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Notes on the courtship behavior of Aplastodiscus arildae (Cruz e Peixoto, 1985)

at an urban forest fragment in southeastern Brazil (Amphibia, Anura, Hylidae)

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 Arquivos do Museu Nacional, Rio de Janeiro, v.64, n.3, p.247-254, jul./set.2006
ISSN 0365-4508

NOTES ON THE COURTSHIP BEHAVIOR OF APLASTODISCUS ARILDAE

(CRUZ & PEIXOTO, 1985) AT AN URBAN FOREST FRAGMENT IN
SOUTHEASTERN BRAZIL (AMPHIBIA, ANURA, HYLIDAE) 1
(With 2 figures)

RONALD R. CARVALHO JR. 2, 3

CONRADO A. B. GALDINO 4
LUCIANA B. NASCIMENTO 2

ABSTRACT: The courtship behavior, advertisement call, and courtship call of Aplastodiscus arildae are
described based on observations realized at Parque das Mangabeiras, Belo Horizonte, Minas Gerais,
Southeastern Brazil. Calling males were observed at leaves above the stream or on the litter near rivulet
banks approximately all year. Female is attracted by the calling male and conducted to the subterranean
nest, a different place from the calling site. The courtship event involves alternated mutual touches by the
couple and calls with higher repetition rate emitted by the male. Aplastodiscus arildae presents reproductive
mode with aquatic eggs deposited in subterranean nests. The advertisement call and courtship call consisted
of a sequence of a unique no pulsed note, but the first presents larger interval among the calls and duration
and higher dominant frequency than the last.
Key words: Hylidae. Aplastodiscus arildae. Courtship behavior. Courtship call. Advertisement call.
RESUMO: Notas sobre o comportamento de corte de Aplastodiscus arildae (Cruz & Peixoto, 1985) em um
fragmento florestal urbano no sudeste do Brasil (Amphibia, Anura, Hylidae).
O comportamento de corte e os cantos de anúncio e de corte de Aplastodiscus arildae são descritos com base
em observações realizadas no Parque das Mangabeiras, Belo Horizonte, Minas Gerais, Sudeste do Brasil.
Machos vocalizam praticamente por todo o ano, utilizando como sítios de vocalização a vegetação marginal
ou o folhedo no barranco na margem de riachos. A fêmea é atraída pelo macho vocalizante, que a leva até o
ninho (uma toca subterrânea), que se situa em local diferente do sítio de vocalização. No processo de condução
ao ninho estão envolvidos toques mútuos entre os indivíduos e emissões de canto de corte emitidas pelo
macho. Aplastodiscus arildae apresenta modo reprodutivo com ovos aquáticos depositados em ninhos
subterrâneos. Os cantos de anúncio e de corte consistem de seqüências de uma única nota não pulsionada,
sendo que o primeiro apresenta maior intervalo entre cantos e duração e freqüência dominante mais elevada
que o segundo.
Palavras-chave: Hylidae. Aplastodiscus arildae. Comportamento de corte. Canto de corte. Canto de anúncio.

INTRODUCTION 2002; LIMA & KELLER, 2003; HARTMANN et al., 2004).

Reproductive modes in amphibians are a
Courtship behavior refers to interactions between combination of ovipositional site, ovum and clutch
males and females to evaluate each other before characteristics, rate and duration of development,
the pair formation and mating, including the use stage, and size of hatchlings and type of parental
of signals by courting males (W ELLS , 1977). care, if any (SALTHE, 1969; SALTHE & DUELLMAN, 1973).
Courtship in frogs involves basically production The greatest variability of reproductive modes in
of advertisement calls by males (DUELLMAN & TRUEB, amphibians is known for the neotropical species
1986). However, literature describing more (DUELLMAN, 1985; HÖDL, 1990) and anurans show
complex signals on courtship of frogs has been more diversity of trends than other amphibians (39
increasing (e.g. H ADDAD & SAWAYA, 2000; LIMA et al., distinct reproductive modes) (HADDAD & PRADO, 2005).

