Herpetology Notes, volume 15: 507-512 (2022) (published online on 19 July 2022)
Aggressive interaction in the Hoogmoed Harlequin Toad
Atelopus hoogmoedi (Anura: Bufonidae), with the description of
its encounter call
André G. Lopes1,2,*, Patrick R. Sanches3, and Carlos E. Costa-Campos4
Atelopus Duméril & Bibron, 1841 comprises 99
species of bufonid toads, mostly diurnal and terrestrial,
distributed from Costa Rica to Bolivia, in northern
Venezuela, and throughout the Guiana Shield and its
surroundings (Frost, 2021). The acoustic knowledge
on this genus is deficient, since vocalisations have
been reported in just over one fifth of its total number
of species. Four main call types have been recognised
within its acoustic repertoire, based on their structure
(pulsed or tonal) and duration: pulsed (92–1240 ms),
pulsed short (5–171 ms), pure tone (159–420 ms), and
pure tone short calls (8–135 ms) (sensu Lötters et al.,
2019; but see updated duration ranges in the caption of
Table 1 herein).
Although there is a consensus that the pulsed call
corresponds to the advertisement call (e.g., Cocroft et
al., 1990; Lötters et al., 2019), the functions associated
to the other call types are partly inconclusive (Jaslow,
1979; Cocroft et al., 1990; Lötters et al., 2002a, 2019). It
has been suggested that the pulsed short call might have
a release (Lötters et al., 2019) or advertisement function
(Granda-Rodríguez et al., 2020), or that it might be
related to some sort of aggressive context (Cocroft et al.,
1990). There is also little agreement on a main specific
1
Laboratório de Taxonomia e Sistemática de Anuros
Neotropicais, Instituto de Ciências Exatas e Naturais do
Pontal, Universidade Federal de Uberlândia, Ituiutaba, Minas
Gerais 38304-402, Brazil.
2
Departamento de Biologia, Faculdade de Filosofia, Ciências
e Letras de Ribeirão Preto, Universidade de São Paulo,
Ribeirão Preto, São Paulo 14040-901, Brazil.
3
Instituto Nacional de Pesquisas da Amazônia, Manaus,
Amazonas 69080-971, Brazil.
4
Laboratório de Herpetologia, Departamento de Ciências
Biológicas e da Saúde, Universidade Federal do Amapá,
Macapá, Amapá 68903-419, Brazil.
*
Corresponding author. E-mail: gomesandrebio@gmail.com
© 2022 by Herpetology Notes. Open Access by CC BY-NC-ND 4.0.
function of the tonal calls; it has been suggested that the
pure tone call might play an aggressive (Jaslow, 1979;
Lötters et al., 2019; Jorge et al., 2020; Rueda-Solano
et al., 2020) or release function (Rueda-Solano et al.,
2020), and that the pure tone short call might play an
aggressive (Cocroft et al., 1990; Lötters et al., 2019;
Rueda-Solano et al., 2020), release (Ibáñez et al., 1995;
Lötters et al., 1999, 2019; Carvajalino-Fernández et
al., 2017; Rueda-Solano et al., 2020) or advertisement
function (Granda-Rodríguez et al., 2020), or that it even
might play a role during the amplexus (Cocroft et al.,
1990; Lötters et al., 2019). Recently, Rueda-Solano et
al. (2020) reported the first case of female vocalisation
in the genus, by describing the release call of a female of
Atelopus laetissimus Ruiz-Carranza et al., 1994, which
consists of short and long unpulsed notes.
Atelopus hoogmoedi Lescure, 1974 is a small-
sized species distributed in the eastern Guiana Shield
encompassing Guiana, Suriname, French Guiana, and
Brazil, and in surrounding Brazilian regions (Frost,
2021). Some studies have reported on its habitat (Kok
and Kalamandeen, 2008; Luger et al., 2009; Ouboter
and Jairam, 2012; Nicolaï et al., 2017), behaviour (Luger
et al., 2009; Almeida-Santos et al., 2016; Nicolaï et al.,
2017; Rössler, 2017; Rößler et al., 2019; Oliveira-Souza
et al., 2020), and vocalisation (Lescure, 1981; Cocroft et
al., 1990; Kok and Kalamandeen, 2008; Costa-Campos
and Carvalho, 2018). So far, the only call type reported
for this species is the pulsed call (i.e., advertisement
call).
In the present study, in order to improve the knowledge
on Atelopus hoogmoedi, we report an unprecedented
aggressive interaction between two males of this
species, and also describe its encounter call. This call
type comprised both pure tone and pure tone short calls,
which are reported for the first time in this species. We
also provide a summary of the call types reported for
each Atelopus species to date.
