglomeration laterally; tooth row formula 2(2)/4(1); jaw sheaths developed, serrated; anterior jaw sheath slightly arched and posterior jaw V-shaped.

In life, dorsal surfaces dark brown marbled with blackish and light brown areas; in front of each eye a golden dot; caudal musculature light brown marbled with golden areas; dorsal and ventral fins dark brown with scattered transparent vermiculations, which are presented as light golden areas. Belly reddish-brown, scattered with golden punctations that are not well distinct; the area over the intestine dark brown, making the coils invisible. Mouth not melanized, with two distinct white dots near the superior lip; spiracle opening bordered by a white aureole; iris golden; pupil black. The color patterns described herein are more evident in earlier stages of development. Early after metamorphosis, the blackish-brown dorsolateral stripe, diagnostic for adult specimens of H. feioi, becomes evident. In preservative, the general color is the same, with the exception of white and golden areas that become pale cream.

Tadpoles of H. feioi were collected in brooks that run over rocky bottoms, with riffles and pools throughout gallery forests, and showed only nocturnal activity. Tadpoles of H. feioi are marbled with blackish and light brown areas, grayish-brown in H. circumdata, H. luctuosa, and H. sazimai and homogeneous black in H. nanuzae; oral disc in H. feioi with four lower rows of teeth, five in H. carvalhoi and H. nanuzae.

Measurements of tadpoles (stage 36; n = 2, in mm).—

Total length 52.2–54.6, body length 19.9–20.0, body width 15.4–16.7, body height 12.2–12.5, dorsal fin height 6.5–7.5, ventral fin height 4.5, tail musculature height 6.5–7.5, interorbital distance 7.7–7.9, internostril distance 0.3–0.7, eye diameter 2.3, nostril diameter 0.6– 0.7, nostril-tip of snout distance 2.3–2.5, eye– nostril distance 2.1–2.3, spiracle-tip of snout distance 6.6–7.0, spiracle-anus distance 7.5–9.0, mouth width 5.5.

Etymology.—

The name is a noun in the genitive case honoring Renato Neves Feio, for his contribution to the knowledge of anurans from Parque Estadual do Ibitipoca and full-time pleasant companionship.

Geographic distribution.—

Hyla feioi is known only from type-locality, in the southern portion of Serra da Mantiqueira (Fig. 5).

Holotype.—

MZUSP 93493, adult male, collected at Campo de Fruticultura da Bocaina (22°38′S, 44°34′W; 510 m altitude), Municipality of São José do Barreiro, State of São Paulo, Brazil, November 1968, by W. C. A. Bokermann, F. M. Oliveira, and O. Oliveira.

Paratopotypes.—

MNRJ 25437–38, adult males, and 25439, adult female, collected with the holotype; MZUSP 93465, 93467, 93469–73, adult males, and 93466, 93468, 93474, adult females, collected November 1965, by F. M. Oliveira, O. Oliveira, C. A. C. Seabra, and W. C. A. Bokermann; MZUSP 93478–79, adult males, and 93480–81, adult females, collected November 1965, by F. M. Oliveira, O. Oliveira, C. A. C. Seabra, and W. C. A. Bokermann; MZUSP 93482–84, adult males, collected November 1967, by C. A. C. Oliveira, F. M. Oliveira, and O. Oliveira; MZUSP 93485–86, 93488–90, 93492, 93495, 93497–98, 93500–501, adult males, and 93487, 93496, adult females, collected with the holotype; MZUSP 93502–503, 93505–508, adult males, and 93504, adult female; MZUSP 93509–12, 93514–20, 93522–25, 93527–28, 93530–35, 93537–40, adult males, 93513, 93521, 93526, 93536, adult females, and 93529, juvenile; MZUSP 93541, adult female; MZUSP 93543, adult male, and 93542, 93544– 45, adult females, collected January 1969, by M. Alvarenga; MZUSP 93546, adult male; MZUSP 93597–98, 93600, adult males, and 93599, adult female, collected November 1965, by F. M. Oliveira, O. Oliveira, C. A. C. Seabra, and W. C. A. Bokermann.

Diagnosis and comparison with other species.—

Species characterized by the following combination of traits: small to medium size (SVL 45.9–56.7 mm in adult specimens); head longer than wide, its width 2.6–3.0 times smaller than SVL; tympanum small, its diameter 1.6–3.1 times smaller than eye diameter; flanks and anterior, superior, and posterior faces of thighs with dark brown transversal stripes, without additional thin stripes between them; in preservative, dorsal surfaces bearing copper or bronze; distinct vocal slits in adult males.

Hyla ahenea is larger than H. astartea, H. feioi, H. ibitipoca, H. nanuzae, H. sazimai (combined SVL 30.1–42.5 mm), and H. ravida (SVL 30.1– 47.6 mm); H. ahenea is smaller then H. carvalhoi, H. gouveai (combined SVL 60.0–70.4 mm), H. circumdata, and H. luctuosa (combined SVL 55.2–71.6 mm). The small tympanum distinguishes H. ahenea (ED/TD 1.6–3.0) from H. izecksohni, H. circumdata, H. luctuosa (ED/TD 1.0–1.5), H. gouveai, and H. ravida (ED/TD 1.2– 1.6). The presence of vocal slits in adult males of H. ahenea distinguish them from H. izecksohni, which lacks such structures. Dorsum always immaculate in H. ahenea, with transverse brown bars in H. circumdata, H. feioi, H. hylax, H. ibitipoca, H. izecksohni, H. luctuosa, H. ravida, and H. sazimai. Anterior and posterior faces of thighs with transverse brown stripes in H. ahenea, immaculate in H. feioi, with thin and fragmented stripes in H. carvalhoi, and with thin stripes among the transverse brown bars in H. hylax.

