Hans Slabbekoorn

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.

Ecology: Birds sing at a higher pitch in urban noise

Great tits hit the high notes to ensure that their mating calls are heard above the citys din.

Birdsong and anthropogenic noise: implications and applications for conservation

The dramatic increase in human activities all over the world has caused, on an evolutionary time scale, a sudden rise in especially low‐pitched noise levels. Ambient noise may be detrimental to birds through direct stress, masking of predator arrival or associated alarm calls, and by interference of acoustic signals in general. Two of the most important functions of avian acoustic signals are territory defence and mate attraction. Both of these functions are hampered when signal efficiency is reduced through rising noise levels, resulting in direct negative fitness consequences. Many bird species are less abundant near highways and studies are becoming available on reduced reproductive success in noisy territories. Urbanization typically leads to homogenization of bird communities over large geographical ranges. We review current evidence for whether and how anthropogenic noise plays a role in these patterns of decline in diversity and density. We also provide details of a case study on great tits (Parus major), a successful urban species. Great tits show features that other species may lack and make them unsuitable for city life. We hypothesize that behavioural plasticity in singing behaviour may allow species more time to adapt to human‐altered environments and we address the potential for microevolutionary changes and urban speciation in European blackbirds (Turdus merula). We conclude by providing an overview of mitigating measures available to abate noise levels that are degrading bird breeding areas. Bird conservationists probably gain most by realizing that birds and humans often benefit from the same or only slightly modified measures.

A behavioural mechanism explaining noise-dependent frequency use in urban birdsong

Acoustic signals are usually very effective in long-distance communication. However, in many habitats animals suffer more and more from signal interference caused by traffic-generated low-frequency noise. Recent observations suggest that birds are able to change the pitch of their song to reduce masking interference, but we still lack experimental evidence. Theoretically, some bird species, when confronted with increased noise levels, may be able to switch to song types in their repertoire with higher frequencies. We tested this hypothesis in the great tit, Parus major, by exposing singing males to low-frequency ‘city’ noise in their natural territories and comparing frequency characteristics of songs before and after song type switching. We also exposed birds to high-frequency, ‘inverse’ city noise, as well as to white noise as a control. Great tits adjusted temporal switching behaviour in response to noise exposure. Song types that were less masked by the noise treatment were sung for longer durations. As a result, all five birds that switched during the low-frequency noise treatment switched to song types with a higher minimum frequency. Similarly, seven of nine birds that switched while exposed to high-frequency noise switched to song types with lower maximum frequencies. These results provide experimental evidence for a short-term behavioural mechanism explaining noise-dependent frequency use in birdsong.

Low-frequency songs lose their potency in noisy urban conditions

Many animal species communicate with their mates through acoustic signals, but this communication seems to become a struggle in urbanized areas because of increasing anthropogenic noise levels. Several bird species have been reported to increase song frequency by which they reduce the masking impact of spectrally overlapping noise. However, it remains unclear whether such behavioral flexibility provides a sufficient solution to noisy urban conditions or whether there are hidden costs. Species may rely on low frequencies to attract and impress females, and the use of high frequencies may, therefore, come at the cost of reduced attractiveness. We studied the potential tradeoff between signal strength and signal detection in a successful urban bird species, the great tit (Parus major). We show that the use of low-frequency songs by males is related to female fertility as well as sexual fidelity. We experimentally show that urban noise conditions impair male–female communication and that signal efficiency depends on song frequency in the presence of noise. Our data reveal a response advantage for high-frequency songs during sexual signaling in noisy conditions, whereas low-frequency songs are likely to be preferred. These data are critical for our understanding of the impact of anthropogenic noise on wild-ranging birds, because they provide evidence for low-frequency songs being linked to reproductive success and to be affected by noise-dependent signal efficiency.

Songs of the city: noise-dependent spectral plasticity in the acoustic phenotype of urban birds

Urbanization leads to homogenization of avian communities through local extinction of rare bird species and increasing numbers of the same common urban bird species over large geographical areas. Successful city birds often persist through some sort of behavioural plasticity that helps them survive and reproduce close to humans, in built-up areas, with all the typical urban feasts and hazards. In this review, I address whether behavioural plasticity of the acoustic phenotype can be an additional factor in explaining which species end up as urban survivors. Anthropogenic noise has been shown to negatively affect avian distribution and reproduction, especially for species that rely on relatively low-frequency songs for mediating territorial conflicts and attracting partners for mating. Spectral differences between songs of city and forest populations of the same species and correlations between individual song frequency use and local noise levels suggest that many successful city species shift song frequency upward under noisy urban conditions. Experimental evidence has confirmed the ability of several species to show rapid spectral adjustments as well as perceptual benefits of singing at higher frequency in noisy habitats. However, empirical evidence of fitness benefits for birds showing the ability and tendency of noise-dependent spectral adjustment is still lacking. Furthermore, depending on the species and the underlying mechanism for spectral change, there may also be fitness costs through a compromise on signal function. These two aspects are only two of many remaining avenues for future studies. The acoustic phenotype of urban birds provides a great model system to study fundamental processes such as causes and consequences of environmentally induced signal changes, ‘cultural assimilation’, and the relationship between phenotypic and genotypic evolution. Furthermore, the current and expected rate of urbanization remains high at a global scale, which will lead to further spread in time and space of artificially elevated noise levels. This should guarantee the continued interest of scientists, politicians and conservationists for many years ahead.

