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Territoriality is a common pattern of space use in animals that has fundamental consequences for ecological processes. In the tropics, all-year resident songbirds usually hold territories throughout the year, whereas most all-year resident temperate species are territorial only during the breeding season. In long-distance migrants, however, the situation is mostly unexplored. Here, we report findings from a Palaearctic-African migrant, the thrush nightingale Luscinia luscina . We found that only a fraction of the males was territorial in their East African winter quarters and that this was related to the stage of their song development. Individuals with full song were territorial towards other full songsters, but not towards birds that sang plastic song (i.e. an earlier stage of song development). Plastic singers were not territorial towards full songsters and often settled closely to territorial males. We suggest that territoriality of thrush nightingales in the winter quarters may be a by-product of rising testosterone levels that trigger song crystallization. Collectively, our study indicates that changes in territoriality can occur rapidly, giving rise to shifting proportions of territorial and non-territorial individuals in a population, which may lead to complex dynamics in settlement patterns and resulting ecological interactions.

The duets of birds have intrigued biologists for a long time, yet much remains unknown about the evolution of these striking collective displays. This is partly because previous studies on duet evolution have been biased to songbirds and neglected other bird groups. In songbirds, the absence of migration has been found to predict the occurrence of duetting, indirectlysupporting the idea that duet communication is linked with pair bonding. Here, we used phylogenetic comparative analyses in a sedentary clade of non-songbirds, the barbets (Capitonidae), to reveal new correlates of duet evolution. We found (i) that duets evolved several times independently in different barbet lineages and (ii) that duetting evolved in association with group living (i.e. the presence of helpers or non-breeding adults during the breeding period), but not with sexual monochromatism or habitat type. Our findings are consistent with a duet function in mate guarding and dominance against subordinate group members as well as joint territory defence. Altogether, the results highlight the importance of the social environment for the evolution of collective signalling.

It has been observed in many songbird species that populations in noisy urban areas sing with a higher minimum frequency than do matched populations in quieter, less developed areas. However, why and how this divergence occurs is not yet understood. We experimentally tested whether chronic noise exposure during vocal learning results in songs with higher minimum frequencies in great tits ( Parus major ), the first species for which a correlation between anthropogenic noise and song frequency was observed. We also tested vocal plasticity of adult great tits in response to changing background noise levels by measuring song frequency and amplitude as we changed noise conditions. We show that noise exposure during ontogeny did not result in songs with higher minimum frequencies. In addition, we found that adult birds did not make any frequency or song usage adjustments when their background noise conditions were changed after song crystallization. These results challenge the common view of vocal adjustments by city birds, as they suggest that either noise itself is not the causal force driving the divergence of song frequency between urban and forest populations, or that noise induces population-wide changes over a time scale of several generations rather than causing changes in individual behaviour., (© 2017 The Author(s).)

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.

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.

Understanding the mechanisms underlying speciation remains a challenge in evolutionary biology. The adaptive radiation of Darwin's finches is a prime example of species formation, and their study has revealed many important insights into evolutionary processes. Here, we report striking differences in mating signals (songs), morphology and genetics between the two remnant populations of Darwin's mangrove finch Camarhynchus heliobates, one of the rarest species in the world. We also show that territorial males exhibited strong discrimination of sexual signals by locality: in response to foreign songs, males responded weaker than to songs from their own population. Female responses were infrequent and weak but gave approximately similar results. Our findings not only suggest speciation in the mangrove finch, thereby providing strong support for the central role of sexual signals during speciation, but they have also implications for the conservation of this iconic bird. If speciation is complete, the eastern species will face imminent extinction, because it has a population size of only 5-10 individuals.

Collective signalling in animals has fascinated biologists for a long time. A recent study on Australian songbirds sheds new light on the function of highly coordinated avian duets.

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.

Vocal amplitude, one of the crucial factors for the exchange of acoustic signals, has been neglected in studies of animal communication, but recent studies on song variation in Common Nightingales Luscinia megarhynchos have revealed new insights into its importance in the singing behavior of territorial birds. In nightingales song amplitude is not maximized per se, but is individually regulated according to the level of masking background noise. Also, birds adjust their vocal intensity according to social variables, as in male-male interactions. Moreover, during such interactions, males exploited the directionality of their songs to broadcast them in the direction of the intended receivers ensuring the most effective signal transmission. Studies of the development of this typical long-range signaling suggest that sound level is highly interrelated with overall developmental progression and learning, and thus should be viewed as an integral part of song ontogeny. I conclude that song amplitude is a dynamic feature of the avian signal system, which is individually regulated according to the ecological demands of signal transmission and the social context of communication.

