Hylodes cardosoi Lingnau, Canedo & Pombal 2008

Diagnosis: This species can be distinguished by the following combination of characters: large, robust body (adult males 35.6-44.1 mm SVL); dorsolateral skin densely granular, with large white tubercles; Toe V with moderate fringing on outer side; advertisement call consisting of 4-8 notes, each with 3-6 frequency-modulated pulses (rapid raise and decay); lack of light dorsolateral stripes; males with external vocal sacs; lack of blue or yellow dorsal spotting. H. asper is similar to H. cardosoi but has a different call (more notes per call and longer duration for H. asper); less frequently has a distinct pair of cloacal tubercles on either side of the vent (cloacal tubercles distinct: 7.9%, weak: 67.3%, lacking: 24.8% of H. asper specimens vs. cloacal tubercles distinct: 89.2%, weak: 8.1%, lacking: 2.7% of H. cardosoispecimens); has longer fringe on Toe V (H. asper); posterior thigh almost always brown with scattered cream blotches in H. asper, vs. brown with anastomosed cream blotches (56% of H. cardosoi specimens), or less frequently, brown with scattered cream blotches (22% of H. cardosoi specimens), or equal brown/cream patches (18.0% of H. cardosoi specimens). Description: The snout-vent length in males is 35.6 mm to 44.1 mm and in females is 36.7 mm to 46.5 mm. The head has a greater length than width. From above, the snout appears rounded and from the lateral view appears protruding, with nostrils directed laterally, a distinct, nearly straight canthus rostralis and a concave loreal region. The tympanum is distinct and is nearly round with a diameter slightly smaller than half the diameter of the eye. A supratympanic fold extends posteriorly from the edge of the orbit and stops before the shoulder. Lateral vocal sacs are well-developed in males. The large tongue is nearly ovoid, and free posteriorly. Vomerine teeth are present in two series between rounded choanae. Maxillary and premaxillary teeth are also present. A row of small, unpigmented tubercles is present on the margin of the upper lip in the male holotype, but never in females. The body is robust, with moderately robust arms and more robust legs. The outer metacarpal tubercle is very large and round, while the inner metacarpal tubercle is small and elliptical. The subarticular tubercles on the fingers are single and round, and the supernumerary tubercles are arranged in rows between fingers. The relative lengths of the fingers are II < I < IV < III. Lateral fringes on the fingers run from the proximal subarticular tubercle to the disc. On the outer margin of finger III is a vestigial fringe, running from the proximal subarticular tubercle to the distal one. The elliptical discs of the fingers are slightly expanded but smaller than the tympanum, with the disc on finger I being the smallest. Well-developed scutes are present on the upper surfaces of discs. The finger discs are as developed as the toe discs. On the legs, the protruding outer metatarsal tubercle is small and round, while the inner metatarsal tubercle is elongated. Subarticular tubercles are single, and can be either rounded or ovoid. The relative lengths of the toes is I < II < V = III < IV. The toes have extensive fringe, and the fringe along the outer margin of toe V extends beyond the end of the subarticular tubercle to a distance almost equal to the tubercle's diameter. Tarsal fringe is also extensive, and continuous distally with the toe fringe of toe I's inner margin on the male holotype. Female paratypes display more poorly developed fringes on the toes, fingers and feet. Slightly expanded toe discs are somewhat elliptical, with a smaller disc on toe V. Well-developed scutes are present on the upper surfaces of discs. The dorsal skin texture of the flanks and thighs are rugose. Ventral surfaces of the thighs and vent are also rugose, but otherwise the venter is smooth. Males lack nuptial asperities or spines on the thumb, but have vocal sacs. Females have weakly developed fringes on fingers, toes, and feet, lack upper lip tubercles, and are slightly larger than males (Lingnau et al. 2008). The overall coloration is brown, with numerous, small, uniformly distributed brown spots on the dorsum. Thighs have two irregular dark brown stripes on the dorsal surfaces, while the tibia has one irregular dark brown and two irregular light brown stripes on the dorsal surface. The pes has two dark brown stripes as well. Two dark brown stripes mark the dorsal surfaces of the brachium. Arms are marked with brown blotches on the anterior and dorsal surfaces. Tubercles on the flanks are white. The head has irregular brown blotches on the sides. Throat, abdomen, and ventral surfaces of the thighs and arms are uniformly cream. Posterodorsal surfaces of the thighs sometimes vary in coloration between specimens; 22.0% had scattered light blotches, 56.0% presented a brown background with anastomosed light blotches, 18.0% displayed equally distributed brown and cream blotches, and 4.0% had brown blotches on a cream background (Lingnau et al. 2008). Hylodes cardosoi was named after the late Dr. Adão José Cardoso and his contributions to the information and knowledge on Brasilian anurans.

Endemic to the Atlantic Forest of southeastern and southern Brazil, Hylodes cardosoi is known from the localities of Marumbi and São João da Graciosa, Municipality of Morrestes, in the State of Paraná, as well as the Municipalities of Apiaí, Capão Bonito and Iporanga in the State of São Paulo (Lingnau et al. 2008).

