Phyllodytes amadoi, Vörös & Dias & Solé, 2017

Vörös, Judit, Dias, Iuri Ribeiro & Solé, Mirco, 2017, A new species of Phyllodytes (Anura: Hylidae) from the Atlantic Rainforest of southern Bahia, Brazil, Zootaxa 4337 (4) : -

Phyllodytes amadoi sp. nov.

Generic placement: We assign the new species to the genus Phyllodytes based on morphological (odontoids on the mandible, carpal, tarsal and ventral tubercules), bioacoustical, and habitat use (bromelids) similarity with other species of the genus.

Holotype. MZUESC 14954 , adult male ( Figs 2 , 3 ), collected on the 7th of October 2015, by Iuri Dias, Leandro O. Santos and Mirco Solé. Type locality: municipality of Una, RPPN Ararauna (15° 18' 38.3" S, 39° 9' 55.9" W, WGS84, 96 m a.s.l.), State of Bahia, Brazil .

Paratypes. SiX adult males ( MZUESC 14950, 14955, 14958, 14960 and MNRJ 91484–85) and two adult females ( MZUESC 14957 and 14959) collected with the holotype; two adult males ( MZUESC 14952–14953) collected on the 3th of September 2015 by Caio Vinícius de Mira Mendes, Iuri R. Dias and Mirco Solé and nine adult males ( MZUESC 14941, 14943–45, 14949, UFBA 14929–30 and HNHMHER 2017.61.1–2.) collected on the 17th of November 2015 by Iuri R. Dias, Andrés Egea Serrano, Mirco Solé and Judit Vörös. All individuals were encountered at the same collection site as the holotype.

Additional specimens. Two juveniles (MZUESC 14946 and 14942) were collected on the 17th of November 2015 by Iuri R. Dias, Andrés Egea Serrano, Mirco Solé and Judit Vörös. The specimens were measured and eXamined for eXternal morphological characters, which further helped to characterize Phyllodytes amadoi . These two specimens were not included in the type series.

Diagnosis. A small species, SVL = 21.7–23.0 mm in females (n = 2) and 15.6–21.5 mm in males (n = 18) characterized by 1) dorsum of body, arms and legs light brown or greenish with small, irregular brown patches; 2) presence of a narrow brown stripe on canthus rostralis; 3) a wide brown stripe eXtending from the corner of the eye to the flanks; 4) snout short, rounded in dorsal view and vertical in profile with a small apical tubercule; 5) two big, widely separated and five small odontoids on both sides of mandible; 6) one–three carpal and one tarsal tubercules; 7) venter with tubercules in siX distinct rows, tubercules in two middle rows significantly greater than in second and third and 8) advertisement call composed by a series for multipulsed notes with 13–17 notes per call with mean of 14.5 ± 4.3 pulses per note, call duration 2.99– 4.106 s and dominant frequency ranging between 3789.8–4306.6 Hz.

Comparisons with other Phyllodytes species. The dorsal colour pattern distinguishes Phyllodytes amadoi from P. maculosus (pale brown with irregular distinctive cream stains), from P. brevirostris , P. edelmoi , P. megatympanum , P. kautskyi and P. luteolus , (immaculate), from P. wuchereri (longitudinal stripes), from P. gyrinaethes (variable marbled dorsal pattern), and from P. punctatus and P. tuberculosus (distinctive brown dots). Size (SVL) of P. amadoi is smaller (21.7–23.0 mm in females and 15.6–21.5 mm in males) than any other Phyllodytes species but the size of bigger individuals overlaps with P. punctatus (18.2–22.8 mm, n = 10; Caramaschi & PeiXoto 2004), P. tuberculosus (24.0–26.0 mm, n = 2; Bokermann 1966 and 21.1–25.8 mm, n = 10; Caramaschi & PeiXoto 2004), P. melanomystax (20.0– 26.6 mm, n = 45; Caramaschi et al. 1992), P. luteolus (15–24 mm, n = 186; Ferreira et al. 2012) and P. megatympanum (21.3–23.7 mm, n = 4; Marciano-Jr et al. 2017). The other species are larger: P. acuminatus (23.0– 24.5 mm, n = 2; Bokermann 1966), P. brevirostris (22.0–24.0, n = 4; PeiXoto & Cruz 1988), P. edelmoi (23.5–28.7 mm, n = 20; PeiXoto et al. 2003), P. gyrinaethes (23.9–27.9 mm, n = 20; PeiXoto et al. 2003), P. maculosus (39.7–48.5 mm, n = 3; Cruz et al. 2006), P. kautskyi (38–43.5 mm; n = not specified; Cruz et al. 2006), and P. wuchereri (25.1–27.1 mm, n = 5; Caramaschi et al. 2004).

