A new species of Phyllodytes (Anura: Hylidae) from the Atlantic Rainforest of southern Bahia, Brazil

Abstract

We describe Phyllodytes amadoi sp. nov., a species of treefrog from the Atlantic Rainforest of southern Bahia, Brazil. The new species can be diagnosed by its small body size (SVL 15.6–23.0 mm) and advertisement call with high dominant frequency (3.789–4.306 Hz). It is known from one locality in the Private Reserve of Natural Heritage Ararauna, in the municipality of Una (15° 18' 38.3" S, 39° 9' 55.9" W, 96 m a.s.l.).

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Accepted by S. Castroviejo-Fisher: 8 Sept. 2017; published: 20 Oct. 2017
ZOOTAXA
ISSN 1175-5326 (print edition)
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Copyright © 2017 Magnolia Press
Zootaxa 4337 (4): 584
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594
http://www.mapress.com/j/zt/
Article
https://doi.org/10.11646/zootaxa.4337.4.9
http://zoobank.org/urn:lsid:zoobank.org:pub:9E156F03-56AB-4541-BACE-A993B89A890C
A new species of Phyllodytes (Anura: Hylidae) from the Atlantic Rainforest of
southern Bahia, Brazil
JUDIT VÖRÖS
1,2,*
, IURI RIBEIRO DIAS
2
& MIRCO SOLÉ
2
1
Department of Zoology, Hungarian Natural History Museum, 1088 Budapest, Baross u. 13., Hungary. E-mail: voros.judit@nhmus.hu
2
Department of Biological Sciences, Universidade Estadual de Santa Cruz, Rodovia Ilhéus-Itabuna, km 16, 45662-900 Ilhéus, Bahia,
Brazil. E-mails: iurirdias@hotmail.com, msole@uesc.br
3
Corresponding author
Abstract
We describe Phyllodytes amadoi sp. nov., a species of treefrog from the Atlantic Rainforest of southern Bahia, Brazil. The
new species can be diagnosed by its small body size (SVL 15.6–23.0 mm) and advertisement call with high dominant fre-
quency (3.789–4.306 Hz). It is known from one locality in the Private Reserve of Natural Heritage Ararauna, in the mu-
nicipality of Una (15° 18' 38.3" S, 39° 9' 55.9" W, 96 m a.s.l.).
Key words: Anura, bioacoustics, Hylidae, Lophiohylinae, morphology, Phyllodytes amadoi
Introduction
The Atlantic Rainforest extends along the Atlantic coast of Brazil and represents one of the richest biodiversity
areas in the world (Myers et al. 2000). In the last 500 years nearly 88 % of the original Atlantic Forest has been lost
and replaced by human-modified landscapes like pastures, croplands or urban areas (Ribeiro et al. 2009; Tabarelli
et al. 2004). Despite its relatively small remaining area (4,000 km
2
), half of the Brazilian amphibian fauna and the
highest endemism rate (85 %) within Brazil can be found in this biome (Cruz & Feio 2007; Haddad et al. 2013).
Species of the hylid frog genus Phyllodytes Wagler, 1830 are endemic to the Atlantic Rainforest of eastern
Brazil. Phyllodytes can be distinguished from other hylids by their odontoids growing on the mandible and strong
affinity to bromelids as environment for living, breeding, egg laying, and larval development (Bokermann 1966;
Caramaschi et al. 2004; Cunha & Napoli 2016; Peixoto 1995). The genus is currently represented by twelve
recognized species (Marciano-Jr et al. 2017; Segalla et al. 2016) out of which, six are known to be found in
southern Bahia—P. luteolus, P. tuberculosus, P. megatympanum, P. melanomystax, P. wuchereri, and P.
maculosus—and, apparently, this diversity is vastly underestimated (Dias et al. 2014a, b; Frost 2017). Even though
P. kautskyi was observed in Serra Bonita (Camacan) and RPPN Estação Veracel (Porto Seguro), both in Bahia,
these records need confirmation. When reporting P. kautskyi from southern Bahia, Simon & Peres (2012) did not
provide diagnosis supporting the differentiation of the studied individuals from P. maculosus despite the
similarities between the two species.
There are attempts to cluster the species belonging to the genus Phyllodytes into specific groups according to
their dorsal colour pattern (Caramaschi et al. 2004; Peixoto et al. 2003) or most recently using bioacoustic
characters, such as the structure of advertisement calls (Roberto & Ávila 2013). However as the phylogenetic
relationships within Phyllodytes are still poorly explored, validity of these groups has not yet been tested in the
context of their evolutionary history.
Herein we describe a new species of Phyllodytes from the Atlantic Rainforest of southern Bahia, based on
morphological and bioacoustic evidences.

