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\begin{document}

\title{INFLUENCE OF CONSPECIFICS AND ROAD NOISE ON THE PROPERTIES OF THE
ADVERTISEMENT CALL OF PHYLLODYTES LUTEOLUS (ANURA, HYLIDAE)}



\author[1]{Leonardo Marques de Abreu}%
\author[2]{Vinicius Guerra Batista}%
\author[3]{Mirco Sole}%
\affil[1]{Universidade Estadual de Santa Cruz}%
\affil[2]{Federal University of Mato Grosso - Campus Sinop}%
\affil[3]{State University of Santa Cruz Department of Biological Sciences}%


\vspace{-1em}



  
  \date{May 25, 2024}


\begingroup
\let\center\flushleft
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\selectlanguage{english}
\begin{abstract}
Acoustic communication in animals can be affected by multiple biotic
(intra and interspecific) and abiotic (e.g., wind and rain) natural
noises. In addition, human beings produce additional novel sources of
noise, which can reduce or inhibit the reception of acoustic signals by
conspecifics, leading to behavioral changes. In this study, we
investigated whether sound of conspecifics and road noise additively
affect the acoustic parameters of the advertisement call of males of a
Yellow Heart-tongued Frog (Phyllodytes luteolus). We hypothesized that
males that vocalize in larger choruses (many males) and in areas close
to highways (anthropic noise) will increase their temporal and spectral
acoustic parameters, respectively, to avoid acoustic signal masking. We
recorded the vocalizations of 38 males in environments close (N=18) to
and distant (N=20) from highways in different social contexts (many or
few individuals in the chorus). Contrary to our expectation, the results
indicated that individuals exposed to road noise had lower dominant
frequency calls than those from natural areas, and that the density of
males in the chorus had no influence on the acoustic parameters.
Furthermore, we found a positive relationship between body size and
intensity, indicating that larger individuals can emit calls that can
reach greater distances. The advertisement call of Phyllodytes luteolus
has a high dominant frequency, with little overlap with the frequency of
anthropic noises (roads), which may explain its presence and
reproductive success of this species in bromeliads from urbanized areas.%
\end{abstract}%



\sloppy


\textbf{INFLUENCE OF CONSPECIFICS AND ROAD NOISE ON THE PROPERTIES OF
THE ADVERTISEMENT CALL OF \emph{PHYLLODYTES LUTEOLUS} (ANURA, HYLIDAE)}

Leonardo Marques de Abreu \textsuperscript{a,c,*}, Vinicius
Guerra\textsuperscript{b}, Mirco Sol\selectlanguage{ngerman}é \textsuperscript{a,c,d}.

\textsuperscript{a} Programa de Pós-graduação em Ecologia e Conservação
da Biodiversidade, Universidade Estadual de Santa Cruz, Ilhéus, Bahia,
Brazil.

\textsuperscript{b} Laboratório de Herpetologia e Comportamento Animal,
Instituto de Ciências Biológicas, Universidade Federal de Goiás,
Goiânia, GO, Brazil.

\textsuperscript{c} Laboratório de Herpetologia Tropical, Universidade
Estadual de Santa Cruz, Ilhéus, Bahia, Brazil

\textsuperscript{d} Herpetology Section, Zoologisches Forschungsmuseum
Alexander Koenig, Adenauerallee 160, Bonn, Germany.

\textbf{*Corresponding author} : Leonardo Marques de Abreu /
leoomarques95@gmail.com



\section*{Abstract}

{\label{abstract}}

Acoustic communication in animals can be affected by multiple biotic
(intra and interspecific) and abiotic (e.g., wind and rain) natural
noises. In addition, human beings produce additional novel sources of
noise, which can reduce or inhibit the reception of acoustic signals by
conspecifics, leading to behavioral changes. In this study, we
investigated whether sound of conspecifics and road noise additively
affect the acoustic parameters of the advertisement call of males of a
Yellow Heart-tongued Frog (\emph{Phyllodytes luteolus} ). We
hypothesized that males that vocalize in choruses (males calling nearby)
and in areas close to highways (anthropic noise) will increase their
temporal and spectral acoustic parameters, respectively, to avoid
acoustic signal masking. We recorded the vocalizations of 38 males in
environments close (N=18) to and distant (N=20) from highways in
different social contexts (many or few individuals calling nearby).
Contrary to our expectation, the results indicated that males calling in
areas close to highways had lower dominant frequency calls than those
from natural areas (far from highways), and that the density of males in
the chorus had no influence on the acoustic parameters. Furthermore, we
found a positive relationship between body size and intensity,
indicating that larger individuals can emit calls that can reach greater
distances. The advertisement call of \emph{Phyllodytes luteolus} has a
high dominant frequency, with little overlap with the frequency of
anthropic noises (roads), which may explain its presence and
reproductive success of this species in bromeliads from urbanized areas.

\textbf{Keywords:} Noise pollution, acoustic communication, anurans,
mate attraction, bromeligens.

\section*{Introduction}

{\label{introduction}}

Acoustic communication is widely used by animals to transmit information
through sounds. These sounds are produced by an emitter (source) and are
propagated in the environment, causing some response in the receivers
(Kime, 2000; Wells, 2000). In environments where acoustic signals
propagate, transmission can be hampered by other different sounds,
causing stress, irritability, reduced fitness, in addition to being
associated with other risk situations (Grenat et al., 2019; Leon et al.,
2019; Troianowski et al., 2017). There are three main types of sound
noise that interfere with the transmission and detection of the species'
acoustic signal: abiotic (environmental), such as the presence of winds,
rains, streams and ocean tides (Caldart et al., 2016); biotic, produced
by intra and interspecific individuals that can form dense social groups
(Lengagne, 2008); and anthropic, which are related to acoustic pollution
caused by humans, such as the flow of automobiles on highways, civil
construction machinery, air transport, ships and boats (Cunnington \&
Fahrig, 2010, 2013). The interference caused by these noises can
negatively influence the fitness of individuals, and consequently affect
populations and communities (Hanna et al., 2014; Hoskin \& Goosem,
2016). Over the last few decades, there has been an increase in studies
on the effects of noise on the acoustic communication of organisms
(Gomes et al., 2022; Grenat et al., 2019). One of the groups of animals
most affected by human noise are anuran amphibians, which use acoustic
signals as their main form of communication (Gomes et al., 2022; Wells,
2007).

During the breeding period, most anurans form dense aggregations in
water bodies (Wells, 2007). Communication between frogs occurs mainly
through the emission of different types of vocalizations (Toledo et al.,
2015), however, the most emitted acoustic signal is the advertisement
call, which has the main function of attracting reproductive partners
and delimiting territories (Guerra et al., 2018; Toledo et al., 2015;
Wells, 2007). The calls of conspecific individuals (and also of other
species) can represent biotic sound noises that interfere in the local
acoustic space. Thus, males in vocalization activity must avoid the
overlapping of these acoustic signals (e.g., temporal and spectral
parameters) in some way (Bittencourt et al., 2016; Herrera-Montes \&
Aide, 2011). Dense choruses of males in vocalizing activity may also
lead to limitations in the ability of females to choose reproductive
partners (Wollerman \& Wiley, 2002).