1
Submitted on July 11, 2005. Accepted on August 22, 2006.
2
Pontifícia Universidade Católica de Minas Gerais, Museu de Ciências Naturais. Av. Dom José Gaspar 500, prédio 40, Coração Eucarístico, 30535-610, Belo
Horizonte, Minas Gerais, Brazil.
3
E-mail: [email protected].
4
Universidade do Estado do Rio de Janeiro, Instituto de Biologia Roberto Alcântara Gomes, Ecologia. Rua São Francisco Xavier 524, Maracanã, 20550-013,
Rio de Janeiro, RJ, Brazil.
248 R.R.CARVALHO JR., C.A.B.GALDINO & L.B.NASCIMENTO

The family Hylidae is one of most diverse families We observed the courtship behavior of A. arildae from
among anurans, with more than 800 species 20:50h to 23:50h on 22 October 2001 and from
(FAIVOVICH et al., 2005), and a broad distribution 20:30h to 21:30h on 23 December 2002. The times
in Brazil. HADDAD & PRADO (2005) recognized 11 described in the text are presented by the interval
reproductive modes to this family. The green 00’ to 180’. Focal-animal and all-occurrence samples
treefrogs of genus Aplastodiscus presents an were used in both observations (LEHNER, 1979), which
apparent synapomorphy related to the were made by a flashlight with a red filter to reduce
reproductive mode, where male constructs a the light interference on the behavior of the treefrogs.
subterranean nest in the muddy side streams or In order to stimulate males we played playback with
ponds. This genus included three species group. advertsiment calls to resident males. The playback
One of them, Aplastodiscus albofrenatus group, was previously recorded using a portable tape
contents the following species: A. albofrenatus recorder Panasonic RQ-L309.
(A.Lutz, 1924); A. arildae (Cruz and Peixoto, 1985); We recorded the advertisement and courtship calls
A. ehrhardti (Müller, 1924); A. eugenioi (Carvalho- with a TASCAM DAP1 recorder and Sennheiser M66
e-Silva and Carvalho-e-Silva, 2005); A. musicus microphone from a male on 2 November 2002. The
(B.Lutz, 1948); and A. weygoldti (Cruz and Peixoto, sonograms were produced by PC computer coupled
1985) (FAIVOVICH et al., 2002; 2005). The species of to the software Avisoft-Sonagraph Light version 2.7.
A. albofrenatus group are distributed within the The oscilogram and power spectrum were obtained
Atlantic Forest domain, from Santa Teresa, State by PC computer coupled to the software Sound
of Espírito Santo, to São Bento do Sul, State of Ruler version 0.941 (G RIDI -P APP , 2003-2004).
Santa Catarina, Brazil (CRUZ & PEIXOTO, 1985). Vocalizations were edited at a sampling frequency
Natural history of these green treefrogs species are of 22 kHz, FFT with 256 points, 16-bit resolution,
nearly unknown, with the exception of A. 50 overlap, and Flap top window.
leucopygius (HADDAD & SAWAYA, 2000), Aplastodiscus ANOVAs and Mann Withney test were performed,
sp. (aff. ehrhardti) (HARTMANN et al., 2004), and to compare the calls parameters, with the software
anecdotal information for some species as A. Statistica for Windows version 5.1 (STATSOFT,
albofrenatus (HARTMANN et al., 2004) and A. eugenioi 1995), as according to the variance assumptions
(CARVALHO-E-SILVA & CARVALHO-E-SILVA, 2005). of homocedasticity and normality. To test these
Aplastodiscus arildae is registered at Serra do Mar, premises it was used Levene and Komogorov-
Serra da Mantiqueira, and Serra do Espinhaço Smirnov’s tests, respectively. The significance index
mountain ranges located in the southeastern was established as 0.05.
Brazilian region (PEDRALLI et al., 2001; FROST, 2004;
NASCIMENTO et al., 2005). The knowledge of natural RESULTS
history of A. arildae is scarce. CRUZ & PEIXOTO (1985)
reported the habitat use for this species and HADDAD Calling males of Aplastodiscus arildae were
& SAZIMA (1992) present anedoctal informations. observed at night during all months in which
Herein, we describe the advertisement call, observations were made, on leaves above streams,
courtship call, and provide observations of or on the litter near rivulet banks in forested areas.
courtship behavior of A. arildae from a secondary In 22 October 2001, two males and one female
forest fragment in Minas Gerais, Brazil. were observed at a rivulet bank, spatially
distributed on the vegetation (Fig.1A). At the
MATERIAL AND METHODS beginning of observation, the two males were 0.3m
and 0.2m above the ground and 1.5m and 2.15m
Observations were made during May 2000 to from the rivulet, respectively. The initial distance
December 2001, November and December 2002 at between males was 0.65m. A female was observed
Parque das Mangabeiras, an urban forest fragment on the ground, 1.5m from the stream. At time 00’
of Belo Horizonte, Minas Gerais, southeastern only male A was emitting advertisement calls on
Brazil (19º55’57”S - 43º56’32”W, at 800-1000m). a low emission rate (not tape recorded). At time
This fragment is located in the Serra do Curral, a 5’, we started the playback from the ground, 0.7m
small mountain range belonging to the Espinhaço of male A, 0.5m of male B and 0.45m of the female,
Mountain Complex, in a transitional region between to stimulate the vocalization activity of the males.
Atlantic Forest and Cerrado domains (sensu Male A answered the playback immediately and
AB’SÁBER, 1977). kept calling until time 10’. Then the female turned

