Dawn singing of the Brownish‐flanked Bush Warbler influences dawn chorusing in a bird community

Despite numerous studies on the function of the avian dawn chorus, few studies have examined whether dawn singing may influence the singing of other species. Here, we built on our previous study which found male Brownish‐flanked Bush Warblers (Horornis fortipes) increase their dawn singing intensity after conspecific playback on the previous day. We reanalyzed those recordings to quantify the start of dawn singing in other nine sympatric songbird species. Ranking‐scaling analyses identified a distinctive sequential pattern of dawn singing among these bird species between the first and the second dawn chorus, and meta‐analysis showed a significant trend to singing earlier in the bird community accompanied by the increase in dawn singing intensity in Brownish‐flanked Bush Warbler. Species with songs most similar to that of the Brownish‐flanked Bush Warbler and species that were phylogenetically distantly related to the Brownish‐flanked Bush Warbler showed a greater shift in the onset of dawn singing. Our study is one of the few studies showing how bird song influences heterospecific singing, and this may influence the temporal organization of song activity in the community, and result in synchronization in singing activities among different species, such as singing in dawn and dusk chorus.     

Early singers attend to conspecific but not heterospecific behavioural cues at dawn

During the dawn chorus songbirds initiate singing activities just prior to sunrise. Environmental factors affect the timing of the dawn chorus, but relatively little is known about how behavioural cues influence chorus timing. We assessed whether early playback of conspecific and heterospecific song influenced chorus start times in two early‐singing temperate species, hermit thrush Catharus guttatus and veery C. fuscescens. We used arrays of GPS time‐synchronized recorders to record the dawn chorus at 30 locations. In each location, thirty minutes prior to natural chorus initiation, we broadcast: 1) an early singing species (hermit thrush or veery), 2) a late singing species (black‐throated green warbler Setophaga virens or Nashville warbler Oreothlypis ruficapilla), 3) a noise control, and 4) also recorded without playback (silent control). Playback treatments were separated by two‐three days at each location. Both hermit thrush and veery sang significantly earlier in response to conspecific playback compared to noise or silent controls. Both species also sang earlier in response to conspecific playback compared to the day before or the day after playback. Neither species sang earlier in response to heterospecific playback (regardless of whether the broadcast species was a naturally early or late song initiator) compared to noise or silent controls. Our results support the theory that the dawn chorus is a communication network where individuals attend primarily to conspecific cues; heterospecific song appears to have minimal impacts on the chorus start times of the two early‐singing species we investigated.     

Eye size in birds and the timing of song at dawn

Why do different species of birds start their dawn choruses at different times? We test the hypothesis that the times at which different species start singing at dawn are related to their visual capability at low light intensities. Birds with large eyes can achieve greater pupil diameters and hence, all other things being equal, greater visual sensitivity and resolution than birds with small eyes. We estimated the maximum pupil diameter of passerine birds by measuring the diameter of the exposed eye surface, and measured the times of the first songs at dawn of songbirds present in different bird communities, and the light intensities at these times. Using phylogenetic comparative analyses, we found that songbirds with large eyes started to sing at lower light intensities (and therefore earlier) than species with smaller eyes. These relationships were stronger when differences in body size were controlled for statistically, and were consistent between two phylogenies and when species were treated as independent data points. Our results therefore provide robust support for the hypothesis that visual capability at low light levels influences the times at which birds start to sing at dawn.     

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.     