Rounded snouth in dorsal view distinguishes Phyllodytes amadoi from P. acuminatus , P. megatympanum , P. luteolus , and P. wuchereri (acuminate) and P. melanomystax (truncate). Truncate snout in profile separates P. amadoi from P. acuminatus , P. brevirostris , P. edelmoi , P. gyrinaethes , P. luteolus , P. megatympanum , P. tuberculosus , and P. wuchereri (acute or protruding). Phyllodytes maculosus , P. kautskyi , P. melanomystax , and P. punctatus also have truncate snout in profile. Phyllodytes amadoi has two large anterior odontoids, similarly to P. acuminatus , P. maculosus , P. megatympanum , P. gyrinaethes , P. luteolus , P. tuberculosus , and P. wuchereri , while P. edelmoi , P.kautskyi , and P. punctatus have only one larger odontoid, and P. brevirostris and P. melanomystax have no large odontoid. The venter with tubercules in distinct rows separates P. amadoi from P. melanomystax , P. kautskyi , and P. maculosus (venter evenly granular without highlighted tubercles).

Holotype description. Head wider than long; snout short, rounded in dorsal view with a small apical tubercule, and vertical in profile; nostril on the tip of the snout, directed anteriorly; canthus rostralis poorly defined; loreal region slightly conveX; eyes directed anterolaterally; eye diameter 32.8 % of head length; interorbital space flat, 30.6 % of head width; tympanum distinct, rounded, diameter of tympanum 53.2 % of eye diameter; supratympanic fold well marked, partially covering the superior margin of tympanum; vocal sac subgular, poorly developed; vocal slits short, near the angle of jaws; tongue large, rounded; vomerine teeth forming straight line behind choanae; each side of mandible with two large anterior odontoids, followed by a series of five small; pupil horizontal.

Forearms slightly wider than upper arms; a row of tubercules along eXternal surface of forearm; length of hand 43.7 % of SVL; fingers short, in order of length I<IV<II<III; subarticular tubercules small and rounded; supernumerary tubercles absent; palmar tubercle well developed and ovoid, thenar tubercle large and elliptical; finger disks well developed, nearly circular; diameter of disk on third finger 36.2 % of eye diameter; webbing absent; males with large nuptial pad.

Legs slender, tibia slightly longer than tigh; sum of tibia and thigh lengths 97.5 % of SVL; a row of weak tubercles along eXternal surface of tarsus; large, spatulate tubercle near tibio-tarsal articulation. Foot with small, round subarticular tubercles; inner metatarsal tubercle large, ovoid, spatulate; outer metatarsal tubercle small, round; toes slender, in order of length, I<V<II<III<IV; toe discs similar size to finger discs; webbing absent between toes I–II; webbing formula I–II 1 ½– 3 III 3– 3 IV 3– 2 V ( Fig. 3 ).

Skin smooth on dorsal surface of body, limbs, and ventral surface of arms and tibiae; throat shagreened; belly granulated with siX rows of round tubercles of which the two middle rows are more evident; ventral surface of thighs with distinct round tubercles of which a pair near thigh insertion are more prominent.

In preservative, dorsum and dorsal surface of limbs cream with small, irregular pale brown patches; wide brown stripe eXtending from the corner of the eye to the flanks; ventral surface of body, limbs and thighs cream; throat, palmar and plantar surfaces light brown.

In life, dorsum and dorsal surface of limbs yellowish or greenish with small, irregular brown patches; wide brown stripe eXtending from the corner of the eye to the flanks; ventral surface of body, limbs and thighs cream; parietal peritoneum white; throat, palmar and plantar surfaces light brown (see paratype on Fig. 4 ).

Variation. Measurements of the 20 specimens composing the type series are shown in Table 1. Dorsal coloration varies from light (MNRJ 91484) to dark yellowish-greenish (MZUESC 14952), while irregular brown patches of dorsal pattern vary from few (MZUESC 14955) to many (MZUESC 14945). The number and size of odontoids are mostly concordant with the holotype but we found a few specimens with irregular pattern such as three-three or three-two enlarged odontoids on each side of mandible. In juveniles enlarged odontoids were absent and one subadult specimen showed only one-one enlarged odontoids. We hypothesize that frogs of Phyllodytes amadoi develop characteristic odontoid structure during adult stage.

