Research PaperAcoustic masking of soniferous species of the St-Lawrence lowlands
Introduction
Communication in the animal kingdom is used by individuals to attract females during the reproductive stage, defend territories and signal food supply to conspecifics (Bradbury & Verenchamps, 2011; Gerhardt & Huber, 2002). Communication can be achieved with visual, olfactory or acoustic signals (Endler, 1993; Rekwot, Ogwu, Oyedipe, & Sekoni, 2001). In noisy environments, acoustic communication can be masked, thus forcing individuals to invest energy into compensatory mechanisms, such as shifting call frequencies (Lengagne, 2008, Slabbekoorn and Peet, 2003), vocalizing louder (Brumm, 2004; Cynx, Lewis, Travel, & Tse, 1998; Penna & Hamilton-West, 2006), or moving away from the noise source (Francis, Ortega, & Cruz, 2011; Goutte, Dubois, & Legendre, 2013). Previous studies have reported examples of adaptations to acoustic masking in anurans (Cunnington & Fahrig, 2010), birds (Hu & Cardoso, 2010; Wood and Yezerinac, 2006) and insects (Morley, Jones, & Radford, 2014; Shieh, Liang, & Chiu, 2012). However, it is not clear if a majority of species in these three taxonomic groups are affected by acoustic masking. For example, a recent meta-analysis by Roca et al. (2016) showed that only small-bodied bird species singing at low frequencies (<3 kHz) increased their dominant frequencies when exposed to anthropogenic noise, whereas anurans were less prone to such shifts.
To estimate the global importance of acoustic masking events, it is important to know the probability of spectral, spatial, and temporal overlap between human-induced and animal-induced sounds. Acoustic masking events occur if the communication signals are emitted i) at the same frequency (spectral overlap), ii) close enough to the noise source (spatial overlap), and iii) concurrently to human-induced sounds (temporal overlap) (Brumm and Slabbekoorn, 2005, Patterson and Green, 1978, Slabbekoorn et al., 2010). Human-induced sounds generally have low dominant frequencies and, as such, should not impact every vocalizing organisms equally (Hu & Cardoso, 2010; Slabbekoorn & Peet,2003). Anurans calls could be more susceptible to acoustic masking due to their spectral overlap with human-induced sounds (e.g., Cunnington & Fahrig, 2010). Hu and Cardoso (2009) similarly argued that birds singing at high frequency should predominate in urban ecosystems because their signaling frequency is out of the range of human-induced sounds.
Acoustic masking can be particularly important in urban settings because species did not coevolve with human-induced sounds (Feng & Schul, 2007). Although, data on the intensity level and the frequency spectrum of urban sounds are relatively easy to obtain (see Wood & Yezerinac, 2006; Wei, van Renterghem, De Coensel, & Botteldooren, 2016), little is known of the amount of temporal overlap between human-induced sounds and animal-induced sounds. Studies that explored the temporal patterns of vocal organisms were mostly conducted in natural ecosystems (e.g., Krause, Gage, & Woo, 2011; Rodriguez et al., 2014) or did not explicitly consider the anthrophony component of the soundscape (e.g., Gage & Axel 2014; Gage, Joo, Kasten, Fox, & Biswas, 2015). Hence, bird and anuran individuals exchanging acoustic signals close to a noise source, and at low frequencies, may avoid acoustic masking by vocalizing when temporal overlap is minimized during the day (Cartwright, Taylor, Wilson, & Chow-Fraser, 2014; Warren, Katti, Ermann, & Brazel, 2006).
The aim of the study was to evaluate the joint probability that acoustic masking could occur for anuran, bird, or stridulating orthopteran species along the spectral, spatial, and temporal dimensions of the acoustic space. We evaluated acoustic overlap in the dominant frequencies, absolute amplitudes (i.e., emitter-receiver distances), and acoustic patterns (i.e., temporal match-mismatch) of human- and animal-induced sounds in three soundscape contexts: urban, peri-urban, and agricultural. We paid particular attention to the daily temporal overlap between human-induced sounds (anthrophony) and animal-induced sounds (biophony), for which no estimates currently exist.
Section snippets
Material and methods
To estimate the likelihood of an acoustic masking event, we independently evaluated the probability of spectral, spatial, and temporal overlap between human-induced sounds (anthrophony) and animal-induced sounds (biophony). We estimated the probability that species of a given taxonomic group t could experience an acoustic masking event when exposed to anthrophony with the following equation:where AM is acoustic masking, SpeO is spectral overlap probability, SpaO is spatial
Spectral overlap
The frequency spectrum of anuran calls showed a large spectral overlap with human-induced sounds, while the spectrum of orthopteran stridulations barely overlapped (Fig. 1). The probability of a spectral overlap between human- and animal-induced sounds was 32%, 26% and 7%, for anuran, bird and orthopteran species, respectively (Table 1). Using the weighted mean frequency did not change the spectral overlap ranking of each taxonomic group, with overlap probabilities ranging from 49% for anurans,
Discussion
An increasing number of studies have raised concerns about the impact of human-induced sounds and acoustic masking events on animal communication and, more generally, on animal behavior (Brumm and Slabbekoorn, 2005, Morley et al., 2014, Roca et al., 2016, Warren et al., 2006). The present study takes a broader perspective and asks whether these examples should be considered the rule or the exception. Our results suggest that acoustic masking per se is unlikely for a majority of species in the
Conclusion
What is the likelihood that a given individual from a regional species pool will experience an event of acoustic masking? Answering this fundamental question is needed to comprehend the impact of human-induced sounds on vocalizing species and to prioritize conservation efforts. We herein provided conservative, upper-bound, estimates of acoustic masking for three taxonomic groups, which suggest that human-induced sounds may not be of major concern in a majority of cases of impaired acoustic
Acknowledgements
The authors acknowledge the financial support provided by the Fonds de recherche du Québec – Nature et technologies (FRQNT) to L.D., the financial support provided by the Natural Sciences and Engineering Research Council of Canada (NSERC) to R.P. and the grants from the Groupe de recherche interuniversitaire en limnologie et en environnement aquatique (GRIL).
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