Territorial behaviour of thrush nightingales outside the breeding season.

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.

Biological sex is binary, even though there is a rainbow of sex roles: Denying biological sex is anthropocentric and promotes species chauvinism: Denying biological sex is anthropocentric and promotes species chauvinism.

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."

Long-term effects of noise pollution on the avian dawn chorus: a natural experiment facilitated by the closure of an international airport.

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 across birds and the conditions for its evolution.

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.

Traffic noise disrupts vocal development and suppresses immune function.

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.

A global analysis of song frequency in passerines provides no support for the acoustic adaptation hypothesis but suggests a role for sexual selection.

Animals use acoustic signals for communication, implying that the properties of these signals can be under strong selection. The acoustic adaptation hypothesis predicts that species in dense habitats emit lower-frequency sounds than those in open areas because low-frequency sounds propagate further in dense vegetation than high-frequency sounds. Signal frequency may also be under sexual selection because it correlates with body size and lower-frequency sounds are perceived as more intimidating. Here, we evaluate these hypotheses by analysing variation in peak song frequency across 5,085 passerine species (Passeriformes). A phylogenetically informed analysis revealed that song frequency decreases with increasing body mass and with male-biased sexual size dimorphism. However, we found no support for the predicted relationship between frequency and habitat. Our results suggest that the global variation in passerine song frequency is mostly driven by natural and sexual selection causing evolutionary shifts in body size rather than by habitat-related selection on sound propagation.

Group living facilitates the evolution of duets in barbets.

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.

Traffic noise exposure depresses plasma corticosterone and delays offspring growth in breeding zebra finches.

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.

Higher songs of city birds may not be an individual response to noise.

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.

Vocal plasticity in a reptile.

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.

Lombard effect onset times reveal the speed of vocal plasticity in a songbird.

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.

Traffic noise drowns out great tit alarm calls.

Anthropogenic noise is one of the fastest growing and most ubiquitous types of environmental pollution and can impair acoustic communication in a variety of animals [1]. Recent research has shown that birds can adjust acoustic parameters of their sexual signals (songs) in noisy environments [2,3], yet we know little about other types of vocalizations. Anti-predator signals contain subtle information that is critical for avoiding predation [4,5], and failure to detect these calls [6,7] as a result of anthropogenic noise pollution could have large fitness consequences by negatively impacting survival. We investigated whether traffic noise impacts both the production and perception of avian alarm calls using a combination of lab and field experiments with great tits (Parus major), a songbird that frequently inhabits noise-polluted environments. In response to experimental noise manipulation in controlled laboratory conditions, great tits increased the amplitude, but not the frequency parameters, of their mobbing alarm calls (hereafter 'alarm calls'). Playback experiments conducted in the wild indicate that current levels of road traffic noise mask alarm calls, impeding the ability of great tits to perceive these critical signals. These results show that, despite the vocal adjustments used to compensate for anthropogenic noise, great tits are not able to restore the active space of their calls in even moderately noisy environments. Consequently, birds are likely to suffer from increased predation risk under noise, with likely effects on their behaviour, populations, and community dynamics in noise-polluted areas.

Airport noise predicts song timing of European birds.

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.

Anthropogenic noise, but not artificial light levels predicts song behaviour in an equatorial bird.

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.

Linking the sender to the receiver: vocal adjustments by bats to maintain signal detection in noise.

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.

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.

Juvenile Galápagos pelicans increase their foraging success by copying adult behaviour.

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.

On the evolution of noise-dependent vocal plasticity in birds.

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.