Sleepless in Town – Drivers of the Temporal Shift in Dawn Song in Urban European Blackbirds

Organisms living in urban environments are exposed to different environmental conditions compared to their rural conspecifics. Especially anthropogenic noise and artificial night light are closely linked to urbanization and pose new challenges to urban species. Songbirds are particularly affected by these factors, because they rely on the spread of acoustic information and adjust their behaviour to the rhythm of night and day, e.g. time their dawn song according to changing light intensities. Our aim was to clarify the specific contributions of artificial night light and traffic noise on the timing of dawn song of urban European Blackbirds (Turdus merula). We investigated the onset of blackbird dawn song along a steep urban gradient ranging from an urban forest to the city centre of Leipzig, Germany. This gradient of anthropogenic noise and artificial night light was reflected in the timing of dawn song. In the city centre, blackbirds started their dawn song up to 5 hours earlier compared to those in semi-natural habitats. We found traffic noise to be the driving factor of the shift of dawn song into true night, although it was not completely separable from the effects of ambient night light. We additionally included meteorological conditions into the analysis and found an effect on the song onset. Cloudy and cold weather delayed the onset, but cloud cover was assumed to reflect night light emissions, thus, amplified sky luminance and increased the effect of artificial night light. Beside these temporal effects, we also found differences in the spatial autocorrelation of dawn song onset showing a much higher variability in noisy city areas than in rural parks and forests. These findings indicate that urban hazards such as ambient noise and light pollution show a manifold interference with naturally evolved cycles and have significant effects on the activity patterns of urban blackbirds.     

Social stimulation of dawn singing in Dusky Flycatchers: a serendipitous experiment

Hypotheses regarding dawn singing in birds remain largely unsupported by quantitative data, especially for suboscine passerines (suborder Tyranni). During a study of the singing behavior of a suboscine, the Dusky Flycatcher (Empidonax oberholseri), in Alberta, Canada, spring storms in 2002 caused the disappearance and presumed mortality of several territorial males and some of their replacements, creating a serendipitous experiment. Overall, 15 males disappeared and, as a result, the number of territorial males in our study area declined from 13 prior to the storms to six afterward. Only one male maintained the same territory throughout the breeding season. During the period of inclement weather and social instability (20 May to 9 June), males sang at high rates during the predawn period. From 10 to 19 June, following this period of unstable weather and turnover in territorial males, males began dawn singing later in the morning, and sang shorter bouts at lower rates. The proportion of males engaging in dawn singing also decreased between the two periods. In contrast, daytime singing activity of paired males was low during both time periods. Playback of dawn songs to territorial males between 20 June and 21 July caused a resumption of some dawn singing. Singing rates were higher on the day of playback than on the day before playback, and the times when dawn singing was initiated were earlier on each of 2 d after playback than on the day before playback. In addition, the proportion of males engaging in dawn singing increased between the day before and the day after playback. Both the decrease in dawn singing when population density was reduced and its partial restoration by playback suggest that dawn singing in this species functions in male–male interactions and support the social‐dynamics hypothesis as an explanation for the dawn chorus in this species.     

Anthropogenic noise reduces bird species richness and diversity in urban parks

Anthropogenic noise is becoming more prevalent in the world and has been shown to affect many animal species, including birds. The impact of such noise was measured in Neotropical urban parks to assess how the noise affects avifauna diversity and species richness. We sampled bird species, and concurrently measured sound pressure (noise) levels (L_eq, equivalent noise levels) in eight urban green areas or parks located in a large city (Belo Horizonte) in south‐eastern Brazil over a 1‐year period. The diversity of sampled points was measured by means of total species richness, Fisher's alpha and Shannon–Wiener diversity indices. Noise levels within all parks were greater than those in natural areas. We found that an increase in noise levels and the area of open habitats surrounding sampling points were negatively related to species richness. Social factors reflecting increased urbanization, such as higher incomes, were also negatively correlated with bird species richness. However, noise was the factor that explained most of the variance. These results suggest that anthropogenic noise can have a significant negative impact on the conservation value of urban parks for bird species.     

