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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.

Biomedical and social scientists are increasingly calling the biological sex into question, arguing that sex is a graded spectrum rather than a binary trait. Leading science journals have been adopting this relativist view, thereby opposing fundamental biological facts. While we fully endorse efforts to create a more inclusive environment for gender-diverse people, this does not require denying biological sex. On the contrary, the rejection of biological sex seems to be based on a lack of knowledge about evolution and it champions species chauvinism, inasmuch as it imposes human identity notions on millions of other species. We argue that the biological definition of the sexes remains central to recognising the diversity of life. Humans with their unique combination of biological sex and gender are different from non-human animals and plants in this respect. Denying the concept of biological sex, for whatever cause, ultimately erodes scientific progress and may open the flood gates to "alternative truths.", (© 2022 The Authors. BioEssays published by Wiley Periodicals LLC.)

The impacts of noise pollution on birdsong have been extensively investigated but potential long-term effects are neglected. Near airports, where noise levels are particularly high, birds start singing earlier in the morning, probably to gain more time of uninterrupted singing before air traffic sets in. In a previous study, we documented this phenomenon in the vicinity of Berlin Tegel airport. In 2020, Tegel airport closed down, giving us the opportunity to investigate potential long-term effects after noise removal and to gain insight into the mechanisms underlying the advancement of dawn singing. We found that several species at the airport shifted their song onset back after the closure and now had similar schedules to their conspecifics at a control site. Some species, however, still sang earlier near the closed airport. While the first suggests plastic adaptation, the latter suggests selection for early singing males in areas with long-lasting noise pollution. Our findings indicate that a uniform behavioural response to anthropogenic change in a community can be based on diverging evolutionary mechanisms. Overall, we show that noise pollution can have long-lasting effects on animal behaviour and noise removal may not lead to immediate recovery in some species.

The broken-wing display is a well-known and conspicuous deceptive signal used to protect birds' broods against diurnal terrestrial predators. Although commonly associated with shorebirds, it remains unknown how common the behaviour is across birds and what forces are associated with the evolution of the display. Here, we use the broken-wing display as a paradigmatic example to study the evolution of a behaviour across Aves. We show that the display is widespread: it has been described in 52 families spread throughout the phylogeny, suggesting that it independently evolved multiple times. Further, we evaluated the association with 16 ecological and life-history variables hypothesized to be related to the evolution of the broken-wing display. Eight variables were associated with the display. We found that species breeding farther from the equator, in more dense environments, with shorter incubation periods, and relatively little nest cover were more likely to perform the display, as were those in which only one parent incubates eggs, species that mob nest predators and species that are altricial or multi-brooded. Collectively, our comprehensive approach identified forces associated with the repeated evolution of this conspicuous display, thereby providing new insights into how deceptive behaviours evolve in the context of predator-prey interactions.

Noise pollution has been linked to learning and language deficits in children, but the causal mechanisms connecting noise to cognitive deficiencies remain unclear because experimental models are lacking. Here, we investigated the effects of noise on birdsong learning, the primary animal model for vocal learning and speech development in humans. We found that traffic noise exposure retarded vocal development and led to learning inaccuracies. In addition, noise suppressed immune function during the sensitive learning period, indicating that it is a potent stressor for birds, which is likely to compromise their cognitive functions. Our results provide important insights into the consequences of noise pollution and pave the way for future studies using birdsong as an experimental model for the investigation of noise-induced learning impairments., (Copyright © 2021 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

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.

The impact of human activity on the acoustic environment is overwhelming, with anthropogenic noise reaching even remote areas of the planet. The World Health Organization has identified noise pollution as one of the leading environmental health risks in humans, and it has been linked to a myriad of short- and long-term health effects in exposed individuals. However, less is known about the health effects of anthropogenic noise exposure on animals. We investigated long- and short-term effects of traffic noise on zebra finches breeding in small communal aviaries, using a repeated measures design. Birds bred in both noise and no-noise conditions, and we measured baseline plasma glucocorticoid levels before, during and after breeding. In addition, we assayed immune function, measured reproductive success and offspring growth and compared rates of extra-pair paternity of breeding adults. Breeding birds had significantly lower baseline plasma corticosterone levels when exposed to traffic noise than when they were not exposed to noise playback. In addition, the nestlings reared during noise exposure were lighter than nestlings of the same parents when breeding in control conditions. Our results suggest that traffic noise poses a more severe hurdle to birds at more vulnerable stages of their life history, such as during reproductive events and ontogeny. While chronic exposure to traffic noise in our birds did not, by itself, prove to be a sufficient stressor to cause acute effects on health or reproductive success in exposed individuals, it did result in disruptions to normal glucocorticoid profiles and delayed offspring growth. However, animals living in urban habitats are exposed to a multitude of anthropogenic disturbances, and it is likely that even species that appear to be thriving in noisy environments may suffer cumulative effects of these multiple disturbances that may together impact their fitness in urban environments., (© The Author(s) 2019. Published by Oxford University Press and the Society for Experimental Biology.)