In addition to biotic noises, more attention has recently been directed
to anthropic noises. This type of noise alters the conditions of the
acoustic environment of many habitats, creating new environmental
pressures that directly affect many animals that communicate
acoustically, including frogs (Barber et al., 2010; Desrochers \&
Proulx, 2017; Knight \& Swaddle, 2011; Sabah et al., 2017), birds
(Bermúdez-Cuamatzin et al., 2009; Herrera-Montes \& Aide, 2011;
Slabbekoorn \& Ripmeester, 2008) and marine mammals (Melcón et al.,
2012; Moore \& Clarke, 2002; Stocker, 2002). Among the anthropic noises,
highways are considered the biggest source of noise pollution, producing
sounds with high energies concentrated in low frequencies (\textless{}5
kHz) (Warren et al., 2006). The urban expansion, and consequently the
road network, not only decreases the availability of habitats but also
increases the amount of human noise, causing negative effects on the
transmission and reception of sound between conspecifics (Bittencourt et
al., 2016; Sun \& Narins, 2005), and may even reduce the chances of
survival of individuals (Gomes et al., 2022; Herrera-Montes \& Aide,
2011). However, the species show several solutions to solve the problems
in the communication limitation imposed by the noises, such as, changing
the temporal and spectral acoustic parameters of the calls to reduce the
noise masking effect (Cunnington \& Fahrig, 2010, 2013; Grenat et al.,
2019).

Among the strategies used by anurans to reduce or avoid the overlap
between biotic and anthropic noises on their calls, there are changes in
amplitude (Halfwerk et al., 2016; Parris et al., 2009; Yi \& Sheridan,
2019), frequency (Caorsi et al., 2017; Cunnington \& Fahrig, 2010),
duration (Zhao et al., 2021) and emission rate (Hanna et al., 2014;
Kaiser \& Hammers, 2009; Legett et al., 2020). These changes can be
advantageous when the individuals are under external influences, since
the acoustic signals indicate the physical condition of the individuals.
Therefore, they must be transmitted in the best possible way in the
environment (Cunnington \& Fahrig, 2010; Kime, 2000), just as the
acoustic adaptation hypothesis predicts (Goutte et al., 2018; Morton,
1975). Thus, changes in parameters of the call may indicate an
adaptation in response to noise, but they may generate additional
fitness costs, negatively affecting survival and reproductive success
(Herrera-Montes \& Aide, 2011). It is often difficult to find evidence
that suggests that changes in the calls of individuals observed in
nature are caused by a single factor (Grenat et al., 2019), as
variations and/or adjustments in acoustic parameters can be influenced
by the environment (abiotic factors; (Kime et al., 2000)), size of
choirs (social factors; (Gambale \& Bastos, 2014; Morais et al., 2012))
and/or level of human noise (Caorsi et al., 2017). Therefore, there may
be confounding factors when trying to explain changes in behavior if the
study does not consider the multiple biological and environmental
aspects to which individuals are exposed.

Biotic factors (body size, weight, predation and abundance of males in
vocalization activity) and abiotic factors (temperature, humidity and
vegetation heterogeneity) influence the anurans vocalizations in
different ways. For example, body size influences the spectral structure
(frequency) of the call, so that larger individuals present calls with
lower frequencies (Kohler et al., 2017). Thus, acoustic signals provide
reliable information about male body size (Bastos et al., 2011; Morais
et al., 2012). The number of individuals in the chorus influences the
intensity of the call as males increase sound pressure to promote
greater attractiveness (Bastos \& Haddad, 2002; Morais et al., 2012). As
frogs are ectothermic animals, temperature influences the metabolic
rate, reflecting changes in the temporal parameters of calls, such as
duration and emission rate (Bastos \& Haddad, 2002; Furtado et al.,
2016). All these aspects must be considered in bioacoustics studies to
avoid bias in the interpretation of results.

Since human activities have impacted the behavior of amphibians in
different ways, in this work we evaluated whether the call of a Yellow
Heart-tongued Frog species is affected by noise pollution produced by
car traffic on highways and by the noise of conspecifics in the chorus.
We hypothesized that (1) males exposed to anthropic noise (road traffic)
will present a higher dominant frequency of the advertisement call to
decrease or avoid signal masking, and that (2) males that vocalize in
conspecific choruses with higher density of individuals will present
higher values in the temporal parameters of the call (e.g., longer call
duration and intensity) to increase the efficiency in signal
transmission (and reduce or avoid overlapping of the call) in the
environment. For this, we compared the acoustic parameters of
advertisement calls of males of \emph{Phyllodytes
luteolus}(Wied-Neuwied, 1821) from natural and urban environments, and
in the presence of loud and quiet choruses. \emph{Phyllodytes luteolus}
is an excellent model organism to test these hypotheses because it is a
common species, forms reproductive choruses, uses acoustic signals as
the main form of communication and is found in bromeliads in natural and
urban environments (Forti et al., 2017; Salles \& Silva-Soares, 2010).

\section*{Materials and methods}

{\label{materials-and-methods}}

\emph{Phyllodytes luteolus} (Wied-Neuwied, 1821) (Figure 1) is a species
of anuran amphibian belonging to the Hylidae family. This species spends
its entire life cycle inside bromeliads and is considered to have a
bromeliad habit (Peixoto, 1995). They prefer bromeliads with greater
complexity of structures and when they form a network connected by
several individuals of the same bromeliad species (Ferreira et al.,
2012; Mageski et al., 2016). Reproduction is prolonged, occurring
throughout the year (Ferreira et al., 2012).

The advertisement call of \emph{P. lutelus} consists of a pulsed call,
with a series of notes without frequency modulation with an average
duration of 5 seconds, and dominant frequencies ranging between 2 and 4
kHz (Cruz et al., 2014; Simon \& Gasparini, 2003). \emph{P.
luteolus}males begin vocalization activity around two hours after sunset
(7 pm). The tadpoles can prey on disease vector mosquito larvae acting
in the biological control of possible diseases transmitted by these
organisms (Salinas et al., 2018). In addition, this species has a diet
specialized in ants with pharmacological potential (Solé \& Loebmann,
2017).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image1/image1}

\end{center}
\end{figure}

\textbf{Figure 1.} Male individual of Phyllodytes luteolus
(Wied-Neuwied, 1821) on the leaves of the bromeliad Aechmea
blanchetiana. Vila do Acu\selectlanguage{ngerman}ípe, municipality of Ilhéus, state of Bahia,
Brazil. Photo: João Victor A. Lacerda.

Field activities were carried out during the month of October 2021 at
five sampling sites in the municipality of Ilhéus, Bahia, Brazil. All
areas have bromeliads used as reproductive sites by \emph{Phyllodytes
luteolus} . Three sites with the presence of vehicle traffic noise were
located next to the BA-001 highway (between 3 and 10 meters away from
the highway), in the stretches between Olivença and Acuípe villages, in
the municipality of Ilhéus. The BA-001 highway runs along the coast of
the state of Bahia and its southern end begins in the city of Mucuri
while the northern end is located on Itaparica Island, in the
Todos-os-Santos Bay of the capital Salvador. The stretch of road on
which it was selected based on the presence of individuals of \emph{P.
luteolus} , at night periods the frequency of vehicles decreased
markedly when compared to daytime periods, but there was the presence of
vehicles in passages. There is no data available from government
agencies regarding the flow of vehicles on this stretch of the highway,
but during our field activities we found that the greatest traffic
occurs between 5 and 7 pm. Two other sampling sites without the presence
of vehicle traffic noise were defined at a minimum distance of two
kilometers from the BA-001 highway (Carr \& Fahrig, 2001) (Figure 2),
where in these environments, as a form of control, there was no other
source of anthropic noise, eventually few noises from other species that
actively communicate acoustically.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image2/image2}

\end{center}
\end{figure}

\textbf{Figure 2.} Breeding sites sampled in the municipality of Ilh\selectlanguage{ngerman}éus,
state of Bahia, northeastern Brazil. Sites A (-15.0844°S, -39.0080°W)
and B (-15.0844°S, -39.0133°W) were two kilometers far from highways
with car traffic (natural environments and no human noise), while sites
C (-14.9398°S, -39.0126 °W), D (-14.9289°S, -39.0177°W) and E
(-15.0866°S, -38.9987°W) were two a ten meters close to highways with
vehicle traffic (noise pollution), the distance from locations C and D
to E was 15 kilometers.

