Another new species (and it’s not over yet) of Phyllodytes Wagler, 1930 (Anura, Hylidae) from the Atlantic Forest of southern Bahia, northeastern Brazil Author Santos, Laisa S. Programa de Pós-Graduação em Zoologia, Departamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Ilhéus, Bahia, Brasil Author Roseno, Rafaella S. 0000-0002-8503-5530 Programa de Pós-Graduação em Zoologia, Departamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Ilhéus, Bahia, Brasil & raf. iroseno 96 @ gmail. com; https: // orcid. org / 0000 - 0002 - 8503 - 5530 raf.iroseno96@gmail.com Author Solé, Mirco 0000-0001-7881-6227 Programa de Pós-Graduação em Zoologia, Departamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Ilhéus, Bahia, Brasil & Herpetology Section, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, D- 53113 Bonn, Germany msole @ uesc. br; https: // orcid. org / 0000 - 0001 - 7881 - 6227 msole@uesc.br Author Dias, Iuri Ribeiro 0000-0002-2825-3494 Programa de Pós-Graduação em Zoologia, Departamento de Ciências Biológicas, Universidade Estadual de Santa Cruz, Ilhéus, Bahia, Brasil & iurirdias @ hotmail. com; https: // orcid. org / 0000 - 0002 - 2825 - 3494 iurirdias@hotmail.com text Zootaxa 2023 2023-11-20 5374 4 519 532 https://mapress.com/zt/article/download/zootaxa.5374.4.4/52304 journal article 10.11646/zootaxa.5374.4.4 1175-5326 10159019 72C9EC0B-F3D1-48F1-9A4C-C9F887C45327 Phyllodytes iuna sp. nov. Phyllodytes sp. 3 — Blotto et al. 2020 Holotype . MZUESC 18950 , an adult male ( Figs. 1 and 2 ) from Estação Ecológica de Wenceslau Guimarães ( - 13.59967 , -39.71880 , 675 m a.s.l. ), municipality of Wenceslau Guimarães , Bahia , Brazil , collected by Iuri R. Dias and Rafael O. de Abreu on 03 February , 2015. Paratopotypes . MZUESC18952 , adult male collected with the holotype; MZUESC 23000 ; 23002, two adult males, and MZUESC 23001 ; 23003, two adult females, collected by Marcos F. Vila Nova in May, 2018 ( Fig. 3 ) . Generic Placement. The new species is assigned to the genus Phyllodytes based on the occurrence of odontoids in the lower jaw. In addition, molecular data support the placement of Phyllodytes sp. nov. as a sister taxon of P. brevirostris + P. edelmoi ( Blotto et al . 2020 ) . Etymology . The specific epithet “iuna ” alludes to a berimbau riff (a musical percussion instrument) played during a Capoeira bout (called “jogo”; “game”). The ‘iuna’ riff represents a rite of passage in Capoeira, marking the moment of transition from student to teacher. When this riff is being played, the game is reserved for teachers and masters. It is also played during posthumous homages to honor the skill of the deceased Capoeira player (capoeirista). Capoeira is an Afro-Brazilian cultural expression, developed by enslaved Africans in Brazil during the colonial era as a means of self-expression, resistance, and community-building. It is recognized as Cultural Heritage of Brazil and Intangible Cultural Heritage of Humanity by UNESCO, symbolizing the black resistance during the slavery period in the country. By adopting the name “iuna ” to honor Capoeira, we are also celebrating the rich Afro-Brazilian heritage and acknowledging the legacy left by Capoeira masters in the resistance, consolidation, and perpetuation of this martial art. Characterization. (1) Medium sized adult males, SVL in males 22.2–25.2 mm (n = 4); SVL in females 24.0– 25.9 mm (n = 2); (2) slender body; (3) head as long as wide (HL 98.7 % of HW); (4) truncated snout with apical tubercle in dorsal view and protuberant in lateral view; (5) nostrils with rounded contours positioned close to the tip of the snout, directed dorsolaterally; (6) canthus rostralis rounded; (7) loreal region concave and straight; (8) tympanum evident; (9) tympanic ring evident; (10) well-marked supratympanic fold from the back of the eyes to the insertion of the arm; (11) prominent eyes, positioned anterolaterally; (12) single sub-gular vocal sac; (13) horizontal pupil; (14) two anterior large, and additional smaller (2–6) subequal odontoids on each side of the mandible; (15) lateral margin of forearms with inconspicuous outer tubercles; (16) palmar tubercle developed and elongated; (17) single and prominent tubercle near the tibiotarsal articulation; (18) brownish yellow coloration; (19) dorsum of body exhibits a coloration with a diffuse band-like pattern from the interocular region to the posterior region of the body; (20) dorsum of arms and legs with small and irregular brown blotches; (21) dark strip from the tip of the snout to the middle of the body; (22) smooth dorsal skin; (23) uniform cream ventral coloration; (24) ventral surface with two rows of central tubercles, with adjacent tubercles bordering without a specific pattern. Comparison with other species . Character states for the other species are shown in parenthesis. Phyllodytes iuna is recovered as the sister taxon of P. brevirostris + P. edelmoi ( Blotto et al ., 2020 ) . Phyllodytes iuna differs from them by having a head that is as wide as it is long, with HL comprising 98.7% of HW (wider than longer; HL 84.0% of HW), truncated snout in dorsal view and protruding in lateral view (rounded snout in dorsal and lateral view), presence of an apical tubercle on the snout (absent), dark stripe between the tip of the snout and the eye, on the canthus rostralis (absent), dorsal coloration with diffuse dark stripes/blotches (immaculate pattern) and single tubercle near the tibiotarsal articulation (several tubercles along the tarsus) ( Fig. 4 ). Phyllodytes iuna is distinguished from P. melanomystax , P. megatympanum , P. magnus , P. maculosus , P. luteolus , P. gyrinaethes , P. kautskyi and P. amadoi by having a head as long as it is wide (wider than long); for having a medium body size ( 22.2–25.9 mm SVL), Phyllodytes iuna is distinguished from P. magnus ( 36.4–41.1 mm SVL), P. kautskyi ( 36.5–43.5 mm SVL) and P. maculosus ( 39.5–43.5 mm SVL); the truncated snout in dorsal view distinguishes Phyllodytes iuna from P. amadoi , P. gyrinaethes , P. maculosus (rounded) and from P. megatympanum , P. kautskyi and P. acuminatus (acute); in lateral view, the snout of Phyllodytes iuna is protuberant, which separates it from P. amadoi , P. maculosus (vertical) and P. melanomystax (rounded); the presence of a tubercle at the tip of the snout separates Phyllodytes iuna from P. praeceptor and P. melanomystax (absent); the tympanum of Phyllodytes iuna is evident, distinguishing it from P. gyrinaethes (hidden); the tympanum size of Phyllodytes iuna is smaller (<7% of the SVL) compared to P. megatympanum (7.73% of the SVL) and P. acuminatus (8.16% of the SVL); Phyllodytes iuna has a well-marked supratympanic fold, distinguishing it from P. gyrinaethes (discreet) and P. wuchereri (weak). The presence of inconspicuous tubercles on the lateral margin of the forearm distinguishes Phyllodytes iuna from P. acuminatus (one well-developed), P. melanomystax and P. tuberculosus (three very evident); Phyllodytes iuna has a round and simple subarticular tubercle, differentiating it from P. maculosus and P. kautskyi (bifid elongated tubercles); the palmar tubercle of Phyllodytes iuna is developed and elongated diverging from P. maculosus , P. wuchereri (ovoid), P. gyrinaethes , P. magnus (round), P. megatympanum (triangular), P. melanomystax (indistinct) and P. punctatus (elliptical); Phyllodytes iuna presents a single tubercle near the tibiotarsal articulation, differentiating it from P. acuminatus , P. kautskyi , P. melanomystax , P. luteolus , P. magnus , P. megatympanum , P. tuberculosus , P. wuchereri (several tubercles along the tarsus), and P. gyrinaethes (two tubercles). Phyllodytes iuna has eye stripes extending from the tip of the snout to halfway down the body, which distinguishes it from P. magnus and P. gyrinaethes (stripes absent); the dorsum of Phyllodytes iuna exhibits a coloration forming a diffuse band-like pattern extending to the posterior region of the body, which differs from P. kautskyi , P. magnus and