Bioacoustics of Bokermannohyla napolii

Description of holotype . Adult male (UFMG 3333; figs. 1, 2(left), and 3). Body robust; head wider than longer. Snout rounded in dorsal and lateral views (figs. 3 A-B, respectively); nostrils directed laterally, much closer to the tip of snout than to the eye; canthus rostralis distinct; loreal region slightly concave; tympanum circular, 83% of the eye diameter; supra-tympanic fold present; pupil horizontal; vocal sac single, subgular, expanded; vocal slits present; tongue round; vomerine teeth in two almost contiguous, slightly arched series between choanae. Crenulated ridge on outer surface of forearms; forearm hypertrophied; finger lengths when put together I<II<IV<III (fig. 3 C); disks of fingers II, III, and IV larger than that of I; disk of finger III 75 % of the tympanum diameter; distal subarticular tubercles of fingers I and II ovoid, those of fingers III and IV bifid/divided; nuptial pads present; pollical tubercle large, elongated; prepollex well-developed, spine curved, completely covered by skin; inner metacarpal tubercle region formed by several smaller granular tubercles; palmar tubercle developed, elongated, divided; supernumerary tubercles present on the palm of hand; hand webbing formula I 2–3 II 2 -–2 2/3 III 2 1/2– 2 IV. Toe lengths when put together I<II<V<III<IV (fig. 3 D); disks of toes III, IV, and V larger than those of I and II; disk of toe IV 68 % of the tympanum diameter; toe webbing formula I 1–2 - II 1–2 III 1–2 + IV 2 +– 1 V ; inner metatarsal tubercle ovoid, outer metatarsal tubercle barely distinct; toes fringed; supernumerary tubercles on the sole of foot; subarticular tubercles of toes rounded; inner tarsal fold weak, extending along full length of the tarsus; outer surface of tarsi. Dorsal surfaces and flanks smooth; belly granular; supra-vent granular ridge defined; discrete pointed dermal protuberance on outer heel present. Measurements of holotype . Morphometric characters (mm) and ratios (%) in relation to SVL ( 52.5 mm ): HL 19.2 (36.6), HW 19.6 (37.3), ED 4.8 (9.1), TD 4.0 (7.6), END 5.1 (9.7), TL 28.5 (54.3), SL 29.0 (55.2), FL 24.5 (46.7), HAL 18.6 (35.4), 3FD 3.0 (5.7), 4TD 2.7 (5.1). Color of holotype in life (fig. 1). Dorsal surfaces uniformly pale cream, with subtly darker transverse stripes; a few white dots and blotches on dorsal surfaces and flanks; background color of the hidden parts of thighs and arms bluish and violet, with 8–9 dark brown vertical bars on posterior and superior surfaces; dorsal surface of shanks with 4–6 dark brown transverse stripes; flanks with subtle violet bars on a yellow marked region; inguinal region tending to the same color (violet) of the hidden parts of thighs, with a few dark brown transverse bars; supra-vent crenulated ridge white; crenulated ridge on outer forearm white; throat, chest, and belly light cream; iris gold, pupil black; tympanum tends to violet. Color of holotype in preservative. Dorsum dark grayish brown with some faded parts on dorsum and limbs, and dark brown transverse stripes barely distinct on dorsum; a few white blotches scattered on dorsum, limbs, and flanks; hidden parts of thighs and arms light gray, their posterior and superior surfaces with 8–9 dark brown vertical bars; anterior surface of thighs immaculate; dorsal surface of shanks with 4–6 dark brown transverse stripes, sometimes fragmented; dark brown transverse stripes on dorsal surface of forearms indistinct; background color of flanks light gray, with a few dark brown reticulations, especially on inguinal region; supra-vent crenulated ridge white; crenulated ridge on outer forearm white; throat, chest, belly, and ventral surfaces of limbs cream, gular region with dark brown punctuations bordering lower jaw, region of tubercles and webbing of hands and feet slightly gray; tympanum brown, slightly lighter than dorsal coloration pattern. Variation. Dorsum varies in color from medium brown (females) to dark gray (males) in ethanol preserved specimens. Number of dark brown vertical bars on dorsal surfaces of forearms (3–4), thighs (6–10), shanks (4–7), and tarsi (3–4). Dark brown transverse stripes on dorsal surfaces may be barely distinct. The male specimen AAG- UFU 0 860 has no distinctive transverse stripes on dorsal surfaces. The presence and extent of white blotches on dorsum and limbs are variable, being completely absent in AAG-UFU 4775. Type series, except the holotype and AAG-UFU 0 860, has a dark brown vertebral stripe, sometimes restricted to the first half of