Description.—

The descriptive statistics are presented in Appendix 1. Medium-sized species (SVL 45.9–56.7 mm). Head length smaller (males 84.8%; females 89.4%), larger (only males, 13.6%), or equal (males 1.5%; females 10.5%) to head width, the latter 2.6–3.0 times smaller than snout–vent length; internarial distance smaller than eye–nostril distance and eye diameter, the latter greater than eye–nostril distance; snout rounded or slightly acuminate in dorsal and lateral outlines; canthus rostralis distinct, loreal region slightly concave to oblique; tympanum small, nearly circular, its width 1.4– 3.1 times smaller than eye diameter in males and 1.6–2.5 times in females; supratympanic fold weakly visible, covering partially the tympanum; nostrils directed laterally and slightly forward; internarial region straight, not furrowed; vomerine teeth in two angular patches between choanae; tongue ovoid or cordiform; vocal sac slightly bilobate, subgular; vocal slits distinct. Forearm hypertrophied in males, with a weakly crenulated dermal fringe; third finger disk diameter equal to or greater than fourth toe disk; subarticular tubercles rounded, distal subarticular tubercle of fourth finger round (90.7%) or bifid (9.3%); supernumerary tubercles distinct; palmar tubercle slightly developed, divided; prepollex large, curved, single (not bifid), and with protruding prepollical spine; modal webbing formula, I 3–3 II 2–3 1/4 III 3⁻– 2⁺ IV. Thigh and tibia with about the same length, their sum nearly equaling the snout– vent length; in males, thigh and tibia lengths 0.42–0.55 of snout–vent length, and 0.44–0.53 in females; subarticular tubercles rounded; supernumerary tubercles distinct; inner metatarsal tubercle ovoid; modal webbing formula, I 2⁻–2⁺ II 1⁺–2 1/2 III 1 1/2–2 1/2 IV 2 1/2–1 1/2 V. Belly glandular; chest, throat, and ventral surfaces of arms smooth; posteroventral surfaces of thighs slightly glandular.

Color in preservative.—

Dorsum bronzed, scattered with dark brown punctations. Posterior faces of flanks with transverse brown stripes (range 0–9; mode 4). Anterior and posterior faces of thighs dark brown, the former immaculate and the latter with 6–10 transverse dark brown stripes, not dichotomized, and without intercalary thin stripes; superior faces of thighs grayish brown, with transverse dark brown stripes. Ventral surfaces of shanks and feet dark brown; thighs and belly cream; chest, arms, hands, and throat, cream with strong melanization. Lorus, superior lip, and snout, blackish brown.

Measurements of holotype.—

SVL 53.1, HL 18.6, HW 18.7, ED 5.6, END 4.8, TD 2.8, UEW 4.0, IOD 5.6, IND 3.3, NSD 2.9, THL 24.6, TL 26.9, FL 37.5, 3FD 2.8, 4TD 2.5.

Sexual dimorphism.—

Adult males of H. ahenea have hypertrophied forearms, enlarged prepollices, and vocal slits, characters absent in adult females. To summarize the degree of morphometric differences between males and females we used principal component analysis. Only adult specimens were sampled. The projection of the individual scores of both sexes resulted in a severe superposition, denoting absence of morphometric differences between males and females.

Geographic distribution.—

Hyla ahenea is known only from the type-locality, in the Serra do Mar (reaching about 1500 m above sea level; Fig. 5), in the Atlantic Forest Morphoclimatic Domain (Ab'Sáber, 1977).

Etymology.—

The specific name, a Latin adjective (aheneus = cooper or bronze), is an allusion to the bronze aspect of dorsal surfaces of preserved specimens.

Comments.—

Hyla ahenea is known only from preserved specimens, lacking any additional information on natural history, advertisement call, and tadpoles.

Advertisement call of Hyla ibitipoca.—

Call with pulsed notes, with unequal duration and intervals, and crescent frequency amplitude (Fig. 3B). Each call comprises one to four notes. Calls with one note have a duration of 0.13–0.20 sec. Calls with two notes have a duration of 0.18– 0.20 sec; duration of the first note nearly 0.09 sec; duration of the second note nearly 0.11 sec. Calls with three notes have a duration of 0.24– 0.26 sec; duration of first note nearly 0.04 sec; duration of second note nearly 0.06 sec; duration of third note nearly 0.11 sec. Calls with four notes have a duration of 0.30–0.36 sec; duration of first note nearly 0.05 sec; duration of second note nearly 0.07 sec; duration of third note nearly 0.06 sec; duration of fourth note nearly 0.11 sec. Dominant frequencies between 0.4–0.6 and 1.7–1.9 kHz. Of 27 notes analyzed, 16 had one pulse group, three had two pulse groups, two had three pulse groups, and six had four pulse groups. Distinct sidebars were present and frequency modulation occurred in the end of the last note.

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The advertisement call of H. ibitipoca is easily distinguished from the syntopic H. feioi by the lower dominant frequency (respectively: 0.4–0.6 kHz and 1.7–1.9 kHz; 1.9–2.4 kHz), and by having calls with one to four notes with duration and arrangement different from that observed in H. feioi (Fig. 3A).