P. megatympanum (immaculate pattern), P. acuminatus , P. luteolus , P. melanomystax and P. tuberculosus (immaculate pattern or with small spots), P. amadoi , P. praeceptor , and P. punctatus (small irregular spots), P. gyrinaethes (marbled pattern), P. maculosus (maculate pattern, but see Novaes-e-Fagundes & Solé 2021), and P. wuchereri (two dorsolateral white stripes outlined by a dark brown to black line with or without irregular patches in dorsum). Phyllodytes iuna has a belly with two rows of central tubercles with adjacent tubercles bordering without showing a pattern, which separates it from P. magnus , P. kautskyi , P. melanomystax , P. gyrinaethes (tubercles not distinct), from P. acuminatus , P. tuberculosus , P. amadoi , P. luteolus , P. wuchereri (six rows of rounded tubercles laterally merging with the granules on the flanks). FIGURE 1. Holotype of Phyllodytes iuna sp. nov. (MZUESC 18950). (A) Dorsal and (B) ventral view of the body. Scale bar = 10 mm Description of the holotype . Adult male in good condition, with a piece of muscle removed from the right thigh for molecular analysis. Detailed measurements are in Table 1 . Head is as long as it is wide (HL 97.4% of HW; HW 35.5% of SVL; HL 34.6% of SVL); snout truncate with apical tubercle in dorsal view and projected slightly forward in lateral view ( Fig. 2 ); nostrils with rounded contours, positioned close to the tip of the snout, directed dorsolaterally; distance between nostrils smaller than the eye-nose distance (IND 57.2% of NDE); large eyes (ED 34.4% of HL; 33.5% of HW; 126.2% of NDE), prominent, positioned anterolaterally; canthus rostralis round; loreal region concave and straight; tympanum evident, relatively large (TD 19.4% of HL), nearly circular; diameter of the tympanum is larger than the distance between nostrils (125% of IND) and smaller than the eye-narrow distance (TD 71.4% of NDE), eye diameter (TD 57.6% of ED), and interorbital distance (TD 28.3% of IOD); diameter of the tympanum is slightly smaller than the width of the discs of the third toe (TD/DF3 = 96.6%) and fourth toe (TD/4TD = 89.9%); tympanic ring evident; the supratympanic fold is well marked from the back of the eyes to near the insertion of the arm and has an arched outline; single vocal sac; vomerine teeth forming two slightly arched series, close to each other arranged horizontally between and behind the coanas; two anterior large, and additional smaller (2–3) subequal odontoids on each side of the mandible; horizontal pupil. FIGURE 2. Holotype of Phyllodytes iuna sp. nov. (MZUESC 18950). (A) Dorsal view of head; (B) Lateral view of head (right side); (C) Palmar view of left hand; (D) Plantar view of left foot. Scale bar = 3 mm Forearm with outer four tubercles, with the most evident ones near the hand. Hands large (HAL 91.5 % of HL), subarticular tubercles simple and round; supernumerary tubercles absent; palmar tubercle elongate and ovoid; fringe between fingers absent; fingers in increasing order of size I <IV <II <III; unpigmented nuptial pad on the internal base of finger I, without dark nuptial excrescence, but showing a different color pattern than the surrounding one with dark spots; hand membrane webbing I 2 - 2 - II 2 + - 2 - III 2 - - 2 + IV. Long hind limbs (THL+TBL+TAL+FL equal 180.3% of SVL), thigh is shorter than tibia (THL 89.7% of TBL; THL 49.3% of SVL; TBL 55.8% of SVL); total of the length of the thigh and tibia greater than the SVL (THL+TBL 104.5% of the SVL); a single tubercle on the inner posteroventral margin of the tarsus, near the junction with the tibia. Tarsus shorter than foot (TAL 66.1% of FL). Foot length shorter than thigh and tibia length (FL 91.2% of THL; FL 81% of TBL). Plantar surface with poorly developed supernumerary tubercles; large elongated internal metatarsal tubercle; small and round external metatarsal tubercle; toes without membranes between the I, II and III toes; foot membrane webbing I 2 - - 2 + II 2 + - 3 III 1 + + 3 IV 1 - 1 + V. Dorsal surfaces