Arq. Mus. Nac., Rio de Janeiro, v.64, n.3, p.247-254, jul./set.2006

 COURTSHIP BEHAVIOR OF APLASTODISCUS ARILDAE 249

and moved toward the playback. After touching courtship calls with the head turned toward the
the recorder, she promptly jumped 0.38m to the female (n=2) or shook the branch, vibrating the
rivulet. We continued the playback and male A perch of female (n=2).
emitted vocalizations on a high emission rate (not The advertisement and courtship call of
tape recorded), attracting the female to him Aplastodiscus arildae are described from the male
(Fig.1B). She climbed a shrub and stopped above recorded on 2 November 2002, at air temperature
male A. At time 50’, she jumped upon male A of 23°C. The adversiment call consisted of a sequence
(Fig.1C), moved to his side, and stayed in this of tonal note; the intervals between consecutive calls
position for 5 minutes. At the time 55’, the couple ranged from 0.85 to 2.83s (x̄ =1.38, SD=0.41, n=83);
alternated mutual touches using their hands the note duration ranged from 0.053 to 0.072s
(Fig.1D). After this sequence of touches, male A (x¯ =0.064, SD=0.003, n=78); the dominant
stopped calling and started moving toward the frequency ranged from 2763.4 to 2870.7Hz
rivulet through the vegetation (Fig.1E). When the (x¯ =2846.8, SD=23.3, n=83) (Figs.2A, B, C).
male A was far from the female, he stopped and The courtship call of A. arildae consisted of a
emitted calls with higher repetition rate, here sequence of tonal note; the intervals between
considered as courtship calls (see H ADDAD, 1995), consecutive calls ranged from 0.464 to 1.154s
until the female approached and touched him (x¯ =0.819, SD=0.156, n=66); the note duration
(Fig.1F). The couple then jumped onto a rock in
ranged from 0.021 to 0.047s (x¯ =0.037, SD=0.004,
the middle of the rivulet, where they alternated
n=70); the dominant frequency ranged from
mutual touches again, sometimes using the side
2843.9 to 3004.9Hz (x¯ =2950.8, SD=23.95, n=70)
of the head (Fig.1G). This sequence was
(Figs.2D, E, F).
performed from 63’ to 73’. The distance covered
by the couple from the beginning of the The advertisement call has larger calls intervals
observations to time 73’ was 2.40m along the (U=357.0; p=0.0) and note duration call
rivulet bank. Male B kept at the same position, (F 2,150=1925.6, p=0.0), and higher dominant
sometimes emitting calls with lower intensity frequency (F2,150=737.9, p=0.0) than courtship call.
than courtship calls, and did not disturb the pair
during the interaction. DISCUSSION
At time 75’, the couple climbed down the rock and
moved 0.75m to a hollow entrance, a small slit Female choice in both events observed in
between rocks at the edge of rivulet, at the water Aplastodiscus arildae seems to be based partially
level. Only the male went into the hollow and the on acoustic signals and the courtship behavior
female stayed at the entrance for approximately observed includes mutual touches and acoustic-
15 minutes (Fig.1H). At time 90’, the female also tactile interactions. Reproductive behavior
went into the hole and we were unable to observe characterized by a stereotyped sequence of
the couple further. Clutch and tadpoles were not mutual touches between both sexes and the male
observed, although we fenced the hollow entrance guiding female to an oviposition site is
at the following morning and monitored it once a characteristic of species which the male
week during the following month. constructs a nest site (e.g. Hylodes phyllodes
During the second observation (23 December Heyer and Crocoft, 1986 – F ARIA et al., 1993);
2002), a female and three calling males were Hylodes asper (Müller, 1924) – H ADDAD & GIARETTA,
observed and the behavior of the female in 1999; A. leucopygius – H ADDAD & SAWAYA, 2000;
choosing the male was the same as described Aplastodiscus sp. (aff. ehrhardti) – HARTMANN et al.,
before. However, it was possible to observe the 2004; A. perviridis A.Lutz, 1950 – H ADDAD et al.,
process by which the male guided the female 2005). In these cases, the final selection of the
to the nest site in more detail. After the female partner still remains with the female, which may
reached the selected male, they exchanged be also based on characteristics of the nest (see
touches for a few minutes. Then the male HADDAD & S AWAYA, 2000).
started to move through the vegetation, followed HARTMANN et al. (2004) observed three stages of
by the female. The male stopped on a branch courtship behavior for Aplastodiscus sp. (aff.
and when the female reached him, they ehrhardti): (1) preliminary female choice, (2)
exchanged mutual touches. When the male interactive courtship, and (3) acceptance or refusal
jumped to another branch, he begun to emit of the male and/or subterranean nest by the female.