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).)

Sophisticated vocal communication systems of birds and mammals, including human speech, are characterized by a high degree of plasticity in which signals are individually adjusted in response to changes in the environment. Here, we present, to our knowledge, the first evidence for vocal plasticity in a reptile. Like birds and mammals, tokay geckos ( Gekko gecko ) increased the duration of brief call notes in the presence of broadcast noise compared to quiet conditions, a behaviour that facilitates signal detection by receivers. By contrast, they did not adjust the amplitudes of their call syllables in noise (the Lombard effect), which is in line with the hypothesis that the Lombard effect has evolved independently in birds and mammals. However, the geckos used a different strategy to increase signal-to-noise ratios: instead of increasing the amplitude of a given call type when exposed to noise, the subjects produced more high-amplitude syllable types from their repertoire. Our findings demonstrate that reptile vocalizations are much more flexible than previously thought, including elaborate vocal plasticity that is also important for the complex signalling systems of birds and mammals. We suggest that signal detection constraints are one of the major forces driving the evolution of animal communication systems across different taxa., (© 2017 The Author(s).)

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., (© 2017. Published by The Company of Biologists Ltd.)

Anthropogenic noise is of increasing concern to biologists and medical scientists. Its detrimental effects on human health have been well studied, with the high noise levels from air traffic being of particular concern. However, less is known about the effects of airport noise pollution on signal masking in wild animals. Here, we report a relationship between aircraft noise and two major features of the singing behavior of birds. We found that five of ten songbird species began singing significantly earlier in the morning in the vicinity of a major European airport than their conspecifics at a quieter control site. As birds at both sites started singing before the onset of air traffic in the morning, this suggests that the birds in the vicinity of the airport advanced their activity to gain more time for unimpaired singing before the massive plane noise set in. In addition, we found that during the day, chaffinches avoided singing during airplane takeoffs, but only when the noise exceeded a certain threshold, further suggesting that the massive noise caused by the airport can impair acoustic communication in birds. Overall, our study indicates that birds may be adjusting their mating signals and time budgets in response to aircraft noise.

Birds in cities start singing earlier in the morning than in rural areas; commonly this shift is attributed to light pollution. Some studies have suggested that traffic noise has a stronger influence on singing activity than artificial light does. Changes in the timing of singing behaviour in relation to noise and light pollution have only been investigated in the temperate zones. Tropical birds, however, experience little seasonal variation in day length and may be less dependent on light intensity as a modifier for reproductive behaviours such as song. To test whether noise or light pollution has a stronger impact on the dawn chorus of a tropical bird, we investigated the singing behaviour of rufous-collared sparrows (Zonotrichia capensis) in Bogota, Colombia at two times during the year. We found that birds in places with high noise levels started to sing earlier. Light pollution did not have a significant effect. Birds may begin to sing earlier in noisy areas to avoid acoustic masking by traffic later in the morning. Our results also suggest that some tropical birds may be less sensitive to variations in day length and thus less sensitive to light pollution.

Short-term adjustments of signal characteristics allow animals to maintain reliable communication in noise. Noise-dependent vocal plasticity often involves simultaneous changes in multiple parameters. Here, we quantified for the first time the relative contributions of signal amplitude, duration, and redundancy for improving signal detectability in noise. To this end, we used a combination of behavioural experiments on pale spear-nosed bats (Phyllostomus discolor) and signal detection models. In response to increasing noise levels, all bats raised the amplitude of their echolocation calls by 1.8-7.9 dB (the Lombard effect). Bats also increased signal duration by 13%-85%, corresponding to an increase in detectability of 1.0-5.3 dB. Finally, in some noise conditions, bats increased signal redundancy by producing more call groups. Assuming optimal cognitive integration, this could result in a further detectability improvement by up to 4 dB. Our data show that while the main improvement in signal detectability was due to the Lombard effect, increasing signal duration and redundancy can also contribute markedly to improving signal detectability. Overall, our findings demonstrate that the observed adjustments of signal parameters in noise are matched to how these parameters are processed in the receiver's sensory system, thereby facilitating signal transmission in fluctuating environments.

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.