The sampling sites were defined as Natural (no traffic noise) or
Highways (with traffic noise). For each recorded individual, we obtained
the social situation in which the male was inserted, being: with chorus
(four or more conspecific individuals in vocalization activity within a
three-meter radius -- CC) or without chorus (four or less conspecific
individuals vocalizing within a radius of three meters -- SC).

Males of \emph{Phyllodytes luteolus} in vocalization activity were found
through visual and auditory searches between 19:00 and 24:00 hrs. The
sampling points were quite representative due to their extensive areas
where they contained many individuals of bromeliads with the presence
of\emph{P. luteolus} . To avoid re-recording the same individual in
vocalization activity, we started the search for \emph{P. luteolus}males
on different sides of the sampling points, thus ensuring that
pseudoreplication biases did not occur, since the individuals were not
marked. After viewing the male, the advertisement calls were recorded
using a Sennheiser ME66 directional microphone coupled to a Marantz
PMD660 hand recorder at a distance of 0.5 meters from the vocalizing
male. As \emph{P. luteolus} males have a long silent interval between
calls, each male was recorded for ten minutes, in order to obtain as
many calls as possible. The recordings were obtained in WAV format, 44
kHz sampling rate and 16-bit resolution. The bioacoustic analyses were
performed with the Raven Pro 1.6 software, using the following
configuration for the spectrogram: type = hann, FFT = 512, overlap =
50\%. Three to five calls were analyzed for each individual. The
acoustic parameters extracted were: call duration (s), number of notes,
number of pulses per call, pulse duration (s), rise time (s), interval
between calls (s), repetition rate (calls/min.) and dominant frequency
(Hz). The acoustic terminologies followed Köhler et al. (2017). The
sound pressure level (SPL; intensity) was obtained for three to five
advertisement calls from each individual using a Minipa MSL-1301 decibel
meter (range: 30-130dB; 125ms, fast) positioned directly in front of and
0.5 meters away from the male in vocalization activity.

Shortly after the recordings were finished, the males were captured for
measurement of the snout-vent length (SVL) with a Western caliper
(accuracy of 0.1 mm). Air temperature and humidity were recorded using a
thermo-hygrometer (TechnoLine WS 9440, 0ºC precision). The variables
were obtained for all recorded individuals. The recordings are deposited
in the Fonoteca Neotropical Jacques Vielliard (catalog number; 0058292
to 0058297).

The noise level present in the environment was measured from the calling
site of each male recorded using a Minipa MSL-1301 decibel meter (range:
30-130dB; 125ms, fast, precision 1.5 dB), positioned 1.5 m above the
ground. The decibel meter was positioned in the four cardinal directions
(north, south, east, west), and for each direction three values of sound
pressure level (SPL; intensity) were obtained, every twenty seconds we
recorded the highest value, so for each one minute we obtained records
of the three highest values, so on for each direction. Subsequently, an
average of all recorded values was extracted to represent the, represent
the environmental noise present during the vocalization activity of
males. To describe the frequency noise felt by the calling males, we
also obtained frequency values (peak and bandwidth 90\% frequency - Hz)
from the recordings of individuals near the road. In the recordings of
males from natural environments, the frequency values of the recordings
were not measured because they were close to zero due to the absence of
noise.

To verify if the environments close to the highway had higher
environmental noise than natural environments, we performed a t-test
using the average SPL obtained from the calling site of each male. To
test whether anthropic (hypothesis 1) and conspecifics (hypothesis 2)
noise alter the acoustic parameters (response variables) of the
advertisement calls of \emph{Phyllodytes luteolus} , we performed
multiple regressions using the type of environment (with and without
traffic noise) and situation (with and without chorus) as explanatory
variables. Since body size and air temperature can influence anurans
calls (Gambale \& Bastos, 2014; Morais et al., 2012; Wells, 2007), these
two factors were used as covariates in the models. We tested the
multicollinearity of the model's independent variables through a
variance inflation factor (VIF) test. We considered variables with VIF
values greater than 3 to be collinear. The VIF did not indicate
collinearity between the variables (environment=2,680, situation=1,063,
SVL=2,520, and temperature=1,088). We evaluated the assumptions of
normal distribution of residuals and homogeneity of variances by graphs
of quantiles and residuals as a function of adjusted values (Zuur et
al., 2009), respectively. Acoustic parameters that did not meet the
assumptions (call duration, number of pulses, and interval between
calls) were logarithmized.

To test significant differences in the set of acoustic parameters
between the environments (with and without traffic noise), a
multivariate analysis of permutational variance (PERMANOVA, Anderson,
2001) based on a Euclidean distance matrix was used. Before this
analysis, we performed a Pearson correlation between the acoustic
parameters and removed the highly correlated ones (number of notes,
number of pulses, rise time and interval between songs). To verify the
homogeneity of the variances we used the PERMDISP (Homogeneity of
Multivariate Dispersion) (Anderson, 2004). For all tests we assigned a
significance level lower than 5\%. The analyses were performed using the
''vegan'', ''lme4'' and ''car'' packages in the R software version 4.1.1
(R Core Team, 2021).

\section*{Results}

{\label{results}}

Recordings of 38 males of \emph{Phyllodytes luteolus} were obtained, 18
in road environments (9 individuals in situation with chorus; and 9
individuals in situation without chorus) and 20 in natural environments
(6 individuals in situation with chorus and 14 individuals in situation
without chorus). The advertisement call structure is represented in
Figure 3. The mean values and standard deviation of each acoustic
parameter of the advertisement call of \emph{P. luteolus} between each
environment and situation are represented in Table 1. The environmental
noise differed significantly between the two environments (t = -9.45, df
= 37, p \textless{} 0.05). Higher noise levels were found in sites close
to highways (average = 56.2 dB) compared to natural areas (45.6 dB)
(Figure 4). Peak and bandwidth 90\% frequencies of recordings from
highway environments averaged 129.199 Hz (86.133--947.461 Hz) and
1143.281 Hz (258.398--4048.242 Hz).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image3/image3}

\end{center}
\end{figure}

\textbf{Figure 3.} Advertisement call structure of Phyllodytes luteolus
(Wied-Neuwied, 1821), spectrogram (above) and oscillogram (below) (air
temperature = 25\selectlanguage{ngerman}°C; body size = 23.7mm), calling in a natural
environment in the municipality of Ilhéus, Bahia, Brazil.