SONG DIVERGENCE BY SENSORY DRIVE IN AMAZONIAN BIRDS

Visual signals are shaped by variation in the signaling environment through a process termed sensory drive, sometimes leading to speciation. However, the evidence for sensory drive in acoustic signals is restricted to comparisons between highly dissimilar habitats, or single‐species studies in which it is difficult to rule out the influence of undetected ecological variables, pleiotropic effects, or chance. Here we assess whether this form of sensory drive—often termed “acoustic adaptation”—can generate signal divergence across ecological gradients. By studying avian communities in two Amazonian forest types, we show that songs of 17 “bamboo‐specialist” bird species differ in predictable ways from their nearest relatives in adjacent terra firme forest. We also demonstrate that the direction of song divergence is correlated with the sound transmission properties of habitats, rather than with genetic divergence, ambient noise, or pleiotropic effects of mass and bill size. Our findings indicate that acoustic adaptation adds significantly to stochastic processes underlying song divergence, even when comparing between habitats with relatively similar structure. Furthermore, given that song differences potentially contribute to reproductive isolation, these findings are consistent with a wider role for sensory drive in the diversification of lineages with acoustic mating signals.

BIRDSONG AND SOUND TRANSMISSION: THE BENEFITS OF REVERBERATIONS

Abstract Animal vocalizations used for long-distance communication are shaped by acoustic properties of the environment. Studies of the relationship between signal design and sound transmission typically focus on habitat-induced limitations due to signal attenuation and degradation. However, signal design may not entirely be explained by habitat limitations, but rather by beneficial consequences of reverberations. Narrow-frequency bandwidth notes (NFB notes) are pure notes that change little in frequency, and are typical for many bird species living in dense tropical forests. In contrast to frequency-modulated notes, we show that reverberations lead to a longer and louder signal after transmission for NFB notes. Furthermore, playback experiments to territorial males of an African passerine indicated that longer notes led to a stronger behavioral response. These results suggest that reverberations may benefit signal efficiency depending on the signal design, and add new insight into the selection pressures imposed on acoustic signals by the environment. Canto de Aves y Transmisión de Sonido: Beneficios de las Reverberaciones Resumen. Las vocalizaciones utilizadas por animales para la comunicación a larga distancia están condicionadas por las propiedades acústicas del entorno. Los estudios sobre la relación entre el diseño de las señales y la transmisión del sonido suelen centrarse en los límites impuestos por el hábitat debido a la atenuación y degradación de la señal. Sin embargo, es posible que el diseño de la señal no esté regido exclusivamente por las limitaciones del habitat, sino por las consecuencias beneficiosas de las reverberaciones. Las notas de frecuencia de banda estrecha (notas NFB) son notas puras que cambian poco de frecuencia y son típicas de varias especies que habitan bosques tropicales densos. Al contrario que en las notas de frecuencia modulada, mostramos que las reverberaciones alargan y aumentan la señal de las notas NFB. Asimismo, experimentos de playback con machos territoriales de un paseriforme africano indican que las notas más largas provocan una mayor respuesta. Estos resultados sugieren que las reverberaciones pueden mejorar la eficiencia de la señal, dependiendo del diseño de la misma, y añaden un nuevo componente a nuestro conocimiento sobre las presiones selectivas impuestas por el entorno sobre las señales acústicas.

Immediate spectral flexibility in singing chiffchaffs during experimental exposure to highway noise

SUMMARY Sound plays an important role in the life of many animals, including many bird species. Typically, male birds sing to defend a territory and to attract mates. Ambient noise may negatively affect the signal efficiency of their songs, which may be critical to reproductive success. Consequently, anthropogenic noise may be detrimental to individual birds and to populations in cities and along highways. Several bird species that are still common in urban areas have been shown to sing at higher frequency at locations where there is more low-frequency traffic noise. Here we show that chiffchaffs along noisy highways also sing with a higher minimum frequency than chiffchaffs nearby at a quiet riverside. Furthermore, through experimental exposure to highway noise we show that these birds are capable of making such adjustments over a very short time scale. The first 10 songs sung during the noise exposure revealed an immediate shift to higher frequencies, with a return to pre-exposure levels in recordings without noise the following day. In a transmission re-recording experiment we tested the impact of a potential measurement artifact by recording playback of the same songs repeatedly under different controlled noise conditions. We found an upward shift in the minimum frequency measurement associated with more noisy recordings of the same song, but this artifact was not of a scale that it could explain the noise-dependent spectral shifts in chiffchaffs.

Chapter 6 – Singing in the wild: The ecology of birdsong

Publisher Summary This chapter focuses on the habitat as a source of selection pressures on sound signals. Despite its advantages for communication over long distances and in low-visibility habitats, sound transmission from sender to receiver in natural environments is by no means without problems. Sound attenuates and degrades with distance, in particular when there are obstacles in the landscape that hinder penetration. Turbulence in the air, because of wind or temperature gradients, causes irregular amplitude fluctuations that may distort a song. Many reflective surfaces, such as trunks, branches, leaves, and water and ground surfaces cause echoes that may interfere with signal perception. Furthermore, in a perfect world, there would be no other sounds to confuse the receiver, but in real life the air is full of sounds day and night, produced by neighboring birds and other animals, and by abiotic factors such as wind or rain. Ambient noise in the background interferes with signal detection and recognition. Thus, in dealing with the ecology of birdsong, one has to consider how sound changes when radiating from sender to receiver as a function of habitat characteristics, and to what extent sound perception is affected by ambient noise that varies with habitat and the animals that live there.