Sensory pollutants alter bird phenology and fitness across a continent.

Expansion of anthropogenic noise and night lighting across our planet1,2 is of increasing conservation concern3-6. Despite growing knowledge of physiological and behavioural responses to these stimuli from single-species and local-scale studies, whether these pollutants affect fitness is less clear, as is how and why species vary in their sensitivity to these anthropic stressors. Here we leverage a large citizen science dataset paired with high-resolution noise and light data from across the contiguous United States to assess how these stimuli affect reproductive success in 142 bird species. We find responses to both sensory pollutants linked to the functional traits and habitat affiliations of species. For example, overall nest success was negatively correlated with noise among birds in closed environments. Species-specific changes in reproductive timing and hatching success in response to noise exposure were explained by vocalization frequency, nesting location and diet. Additionally, increased light-gathering ability of species' eyes was associated with stronger advancements in reproductive timing in response to light exposure, potentially creating phenological mismatches7. Unexpectedly, better light-gathering ability was linked to reduced clutch failure and increased overall nest success in response to light exposure, raising important questions about how responses to sensory pollutants counteract or exacerbate responses to other aspects of global change, such as climate warming. These findings demonstrate that anthropogenic noise and light can substantially affect breeding bird phenology and fitness, and underscore the need to consider sensory pollutants alongside traditional dimensions of the environment that typically inform biodiversity conservation.

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.

A stochastic simulation model for assessing the masking effects of road noise for wildlife, outdoor recreation, and bioacoustic monitoring.

Traffic noise is one of the leading causes of reductions in animal abundances near roads. Acoustic masking of conspecific signals and adventitious cues is one mechanism that likely causes animals to abandon loud areas. However, masking effects can be difficult to document in situ and the effects of infrequent noise events may be impractical to study. Here, we present the Soundscapes model, a stochastic individual-based model that dynamically models the listening areas of animals searching for acoustic resources ("searchers"). The model also studies the masking effects of noise for human detections of the searchers. The model is set in a landscape adjacent to a road. Noise produced by vehicles traveling on that road is represented by calibrated spectra that vary with speed. Noise propagation is implemented using ISO-9613 procedures. We present demonstration simulations that quantify declines in searcher efficiency and human detection of searchers at relatively low traffic volumes, fewer than 50 vehicles per hour. Traffic noise is pervasive, and the Soundscapes model offers an extensible tool to study the effects of noise on bioacoustics monitoring, point-count surveys, the restorative value of natural soundscapes, and auditory performance in an ecological context.

Long-term noise pollution affects seedling recruitment and community composition, with negative effects persisting after removal.

Noise pollution can affect species' behaviours and distributions and may hold significant consequences for natural communities. While several studies have researched short-term effects of noise, no long-term research has examined whether observed patterns persist or if community recovery can occur. We used a long-term study system in New Mexico to examine the effects of continuous natural gas well noise exposure on seedling recruitment of foundational tree species (Pinus edulis, Juniperus osteosperma) and vegetation diversity. First, we examined seedling recruitment and vegetation diversity at plots where current noise levels have persisted for greater than 15 years. We then examined recruitment and diversity on plots where noise sources were recently removed or added. We found support for long-term negative effects of noise on tree seedling recruitment, evenness of woody plants and increasingly dissimilar vegetation communities with differences in noise levels. Furthermore, seedling recruitment and plant community composition did not recover following noise removal, possibly due in part to a lag in recovery among animals that disperse and pollinate plants. Our results add to the limited evidence that noise has cascading ecological effects. Moreover, these effects may be long lasting and noise removal may not lead to immediate recovery.

Light pollution affects West Nile virus exposure risk across Florida.

Emerging infectious diseases (EIDs) present global health threats, and their emergences are often linked to anthropogenic change. Artificial light at night (ALAN) is one form of anthropogenic change that spans beyond urban boundaries and may be relevant to EIDs through its influence on the behaviour and physiology of hosts and/or vectors. Although West Nile virus (WNV) emergence has been described as peri-urban, we hypothesized that exposure risk could also be influenced by ALAN in particular, which is testable by comparing the effects of ALAN on prevalence while controlling for other aspects of urbanization. By modelling WNV exposure among sentinel chickens in Florida, we found strong support for a nonlinear relationship between ALAN and WNV exposure risk in chickens with peak WNV risk occurring at low ALAN levels. Although our goal was not to discern how ALAN affected WNV relative to other factors, effects of ALAN on WNV exposure were stronger than other known drivers of risk (i.e. impervious surface, human population density). Ambient temperature in the month prior to sampling, but no other considered variables, strongly influenced WNV risk. These results indicate that ALAN may contribute to spatio-temporal changes in WNV risk, justifying future investigations of ALAN on other vector-borne parasites.