\textbf{Table 1.} Mean and standard deviation values of each acoustic
parameter of the advertisement call of \emph{Phyllodytes
luteolus}between environments and situations in which males were
exposed.\selectlanguage{english}
\begin{longtable}[]{@{}llllll@{}}
\toprule
\begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Acoustic parameters}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Environments}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Environments}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Environments}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Environments}\strut
\end{minipage} & \begin{minipage}[b]{0.14\columnwidth}\raggedright\strut
\textbf{Environments}\strut
\end{minipage}\tabularnewline
\midrule
\endhead
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Highway (n = 19)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Highway (n = 19)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Natural (n = 20)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Natural (n = 20)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Natural (n = 20)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
With chorus (n = 9)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Without chorus (n = 10)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
With chorus (n = 6)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
With chorus (n = 6)\strut
\end{minipage} & \begin{minipage}[t]{0.16\columnwidth}\raggedright\strut
Without chorus (n = 14)\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Dominant frequency (kHz)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3473.3 -- 345.7\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3460.4 -- 389.8\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3874.5 -- 298\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3874.5 -- 298\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3889.9 -- 314.1\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Call duration (s)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
5.554 -- 0.803\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
5.713 -- 1.712\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
5.751 -- 1.736\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
5.751 -- 1.736\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
5.127 -- 0.719\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Rise Time (s)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.020 -- 0.630\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.162 -- 1.170\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
2.902 -- 0.822\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
2.902 -- 0.822\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.365 -- 0.881\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Interval between calls (s)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
75 -- 31.7\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
69.3 -- 18.4\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
79.9 - 15\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
79.9 - 15\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
78.8 -- 27.9\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Notes number\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
18.7 -- 2.948\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
17.9 -- 2.514\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
19.3 -- 2.658\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
19.3 -- 2.658\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
18.2 -- 3.445\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Pulse number\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
806.9 -- 239.3\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
1016.7 -- 483.7\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
884.6 -- 71.2\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
884.6 -- 71.2\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
771.5 -- 154.9\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Pulse duration (s)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.0028 -- 0.0005\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.0026 -- 0.0007\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.0024 -- 0.0001\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.0025 -- 0.0004\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
0.0025 -- 0.0004\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Intensity (dB)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
71.1 -- 4.22\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
69.9 -- 4.06\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
71.7 -- 7.20\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
70.8 -- 6.14\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
70.8 -- 6.14\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Advertisement call rate (calls/5min)\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.6 -- 1.414\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.9 -- 1.100\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.3 -- 1.032\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.7 -- 1.138\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
3.7 -- 1.138\strut
\end{minipage}\tabularnewline
\begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
Advertisement call total\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
7.5 -- 3.16\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
7.1 -- 1.72\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
6.6 -- 1.21\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
7.1 -- 2.21\strut
\end{minipage} & \begin{minipage}[t]{0.14\columnwidth}\raggedright\strut
7.1 -- 2.21\strut
\end{minipage}\tabularnewline
\bottomrule
\end{longtable}

The dominant frequency of the advertisement call differed significantly
between the two environments (E = -358.4, df = 33, p = 0.048), so that
males from the sites in the natural areas (without road noise) showed
higher values of the dominant frequency (Figure 5). No other acoustic
parameter showed a significant difference between the environments in
relation to the presence of traffic or situation (with or without
chorus) (Table 2). Additionally, we found a significant positive
relationship between the intensity of the calls and the snout-vent
length (E = 3.302, df = 33, p = 0.003) (Table 2), that is, the larger
the individual the higher the signal intensity (Figure 6). Considering
the set of acoustic parameters, there was no influence of the
environment or the situation on the acoustic behavior of \emph{P.
luteolus} (F = 0.005, df = 36, p = 0.942).\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image4/image4}

\end{center}
\end{figure}

\textbf{Figure 4.} Boxplot representing the differences in intensity
(dB; sound pressure level) recorded between environments (Natural and
Highway), in the municipality of Ilh\selectlanguage{ngerman}éus, Bahia, Brazil. The boxplots
show the median (middle line), first and third quartiles (upper and
lower bounds of the box), the points presented in the boxplot represent
each noise value in decibels for each individual and the extreme lines
indicate the non-outlier range. The red points represent the mean
values.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image5/image5}

\end{center}
\end{figure}

\textbf{Figure 5.} Boxplot representing the differences in the dominant
frequency of the advertisement call of Phyllodytes luteolus between
environments (Natural and Highway) in the municipality of Ilh\selectlanguage{ngerman}éus, Bahia,
Brazil. The boxplots show the median (middle line), first and third
quartiles (upper and lower bounds of the box), and the extreme lines
indicate the non-outlier range. The red points represent the mean
values.\selectlanguage{english}
\begin{figure}[H]
\begin{center}
\includegraphics[width=0.70\columnwidth]{figures/image6/image6}

\end{center}
\end{figure}

\textbf{Figure 6.} Positive relationship between advertisement call
intensity (dB; sound pressure level) and body size of males of
Phyllodytes luteolus. The line represents the estimated value from the
linear regression analysis.

\textbf{Table 2.} Results of multiple regression analysis between the
acoustic parameters of the advertisement call of Phyllodytes luteolus on
the independent variables recorded in the municipality of Ilh\selectlanguage{ngerman}éus, Bahia,
Brazil. In bold, significant p-values (\textless{}0.05).\selectlanguage{english}
\begin{longtable}[]{@{}lllll@{}}
\toprule
\textbf{Acoustic parameters} & \textbf{Independent variables} &
\textbf{Coeficients} & \textbf{Standard error} & P\tabularnewline
\midrule
\endhead
Dominant frequency (kHz) & Environments & -358.458 & 174.900 &
\textbf{0.048}\tabularnewline
& Situation & -7.457 & 119.583 & 0.950\tabularnewline
& SVL & -33.771 & 74.628 & 0.653\tabularnewline
& Temperature & -2.960 & 49.657 & 0.952\tabularnewline
Call duration (s) & Environments & 0.469 & 0.635 & 0.465\tabularnewline
& Situation & -0.170 & 0.434 & 0.698\tabularnewline
& SVL & -0.114 & 0.271 & 0.675\tabularnewline
& Temperature & -0.170 & 0.180 & 0.352\tabularnewline
Rise Time (s) & Environments & -0.207 & 0.459 & 0.655\tabularnewline
& Situation & 0.249 & 0.314 & 0.432\tabularnewline
& SVL & 0.079 & 0.196 & 0.686\tabularnewline
& Temperature & 0.169 & 0.130 & 0.203\tabularnewline
Interval between calls (s) & Environments & -16.908 & 12.708 &
0.192\tabularnewline
& Situation & -1.433 & 8.688 & 0.870\tabularnewline
& SVL & 4.442 & 5.422 & 0.419\tabularnewline
& Temperature & -0.028 & 3.608 & 0.994\tabularnewline
Notes number & Environments & -0.013 & 0.116 & 0.906\tabularnewline
& Situation & -0.056 & 0.079 & 0.477\tabularnewline
& SVL & -0.004 & 0.049 & 0.922\tabularnewline
& Temperature & 0.005 & 0.033 & 0.856\tabularnewline
Pulse number & Environments & 86.76 & 0.597 & 0.554\tabularnewline
& Situation & 45.42 & 99.43 & 0.650\tabularnewline
& SVL & 15.09 & 62.05 & 0.809\tabularnewline
& Temperature & 74.40 & 41.29 & 0.080\tabularnewline
Pulse duration (s) & Environments & 1.077 & 2.761 & 0.416\tabularnewline
& Situation & -1.824 & 1.888 & 0.924\tabularnewline
& SVL & 6.487 & 1.178 & 0.586\tabularnewline
& Temperature & -8.021 & 7.838 & 0.314\tabularnewline
Intensity (dB) & Environments & -27.156 & 34.011 & 0.430\tabularnewline
& Situation & -0.453 & 1.685 & 0.789\tabularnewline
& SVL & 3.302 & 1.051 & \textbf{0.003}\tabularnewline
& Temperature & 0.722 & 0.700 & 0.309\tabularnewline
Call rate (call/5min) & Environments & 0.299 & 0.266 &
0.261\tabularnewline
& Situation & 0.061 & 0.179 & 0.731\tabularnewline
& SVL & -0.123 & 0.114 & 0.282\tabularnewline
& Temperature & -0.062 & 0.073 & 0.393\tabularnewline
Advertisement call total & Environments & 1.194 & 1.074 &
0.274\tabularnewline
& Situation & -0.110 & 0.734 & 0.881\tabularnewline
& SVL & -0.403 & 0.458 & 0.385\tabularnewline
& Temperature & -0.443 & 0.305 & 0.155\tabularnewline
\bottomrule
\end{longtable}