The aggressive interaction occurred on 14 February
508 André G. Lopes et al.

Table 1. Call types reported for Atelopus species. * Described in this study. 1 Other calls were reported for this species. 2 Referred
to as “partially pulsed” by Cocroft et al. (1990). Duration ranges: pulsed calls (92–2800 ms); pulsed short calls (20–171 ms); pure
tone calls (137–420 ms); pure tone short calls (5–165 ms). Table and duration ranges modified from Lötters et al. (2019).

Species Pulsed Pulsed Pure tone Pure tone References

call
A. barbotini1 Lescure, 1981
A. carbonerensis1 Rivero, 1974
A. certus Barbour, 1923
A. chiriquiensis Shreve, 1936
A. cruciger1 (Lichtenstein & Martens, 1856)
A. exiguus (Boettger, 1892)
A. flavescens1 Duméril & Bibron, 1841
A. franciscus Lescure, 1974
A. fronterizo Veselý & Batista, 2021
A. glyphus Dunn, 1931
A. hoogmoedi Lescure, 1974
A. laetissimus1 Ruiz-Carranza et al., 1994
A. limosus Ibáñez et al., 1995
A. manauensis Jorge et al., 2020
A. minutulus1 Ruiz-Carranza et al., 1988
A. mucubajiensis Rivero, 1974
A. nahumae Ruiz-Carranza et al., 1994
A. nicefori Rivero, 1963
A. peruensis Gray & Cannatella, 1985
A. pulcher (Boulenger, 1882)
A. reticulatus1 Lötters et al., 2002
A. senex Taylor, 1952
A. spumarius Cope, 1871
A. tamaense La Marca et al., 1990
A. tricolor Boulenger, 1902
A. varius (Lichtenstein & Martens, 1856)
Atelopus cf. loettersi De la Riva et al., 2011
Atelopus sp. 'Itaya'
short call call short call
X Lescure (1981); Lescure and Marty (2000) (as A. spumarius
barbotini)
X X X X Lötters et al. (2019)
X Veselý and Batista (2021)
X X X Jaslow (1979); Lötters et al. (1999)
X X2 X Cocroft et al. (1990)
X Coloma et al. (2000)
X Lescure (1981); Lescure and Marty (2000)
X Lescure (1981); Lescure and Marty (2000); Boistel et al. (2011)
X X2 Cocroft et al. (1990) (as Atelopus sp.); Veselý and Batista (2021)
X Veselý and Batista (2021)
X X* X* Lescure (1981) (as A. spumarius hoogmoedi); Cocroft et al. (1990)
(as A. s. hoogmoedi); Kok and Kalamandeen (2008); Costa-Campos
and Carvalho (2018); present study
X X X X Granda-Rodríguez et al. (2020); Rueda-Solano et al. (2020)
X X Ibáñez et al. (1995)
X X X X Jorge et al. (2020)
X X2 X Cocroft et al. (1990)
X X Lötters et al. (2019)
X Carvajalino-Fernández et al. (2017)
X Cocroft et al. (1990)
X Lötters et al. (1999)
X Lötters et al. (2002b)
X X X Lötters et al. (2002a)
X Cocroft et al. (1990)
X Lescure (1981) (as A. spumarius spumarius)
X Lötters et al. (2019)
X X Lötters et al. (1999)
X X2 X Cocroft et al. (1990) (in part, also as A. zeteki)
X2 Cocroft et al. (1990) (as A. spumarius spumarius)
X Asquith and Altig (1987) (as. A. spumarius)

2015, at ca. 09:54 h, in a terra-firme forest next to a
10 m wide stream, located in the Colônia de Água
Branca do Amapari, municipality of Serra do Navio,
Amapá state, Brazil (0.9410°N, 051.9945°W; datum
WGS84). A video of the interaction was recorded
during 02:03 minutes, with a Sony HDR-CX 440 Full
HD camera. Specimens were collected, euthanised with
5% lidocaine, fixed in 10% formalin, preserved in 70%
ethanol, and deposited in the Herpetological Collection
of the Universidade Federal do Amapá (accession
numbers: CECC 3209, CECC 3210). The video file
was deposited in this same collection (label: Atelopus_
hoogmoedi_3209_3210) and also in the Fonoteca
Neotropical Jacques Vielliard (access code: ZUEC-
VID 977). The audio from the video was extracted and
converted to a readable audio file (.wav) (sampling
rate of 48.0 kHz; 16 bits resolution), for the purpose
of analysing the calls emitted by the individuals. Prior
to the analysis, the audio file was filtered to reduce
background noise and then normalised (peak -1.0 dB)
in Audacity v. 2.2.2 software (Audacity Team, 2021).
