Henrik Brumm

Acoustic Communication in Noise

Publisher Summary Environmental noise can affect acoustic communication through limiting the broadcast area, or active space, of a signal by decreasing signal-to-noise ratios at the position of the receiver. At the same time, noise is ubiquitous in all habitats and is, therefore, likely to disturb animals, as well as humans, under many circumstances. However, both animals and humans have evolved diverse solutions to the background noise problem, and this chapter reviews recent advancements in studies of vocal adaptations to interference by background noise and relate these to fundamental issues in sound perception. The chapter starts with the discussion of senders side by considering potential evolutionary shaping of species-specific signal characteristics and individual short‐term adjustments of signal features. Subsequently, it focuses on the receivers of signals and reviews their sensory capacities for signal detection, recognition, and discrimination and relates these issues to auditory scene analysis and the ecological concept of signal space. The data from studies on insects, anurans, birds, and mammals, including humans, and to a lesser extent available work on fish and reptiles is also discussed in the chapter.

Acoustic communication in noise: regulation of call characteristics in a New World monkey

SUMMARY This study on common marmosets Callithrix jacchus is the first to examine noise-dependent mechanisms of vocal plasticity in a New World monkey. Since acoustic communication can be considerably impaired by environmental noise, some animals have evolved adaptations to counteract its masking effects. The studied marmosets increased the sound level of their spontaneous calls in response to increased levels of white noise broadcast to them. Possibly, such noise-dependent adjustment of vocal amplitude serves to maintain a specific signal-to-noise ratio that is favourable for signal production. Concurrently, the adjustment of vocal amplitude can maintain a given active space for communication. In contrast to some bird species, no noise-induced increase in the number of syllables per call series could be found, showing that an increased serial redundancy of vocal signals was not used to communicate under noisy conditions. Finally, we examined a possible noise-dependent prolongation of vocal signals. This approach was guided by the findings of perceptional studies, which suggest an increased detection probability of prolonged signals in noise by temporal summation. Marmosets indeed increased the duration of their call syllables along with increasing background noise levels. This is the first evidence of such mechanism of vocal plasticity in an animal communication system.

Noise-dependent song amplitude regulation in a territorial songbird

Abstract Some animals that use sound to communicate compensate for interference from background noise by adjusting the amplitude of their vocalizations as environmental noise levels vary. Territorial songbirds may have evolved a different strategy, since they can be expected to benefit from maximizing the amplitude of their songs to defend territories and attract females. We tested this hypothesis with calibrated measurements of the song level of male nightingales, Luscinia megarhynchos . All birds increased the sound level of their songs in response to an increase in white noise broadcast to them. A second experiment revealed that noise in the spectral region of their own songs was most effective in inducing the birds to increase vocal intensity. These findings show that nightingales do not maximize song amplitude but regulate vocal intensity dependent on the level of masking noise. The adjustment of vocal amplitude may serve to maintain a specific signal-to-noise ratio that is favourable for signal production. Concurrently, increasing the intensity of songs can maintain a given active space for communication. Thus, vocal amplitude in a territorial songbird can be interpreted as a flexible trait, which is individually regulated according to ecological demands from signal transmission. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved .

Blackbirds sing higher-pitched songs in cities: adaptation to habitat acoustics or side-effect of urbanization?

When animals colonize cities they often have to adapt their physiology, life history and behaviour to the novel environment. Songbirds rely on acoustic communication for reproduction, and recent studies indicate that songs vary between urban and nonurban habitats. In cities, birds sing louder or use higher frequencies compared to their conspecifics in forests. These habitat-specific differences in song have been interpreted as an adaptation of the city birds to mitigate acoustic masking by low-frequency traffic noise. We compared the songs of blackbirds, Turdus merula, from the city centre of Vienna and the Vienna Woods and found that forest birds sang at lower frequencies and with longer intervals between songs. This difference in song pitch might reflect an adaptation to urban ambient noise. However, the song divergence could also be the result of more intense vocal interaction in the more densely populated city areas or a side-effect of physiological adaptation to urban habitats. We emphasize the need for experimental studies in blackbirds, but also in other species, to clarify a possible causal link between urban acoustics and song characteristics of city birds.

Birds and Anthropogenic Noise: Are Urban Songs Adaptive?

In cities with intense low‐frequency traffic noise, birds have been observed to sing louder and at a higher pitch. Several studies argue that higher song pitch is an adaptation to reduce masking from noise, and it has even been suggested that the song divergence between urban and nonurban songs might lead to reproductive isolation. Here we present models of signal transmission to compare the benefits of raised song amplitude and song pitch in terms of sound transmission. We chose two bird species that sing with higher pitch in urban areas, the great tit (Parus major) and the blackbird (Turdus merula). For both species, we calculated communication distances in response to different levels of urban noise and in their natural forest habitats. We found that an increase in vocal pitch increased communication distance only marginally. In contrast, vocal amplitude adjustments had a strong and significantly larger effect. Our results indicate that frequency changes of urban songs are not very effective in mitigating masking from traffic noise. Increased song pitch might not be an adaptation to reduce signal masking but a physiological side effect of singing at high amplitudes or an epiphenomenon of urbanization that is not related to signal transmission.

