Bioacoustics of Proceratophrys goyana

Description. Based on adult male specimens (N = 18). SVL: 38.8–46.5 mm (measurements in Table 1 ). General aspect of body ovoid, stout and warty. Head wider than long (Head length/width = 83.3–88.9%), its length 37.2–39.6% and width 42.5–46.0% SVL. Snout outline rounded in dorsal view, near vertical to obtuse in profile. Nostrils elliptical, with slightly prominent borders. Eyes anterolaterally directed. Preocular cutaneous crest not evident. Canthus rostralis highlighted by the presence of a moderate canthal crest. Loreal region slightly concave, tending to plan. Region between eyes and nostrils concave. An enlarged tubercle posterior and ventral to mouth commissure. An oblique series of warts from below eye to mouth commissure. Upper eyelids (21.2–24.2% head width) triangular, bordered by warts, with a fleshy and slightly expanded tip; warts covering upper eyelids somewhat arranged in rows, a distinct row in the anterior half of each eyelid directed to the interocular region, not contacting each other by the space of one–two warts. Frontoparietal crests visible as slight elevations, delimiting a shallow depression between them. Tympanum indistinct. A symmetrical pair of ridges on dorsum, initially converging from the posterior border of upper eyelids to around the middle of dorsum, then diverging and converging again in the sacral region to meet each other above coccyx, forming a lozenge in the posterior half of dorsum; warts in a regular, well-defined series all along the extent of symmetrical dorsal ridges. Arms relatively short and stout; upper arm enclosed by a patagium; forearm with a well-defined ventrolateral series of enlarged tubercles, its length 21.6–23.3% SVL. Hands large (26.7–28.1% SVL), finger webbing and nuptial pads absent; outer metacarpal tubercle bifid, the two parts similar sized; inner metacarpal tubercle enlarged, oval; enlarged subarticular tubercles; other smaller, supernumerary tubercles scattered along the ventral surface of hand; sides of fingers ridged; tip of fingers keratinized; length of fingers: III>I>II>IV. Legs relatively short and stout; thigh length 35.8–37.9% SVL, tibia length 36.3–37.4% SVL; tarsus with a well-defined ventromedial series of enlarged tubercles. Feet large (41.1–42.5% SVL), toe webbing poorly developed; outer metatarsal tubercle barely distinct; inner metatarsal tubercle enlarged, strongly keratinized, spade-like, with a series of tubercles lying on its inner basis; enlarged subarticular tubercles; other smaller, supernumerary tubercles scattered along the ventral surface of foot; sides of toes ridged; tip of toes keratinized; length of toes: IV>III>II≈V>I. Skin shagreened, covered by warts variable in size (except ventral surfaces), 1–2 relatively well-defined oblique rows of warts on the sides of trunk. A pair of enlarged tubercles below and lateral to cloacae. Vocal sac single, not or poorly expanded externally. Tongue cordiform, free posteriorly. Vomerine teeth lying in two patches between choanae. Vocal slits present. Female specimens (N = 4) basically share the features of males, except by the larger size ( Table 1 ) and absence of vocal slits. Color. In preservative (70% ethanol), browned background, somewhat grayish or reddish in some specimens. Irregular blotches, slightly lighter or darker than the background may or not be present on dorsum and flanks. The area delimited by the pair of symmetrical ridges can be blotched or have a uniform color pattern. In all specimens, the symmetrical dorsal ridges are bordered laterally by dark brown undulations, less conspicuous in sacral region. Sides of the snout and dorsal surfaces of arms, legs, fingers and toes with alternated light and dark brown bars. Ventral surfaces tending from cream to light brown, with small dark spots on belly and chest, at least in some extent. In males, the throat color is darker than the rest of ventral background. In life, the color pattern was the same ( Figure 1 ). Natural history. Specimens of P. goyana were found at Vale da Lua, an area at the Chapada dos Veadeiros characterized by a large conglomerate of rocks. The breeding site of this species ( Figure 4 ) consisted of streamlets running over rocks, surrounded by grasses and shrubs. Calling specimens were found at night, hidden under rocks and grass bushes or completely exposed. We have just visited this site during the rainy season ( November 2011 and 2012), so we don’t have further information about the breeding season of P. goyana . Geographic distribution. Presently, P. goyana is thought to have a widespread distribution, mainly throughout the Cerrado biome in central Brazil . A distribution map for the species was recently published by Teixeira Jr. et al. (2012).

Diagnosis. In the genus Proceratophrys by lacking nuptial pads on thumb, body without enlarged glands, toes not webbed and ridged laterally, supernumerary tubercles present on hands and feet, dorsal surfaces of fingers and toes wrinkled, and by presenting a pair of symmetrical ridges on dorsum, extending from upper eyelids to sacral region. In the P. cristiceps group by lacking long palpebral appendages and postocular swellings. This species is characterized by: (i) SVL = 38.8–46.5 mm in males and 55.8–62.6 in females; (ii) a symmetrical pair of dorsal ridges extending sinuously and continuous from the upper eyelids to coccyx; (iii) upper eyelids triangular, with a fleshy, slightly expanded tip; (iv) frontoparietal crests visible as slight elevations, delimiting a shallow depression between them; (v) overall color pattern browned, with the symmetrical dorsal ridges bordered laterally by dark brown undulations; (vi) advertisement call emitted in a multi-note pattern, each noted pulsed, the first and last notes differing from each other and from those amidst the call in temporal features. Comparison with other species. Proceratophrys goyana can be distinguished from the other species of the P. cristiceps group, and from those not associated to any group, mainly by the following combination of characters (data for species in comparison in parenthesis): in size (male SVL = 38.8–46.5 mm in P. goyana ), from P. huntingtoni (SVL = 31.8–38.7 mm ), P. minuta (SVL = 20.0– 25.2 mm ), P. moratoi (24.7–31.0 mm), P. redacta ( 26.9–29.7 mm ) and P. rondonae ( 62.9 mm – only the holotype measured); from P. aridus , P. carranca and P. cristiceps (frontoparietal crests not developed) by the frontoparietal crests visible as slight elevations, delimiting a shallow depression between them; from P. a r i d u s , P. caramaschii , P. carranca , P. cristiceps , P. cururu , P. moratoi , P. redacta , P. vielliardi (upper eyelids rounded, not expanded), P. minuta , P. rondonae and P. s c h i rc h i (upper eyelids with multiple short expansions/appendages) by the upper eyelids triangular, with a fleshy, slightly expanded tip; from P. concavitympanum and P. strussmannae (tympanum visible as a depression in the skin) by the tympanum not visible externally; from P. carranca , P. cururu , P. huntingtoni , P. moratoi and P. vielliardi by the symmetrical pair of dorsal ridges extending sinuously and continuous from the upper eyelids to coccyx (interrupted in the sacral region or completely absent in these species); from P. moratoi and P. vielliardi (overall color mostly in tones of gray, with rounded dark blotches) by the overall color pattern browned, with the symmetrical dorsal ridges bordered laterally by dark brown undulations; from P. concavitympanum , P. cristiceps , P. cururu , P. huntingtoni and P. moratoi by the multi-note advertisement call pattern (single note).