Calls were analysed in Raven Pro v. 1.5 software (K.
Lisa Yang Center for Conservation Bioacoustics,
2022) with the following settings: window size = 1024
samples; 3 dB filter bandwidth = 67.4 Hz; window
type = Hann; overlap = 90% (locked); hop size = 2.13
ms; DFT size = 1024 samples; grid spacing = 46.9 Hz.
Acoustic terminology and definitions followed Köhler
et al. (2017). Dominant, maximum, and minimum
frequency values were respectively obtained with the
“Peak Frequency”, “Frequency 95%”, and “Frequency
5%” functions. Frequency modulation was calculated as
“dominant frequency of the final 10% portion of the call
- dominant frequency of the initial 10% portion”. Sound
Aggressive interaction in the Hoogmoed Harlequin Toad 509

figures were generated in R platform v. 3.6.2 (R Core
Team, 2019) with seewave v. 2.1.6 (Sueur et al., 2008)
and tuneR v. 1.3.3 (Ligges et al., 2018) packages, under
the following settings: window = Hanning; overlap =
85%; FFT = 1024.
The two males of Atelopus hoogmoedi were found in
the leaf litter aggressively interacting at a very close-
range. First, males were facing each other ca. 15 cm
apart, with only male 1 (CECC 3209; snout-vent length
of 22.90 mm) emitting encounter calls. After the video
recording started, male 1 emitted four calls (Fig. 1A)
and took a step towards male 2 (CECC 3210; snout-vent
length of 30.20 mm), emitted two more calls, slightly
moved its left front limb forward and emitted another
call. Then male 2 took a jump away from male 1 (Fig.
1B), but this latter took a short jump in its direction.
Immediately, male 2 took another jump away from male
1, and the latter chased it by taking another jump, but
male 2 promptly jumped away again. At this moment
the individuals were ca. 50 cm apart; male 1 directed
its body to male 2, emitted two calls and took a step
forward, then emitted another call and took a jump
forward. Then, male 1 emitted another call, took a
step forward and emitted five calls (Fig. 1C), being
that during the emission of its fifth call, male 2 started
vocalising. In this moment male 1 stopped vocalising,
while male 2 emitted three consecutive encounter
calls (only two were analysed here, since the first one
overlapped with a call of male 1). Immediately after
male 2 emitted the third call, male 1 took a jump in its
direction and emitted two calls, and then male 2 stopped
calling. From then on, male 1 stood directed to male 2
and started a continuous emission of advertisement calls
(i.e., pulsed call) for 01:23 minute (Fig. 1D). Then male
2 moved away without calling, and both individuals
were collected.
The encounter calls emitted by male 1 correspond to
pure tone calls (Fig. 2A). This call type (n = 16 calls)

Figure 1. Aggressive interaction between male 1 (CECC 3209; white arrow) and male 2 (CECC 3210; red arrow) of Atelopus
hoogmoedi. (A) Individuals in close-range interaction, with male 1 emitting encounter calls. (B) Moment in which male 2 took
the first jump away from male 1. (C) Male 1 emitting encounter calls, with its body directed to male 2. (D) Male 1 emitting
advertisement calls, with its body directed to male 2. These figures are frames extracted from the video recording of the interaction
(video access code: ZUEC-VID 977; see text for details). Video recorded by Rubenilson Oliveira Pinto.
510
consists of a single tonal note which sounds like a soft
whistle, has duration of 255 ± 33 ms (190–316 ms)
and inter-call interval of 1922 ± 1134 ms (924–5220
ms). Calls have a slight upward frequency modulation
over their duration, followed by a minor downward
modulation at their very final portion; in most calls (n
= 11) there is also a slight downward modulation at
the very onset before the upward modulation. The call
frequency modulation is of 70 ± 82 Hz (-94–188 Hz).
Calls have the dominant frequency at 2150 ± 23 Hz
(2109–2203 Hz), minimum frequency at 2051 ± 50 Hz
(1969–2156 Hz), and maximum frequency at 2203 ± 17
Hz (2156–2250 Hz).
The encounter calls emitted by male 2 correspond
to pure tone short calls (Fig. 2B). This call type (n =
2 calls) consists of a single tonal note which sounds
like a short peep, has duration of 137 ± 11 ms (129–
144 ms) and inter-call interval of 596 ± 0 ms. Calls are
non-frequency modulated. Calls have the dominant
frequency at 2344 ± 0 Hz, minimum frequency at 2273
± 33 Hz (2250–2297 Hz), and maximum frequency at
2391 ± 0 Hz.