Ambient noise, motor fatigue, and serial redundancy in chaffinch song

Many animal signals are performed in a highly redundant manner as in some bird species where males sing several renditions of one song type before switching to another. However, differences in signal redundancy between contexts and between individuals are only poorly understood. We found that chaffinches (Fringilla coelebs) in noisier areas (i.e., close to waterfalls and torrents) sang longer bouts of the same song type before switching to a new type, suggesting that they use increased serial redundancy to get the message across in noisy conditions. This is the first evidence of a noise-dependent adjustment of signal redundancy in a songbird. In addition, we found that song types with faster trills were sung in shorter bouts suggesting that the performance of highly redundant song series is probably limited by motor constraints. Thus, in noisy environments, serial redundancy in bird song may reflect a trade-off between successful signal transmission and preventing motor fatigue.

On the relationship between, and measurement of, amplitude and frequency in birdsong

A growing number of studies ask whether and how bird songs vary between areas with low versus high levels of anthropogenic noise. Across numerous species, birds are seen to sing at higher frequencies in urban versus rural populations, presumably because of selection for higher-pitched songs in the face of low-frequency urban noise, or because birds can avoid masking directly by shifting to higher-frequency sounds (Fernandez-Juricic et al. 2005; Slabbekoorn & den Boer-Visser 2006; Nemeth & Brumm 2009; Gross et al. 2010; Potvin et al. 2010). In addition to changing song frequency, birds are also reported to respond to increased background noise by singing at higher amplitudes (Brumm & Zollinger 2011). Nightingales, Luscinia megarhynchos, for example, sing with a higher sound pressure level in areas with intense traffic noise as compared to quieter locations (Brumm 2004). While frequencyand amplitude-based responses to ambient noise are often considered independently, the twomight also vary in tandem

Signalling through acoustic windows: Nightingales avoid interspecific competition by short-term adjustment of song timing.

The function of bird song is closely linked to sexual selection through female choice and male–male competition, and thus variation in communication success is likely to have major fitness consequences for a singing male. A crucial constraint on signal transmission is imposed by background noise, which may include songs from other species. I investigated whether singing nightingales (Luscinia megarhynchos) avoid temporal overlap with the songs of other bird species in a playback experiment. I analysed the temporal song patterns of six males, each of which were exposed to songs of other species. The nightingales significantly avoided overlapping their songs with the playback songs, and started singing preferentially during the silent intervals between the heterospecific songs. This timing of song onset produced a greater variability in pause duration compared to the nightingales’ undisturbed solo singing. These findings suggest that birds adjust their song timing to avoid acoustic interference on short temporal scales, and thus are able to improve the efficiency of acoustic communication in complex sonic environments. Moreover, the results indicate that temporal song patterns can be affected by the songs of other species, and thus such influences should be taken into account when studying bird song in the field.

Bird song and anthropogenic noise: vocal constraints may explain why birds sing higher-frequency songs in cities

When animals live in cities, they have to adjust their behaviour and life histories to novel environments. Noise pollution puts a severe constraint on vocal communication by interfering with the detection of acoustic signals. Recent studies show that city birds sing higher-frequency songs than their conspecifics in non-urban habitats. This has been interpreted as an adaptation to counteract masking by traffic noise. However, this notion is debated, for the observed frequency shifts seem to be less efficient at mitigating noise than singing louder, and it has been suggested that city birds might use particularly high-frequency song elements because they can be produced at higher amplitudes. Here, we present the first phonetogram for a songbird, which shows that frequency and amplitude are strongly positively correlated in the common blackbird (Turdus merula), a successful urban colonizer. Moreover, city blackbirds preferentially sang higher-frequency elements that can be produced at higher intensities and, at the same time, happen to be less masked in low-frequency traffic noise.

Chapter 1 Environmental Acoustics and the Evolution of Bird Song

Abstract Any signal must get from a sender to a receiver if information is to be transmitted. In the case of bird song, the acoustic properties of the habitat may hinder this being achieved. However, birds as senders and receivers have evolved numerous adaptations to overcome the problem of getting the message across. In this chapter, we explore habitat-dependent patterns of sound transmission, the effects of noise, signal perception, and signal interpretation such as auditory distance assessment with a specific focus on the solutions that selection has generated. We argue that along with other possible selective forces, such as sexual selection, the combination of environmental constraints on signal transmission, noise levels, and the use of signal degradation as a distance cue need mutual consideration to gain a more thorough understanding of the astounding variety of avian song and the many different ways in which birds use it.