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Territoriality is a common pattern of space use in animals that has fundamental consequences for ecological processes. In the tropics, all-year resident songbirds usually hold territories throughout the year, whereas most all-year resident temperate species are territorial only during the breeding season. In long-distance migrants, however, the situation is mostly unexplored. Here, we report findings from a Palaearctic–African migrant, the thrush nightingale Luscinia luscina. We found that only a fraction of the males was territorial in their East African winter quarters and that this was related to the stage of their song development. Individuals with full song were territorial towards other full songsters, but not towards birds that sang plastic song (i.e. an earlier stage of song development). Plastic singers were not territorial towards full songsters and often settled closely to territorial males. We suggest that territoriality of thrush nightingales in the winter quarters may be a by-product of rising testosterone levels that trigger song crystallization. Collectively, our study indicates that changes in territoriality can occur rapidly, giving rise to shifting proportions of territorial and non-territorial individuals in a population, which may lead to complex dynamics in settlement patterns and resulting ecological interactions. [ABSTRACT FROM AUTHOR]

It has been observed in many songbird species that populations in noisy urban areas sing with a higher minimum frequency than do matched populations in quieter, less developed areas. However, why and how this divergence occurs is not yet understood. We experimentally tested whether chronic noise exposure during vocal learning results in songs with higher minimum frequencies in great tits (Parus major), the first species for which a correlation between anthropogenic noise and song frequency was observed. We also tested vocal plasticity of adult great tits in response to changing background noise levels by measuring song frequency and amplitude as we changed noise conditions. We show that noise exposure during ontogeny did not result in songs with higher minimum frequencies. In addition, we found that adult birds did not make any frequency or song usage adjustments when their background noise conditions were changed after song crystallization. These results challenge the common view of vocal adjustments by city birds, as they suggest that either noise itself is not the causal force driving the divergence of song frequency between urban and forest populations, or that noise induces population-wide changes over a time scale of several generations rather than causing changes in individual behaviour. [ABSTRACT FROM AUTHOR]

Animals that use vocal signals to communicate often compensate for interference and masking from background noise by raising the amplitude of their vocalisations. This response has been termed the Lombard effect. However, despite more than a century of research, little is known how quickly animals can adjust the amplitude of their vocalisations after the onset of noise. The ability to respond quickly to increases in noise levels would allow animals to avoid signal masking and ensure their calls continue to be heard, even if they are interrupted by sudden bursts of high-amplitude noise. We tested how quickly singing male canaries (Serinus canaria) exhibit the Lombard effect by exposing them to short playbacks of white noise and measuring the speed of their responses. We show that canaries exhibit the Lombard effect in as little as 300 ms after the onset of noise and are also able to increase the amplitude of their songs mid-song and mid-phrase without pausing. Our results demonstrate high vocal plasticity in this species and suggest that birds are able to adjust the amplitude of their vocalisations very rapidly to ensure they can still be heard even during sudden changes in background noise levels. [ABSTRACT FROM AUTHOR]

In birdsong, and in most commonly studied acoustic communication systems, research has often focused on temporal and frequency-related signal parameters. However, although variations in amplitude are often overlooked and seldom measured, they are just as critical in communication. Recent studies have demonstrated that vocal amplitude plays an important role in both territorial behaviours and mate choice in birds. Several songbird species have been shown to produce low-amplitude songs, used primarily in aggressive encounters between males. On the other hand, loud songs may be an honest signal of current condition in males and recent studies have shown that females may prefer high-amplitude songs. Although it is generally assumed that louder song is more costly to produce, there is little empirical evidence to support this assumption. Here we review data on the metabolic costs of singing at different vocal amplitudes, and discuss recent studies from our laboratory showing that louder songs elicit stronger aggressive responses from territorial males. Together, these findings suggest that while the energetic costs of singing loudly are negligibly small, social aggression may be a key constraint that limits the upper amplitude of vocal signals. Finally we discuss future directions that can increase our understanding of the role that amplitude plays in acoustic communication in animals. [ABSTRACT FROM AUTHOR]

Several recent studies have tested the hypothesis that song quality in adult birds may reflect early developmental conditions, specifically nutritional stress during the nestling period. Whilst all of these earlier studies found apparent links between early nutritional stress and song quality, their results disagree as to which aspects of song learning or production were affected. In this study, we attempted to reconcile these apparently inconsistent results. Our study also provides the first assessment of song amplitude in relation to early developmental stress and as a potential cue to male quality. We used an experimental manipulation in which the seeds on which the birds were reared were mixed with husks, making them more difficult for the parents to obtain. Compared with controls, such chicks were lighter at fledging; they were thereafter placed on a normal diet and had caught up by 100 days. We show that nutritional stress during the first 30 days of life reduced the birds’ accuracy of song syntax learning, resulting in poorer copies of tutor songs. Our experimental manipulations did not lead to significant changes in song amplitude, song duration or repertoire size. Thus, individual differences observed in song performance features probably reflect differences in current condition or motivation rather than past condition. [ABSTRACT FROM AUTHOR]

Recent studies have shown that territorial songbirds do not maximize vocal amplitude. Instead, song intensity appears to be a flexible trait that is individually regulated. Given the benefits of singing loudly with regard to signal transmission in the context of territory defence and mate attraction, we investigated whether songbirds adjust the sound level of their territorial songs depending on social influences. While interacting with the playback of a simulated conspecific rival, seven male nightingales, Luscinia megarhynchos, increased the sound level of their songs, on average, by more than 5 dB. The nightingales increased their song level to significantly lower values (about 1 dB) in response to control songs of other species, so the revealed increase in vocal intensity during interactions cannot be fully explained by the acoustical masking of the interfering conspecific songs. Thus, the birds improved signal transmission to a higher degree during conspecific male–male interactions. [Copyright &y& Elsevier]

Investigates noise-dependent song amplitude in a territorial songbird. Ability of songbirds to maximize the amplitude of songs to defend territories and attract females; Favorability of the adjustment of vocal amplitude for signal production; Interpretation of the vocal amplitude in a territorial songbird.