\section*{Discussion}

{\label{discussion}}

Males of \emph{Phyllodytes luteolus} presented advertisement calls with
a higher dominant frequency in natural environments when compared to
highway environments, and there is no evidence that chorus noise alters
the acoustic parameters of this species. Therefore, \emph{P.
luteolus}does not seem to be affected by traffic noise, which may
explain its presence in anthropized areas. In addition, we found a
positive relationship between advertisement call intensity and male body
size.

Contrary to what was expected by our hypothesis (1), males
of\emph{Phyllodytes luteolus} from natural areas presented higher
dominant frequencies of the advertisement call compared to individuals
from environments with road noise. Although some anuran species can
increase the dominant frequency of calls to avoid overlapping with the
noise frequency (e.g., \emph{Scinax nasicus} , Leon et al., 2019;
\emph{Litoria ewingii} , Parris et al., 2009; \emph{Rana clamitans} and
\emph{Rana pipiens} , Cunnington \& Fahrig 2010, and \emph{Amolops
torrents} , Zhao et al., 2018), others can decrease the dominant
frequency (e.g.,\emph{Boana bischoffi} , Caorsi et al., 2017). In an
experimental study, Caorsi et al. (2017) observed that \emph{Boana
leptolineata} did not change the dominant frequency when exposed to
different levels of anthropic noise. Like \emph{B. leptolineata} , males
of \emph{Phyllodytes luteolu} s have calls with an average dominant
frequency (3.46 kHz) above the mean energy concentration of road noise
(below 1 kHz) (Cunnington \& Fahrig, 2010; Warren et al., 2006).
Therefore, species that have calls with higher frequencies have little
or no acoustic overlap with anthropic noise. In addition, the frequency
of ambient noise in highway areas was very low and may not represent
sufficient noise to disrupt\emph{P. luteolus} vocalizations. The
dominant frequency of the call is usually a static property (little
intra-individual variation) as it is related to morphological
characteristics of males (Gerhardt, 1991; K\selectlanguage{ngerman}öhler et al., 2017), serving
as an indicator of the quality of males (Bastos et al., 2011; Gingras et
al., 2013), being an important parameter for the sexual selection system
of a population (specific recognition and mate selection by females).
Thus, dominant frequency values may be stabilized across populations
(Friedl, 2006, Andreani et al., 2020). Future studies should investigate
possible causes for the difference in call frequency of different anuran
populations, especially those exposed to anthropic noise.

The advertisement call of \emph{P. luteolus} has a narrow bandwidth in
which the energy distributions around the dominant frequency are short.
Some studies suggest that acoustic signals with higher bandwidths may be
less detectable against noise since the signal energy is spread over a
wide range of frequencies (Parris et al., 2009; Warren et al., 2006),
therefore \emph{P. luteolus} would have an advantage in noisier
environments. Many altered habitats present a constant noise pattern in
certain frequency bands and for a long period of time, acting as a
selection pressure (Warren et al., 2006). In this sense, more studies
need to be carried out to assess how different types of noise, and/or
even anthropogenic vibrations (e.g., Caorsi et al., 2020) can influence
the reproductive behavior of anuran amphibians.

There was no influence of road noise or situation (with or without
chorus) on the temporal parameters of the advertisement call of \emph{P.
luteolus} , also not corroborating our hypothesis (II). The
advertisement call of \emph{P. luteolus} has a long duration (around 5
sec.) and a series of identical notes without frequency modulation. The
temporal structure of the call can influence the active space of the
signal when propagated in the environment, increasing the fidelity of
the transmitted information, and, therefore, decreasing the probability
of acoustic overlap (Ey \& Fischer, 2009; Warren et al., 2006), an
attribute that is advantageous in environments with high noise levels.
In general, the social context in which males are inserted in the
reproductive site influences the change in call parameters (Bosch \& De
La Riva, 2004; Toledo et al., 2015). Furthermore, males of \emph{P.
luteolus} begin to vocalize after the period of greatest road traffic
(between 5 and 7 pm), and therefore, the lower flow of vehicles at night
may not represent a source of noise that modifies the acoustic
parameters of the species. It is necessary to investigate whether other
factors are more important to determine changes in the acoustic
parameters of \emph{P. luteolus} , such as the density of conspecific
individuals in the choirs. Experimental studies on the immediate
response of males are important for a better understanding of the
behavioral strategies adopted by \emph{P. luteolus} on the balance of
costs and benefits determined by changes in acoustic parameters when
they are subjected to different noise levels.

We found a positive relationship between advertisement call intensity
and male body size. In general, morphological characteristics of males
can determine the temporal parameters of the call (Kohler et al., 2017;
Wells, 2007). For example, larger males have larger lungs and,
therefore, greater energy reserves, being able to emit calls with
greater intensity (Wells, 2007; Wells \& Schwartz, 2006). Possibly, even
in noisy scenarios, larger males are able to increase the intensities of
their calls to avoid signal masking and promote greater signal
detection, localization and discrimination (Halfwerk et al., 2016; Yi \&
Sheridan, 2019). Therefore, the intensity of the call is an important
parameter for sexual selection (Gerhardt \& Huber, 2002; Wells, 2007)
since the sounds with higher intensities can propagate over long
distances, potentially attracting a greater number of females and thus
obtaining greater reproductive success (Bastos et al., 2011; Kime, 2000;
Penna \& Solís, 1998).

The Yellow Heart-tongued Frog (\emph{Phyllodytes luteolus} ) has
advantageous acoustic characteristics in conditions where anthropogenic
noise is present. Therefore, even in noisy environments, individuals of
this species manage to recognize and discriminate in the conspecific
chorus. This fact may explain the successful occupation of areas outside
its original range, having already been reported as an invasive species
(Forti et al., 2017). Invasive species may impose limits on reproductive
success and survival of other frog species that live in the same
habitat. For example, males of \emph{P. luteolus} calling in the same
frequency range as the species \emph{Ischnocnema} sp. The acoustic noise
caused by the vocalization of an invasive species can harm the acoustic
communication system in native species as it can influence the ability
of females to locate males in the reproductive environment (Forti et
al., 2017). This is the first study evaluating the influence of sound
noise on a bromeliad anuran. Future studies may increase our knowledge
on the effects of anthropogenic sound and serve as a subsidy for
conservation actions, especially those aimed at acoustic monitoring in
noisy environments.