Long-term influence of simulated territorial intrusions on dawn and dusk singing in the Winter Wren: spring versus autumn

Males of many songbird species have peaks of singing activity at dawn and dusk. Singing during those twilight periods can function in territory proclamation, and males are suggested to adjust song output to the level of intruder pressure. We used song playback during the breeding season to simulate intrusions into territories of male Winter Wrens (Troglodytes troglodytes) shortly after dawn. We then compared male singing behaviour during the dawn and dusk chorus before and 1 day after the simulated intrusion. One day after the playback, male Wrens increased their song output before sunrise, which confirms our results from a previous study on dawn singing in autumn territories. At dusk, on the evening following the playback, males slightly increased song output after sunset, but singing activity at dusk was generally very low. We found no significant changes of song output after sunrise, before sunset, and between 2 days of control without playback. These results are consistent with the hypothesis that dawn and dusk singing is important for territory defence in spring. Unlike in autumn, however, increased singing in spring at dawn and dusk could also serve to defend other resources such as fertile mates or to strengthen the pair bond after a territorial challenge. In comparison with the results on autumnal singing, male Wrens started singing earlier at dawn during the breeding season, and they generally sang more songs at dawn and immediately after playback. The increase in absolute numbers of songs sung in the morning after playback seemed greater in spring than in autumn; however, the proportional increase relative to overall song output was similar in both seasons.     

Vocal traits and diet explain avian sensitivities to anthropogenic noise

Global population growth has caused extensive human‐induced environmental change, including a near‐ubiquitous transformation of the acoustical environment due to the propagation of anthropogenic noise. Because the acoustical environment is a critical ecological dimension for countless species to obtain, interpret and respond to environmental cues, highly novel environmental acoustics have the potential to negatively impact organisms that use acoustics for a variety of functions, such as communication and predator/prey detection. Using a comparative approach with 308 populations of 183 bird species from 14 locations in Europe, North American and the Caribbean, I sought to reveal the intrinsic and extrinsic factors responsible for avian sensitivities to anthropogenic noise as measured by their habitat use in noisy versus adjacent quiet locations. Birds across all locations tended to avoid noisy areas, but trait‐specific differences emerged. Vocal frequency, diet and foraging location predicted patterns of habitat use in response to anthropogenic noise, but body size, nest placement and type, other vocal features and the type of anthropogenic noise (chronic industrial vs. intermittent urban/traffic noise) failed to explain variation in habitat use. Strongly supported models also indicated the relationship between sensitivity to noise and predictive traits had little to no phylogenetic structure. In general, traits associated with hearing were strong predictors – species with low‐frequency vocalizations, which experience greater spectral overlap with low‐frequency anthropogenic noise tend to avoid noisy areas, whereas species with higher frequency vocalizations respond less severely. Additionally, omnivorous species and those with animal‐based diets were more sensitive to noise than birds with plant‐based diets, likely because noise may interfere with the use of audition in multimodal prey detection. Collectively, these results suggest that anthropogenic noise is a powerful sensory pollutant that can filter avian communities nonrandomly by interfering with birds' abilities to receive, respond to and dispatch acoustic cues and signals.     

Conveying information with one song type: changes in dawn song performance correspond to different female breeding stages

Many bird species participate in dawn singing, a behaviour categorized by intensive singing at dawn; however, many of these species deliver only one song type at dawn. While there are many proximate and ultimate hypotheses for why birds sing at dawn, little is known about whether males are able to vary one simple song to convey different information. We used autonomous acoustic recorders to record dawn songs of field sparrows and quantified three parameters of singing performance: 1) bout length, 2) song rate, and 3) song complexity. We found that males sang the longest dawn bouts during their mate's fertile period, the highest song rates during the post-fertile period, and the most complex songs during the pre-fertile period. The change in dawn singing behaviour with their mate's breeding stage suggest the purpose of dawn song may be context dependent. Our results demonstrate that male field sparrows, while only having a single song type sung at dawn, may convey information for both intra- and intersexual purposes. While it is generally assumed that dawn song has a specific function, the variability in the duration, rate, and complexity of dawn song in field sparrows suggests that they are conveying different information and that dawn song likely has multiple functions.     