The species tends to act warily, and reacts to disturbances by jumping into water, but returning to the original location a few minutes later (Lingnau et al. 2008). Males have been observed performing visual displays of body-raising and foot-flagging, and sometimes simultaneously emitting an advertisement call (Lingnau et al. 2008). This behavior has been similarly observed in other species of the Hylodes nasus group, such as Hylodes asper (Haddad and Giaretta 1999). The call is given sporadically, and consists of 4-8 notes of 3-6 frequency-modulated pulses each, at a dominant frequency of 3.9-5.8 kHz. Generally the dominant frequency is associated with the third harmonic, but sometimes the second and third harmonics have equal amounts of energy, making it difficult to distinguish the dominant frequency (Lingnau et al. 2008). Hylodes cardosoi males and females appear to practice some site fidelity (Lingnau et al. 2008). Hylodes cardosoi is sometimes sympatric with Hylodes heyeri in stream habitats (Lingnau et al. 2008).

Diagnosis Densely granular dorsolateral surfaces; large size (adult male snout-vent length 35.6-44.1 mm); robust body; light dorsolateral stripes absent; moderate-sized fringe on outer side of toe V; advertisement call with 4-8 notes, each note with 3-6 pulses in rapid raise and decay frequency modulation.

Known from Marumbi and São João da Graciosa, both localities in the Municipality of Morretes, State of Paraná, southern Brazil, and from the Municipalities of Apiaí, Capão Bonito, and Iporanga, State of São Paulo, southeastern Brazil.

Description of holotype Body robust; head longer than wide; snout protruding in lateral view, round in dorsal view; nostrils not protuberant, directed laterally; canthus rostralis distinct, almost straight; loreal region concave; tympanum distinct, nearly round, its diameter slightly smaller than half of eye diameter; supratympanic fold starting at posterior corner of eye, not reaching shoulder; vocal sacs lateral, well developed; tongue nearly ovoid, large, free posteriorly; vomerine teeth in two series between choanae; choanae round; maxillary and premaxillary teeth present; a row of unpigmented and small tubercles on margin of upper lip. Arms moderately robust; thumb without nuptial asperities or spines; outer metacarpal tubercle very large, nearly round; inner metacarpal tubercle small, elliptical; subarticular tubercles single, round; number of subarticular tubercles on fingers (I to IV) 1-1-2-2; supernumerary tubercles arranged in rows between fingers II, III, and IV; relative lengths of fingers II , I , IV , III; fingers bearing lateral fringes from proximal subarticular tubercle to disc; finger III bearing vestigial fringe on outer margin from proximal subarticular tubercle to the distal one; finger discs elliptical, slightly expanded, smaller than tympanum; disc of finger I smaller than other discs; upper surfaces of discs with well developed scutes; finger discs as developed as toe discs. Legs robust; outer metatarsal tubercle small, round, protruding; inner metatarsal tubercle elongated; subarticular tubercles single, rounded or ovoid; number of subarticular tubercles in toes (I to V) 1-1-2-3-2; relative lengths of toes I , II , V . III , IV; toes extensively fringed, fringe on outer margin of toe V exceeds the end of subarticular tubercle in a distance nearly equal the diameter of a subarticular tubercle; tarsal fringe extensive, continuous distally with toe fringe on inner margin of toe I; toe discs slightly expanded, nearly elliptical; toe disc I in the left foot malformed; disc of toe V smaller than other discs; upper surfaces of discs with developed scutes. Dorsal skin texture rugose, mainly in flanks, thighs and around vent; undersurfaces smooth; skin fold across abdomen near inguinal region (probably caused by preservation); skin texture rugose around vent and on ventral surfaces of thighs. Measurement of the holotype: Snout-vent length 41.5; head length 15.3; head width 13.0; tympanum diameter 2.6; eye diameter 5.5; interorbital distance 3.8; eye-nostril distance 3.3; internarinal distance 4.9; snout to nostril distance 2.1; thigh length 21.0; tibia length 22.3; foot length 20.4. Coloration Overall color brown; dorsum with numerous uniformly distributed small brown spots; dorsum of thighs with two irregular dark brown stripes; dorsum of tibia with three irregular dark brown and two brown stripes; dorsum of pes with two dark brown stripes. Dorsum of brachium with two dark brown stripes; anterior and dorsal surfaces of arms with brown blotches; white tubercles on flanks; lateral aspect of head with irregular brown blotches; abdomen and throat uniformly cream; ventral surfaces of thighs and arms uniformly cream. Variation Females have poorly developed fringes on toes, fingers, and feet; no tubercles on upper lip; and are slightly larger than males. The fringe on the inner margin of finger III (n=42) may be: complete or nearly complete (73.8%), vestigial (23.8%), or absent (2.4%). The fringe on the outer margin of finger III may be: complete or nearly complete (4.8%), vestigial (66.7%), or absent (28.6%). The fringe on the inner margin of finger IV may be: complete (19.1%), vestigial (76.2%), or absent (4.8%). Specimens (n 5 50) also vary in color pattern of posterodorsal surface of thighs: 22.0% present scattered light blotches, 56.0% present brown background with anastomosed light blotches, 18.0% present brown and cream blotches equally distributed, and 4.0% present cream background with brown blotches.

NE.Not evaluated.