\section*{Conclusion}

{\label{conclusion}}

This work investigated aspects of the environmental condition, social
context and morphological factors on the acoustic parameters of the
Yellow Heart-tongued Frog (\emph{Phyllodytes luteolus} ) advertisement
call when exposed to anthropic road noise. The results show that the
dominant frequency of calls is higher in environments without traffic
noise. In addition, we also found that larger males can produce signals
with greater intensity. As the emission of calls is the main form of
communication in anurans, we concluded that \emph{P. luteolus} presented
characteristics in the structure of the advertisement call that may
represent advantages in noisy anthropic environments. Future studies are
important to investigate how different sources of noise can affect in
different ways the acoustic communication and population dynamics of
organisms that use the emission of acoustic signals to communicate.

\section*{Acknowledgments}

{\label{acknowledgments}}

We thank E. Neto and M. Vaz-Ribeiro for their assistance in the
fieldwork. The research was funded by the Fundação de Amparo à Pesquisa
do Estado da Bahia (FAPESB) through a master's scholarship to L.
Marques; by the Conselho Nacional de Desenvolvimento Científico e
Tecnológico (CNPq) with the availability of funds for the execution of
the fields and a research grant (309365/2019-8) to M. Solé; the
Coordenação de Aperfeiçoamento de Pessoal de Nível Superior (CAPES) for
providing the PROAP budget for carrying out the field work; and SISBio -
ICMBio for granting the collection license/authorization (77736) of
biological material. V. Guerra receives scholarship from CNPq (DTI-B -
process n\selectlanguage{english}* 380073/2023-4). This paper is developed in the context of
National Institutes for Science and Technology (INCT) in Ecology,
Evolution and Biodiversity Conservation, supported by MCTIC/CNpq (proc.
465610/2014-5) and FAPEG (proc. 201810267000023). We thank Fonoteca
Neotropical Jacques Vielliard (FNJV) for depositing the vocalization
recordings in its collection.

\textbf{Author contributions}

\textbf{Leonardo} : Conceptualization, Writing -- original draft,
Methodology, Data curation, Investigation,
Visualization.\textbf{Vinicius} : Methodology, Formal analysis, Writing
-- Review \& Editing. \textbf{Mirco} : Writing -- Review \& Editing,
Supervision, Resources, Project administration.

\textbf{Declaration Conflict of Interest}

The authors declare that they have no known competing financial
interests or personal relationships that could have appeared to
influence the work reported in this paper.

\textbf{Data Availability Statement}

The data source used for morphological, climatic, social values and
acoustic parameters of the advertisement call are available as
supplementary material. All files mentioned above will be also available
on Dryad Repository upon acceptance.

\section*{References}

{\label{references}}

Anderson, MJ, 2001. A new method for non\selectlanguage{english}-parametric multivariate
analysis of variance.~Austral ecology. 26, 32--46,
https://doi.org/10.1111/j.1442-9993.2001.01070.pp.x.

Anderson, MJ, 2004. PERMANOVA 2factor: a FORTRAN computer program for
permutational multivariate analysis of variance using permutation
tests.~University of Auckland, Auckland.

Andreani, TL, Bastos, RP, Dias, TM, Prado, CPA, Morais, AR, 2021.
Acoustic variability among male gladiator frogs, \emph{Boana
goiana}(Lutz, 1968) (Anura: Hylidae): an 18-year analysis across several
reproductive seasons. Amphibia-Reptilia. 42:43-57.
http://dx.doi.org/10.1163/15685381-bja10029

Barber, JR., Crooks, KR, Fristrup, KM, 2010. The costs of chronic noise
exposure for terrestrial organisms. Trends in Ecology and Evolution. 25,
180--189, https://doi.org/10.1016/j.tree.2009.08.002.

Bastos, RP., Alcantara, B, Morais, AR, Lingnau, R, Signorelli. L, 2011.
Vocal behaviour and conspecific call response in \emph{Scinax centralis}
. Herpetological Journal, 21, 43--50.

Bastos, RP, Haddad CFB, 2002. Acoustic and aggressive interactions
in\emph{Scinax rizibilis} (Anura: Hylidae) during the reproductive
activity in southeastern Brazil. Amphibia-Reptilia. 23, 97--104.

Bermudez-Cuamatzin, E, Rios-Chelen, A, Gil, D, Garcia, CM, 2009.
Strategies of song adaptation to urban noise in the house finch:
syllable pitch plasticity or differential syllable use? Behaviour. 146,
1269--1286, https://doi.org/10.1163/156853909X423104.

Bertoluci, J, Brassaloti, RA, Junior, JWR, Vilela, VMF, Sawakuchi, HO,
2007. Species composition and similarities among anuran assemblages of
forest sites in southeastern brazil. Sci. Agric. (Piracicaba). 64,
364--374, https://doi.org/10.1590/S0103-90162007000400007.

Bittencourt, L, Barbosa, M, Secchi, E, Lailson-Brito, J, Azevedo, A,
2016. Acoustic habitat of an oceanic archipelago in the Southwestern
Atlantic. Deep-Sea Research Part I: Oceanographic Research Papers. 115,
103--111, https://doi.org/10.1016/j.dsr.2016.06.001.

Bosch, J, De La Riva, I, 2004. Are frog calls modulated by the
environment? An analysis with anuran species from Bolivia. Canadian
Journal of Zoology. 82, 880--888, https://doi.org/10.1139/z04-060.

Caldart, VM, Iop, S, Lingnau, R, Cechin, SZ, 2016. Communication in a
noisy environment: short-term acoustic adjustments and the underlying
acoustic niche of a Neotropical stream-breeding frog. Acta Ethologica.
19, 151--162, https://doi.org/10.1007/s10211-016-0235-2.

Carr, L.W., Fahrig, L., 2001. Effect of road traffic on two amphibian
species of differing vagility. Conserv. Biol. 15, 1071--1078,
https://doi.org/10.1046/j.1523-1739.2001.0150041071.x.

Caorsi, VZ, Both, C, Cechin, S, Antunes, R, Borges-Martins, M, 2017.
Effects of traffic noise on the calling behavior of two Neotropical
hylid frogs. PLoS ONE. 12, 1--14,
https://doi.org/10.1371/journal.pone.0197632.

Cruz, D., Marciano-jr, E, Napoli, MF, 2014. Advertisement and courtship
calls of \emph{Phyllodytes wuchereri} (Peters, 1873) (Anura: Hylidae).
Zootaxa. 3774, 97--100, https://doi.org/10.11646/zootaxa.3774.1.8.

Cunnington, GM, Fahrig, L, 2010. Plasticity in the vocalizations of
anurans in response to traffic noise. Acta Oecologica. 36, 463--470,
https://doi.org/10.1016/j.actao.2010.06.002.

Desrochers, L, Proulx, R, 2017. Acoustic masking of soniferous species
of the St-Lawrence lowlands. Landscape and Urban Planning. 168, 31--37,
https://doi.org/10.1016/j.landurbplan.2017.09.015.