Etymology. The name is a patrimony for Jorge Amado, a Brazilian modernist writer who had an enormous influence on Brazilian literature. He lived in the same region where the new species was discovered and he adored frogs and enjoyed collecting all kinds of objects that were related to them. As a well-travelled man, he grew a big collection of these "frog-souvenirs" from all over the world, which are partially on display in his home in Salvador, Brazil.

Natural History. Specimens were always spotted during the night, and males called from terrestrial and small epiphytic bromelids at 1.5–2.0 m height on the trunks of large trees in a regenerating forest along a small stream known as Rio Pimenta. The species appeared to be locally abundant during the three visits to the area.

The type locality in the municipality of Una is located inside Ararauna farm, which covers an area of 100 hectares, of which 40 have been converted into a private reserve of natural heritage. The Biological Reserve of Una, with more than 18,000 hectares, one of the largest conservation units in the region, is located nearby in the same municipality, but to date the new species has not been reported from this reserve.

Advertisement call. A total of 17 calls from two specimens (MZUESC 14954 holotype; SVL = 20 mm and MZUESC 14955, SVL = 20 mm) were recorded on 7th of October 2015 at 19 h 30 min with air temperature 22.6°C. The advertisement call of Phyllodytes amadoi is composed of 13–17 (14.5 ± 1; n = 17) pulsed notes ( Fig. 5 ). Call duration is 2.99–4.106 s (3.41 ± 0.28; n = 17) and it is emitted in intervals of 41–68 s (53.4 ± 8.5; n = 15), resulting in a call rate of 1.19 call/min. Notes last 0.008–0.119 s (0.043 ± 0.021; n = 247) with 3–21 (14.5 ± 4.3; n = 247) pulses per note and internote intervals of 0.137–0.285 (0.204 ± 0.02; n = 230). Pulse repetition rate is of 368.37 ± 86.6 pulses/s (151–625; n = 247).

As a general pattern, an increase in pulse number in each note can be perceived through each call, the first notes always being shorter than the last ones. During the second half of the call (5–7 final notes) the 1–4 initial pulses are more interspaced from the rest of the note ( Fig. 5C ).

The dominant frequency of the call is 3789.8–4306.6 Hz (3962 ± 192.6; n=17) and of notes 2411.7–4306.6 Hz (3758.4 ± 405.7, n=247). The dominant frequency along the notes varies, but the lowest values (around 2411.7– 2756.2 Hz) are in the first three notes.

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Bioacoustic comparisons with other species. Of the 12 recognized species in the genus, only three have no available bioacoustic data ( Phyllodytes brevirostris , P. maculosus and P. punctatus ). The advertisement call of P. amadoi is more similar to the call of P. wuchereri ( Cruz et al. 2014; Magalhães et al. 2015), P. tuberculosus ( Juncá et al. 2012) , P. edelmoi ( Lima et al. 2008) , P. gyrinaethes ( Roberto & Ávila 2013) , and P. luteolus ( Weygoldt 1981) because all of them consist of a series of pulsed notes. The advertisement call of P. amadoi differs from the call of these other species due to its higher dominant frequency (3.789–4.306 Hz in P. amadoi vs 1.490–3.520 Hz in the other species), eXcept for P. luteolus , with a dominant frequency between 2 to 6 kHz according to literature ( Weygoldt 1981). Call duration, between 2.9– 4.1 s in P. amadoi is shorter than in P. tuberculosus , P. edelmoi and P. luteolus (4.28– 9.35 s) and longer than in P. gyrinaethes (1.3– 2.3 s). The advertisement calls of P. acuminatus ( Campos et al. 2014) , P. kautskyi ( Simon & Gasparini, 2003) , P. megatympanum ( Marciano-Jr et al. 2017) and P. melanomystax ( Nunes et al. 2007) are unpulsed.

Figures

FIGURE 2. Phyllodytes amadoi SP. nov. in preservative (holotype, MZUESC 14954, SVL = 20 mm).

FIGURE 3. Phyllodytes amadoi SP. nov., holotype MZUESC 14954, ventral views of hand (A) and foot (B); dorsal (C) and lateral (D) view of the head. Drawing courtesy of Viktória Szőke (HNHM).

FIGURE 4. Photos of Phyllodytes amadoi SP. nov., paratype MZUESC 14995 in life. Photos: Mirco Solé.

FIGURE 5. (A) Oscillogram and corresponding spectrogram of the advertisement call of Phyllodytes amadoi SP. nov. (holotype). Detailed example of firsts notes of the call being shorter (B) and final notes longer and with initial pulses interspaced from the rest of the note (C). Recorded on 7 th of October 2015 at 19 h 30 min. Air temperature during recording = 22.6 ° C.