Patterns of dawn singing by Buff-breasted Flycatchers

ABSTRACT.   Many passerine species exhibit a "dawn chorus"—a bout of intense singing activity before or at dawn, but our understanding of this phenomenon is poor. Tyrant flycatchers (Tyrannidae) exhibit pronounced daily bouts of dawn singing. I documented this behavior in several populations of Buff-breasted Flycatchers ( Empidonax fulvifrons ) in Arizona, tape recording >30,000 songs of 23 individuals during dawn singing. Individual males sang a dawn bout each morning, even during breeding phases when daytime song was almost completely absent. Dawn bouts began 5–10 min before local civil twilight and continued for 25–30 min. Each male possessed two song types delivered at high rates during dawn singing (four times the rate during sustained daytime singing). Song rate varied significantly over the course of dawn bouts, increasing to 55 songs / min at mid-bout, then declining to the end of the bout. Type 2 songs comprised about 30% of songs during dawn singing, and decreased significantly in proportion during the final 10 min of the bout. Songs of the two types were delivered in a nonrandom fashion. Males sang at locations near territory boundaries and pairs of neighbors engaged in counter-singing from the same locations each morning. A number of dawn singing bouts ended with attempted or successful copulations. These observations are consistent with the social dynamics hypothesis for the functional significance of dawn singing in this species.     

Sunrise in the city: disentangling drivers of the avian dawn chorus onset in urban greenspaces

Urban systems are known to have a number of effects on avian richness, density, and morphological and behavioral traits. However, no study to date has simultaneously examined the wide range of urban variables in relation to the avian dawn chorus, a complex behavioral phenomenon. Previous studies investigating adjustments of the dawn chorus onset in urban settings have mainly been confined to relationships with noise and light levels. In addition to noise and light levels, in this study we included other potentially related environmental characteristics describing vegetation structure, urban infrastructure, and human activity, all of which have been shown to be drivers of bird diversity in urban areas. We conducted dawn chorus surveys at 38 Los Angeles urban greenspaces and used a classification and regression tree analysis to identify specific urban scenarios that best explained timing differences in the dawn chorus onset. Our results show that light level was the most important variable related to the dawn chorus onset time, in which, counter‐intuitively, bird communities in greenspaces with higher light levels had later onsets. In addition, noise was an important factor for the chorus onset in greenspaces with higher light levels. Although our results differ from those of previous studies, these findings highlight the importance of noise and light levels in explaining dawn chorus onset variation, indicating the need for further research in untangling this complex and ecologically important phenomenon.     

Experimentally increased noise levels change spatial and singing behaviour

The reasons why animal populations decline in response to anthropogenic noise are still poorly understood. To understand how populations are affected by noise, we must understand how individuals are affected by noise. By modifying the acoustic environment experimentally, we studied the potential relationship between noise levels and both spatial and singing behaviour in the European robin (Erithacus rubecula). We found that with increasing noise levels, males were more likely to move away from the noise source and changed their singing behaviour. Our results provide the first experimental evidence in a free ranging species, that not merely the presence of noise causes changes in behaviour and distribution, but that the level of noise pollution plays a crucial role as well. Our results have important implications for estimating the impact of infrastructure which differs in the level of noise produced. Thus, governmental planning bodies should not only consider the physical effect on the landscape when assessing the impact of new infrastructure, but also the noise levels emitted, which may reduce the loss of suitable habitats available for animals.     

Dawn Singing Intensity of the Male Brownish‐Flanked Bush Warbler: Effects of Territorial Insertions and Number of Neighbors

The dawn chorus is a period of peak singing activity of many songbirds. Numerous studies have sought to understand this widespread phenomenon, and many hypotheses have been proposed to explain the dawn chorus. The social dynamics hypothesis proposes that dawn singing plays an important role in the announcement of territorial occupancy and the regulation of social relationships between males; it predicts that the dawn chorus vocal behavior varies with changes in social relationships. In this study, we tested the influence of territorial insertions and the number of neighbors, on the intensity of the brownish‐flanked bush warbler (Cettia fortipes)'s dawn singing. We found that simulated territorial insertions (playback) caused the males to increase their dawn singing significantly the next day, and males that had many neighbors exhibited more intense dawn singing than did males with few neighbors. Our study provides evidence that dawn singing plays an important role in the announcement of territorial occupancy and the regulation of the social relationships between the males.     