Eterovick, PC, 2008. Use and sharing of calling and retreat sites
by\emph{Phyllodytes luteolus} in a modified environment. Journal of
Herpetology. 33, 17--22, https://doi.org/10.2307/1565538.

Ey, E, Fischer, J, 2009. The ``Acoustic Adaptation Hypothesis'' - A
review of the evidence from Birds, Anurans and Mammals. Journal of
Bioacoustics. February 21--48,
https://doi.org/10.1080/09524622.2009.9753613.

Ferreira, RB, Schineider, JAP, Teixeira, RL, 2012. Diet, fecundity, and
use of Bromeliads by \emph{Phyllodytes luteolus} (Anura: Hylidae) in
Southeastern Brazil. Journal of Herpetology. 46, 19--24,
https://doi.org/10.1670/09-040.

Forti, LR, Becker, G, Tacioli, L, Pereira, VR, Santos, AC, Oliveira, I,
Haddad, CFB, Toledo, LF, 2017. Perspectives on invasive amphibians in
Brazil. PLoS ONE. 12, 1--22,
https://doi.org/10.1371/journal.pone.0184703.

Friedl, TWP, 2006. Individual male calling pattern and male mating
success in the European Treefrog (\emph{Hyla arborea} ): Is there
evidence for directional or stabilizing selection on male calling
behaviour? Ethology, 112, 116--126.
https://doi.org/10.1111/j.1439-0310.2005.01132.x

Furtado, R, Santos, S, Dias, TM, Bastos, RP, Nomura, F, 2016. Vocal
repertoire during reproductive and aggressive contexts of three
Brazilian Tree Frogs: \emph{Bokermannohyla sapiranga, Hypsiboas
albopunctatus and H. Goianus} (Anura: Hylidae). South American Journal
of Herpetology. 11, 136--147, https://doi.org/10.2994/SAJH-D-16-00002.1.

Gambale, PG, Bastos, RP, 2014. Vocal repertoire and bioacoustic analyses
in \emph{Physalaemus cuvieri} (Anura, Leptodactylidae) from southern
Brazil. Herpetological Journal, 24, 31--39.

Gerhardt, HC, Tanner, SD, Corrigan, CM, Walton, H, 2000. Female
preference functions based on call duration in the gray tree frog
(\emph{Hyla versicolor} ). Behavioral Ecology. 11, 663--669,
https://doi.org/10.1093/beheco/11.6.663.

Gerhardt, HC, Huber, F, 2002 Acoustic communication in insects and
anurans: common problems and diverse solutions. 1. ed. Chicago:
University Of Chicago Press.

Gingras, B, Boeckle, M, Herbst, CT, Fitch WT, 2013. Call acoustics
reflect body size across four clades of anurans. Journal of Zoology.
289, 143--150, https://doi.org/10.1111/j.1469-7998.2012.00973.x.

Gomes, L, Sole, M, Sousa-Lima, RS, Baumgarten, JE, 2022. Influence of
anthropogenic sounds on insect , anuran and bird acoustic signals\selectlanguage{english} : A
Meta-Analysis. Frontiers in Ecology and Evolution. 10, 1--10,
https://doi.org/10.3389/fevo.2022.827440.

Goutte, S, Dubois, A, Howard, SD, Marquez, R, Rowley, JJL, Dehling, J.
M., Grandcolas, P., Xiong, R. C., Legendre, F. 2018. How the environment
shapes animal signals\selectlanguage{english} : a test of the acoustic adaptation hypothesis in
frogs. Journal of Evolutionary Biology. 31, 148--158,
~https://doi.org/10.1111/jeb.13210.

Grenat, PR, Pollo, FE, Ferrero, MA, Martino, AL, 2019. Differential and
additive effects of natural biotic and anthropogenic noise on call
properties of \emph{Odontophrynus americanus} (Anura, Odontophryinidae):
Implications for the conservation of anurans inhabiting noisy
environments. Ecological Indicators. 99, 67--73,
https://doi.org/10.1016/j.ecolind.2018.12.014.

Guerra, V, Llusia, D, Gambale, PG, Morais, AR, Marquez, R, Bastos, RP,
2018. The advertisement calls of Brazilian anurans: Historical review,
current knowledge and future directions. PLoS ONE. 13, 1--22,
https://doi.org/10.1371/journal.pone.0191691.

Halfwerk, W, Lea, AM, Guerra, MA, Page, RA, Ryan, MJ, 2016. Vocal
responses to noise reveal the presence of the Lombard effect in a frog.
Behavioral Ecology. 27, 669--676, https://doi.org/10.1093/beheco/arv204.

Hanna, DEL, Wilson, DR, Blouin-Demers, G, Mennill, DJ, 2014. Spring
peepers \emph{Pseudacris crucifer} modify their call structure in
response to noise. Current Zoology. 60, 438--448,
https://doi.org/10.1093/czoolo/60.4.438.

Herrera-Montes, MI, Aide, TM, 2011. Impacts of traffic noise on anuran
and bird communities. Urban Ecosystems. 4, 15--427,
https://doi.org/10.1007/s11252-011-0158-7.

Hoskin, CJ, Goosem, MW, 2016. Road impacts on abundance, call traits,
and body size of Rainforest Frogs in Northeast Australia. Ecology and
Society. 15, 15, DOI:~10.5751/ES-03272-150315.

Kaiser, K, Hammers, JL, 2009. The effect of anthropogenic noise on male
advertisement call rate in the neotropical treefrog, \emph{Dendropsophus
triangulum} . Behaviour. 146, 1053--1069,
https://doi.org/10.1163/156853909X404457.

Kight, CR, Swaddle, JP, 2011. How and why environmental noise impacts
animals: An integrative, mechanistic review. Ecology Letters. 04,
1052--1061,~ https://doi.org/10.1111/j.1461-0248.2011.01664.x.

Kime, NM, Turner, WR, Ryan, MJ, 2000. The transmission of advertisement
calls in Central American frogs. Behavioral Ecology. 11, 1--83,
https://doi.org/10.1093/beheco/11.1.71.

Kohler, J, Jansen, M, Rodriguez, A, Kok, PJR, Toledo, LF, Emmerich, M,
Glaw, F, Haddad, CFB, Rodel, MO, Vences, M, 2017. The use of
bioacoustics in anuran taxonomy: Theory, terminology, methods and
recommendations for best practice. Zootaxa. 4251, 1-124,
https://doi.org/10.11646/zootaxa.4251.1.1.

Legett, HD, Madden, RP, Aihara, I, Bernal, XE, 2020. Traffic noise
differentially impacts call types in a Japanese treefrog
(\emph{Buergeria japonica} ). Ethology. 126, 76--583,
https://doi.org/10.1111/eth.13009.

Lengagne, T, 2008. Traffic noise affects communication behaviour in a
breeding anuran, \emph{Hyla arborea} . Biological Conservation. 141,
2023--2031, https://doi.org/10.1016/j.biocon.2008.05.017.

Leon, E, Peltzer, PM, Lorenzon, R, Lajmanovich, RC, Beltzer, AH, 2019.
Effect of traffic noise on \emph{Scinax nasicus} advertisement call
(Amphibia, Anura). Ilheringia. 109, 1--8,
https://doi.org/10.1590/1678-4766e2019007.