The observed interaction revealed an escalated
aggressive behaviour in Atelopus hoogmoedi, beginning
with encounter call emissions, and escalating to
approaching attempts and chasing. Although we did
not observe what initially triggered the interaction,
we cogitate that male 2 invaded the territory of male
1. While male 1 showed a more combative behaviour
by vocalising most of the time and chasing male 2, the
Figure 2. Spectrograms (above) and respective oscillograms (below) of the encounter calls emitted by the two individuals of
Atelopus hoogmoedi during the aggressive interaction. (A) A pure tone call emitted by the individual CECC 3209. (B) A pure tone
short call emitted by the individual CECC 3210.
André G. Lopes et al.
latter just tried to escape and emitted only three calls.
Such behaviours are in line with the previously reported
in other agonistic interactions in anurans (e.g., Martins
et al., 1998; Bastos and Haddad, 2002; Guimarães and
Bastos, 2003), supporting the hypothesis that male 1
was the resident and male 2 the intruder.
The pure tone call and the pure tone short call are
herein reported for the first time in Atelopus hoogmoedi.
Since these two call types were emitted by males during
a close-range aggressive interaction, it can be reliably
assumed that both compose the encounter call of the
species, thus playing an aggressive function. Some
previous studies on vocalisations of Atelopus species
also assigned an aggressive function to pure tone
(Jaslow, 1979; Lötters et al., 2019; Jorge et al., 2020;
Rueda-Solano et al., 2020) and pure tone short calls
(Cocroft et al., 1990; Lötters et al., 2019; Rueda-Solano
et al., 2020).
In addition to a slight audible difference, these tonal
calls of Atelopus hoogmoedi can be distinguished from
each other mainly because the pure tone call is longer
and has lower values for spectral traits than the pure
tone short call. Besides, both calls allow to distinguish
A. hoogmoedi from its congeners. Pure tone calls of A.
hoogmoedi differ from those of congeners in dominant
frequency (lower than in A. cruciger, A. manauensis, A.
minutulus, A. mucubajiensis, A. reticulatus; higher than
in A. carbonerensis, A. chiriquiensis, A. laetissimus),
minimum frequency (lower than in A. manauensis;
higher than in A. carbonerensis, A. laetissimus),
Aggressive interaction in the Hoogmoed Harlequin Toad
maximum frequency (lower than in A. manauensis and A.
mucubajiensis), and frequency modulation (lower than
in A. cruciger and A. minutulus). Pure tone short calls of
A. hoogmoedi differ from those of congeners in duration
(longer than in A. carbonerensis, A. chiriquiensis, A.
laetissimus, A. limosus, A. mucubajiensis, A. peruensis,
A. reticulatus, A. tamaense, A. tricolor, A. varius),
dominant frequency (lower than in A. manauensis, A.
mucubajiensis, A. reticulatus, A. tamaense; higher than
in A. carbonerensis, A. chiriquiensis, A. laetissimus,
A. limosus, A. nahumae, A. peruensis, A. varius),
minimum frequency (lower than in A. manauensis;
higher than in A. carbonerensis, A. laetissimus, A.
limosus, A. mucubajiensis), and maximum frequency
(lower than in A. manauensis, A. tamaense; higher
than in A. carbonerensis, A. laetissimus). See Table 1
for a summary of the call types reported for Atelopus
species.
This is the first report of aggressive interaction in
males of Atelopus hoogmoedi, thus contributing to
the knowledge on the behavioural repertoire of this
species. The study also adds new data to the acoustic
repertoire of A. hoogmoedi, and provide evidence that
may be relevant towards a better understanding on
the actual function of the tonal calls in the genus. The
present contributions may help to build up a framework
for future studies that aim to understand the evolution
of the different call types in Atelopus, as well as their
respective underlying functions.
Acknowledgments. We thank Albertina Pimentel Lima for
the pre-peer review on the original version of this study, and
an anonymous reviewer for making valuable comments and
suggestions. We thank Christoph Jaster (PARNA Montanhas do
Tumucumaque) for the logistical support during the field­work,
and Rubenilson O. Pinto for kindly providing its recording of the
interaction. We thank Simone Dena for the help in depositing the
video in the Fonoteca Neotropical Jacques Vielliard. Conselho
Nacional de Desenvolvimento Científico e Tecnológico (CNPq)
granted a master’s fellowship to AGL (process n. 130380/2020-
2). The Cornell Lab of Ornithology (Center for Conservation
Bioacoustics) provided a free license of Raven Pro 1.5 software
to AGL. Specimens were collected under the collection
permits provided by Instituto Chico Mendes de Conservação
da Biodiversidade/Sistema de Autorização e Informação em
Biodiversidade (process n. 32651-1).
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Accepted by Fabrício Hiroiuki Oda