Artificial night light and anthropogenic noise interact to influence bird abundance over a continental scale.

The extent of artificial night light and anthropogenic noise (i.e., "light" and "noise") impacts is global and has the capacity to threaten species across diverse ecosystems. Existing research involving impacts of light or noise has primarily focused on noise or light alone and single species; however, these stimuli often co-occur and little is known about how co-exposure influences wildlife and if and why species may vary in their responses. Here, we had three aims: (1) to investigate species-specific responses to light, noise, and the interaction between the two using a spatially explicit approach to model changes in abundance of 140 prevalent bird species across North America, (2) to investigate responses to the interaction between light exposure and night length, and (3) to identify functional traits and habitat affiliations that explain variation in species-specific responses to these sensory stimuli with phylogenetically informed models. We found species that responded to noise exposure generally decreased in abundance, and the additional presence of light interacted synergistically with noise to exacerbate its negative effects. Moreover, the interaction revealed negative emergent responses for several species that only reacted when light and noise co-occurred. Additionally, an interaction between light and night length revealed 47 species increased in abundance with light exposure during longer nights. In addition to modifying behavior with optimal temperature and potential foraging opportunities, birds might be attracted to light, yet suffer inadvertent physiological consequences. The trait that most strongly related to avian response to light and noise was habitat affiliation. Specifically, species that occupy closed habitat were less tolerant of both sensory stressors compared to those that occupy open habitat. Further quantifying the contexts and intrinsic traits that explain how species respond to noise and light will be fundamental to understanding the ecological consequences of a world that is ever louder and brighter.

Chronic anthropogenic noise disrupts glucocorticoid signaling and has multiple effects on fitness in an avian community.

Anthropogenic noise is a pervasive pollutant that decreases environmental quality by disrupting a suite of behaviors vital to perception and communication. However, even within populations of noise-sensitive species, individuals still select breeding sites located within areas exposed to high noise levels, with largely unknown physiological and fitness consequences. We use a study system in the natural gas fields of northern New Mexico to test the prediction that exposure to noise causes glucocorticoid-signaling dysfunction and decreases fitness in a community of secondary cavity-nesting birds. In accordance with these predictions, and across all species, we find strong support for noise exposure decreasing baseline corticosterone in adults and nestlings and, conversely, increasing acute stressor-induced corticosterone in nestlings. We also document fitness consequences with increased noise in the form of reduced hatching success in the western bluebird (Sialia mexicana), the species most likely to nest in noisiest environments. Nestlings of all three species exhibited accelerated growth of both feathers and body size at intermediate noise amplitudes compared with lower or higher amplitudes. Our results are consistent with recent experimental laboratory studies and show that noise functions as a chronic, inescapable stressor. Anthropogenic noise likely impairs environmental risk perception by species relying on acoustic cues and ultimately leads to impacts on fitness. Our work, when taken together with recent efforts to document noise across the landscape, implies potential widespread, noise-induced chronic stress coupled with reduced fitness for many species reliant on acoustic cues.

Direct and indirect effects of noise pollution alter biological communities in and near noise-exposed environments.

Noise pollution is pervasive across every ecosystem on Earth. Although decades of research have documented a variety of negative impacts of noise to organisms, key gaps remain, such as how noise affects different taxa within a biological community and how effects of noise propagate across space. We experimentally applied traffic noise pollution to multiple roadless areas and quantified the impacts of noise on birds, grasshoppers and odonates. We show that acoustically oriented birds have reduced species richness and abundance and different community compositions in experimentally noise-exposed areas relative to comparable quiet locations. We also found both acoustically oriented grasshoppers and odonates without acoustic receptors to have reduced species richness and/or abundance in relatively quiet areas that abut noise-exposed areas. These results suggest that noise pollution not only affects acoustically oriented animals, but that noise may reverberate through biological communities through indirect effects to those with no clear links to the acoustic realm, even in adjacent quiet environments.