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Material and methods
Between the years 2015–2016 twenty-two specimens of the new species were collected in the Private Reserve of
Natural Heritage Ararauna in the municipality of Una (Fig. 1).
Adult and juvenile specimens were collected manually and transferred to the laboratory of the Universidade
Estadual de Santa Cruz (UESC) for further processing. Lidocaine was used for anasthetizing and killing the
specimens. Specimens were fixed overnight in 10 % formaline and preserved in 70 % ethanol.
FIGURE 1. Type locality of P. amadoi sp. nov. in Southern Bahia, Brazil. Atlantic Forest remnants and Conservation priority
according to SOS Mata Atlântica (2014) and Ministério do Meio Ambiente, MMA (2000), respectively.
Specimens were deposited at Collection of Amphibians and Reptiles of Museu de Zoologia da Universidade
Estadual de Santa Cruz, Bahia, Brazil (MZUESC), Collection of Amphibians and Reptiles of the Hungarian
Natural History Museum (HNHM), Museu de Zoologia of the Universidade Federal da Bahia (MZUFBA), and
Museu Nacional, Rio de Janeiro (MNRJ).
Morphological measurements were obtained with a digital caliper (0.1 mm precision) using an ocular
micrometer in a Leica stereomicroscope. The following measurements were obtained from 20 adult specimens
following Duelmann (1970) and Heyer et al. (1990): snouth–vent lenght (SVL), head length (HL), head width
(HW), inter–narine distance (IND), eye–narine distance (END), eye diameter (ED), upper eyelid width (UEW),
inter–orbital distance (IOD), tympanum diameter (TD), hand length (HL), thigh length (THL), tibia length (TL),
femur length (FL), finger disc diameter (FD). Webbing formula notation follows Myers & Duellman (1982).
We recorded advertisement calls with a Sennheiser ME45 unidirectional microphone attached to a Tascam
DR1 digital recorder. The microphone was placed at an approximate distance of 10–20 cm from the calling site
inside bromeliads of the recorded specimen. Calls were analyzed using the program Raven Pro 1.4 at 44.1 kHz

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with 16 bit resolution. Waveforms and audiospectrograms were produced according to the following parameters:
72 points FFT, hann window, with 50 % overlap. Advertisement call terminology follows Duellman & Trueb
(1986).
Colour in life was described based on field observations and photographs of live specimens. Sex was
determined by secondary sexual characters such as nuptial pads and vocal slits.
Results
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.
FIGURE 2. Phyllodytes amadoi sp. nov. in preservative (holotype, MZUESC 14954, SVL = 20 mm).
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.

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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.
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

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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. k a ut sk yi , 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).
Vari a t i o n. 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.

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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.
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 7th of October 2015 at 19 h 30 min. Air temperature during recording =
22.6°C.

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0=8(6& )              
0=8(6& 0              
0=8(6& )              
0=8(6& 0              
+1+0+(5

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+1+0+(5

0              
8)%$ 0              
8)%$ 0              
015- 0              
015- 0              
  
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±
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±


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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.
Discussion
Phyllodytes is one of the frog genera endemic to the Atlantic Rainforest where the number of new species
discoveries expanded recently; six (50 %) out of the twelve species have been described in the last 15 years. The
descriptions are based on morphological characters and bioacoustic comparisons; no comprehensive molecular
taxonomical investigation has been conducted within the group yet. In the lack of available reference sequences
(only sequences of P. luteolus gene fragments can be found when searching NCBI GenBank Database) our
description of P. amadoi relies on the sharp differences in morphology and advertisement call compared to the
already recognized species. The small body size (smallest species within the genus) and the high dominant
frequency of the advertisement call make the species easily distinguishable within Phyllodytes. The number of
candidate species in the genera is expected to expand further via monitoring programs, environmental assessments
and possibly molecular phylogenetic studies (Dias et al. 2014b).
Phyllodytes amadoi is known only from one locality, but its occurance is probably more widespread. This
habitat is part of a private reserve of natural heritage and is probably not threatened by direct deforestation. Most of
the Phyllodytes species—except for P. luteolus, which occurs from the State of Pernambuco to southern Espírito
Santo—are known from a relatively narrow distribution range or are data deficient (Amphibiaweb 2016).
Therefore they might be extremely affected by factors causing amphibian decline including habitat loss (Eterovick
et al. 2005), habitat fragmentation (Dixo et al. 2009), infectuous diseases such as the amphibian chytrid fungus
Batrachochytrium dendrobatidis (Carnaval et al. 2006; James et al. 2015; Rodriguez et al. 2014), natural disasters
(Papp & Papp 2000) or the use of pesticides (Egea-Serrano & Solé 2017). The amount of cryptic diversity among
Phyllodytes should be screened before it could be lost.
Acknowledgements
We thank Tadeu Teixeira Medeiros, Marcos Vila Nova, Daniel Grundmann, Rose Oliveira, and Elson Rios for
technical help in the collection, Victor Goyannes Dill Orrico for encouraging discussions, and Andrés Egea
Serrano for help in the field. We are thankful to Julia and Nora Nüscheler for opening the doors of RPPN Ararauna
and Wesley DaRocha for logistical support during fieldwork. Special thanks to Tamás Németh for his contribution
to the photographs of holotype and Eszter Vörös for English corrections. Drawings of holotype were made by
Viktória Szőke (HNHM). We are grateful to the colleagues Caio Vinícius de Mira Mendes and Leandro O. Santos
for collaboration during field activities. Specimens were collected under the permit number #13708–1 issued by the
Intituto da Conservação da Biodiversidade Chico Mendes–ICMBio. JV was supported by the Hungarian Eötvös
State Fellowship. Scholarships were provided by the Coordenação de Aperfeiçoamento de Pessoal de Nível
Superior (CAPES) and Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq–Project: 404417/
2015–9, Process: 150372/2016–7) to IRD. MS was funded by a „Bolsa de Produtividade em Pesquisa” (CNPq–
304999/2015–6), a “Universal” project (CNPq–449930/2014–9) and a “PROTAX” project (CNPq–440615/2015–
1). Fundação O Boticário de Proteção à Natureza (0991_20132) partially funded this study.

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