Arq. Mus. Nac., Rio de Janeiro, v.64, n.3, p.247-254, jul./set.2006

250 R.R.CARVALHO JR., C.A.B.GALDINO & L.B.NASCIMENTO

Fig.1- Schematic diagram showing the reproductive behavior of Aplastodiscus arildae at Parque das Mangabeiras, Belo
Horizonte, Minas Gerais, Brazil: (A) individuals of A. arildae at the beginning of observations; (B) female closed to male A;
(C) female above the male A; (D) female touching the dorsum of male; (E) male guiding the female; (F) female touching
male during the trajectory; (G) mutual touches by the couple using the lateral part of heads; (H) female at the hollow
entrance. Drawings based on the narrative recording during field observations and photos.

Arq. Mus. Nac., Rio de Janeiro, v.64, n.3, p.247-254, jul./set.2006

 COURTSHIP BEHAVIOR OF APLASTODISCUS ARILDAE 251

A D

B E

C F

Fig.2- Advertisement call (A) oscilogram; (B) sonogram of sequence of two calls; (C) power spectrum and courtship call (D)
oscilogram; (E) sonogram of sequence of two calls; (F) power spectrum of Aplastodiscus arildae at Parque das Mangabeiras,
Belo Horizonte, Minas Gerais, Brazil (air temperature 23ºC).