dorsal length. One female (AAG-UFU 4773) has distal subarticular tubercles of finger III divided; one male (AAG-UFU 4774) has distal subarticular tubercles fingers III and IV cordiform. Two male specimens (AAG-UFU 4775 and ZUEC 16610) have distal subarticular tubercles of finger I divided. Natural History. Despite extensive fieldworks since 2001, this species was only heard (found) in 2009, during an explosive breeding event. Afterwards, a few males were heard once at the end of 2011. Males started calling late at night (around 01:00 AM), and a few males were heard still calling at nearly midday on the next day. They gathered around a muddy slow-watered section of a stream at the forest border. Males (N = 10) were calling from the ground or perched on low (< 1 m high) vegetation. Egg clutches (N = 4; fig. 4) were laid 5–10 cm deep and hidden among dead broad leaves under water. Eggs were adhered to each other so that the entire egg mass (fresh, <6 hours) could be removed from water using both hands to scoop it up. Number of eggs per clutch ranged from 373 to 742 (mean 637, SD = 176, N = 4 egg clutches); yolk diameter ranged from 1.2 to 2.3 mm (mean 1.8, SD = 0.3, N = 3 egg clutches); gelatinous capsules ranged from 3.8 to 5.4 mm (mean 4.4, SD = 0.6, N = 4 egg clutches). Three collected females released ovocytes inside plastic bags: yolk diameter ranged from 1.5 to 2.2 (mean 2.0, SD = 0.6); gelatinous capsules ranged from 3.1 to 5.4 mm (mean 4.2, SD = 0.5). Some males had their backs scratched (fig. 1), probably as a result of agonistic interactions. The new species occurs simpatrically with Bokermannohyla sazimai , Hypsiboas faber (Wied-Neuwied) , Hypsiboas goianus (Lutz) , Hypsiboas lundii (Burmeister) , and Scinax canastrensis (Cardoso & Haddad) . Geographic distribution. Bokermannohyla napolii sp. nov. is known only from the type locality. (fig. 5). Etymology. The name is a noun in the genitive case honoring Marcelo F. Napoli for his extensive contribution to the knowledge of the genus Bokermannohyla . Vocalizations of Bokermannohyla napolii sp. nov. Six males recorded (N = 76 advertisement call samples; N = 43 aggressive call samples). Quantitative traits are summarized in Table 2 . Advertisement call is composed of two note types (referred herein to as note types A and B; figs. 6 A-B) emitted consecutively at a rate of 1–14 calls/minute (mean 6.7, SD = 4.6). Additional notes B may eventually be emitted just after a typical A/B sequence though. Advertisement call duration (notes A + B) was 460–807 ms (mean 619.7, SD = 71.7). Note A is a short note (53– 156 ms; mean 93.4, SD = 15.6) with 5–6 harmonics. Dominant frequencies from 0.39–0.56 kHz (mean 0.43, SD = 0.07), from 0.73–0.90 kHz (mean 0.77, SD = 0.06), from 1.34–1.51 kHz (mean 1.47, SD = 0.05), and from 1.68– 1.85 kHz (mean 1.82, SD = 0.08), which correspond to the first four harmonics. Note B is a long note (242–550 ms; mean 399.5, SD = 77.4) with a harmonic structure (5–6 harmonics; sometimes ill-defined), and sound intensity concentrated the most on the first part of the note, and sound intensity modulations forming 1–5 disjunctive or juxtaposed harmonic groups (mean 2.5, SD = 1.2) along its extent, or with ill-defined harmonic structure (N = 8 note samples). A/B internote interval from 82–167 ms (mean 133.3, SD = 17.2). Dominant frequencies from 0.47–0.65 kHz (mean 0.51, SD = 0.06), from 0.82–1.34 kHz (mean 1.17, SD = 0.14), from 1.42–1.68 kHz (mean 1.57, SD = 0.07), and from 2.02–2.11 kHz (mean 2.05, SD = 0.04), which correspond to the first four harmonics. We have also identified a third note type (referred herein to as note type C; fig. 7 A) that should possibly be attributed to a kind of aggressive call ( sensu Wells 2007 ) because it has an ill-defined lower sound intensity structure with remarkable differences in temporal traits (longer duration) in comparison with note B, yet similar in spectral traits (dominant frequencies). Besides, this aggressive call has no distinguishable harmonic structure as in notes A and B of the advertisement call. Note C is emitted at a rate of 12–26 notes/minute (mean 20.0, SD = 5.7) alone or just after a note A. The emission rate of advertisement calls of a given male was inversely proportional to the emission rate of aggressive calls. Note C duration was 437–1,120 ms (mean 763.9, SD = 210.4) with A/C internote interval from 90–128 ms (mean 115.3, SD = 14.6). Despite the absence of well-defined harmonic structure, dominant frequencies coincided with those of note type A.