Anthropogenic noise decreases urban songbird diversity and may contribute to homogenization

More humans reside in urban areas than at any other time in history. Protected urban green spaces and transportation greenbelts support many species, but diversity in these areas is generally lower than in undeveloped landscapes. Habitat degradation and fragmentation contribute to lowered diversity and urban homogenization, but less is known about the role of anthropogenic noise. Songbirds are especially vulnerable to anthropogenic noise because they rely on acoustic signals for communication. Recent studies suggest that anthropogenic noise reduces the density and reproductive success of some bird species, but that species which vocalize at frequencies above those of anthropogenic noise are more likely to inhabit noisy areas. We hypothesize that anthropogenic noise is contributing to declines in urban diversity by reducing the abundance of select species in noisy areas, and that species with low‐frequency songs are those most likely to be affected. To examine this relationship, we calculated the noise‐associated change in overall species richness and in abundance for seven common songbird species. After accounting for variance due to vegetative differences, species richness and the abundance of three of seven species were reduced in noisier locations. Acoustic analysis revealed that minimum song frequency was highly predictive of a species' response to noise, with lower minimum song frequencies incurring greater noise‐associated reduction in abundance. These results suggest that anthropogenic noise affects some species independently of vegetative conditions, exacerbating the exclusion of some songbird species in otherwise suitable habitat. Minimum song frequency may provide a useful metric to predict how particular species will be affected by noise. In sum, mitigation of noise may enhance habitat suitability for many songbird species, especially for species with songs that include low‐frequency elements.     

Traffic noise affects forest bird species in a protected tropical forest

The construction of roads near protected forest areas alters ecosystem function by creating habitat fragmentation and through several direct and indirect negative effects such as increased pollution, animal mortality through collisions, disturbance caused by excessive noise and wind turbulence. Noise in particular may have strong negative effects on animal groups such as frogs and birds, that rely on sound for communication as it can negatively interfere with vocalizations used for territorial defense or courtship. Thus, birds are expected to be less abundant close to the road where noise levels are high. In this study, we examined the effects of road traffic noise levels on forest bird species in a protected tropical forest in Costa Rica. Data collection was conducted in a forest segment of the Carara National Park adjacent to the Coastal Highway. We carried out 120 ten minute bird surveys and measured road noise levels 192 times from the 19th to the 23rd of April and from the 21st to the 28th of November, 2008. To maximize bird detection for the species richness estimates we operated six 12 m standard mist nets simultaneously with the surveys. The overall mist-netting effort was 240 net/h. In addition, we estimated traffic volumes by tallying the number of vehicles passing by the edge of the park using 24 one hour counts throughout the study. We found that the relative abundance of birds and bird species richness decreased significantly with the increasing traffic noise in the dry and wet season. Noise decreased significantly and in a logarithmic way with distance from the road in both seasons. However, noise levels at any given distance were significantly higher in the dry compared to the wet season. Our results suggest that noise might be an important factor influencing road bird avoidance as measured by species richness and relative abundance. Since the protected forest in question is located in a national park subjected to tourist visitation, these results have conservation as well as management implications. A decrease in bird species richness and bird abundance due to intrusive road noise could negatively affect the use of trails by visitors. Alternatives for noise attenuation in the affected forest area include the enforcement of speed limits and the planting of live barriers.     