Arq. Mus. Nac., Rio de Janeiro, v.64, n.3, p.247-254, jul./set.2006

252 R.R.CARVALHO JR., C.A.B.GALDINO & L.B.NASCIMENTO

Although we did not observe any kind of choice ehrhardti), in A. arildae, as in A. leucopygius, courtship
related to nest characteristics for A. arildae female, involves calls and tactile signals. Mechanical vibration
we hypothesized that female choice relied on of the vegetation branches could be an additional
acoustic selection at the beginning of courtship and way to male conduct the female in A. arildae.
not at the end of the whole process. The elaborate HÖDL & AMÉZQUITA (2001) indicated that some special
courtship behavior observed, the long distance from ecological conditions favored the evolution of visual
vocal site to the nest, the likelihood of a predator signals. These ecological conditions are displaying
encounter, and the risks for the female to be at elevated perches, diurnality, aposematism and
intercepted by a satellite males during the trajectory, displaying at continuos high environment noise
could make the choice after the inspection of the levels. Except by the period of activity, Aplastodiscus
nest costly for both males and females. Hence, the arildae at Parque das Mangabeiras are under these
final selection on the basis of the nest characteristics ecological conditions. Therefore, it is essencial to
(HADDAD & SAWAYA, 2000; HARTMANN et al., 2004) might have more observations to conclude that this species
be cautiously invoked and so more detailed studies do not present visual communication. In spite of
are needed to evaluate these hypothesis. insufficently observations, we suggested that A.
Although the function of visual cues has only arildae could present an additional way of
ocassionally been tested experimentally, the results communication, a mechanical vibration of the
suggested that visual signaling is a significant mode substrate.
of communication in a few anuran species (HÖDL & Calling sites that differ from oviposition sites also
AMÉZQUITA, 2001). This kind of communication is are found in other tropical hylids species (e.g., D.
better known for diurnal species (e.g., Colostethus elegans Wied-Neuwied, 1824 – BASTOS & HADDAD,
trinitatis Garman, 1888 – W E L L S , 1980; 1996; A. leucopygius – H ADDAD & SAWAYA, 2000;
Brachycephalus ephippium Spix, 1824 – POMBAL et Aplastodiscus sp. (aff. ehrhardti) – HARTMANN et al.,
al., 1994; Hylodes asper – HADDAD & GIARETTA, 1999). 2004). H ADDAD & SAWAYA (2000) showed for A.
O W A S K A & R AND (2001) pointed out that leucopygius and H A R T M A N N et al. (2004) for
Eleutherodactylus diastema Stejneger, 1904, a Aplastodiscus sp. (aff. ehrhardti) that call sites are
nocturnal species, probably uses visual cues during on leaves under water and the oviposition site are
reproductive display, as is known for Phyllomedusa subterranean nests. Aplastodiscus arildae males
distincta Lutz, 1950 (CASTANHO, 1994). Aplastodiscus used leaves above the water as calling sites too.
sp. (aff. ehrhardti) presents a visual signaling and a Although we did not observe the direct events of
diverse repertoire of limb movements exhibited by oviposition, we considered the courtship behavior
both sexes. Male and female show a stereotyped described for A. arildae as evidence that this species
behavior, moving the limbs up and down, alternating deposits eggs in subterranean nests.
positions, and sometimes being face to face, The reproductive behavior of A. arildae is similar to
sometimes side by side (HARTMANN et al., 2004). that described for A. leucopygius (see HADDAD & SAWAYA,
Courtship in this species has greater duration 2000) and Aplastodiscus sp. (aff. ehrhardti) (HARTMANN
compared to other amphibians and this long et al., 2004). Thus, the reproductive mode for A.
interaction between males and females may have at arildae consisted of: aquatic eggs; eggs and early larval
least three purposes: (1) to evaluate the reproductive stages in subterranean constructed nests;
condition of the mate, because elaborate behavior subsequent to flooding, exotrophic tadpoles in ponds
may indicated physiological condition and individual or streams (mode 5, sensu HADDAD & PRADO, 2005).
attributes, influencing acceptance or refusal of the Further studies should be addressed to precisely
mate; (2) to stimulate ovulation, as relative long evaluated this aspect. Therefore, the evidences of
periods of courtship may be necessary to trigger equal events on courtship behavior and reproductive
ovulation; and/or (3) to lead the female from the mode for A. arildae, A. leucopygius (see HADDAD &
calling site to the nest passing obstacles such as SAWAYA, 2000) and Aplastodiscus sp. (aff. erhardti) (see
leaves, trunks, and roots that could obstruct HARTMANN et al., 2004), even though to A. perviridis
progress to the nest (see HARTMANN et al., 2004). (H ADDAD et al., 2005) may have confirm the
Inasmuch as A. arildae is nocturnal, we believe that monophyletism of this group as suggested by HADDAD
the decreased moonlight reduces the probability of et al. (2005) and proposed by FAIVOVICH et al. (2005).
visual communication, based on the fact that the male The advertisement call here described presents
emitted courtship calls and waited for the female to differences for dominant frequency and interval
touch him. In contrast with Aplastodiscus sp. (aff. among calls from that reported by HEYER et al.