Machado, RA, Bernarde, PS, Morato, SAA, Anjos, L, 1999. Analise
comparada da riqueza de anuros entre duas areas com diferentes estados
de conservacao no municipio de Londrina, Parana, Brasil (Amphibia,
Anura). Revista Brasileira de Zoologia. 16, 997--1004,
https://doi.org/10.1590/S0101-81751999000400009..

Mageski, MM, Ferreira, RB, Beard, KH, Costa, LC, Jesus, PR, Medeiros,
CC, Ferreira, PD, 2016. Bromeliad selection by \emph{Phyllodytes
luteolus} (Anura, Hylidae): the influence of plant structure and water
quality factors. Journal of Herpetology. 50, 108--112,
https://doi.org/10.1670/14-166.

Melcon, ML, Cummins, AJ, Kerosky, SM, Roche, LK, Wiggins, SM,
Hildebrand, JA. 2012. Blue whales respond to anthropogenic noise. PLoS
ONE. 7, 1--6, https://doi.org/10.1371/journal.pone.0032681.

Moore, SE, Clarke, JT, 2002. Potential impact of offshore human
activities on gray whales. J. Cetacean Res. Manage. 4, 9--25.

Morais, AR, Batista, VG, Gambale, PG, Signorelli, L, Bastos, RP, 2012.
Acoustic communication in a Neotropical frog (\emph{Dendropsophus
minutus} ): vocal repertoire, variability and individual discrimination.
Herpetological Journal. 22, 249-257.

Morton, E. S. Ecological sources of selection on avian sounds. 1975. The
American Naturalist, 109, 17--34.

Parris, KM, Velik-Lord, M, North, JMA, 2009. Frogs call at a higher
pitch in traffic noise. Ecology and Society. 14, 15,
http://www.ecologyandsociety.org/vol14/iss1/art25/.

Penna, M, Solis, R, 1998. Frog call intensities and sound propagation
iin the South American temperate forest region. Behavioral Ecology and
Sociobiology. 42, 371--381, https://doi.org/10.1007/s002650050452.

Peixoto, O. L., 1995. Associacao de anuros a bromeliaceas na Mata
Atlantica. Revista Universidade Rural: Serie Ciencias da Vida.
Seropedica 17: 75-83

R Core Team, 2016. R: A language and environment for statistical
computing. R Foundation for Statistical Computing, Vienna, Austria.
Available from: http://www. R-project.org/.

Sabagh, LT, Ferreira, RB, Rocha, CFD, 2017. Host bromeliads and their
associated frog species: further considerations on the importance of
species interactions for conservation. Symbiosis. 73, 201--211,
https://doi.org/10.1007/s13199-017-0500-9.

Salinas, A. S., Costa, RN, Orrico, VGD, Sole, M, 2018. Tadpoles of the
bromeliad-dwelling frog \emph{Phyllodytes luteolus} are able to prey on
mosquito larvae. Ethology Ecology \& Evolution. 30, 85--496,
https://doi.org/10.1080/03949370.2018.1438518.

Salles, ROL, Silva-Soares, T, 2010. \emph{Phyllodytes luteolus} (Anura,
Hylidae) as an Alien Species in the Rio de Janeiro municipality, State
of Rio de Janeiro, Southeastern Brazil. Herpetology Notes. 3, 57--258.

Simon, JE, Gasparini, JL, 2003. Descricao da vocalizacao
de\emph{Phyllodytes kautskyi} Peixoto e Cruz, 1988
(Amphibia,Anura,Hylidae). Biologia, 1988, 7--54.

Slabbekoorn, H, Ripmeester, EAP, 2008. Birdsong and anthropogenic noise:
Implications and applications for conservation. Molecular Ecology. 17,
72--8, https://doi.org/10.1111/j.1365-294X.2007.03487.x.

Sole, M., Loebmann, D. 2017. Disgusting or delicious? Predatory behavior
of the hylid frog \emph{Phyllodytes luteolus} on sympatric ants.
Ilheringia, Serie Zoologia. 107. DOI: 10.1590/1678-4766e2017030.

Stocker, M. 2002. Fish, mollusks and other sea animals' use of sound,
and the impact of anthropogenic noise in the marine acoustic
environment. The Journal of the Acoustical Society of America. 112,
2431--2431.

Sun, JWC, Narins, PM, 2005. Anthropogenic sounds differentially affect
amphibian call rate. Biological Conservation. 121, 419--427,
https://doi.org/10.1016/j.biocon.2004.05.017.

Toledo, LF, Martins, IA, Bruschi, DP, Passos, MA, Alexandre, C, Haddad,
CFB, 2015. The anuran calling repertoire in the light of social context.
Acta Ethologica, 18, 87--99, https://doi.org/10.1007/s10211-014-0194-4.

Toledo, LF, Carvalho-e-Silva, SP, Sanchez, C, Almeida, MA, Haddad, CFB,
2010. A revisao do Codigo Florestal Brasileiro: impactos negativos para
a conservacao dos anfibios. Biota Neotrop. 10, 35--38,
https://doi.org/10.1590/S1676-06032010000400003.

Troianowski, M, Mondy, N, Dumet, A, Arcanjo, C, Lengnage, T, 2017.
Effects of traffic noise on tree frog stress levels, immunity, and color
signaling. Conservation Biology. 31, 132--1140,
https://doi.org/10.1111/cobi.12893.

Warren, P. S., Katti, M, Ermann, M, Brazel, A, 2006. Urban bioacoustics:
It's not just noise. Animal Behaviour. 71, 491--502,
https://doi.org/10.1016/j.anbehav.2005.07.014.

Wells, K. D. 2007. The Ecology and Behavior of Amphibians, The
University of Chicago Press. Chicago and London.

Wells, KD, Schwartz, JJ, 2006. The behavioral ecology of Anuran
communication. In: Narins, PM. et al. (Eds.). Hearing and sound
communication in Amphibians. New York: Springer Handbook of Auditory
Research, 44--86.

Wollerman, L, Wiley, RH, 2002. Background noise from a natural chorus
alters female discrimination of male calls in a neotropical frog. Animal
Behaviour. 63, 15--22, https://doi.org/10.1006/anbe.2001.1885.

Yi, YZ, Sheridan, JA, 2019. Effects of traffic noise on vocalisations of
the rhacophorid tree frog \emph{Kurixalus chaseni} (Anura:
Rhacophoridae) in Borneo. Raffles Bulletin of Zoology. 67, 77--8, DOI:
10.26107/RBZ-2019-0007.

Zhao, L, Sun, X, Chen, Q, Yang, Y, Wang, J, Ran, J, Brauth, SE, Tang, Y,
Cui, J, 2018 Males increase call frequency, not intensity, in response
to noise, revealing no Lombard effect in the little torrent frog.
Ecology and Evolution. 8, 11733--11741,

Zhao, L, Wang, T, Guo, R, Zhai, X, Zhou, L, Cui, J, Wang, J, 2021.
Differential effect of aircraft noise on the spectral-temporal acoustic
characteristics of frog species. Animal Behaviour. 182, 9--18, DOI:
10.1016/j.anbehav.2021.09.014.

Zuur, AF, Ieno, EN, Walker, NJ, Saveliev, AA, Smith, GM, 2009.~Mixed
effects models and extensions in ecology with R~(Vol. 574). New York:
Springer.\selectlanguage{english}
\begin{figure}[H]
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