smooth; calcar appendix and supracloacal crest absent. Ventral surface of thighs with tubercles; ventral tubercles reaching to pectoral; mesal pericloacal tubercles absent. FIGURE 3. Paratopotypes of Phyllodytes iuna sp. nov. MZUESC 18952 (A and F); MZUESC 23000 (B and G); MZUESC 23001 (C and H); MZUESC 23002 (D and I); MZUESC 23003 (E and J). Scale bar = 10 m Color of the holotype in preservative . In preservative it has a yellowish-brown coloration, a dorsal coloration presenting a diffuse band-like pattern extending halfway down the body and a dark band extending from the eyes to halfway down the body; the entire ventral surface is cream colored. FIGURE 4. Variation pattern of the dorsal coloration of P. iuna sp. nov. and its sister clade P. edelmoi and P. brevirostris . Paratopotypes of Phyllodytes iuna sp. nov. (MZUESC 18952, A and MZUESC 23001, B, Photos by Rafael O. de Abreu and Marcos Vila Nova, respectively); Phyllodytes edelmoi (C, Reserva Biológica de Saltinho, Tamandaré—PE, Photo by Pedro Ivo Simıes) and Phyllodytes brevirostris (D, Cruz do Espírito Santo—PB, Photo by Washington Luiz S. Vieira). Color in life . Based on digital photographs of paratopotypes ( Fig. 4A and B ). In life it is yellow with shades of dark brown in the snout region, but this color smooths throughout the body ending in light yellow in the cloacal region. Dorsum coloration forming a diffuse band-like pattern extending halfway down the body. Dark brown stripe extending from the eyes halfway down the body. Belly uniformly yellowish. Variation. The type series is morphologically consistent with the holotype . The measurements of the type series are summarized in Table 1 . However, MZUESC 18952 and MZUESC 23001 in lateral view have a truncated snout; MZUESC 23003 has poorly developed tubercles along the tarsus with a more developed one in the tibiotarsal region; MZUESC 18952 has a dorsal coloration pattern with fewer blotches than the rest of the type-series ( Fig. 3 ). TABLE 1. Measurements (mm) of the type series of Phyllodytes iuna sp. nov.

Males	Females
Holotype MZUESC 18950	Paratype MZUESC 18952	Paratype MZUESC 23000	Paratype MZUESC 23002	Min/Max	Mean ± SD	Paratype MZUESC 23001	Paratype MZUESC 23003
SVL	22.2	23.2	25.2	23.3	22.2–25.2	23.4 ± 1.08	24.0	25.9
HW	7.9	7.7	7.9	7.9	7.7–7.9	7.8 ± 0.09	8.1	8.6
HL	7.7	8.2	8.9	8.2	7.7–8.9	8.2 ± 0.43	8.1	8.3
IND	1.2	1.2	1.6	1.5	1.2–1.6	1.3 ± 0.18	1.5	1.5
IOD	5.3	5.4	5.8	5.7	5.3–5.8	5.5 ± 0.21	5.3	5.7
END	2.1	2.3	2.6	2.9	2.1–2.9	2.4 ± 0.3	2.4	2.9
ED	2.3	2.5	2.6	2.6	2.3–2.6	2.5 ± 0.12	2.4	2.6
TD	1.5	1.7	1.6	1.5	1.5–1.7	1.5 ± 0.08	1.4	1.5
THL	10.9	11.1	11.0	10.5	10.5–11.1	10.8 ± 0.23	10.0	11.5
TBL	12.4	12.8	13.1	11.7	11.7–13.1	12.5 ± 0.52	12.5	13.4
AL	6.6	6.4	7.1	6.7	6.4–7.1	6.7 ± 0.25	6.9	7.3
FL	10.1	9.8	10.0	9.9	9.8–10.1	9.9 ± 0.11	9.8	11.0
HAL	7.1	6.8	7.7	7.0	6.8–7.7	7.1 ± 0.34	7.0	8.1
DF3	1.4	1.3	1.3	1.1	1.1–1.4	1.2 ± 0.11	1.4	1.3
4TD	1.3	1.3	1.3	1.2	1.2–1.3	1.2 ± 0.04	1.3	1.2

Characters presented as informed in M&M section. SD = standard deviation. Natural history and geographic distribution. Specimens were sighted at night and males vocalized in terrestrial bromeliads within well-preserved forests between 650 and 800 m a.s.l. Phyllodytes iuna is currently known only from Estaç ã o Ecológica de Wenceslau Guimar ã es (EEWG), a conservation unit managed by the state of Bahia , located in the north of the Central Corridor of the Atlantic Forest with a total area of ~ 24 km ² ( Fig. 5 ). This area was previously exploited for logging and cacao ( Theobroma cacao ) cultivation; however, many areas have remained untouched, especially near streams and upland areas (Rodrigues et al . 2013). FIGURE 5. Geographic distribution of Phyllodytes iuna s p. nov. Genetic distance . Genetic divergence with uncorrected p-distance between Phyllodytes iuna and the species that form its sister clade ( P. brevirostris and P. edelmoi ) were 4.4–5.0% for the partial 16S rDNA, whereas for the other species of the genus varied among 5.8–14.2% ( Table 2 ).