Arq. Mus. Nac., Rio de Janeiro, v.64, n.3, p.247-254, jul./set.2006

 COURTSHIP BEHAVIOR OF APLASTODISCUS ARILDAE 253

(1990) for A. arildae at Boracéia, São Paulo, Brazil. CRUZ, C.A.G. & PEIXOTO, O.L., 1985. Espécies verdes
The maximum frequency in their sonogram was de Hyla: o complexo “albofrenata” (Amphibia, Anura,
approximately 4000Hz and the calls intervals Hylidae). Arquivos da Universidade Federal Rural do
ranged from 0.7 to 1.7s (21.8ºC air temperature). Rio de Janeiro, 7(1-2):59-70.
The advertisement and courtship calls from the DUELLMAN, W.E., 1985. Reproductive modes in
present study were recorded from the same male, anuran amphibians: phylogenetic significance of
at the same night, by the same air temperature, adaptive strategies. South African Journal of Science,
but over different behaviors. The advertisement 81:174-178.
calls was emitted at the absence of female and
the courtship calls were only emitted when the DUELLMAN, W.E. & TRUEB, L., 1986. Biology of
Amphibians. New York: McGraw-Hill. 670p.
male and female begun the trajectory to the
oviposition site. FAIVOVICH, J.; CRUZ, C.A.G. & PEIXOTO, O.L., 2002.
The identity of Hyla ehrhardti Muller, 1924 (Anura:
Hylidae). Journal of Herpetology, 36(2):325-327.
ACKNOWLEDGEMENTS
FAIVOVICH, J.; HADDAD, C.F.B.; GARCIA, P.C.A.;
We thank administrative and security of Parque FROST, D.R.; CAMPBELL, J.A. & WHEELER, W.C., 2005.
das Mangabeiras which facilitated our work in Systematic review of the frog family Hylidae, with special
the field. C.A.G.Cruz, U.Caramaschi (Museu reference to Hylinae: phylogenetic analysis and
Nacional, Rio de Janeiro), C.F.B.Haddad taxonomic revision. Bulletin of the American Museum
(Universidade Estadual Paulista, Rio Claro, SP), of Natural History, 294:1-240.
A.S.Rand (in memoriam) and an three
FARIA, D.M.; CASAIS SILVA, L.L. & RODRIGUES, M.T.,
anonymous reviewers kindly reviewed the 1993. Nota sobre reprodução de Hylodes phyllodes (Anura,
manuscript offering helpful suggestions. We Leptodactylidae). In: CONGRESSO LATINO AMERICANO
acknowledge G.C.Zorzin, for the illustration, DE HERPETOLOGIA, 3., 1993, Campinas. Resumos...
F.M.H.Nunes, for helping in fieldwork, and Campinas: Universidade Estadual de Campinas. p.150.
H.Paprocki (PUC Minas Gerais) for the english
review. C.A.B. Galdino received PhD grant from FROST, D.R., 2004. Amphibian Species of the World –
an on line reference. Disponível em: <http://
Coordenação de Aperfeiçoamento de Pessoal de
research.amnh.org/herpetology/amphibia/index.html>.
Nível Superior (CAPES) and L.B.Nascimento is
Acesso em: 10 jun. 2005.
grateful to Fundo de Incentivo à Pesquisa (FIP)/
PUC Minas for the financial support. GRIDI-PAPP, M. 2003-2004. Sound Ruler: Acoustic
Analysis, versão 0.941. Disponível em: <http://
soundruler.sourceforge.net/oldsite/index-br.htm>.
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