Signal plasticity is considered an important step in the evolution of animal communication. In acoustic communication, signal transmission is often constrained by background noise. One adaptation to evade acoustic signal masking is the Lombard effect, in which an animal increases its vocal amplitude in response to an increase in background noise. This form of signal plasticity has been found in mammals, including humans, and some birds, but not frogs. However, the evolution of the Lombard effect is still unclear. Here we demonstrate for the first time the Lombard effect in a phylogentically basal bird species, the tinamou Eudromia elegans. By doing so, we take a step towards reconstructing the evolutionary history of noise-dependent vocal plasticity in birds. Similar to humans, the tinamous also raised their vocal pitch in noise, irrespective of any release from signal masking. The occurrence of the Lombard effect in a basal bird group suggests that this form of vocal plasticity was present in the common ancestor of all living birds and thus evolved at least as early as 119 Ma.

Social learning is the building block of culture and traditions in humans and nonhuman animals, and its study has a long history. Most investigations have addressed either the causation or the function of social learning. Though much is known about the underlying mechanisms of social learning, demonstrations of its adaptive value in a natural setting are lacking. Here we show that juvenile brown pelicans (Pelecanus occidentalis) can increase their foraging efficiency by copying adult diving behaviour, suggesting that social learning helps juveniles to find profitable food patches. Our findings demonstrate the potential fitness consequences of behavioural copying and thus highlight the possible adaptive importance of social learning.

The evolution of mating signals is closely linked to sexual selection. Acoustic ornaments are often used as secondary sexual traits that signal the quality of the signaller. Here we show that song performance reflects age and reproductive success in the rock sparrow (Petronia petronia). In an Alpine population in south-east France, we recorded the songs of males and assessed their genetic breeding success by microsatellite analysis. In addition to temporal and spectral song features, we also analysed for the first time whether the sound pressure level of bird song reflects reproductive success. Males with higher breeding success sang at a lower rate and with a higher maximum frequency. We found also that older males gained more extra-pair young and had a higher overall breeding success, although they also differed almost significantly by having a higher loss of paternity in their own nests. Older males could be distinguished from yearlings by singing at lower rate and higher amplitudes. Our findings suggest that song rate may be used as a signal of age and together with song pitch as a signal of reproductive success in this species. Alternatively, younger and less successful males might try to compensate their inferior status by increased song rates and lower pitch. Independent of age and quality, high-amplitude songs correlated with paternity loss in the own nest, suggesting that in this species song amplitude is not an indicator of male quality but high-intensity songs may be rather a response to unfaithful social mates.

Bird song is a widely used model in the study of animal communication and sexual selection, and several song features have been shown to reflect the quality of the singer. Recent studies have demonstrated that song amplitude may be an honest signal of current condition in males and that females prefer high amplitude songs. In addition, birds raise the amplitude of their songs to communicate in noisy environments. Although it is generally assumed that louder song should be more costly to produce, there has been little empirical evidence to support this assumption. We tested the assumption by measuring oxygen consumption and respiratory patterns in adult male zebra finches (Taeniopygia guttata) singing at different amplitudes in different background noise conditions. As background noise levels increased, birds significantly increased the sound pressure level of their songs. We found that louder songs required significantly greater subsyringeal air sac pressure than quieter songs. Though increased pressure is probably achieved by increasing respiratory muscle activity, these increases did not correlate with measurable increases in oxygen consumption. In addition, we found that oxygen consumption increased in higher background noise, independent of singing behaviour. This observation supports previous research in mammals showing that high levels of environmental noise can induce physiological stress responses. While our study did not find that increasing vocal amplitude increased metabolic costs, further research is needed to determine whether there are other non-metabolic costs of singing louder or costs associated with chronic noise exposure.

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.

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.

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.

Birds adjust their songs to make themselves heard in the presence of ambient noise. New research comparing songs of great tits across Europe shows how animals adapt their signals to the urban din.

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.

Primates acquire knowledge about relationships of third parties and group structure by monitoring their conspecifics. We show that Barbary macaques (Macaca sylvanus) utter specific vocalizations while monitoring interactions of other group members. As they did not direct other behaviours to the interacting group members, we provisionally termed these vocalizations 'vocal comments'. We investigated the acoustic properties of these comments and the social contexts in which they occurred. Most adult males and females of two studied groups produced low-amplitude calls when observing close contact interactions of other group members. The acoustic features of these calls varied with characteristics of the commented situation. Our results suggest that such calls might not be directed towards the agents of the commented situation, but towards other group members. The vocal comments may signal the caller's awareness of the observed interaction and possibly attract the attention of others to the situation.

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.

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.