Understanding the function of nocturnal song in ovenbirds: males do not countersing at night

Many diurnal bird species vocalize at night, however the function of nocturnal song is, generally, still poorly understood. Previous research has suggested that nocturnal song may serve a social function and is influenced by environmental factors. To test whether males attend to the nocturnal song of conspecifics, we experimentally exposed ovenbirds Seiurus aurocapilla to nocturnal flight songs, and recorded their response both during the night and during the following dawn chorus. We compared latency to song and vocal output before and after playback exposure to determine if males altered their vocalizations in response to exposure to flight songs from an unknown male. We found no evidence of counter singing or change in nocturnal song output, nor a change in vocal output during the dawn chorus following playback exposure. Our results suggest that, in ovenbirds, nocturnal song does not serve as an intraspecific social function. Nocturnal song, through rare, may be significant in the mating systems of some diurnal bird species, and requires additional study.     

Interspecific Response to Playback of Bird Song

Responses to bird song have usually only been studied at the intraspecific level. I experimentally tested whether playback of the song of the black wheatear Oenanthe leucura in an area in S Spain resulted in responses from conspecifics as well as heterospecific birds by comparing the numbers of individuals singing before and after playback. The number of singing male black wheatears increased considerably, but also the number of singing males of five other passerine species increased significantly. The heterospecific response to playback may be due (1) to interspecific territoriality, (2) to black wheatear song signalling the absence of predators, or (3) to heterospecifics confusing the species-identity of the singer. The second alternative is considered more likely, since an ecologically wide array of species increased their song rate following playback. The conspicuous dawn (and dusk) chorus of bird song may be augmented by social facilitation due to the singing of conspecifics as well as heterospecifics.     

Variation in singing style use in the reed bunting Emberiza schoeniclus: influencing factors and possible functions

The two main functions of bird song are territory defence and mate attraction. Considerable progress has been made in understanding how species adjust the use of songs to serve these and other (presumed) functions of bird song, but the striking variety of singing behavior observable in wild birds remains enigmatic. Some species make do with simple songs and small repertoires, while others show large, complex repertoires and still others have evolved several distinct singing styles. In most species with distinct singing styles, however, the functions of singing styles are poorly understood. Two distinct singing styles (type I and II, respectively) have long been known in the reed bunting Emberiza schoeniclus, while a new third one has recently been reported to exist. We first quantitatively investigated the evidence for the existence of three singing styles. Then, we tested predictions of the mate attraction hypothesis, the mate guarding hypothesis and the territory defence hypothesis by examining the relations between singing style use with social and temporal factors. Cluster and discriminant analyses supported the existence of three (instead of two) singing styles, which could be differentiated based on four variables referring to song structure and complexity. Use of singing styles was related to male mating status (consistent with the mate attraction hypothesis), but not to female breeding stage (no support for the mate guarding hypothesis). Finally, use of singing styles differed in relation to time of day, with the dawn chorus of paired reed buntings consisting almost exclusively of songs of the recently discovered type III singing style and daytime singing primarily consisting of songs of long‐known type I (in unpaired males) or II singing styles (in paired males). Our findings suggest that one singing style (type I) primarily serves to attract a social mate, although an additional territorial function of this singing style cannot be dismissed. The function(s) of the other two singing styles, both only sung by paired males, are not related to attraction of a social mate or to the own female's fertility, but appear to be important in the context of territory defence and extra‐pair matings.     

Stadtamseln sind anders

City blackbirds are different Abiotic factors such as traffic noise and especially artificial light do not just influence species in cities but also change them. The example of the European blackbird (Turdus merula) illustrates how adaptions to city environments can initiate micro‐evolutionary changes. Artificial light clearly alters daily activity patterns and leads to changes in reproductive behaviour: blackbirds sing earlier in the morning, are awake for longer and start reproduction earlier in the year. Traffic noise also changes their song. In order to compensate for interfering traffic noise, city blackbirds sing louder and at higher frequencies. In addition, blackbirds in cities are less curious but also show less fear towards humans than their forest counterparts. Cities have the potential to become a home for some species, but they also change them. One of the big questions of the future will be if, in the globally expanding city environments, this process will lead to the evolution of adapated city species.