The phantom chorus: birdsong boosts human well-being in protected areas.

Spending time in nature is known to benefit human health and well-being, but evidence is mixed as to whether biodiversity or perceptions of biodiversity contribute to these benefits. Perhaps more importantly, little is known about the sensory modalities by which humans perceive biodiversity and obtain benefits from their interactions with nature. Here, we used a 'phantom birdsong chorus' consisting of hidden speakers to experimentally increase audible birdsong biodiversity during 'on' and 'off' (i.e. ambient conditions) blocks on two trails to study the role of audition in biodiversity perception and self-reported well-being among hikers. Hikers exposed to the phantom chorus reported higher levels of restorative effects compared to those that experienced ambient conditions on both trails; however, increased restorative effects were directly linked to the phantom chorus on one trail and indirectly linked to the phantom chorus on the other trail through perceptions of avian biodiversity. Our findings add to a growing body of evidence linking mental health to nature experiences and suggest that audition is an important modality by which natural environments confer restorative effects. Finally, our results suggest that maintaining or improving natural soundscapes within protected areas may be an important component to maximizing human experiences.

Macroevolutionary Patterning in Glucocorticoids Suggests Different Selective Pressures Shape Baseline and Stress-Induced Levels.

Glucocorticoid (GC) hormones are important phenotypic mediators across vertebrates, but their circulating concentrations can vary markedly. Here we investigate macroevolutionary patterning in GC levels across tetrapods by testing seven specific hypotheses about GC variation and evaluating whether the supported hypotheses reveal consistent patterns in GC evolution. If selection generally favors the "supportive" role of GCs in responding effectively to challenges, then baseline and/or stress-induced GCs may be higher in challenging contexts. Alternatively, if selection generally favors "protection" from GC-induced costs, GCs may be lower in environments where challenges are more common or severe. The predictors of baseline GCs were all consistent with supportive effects: levels were higher in smaller organisms and in those inhabiting more energetically demanding environments. During breeding, baseline GCs were also higher in populations and species with fewer lifetime opportunities to reproduce. The predictors of stress-induced GCs were instead more consistent with the protection hypothesis: during breeding, levels were lower in organisms with fewer lifetime reproductive opportunities. Overall, these patterns indicate a surprising degree of consistency in how some selective pressures shape GCs across broad taxonomic scales; at the same time, in challenging environments selection appears to operate on baseline and stress-induced GCs in distinct ways.

No evidence that warmer temperatures are associated with selection for smaller body sizes.

Reductions in animal body size over recent decades are often interpreted as an adaptive evolutionary response to climate warming. However, for reductions in size to reflect adaptive evolution, directional selection on body size within populations must have become negative, or where already negative, to have become more so, as temperatures increased. To test this hypothesis, we performed traditional and phylogenetic meta-analyses of the association between annual estimates of directional selection on body size from wild populations and annual mean temperatures from 39 longitudinal studies. We found no evidence that warmer environments were associated with selection for smaller size. Instead, selection consistently favoured larger individuals, and was invariant to temperature. These patterns were similar in ectotherms and endotherms. An analysis using year rather than temperature revealed similar patterns, suggesting no evidence that selection has changed over time, and also indicating that the lack of association with annual temperature was not an artefact of choosing an erroneous time window for aggregating the temperature data. Although phenotypic trends in size will be driven by a combination of genetic and environmental factors, our results suggest little evidence for a necessary ingredient-negative directional selection-for declines in body size to be considered an adaptive evolutionary response to changing selection pressures.

Sound settlement: noise surpasses land cover in explaining breeding habitat selection of secondary cavity-nesting birds.

Birds breeding in heterogeneous landscapes select nest sites by cueing in on a variety of factors from landscape features and social information to the presence of natural enemies. We focus on determining the relative impact of anthropogenic noise on nest site occupancy, compared to amount of forest cover, which is known to strongly influence the selection process. We examine chronic, industrial noise from natural gas wells directly measured at the nest box as well as site-averaged noise, using a well-established field experimental system in northwestern New Mexico. We hypothesized that high levels of noise, both at the nest site and in the environment, would decrease nest box occupancy. We set up nest boxes using a geospatially paired control and experimental site design and analyzed four years of occupancy data from four secondary cavity-nesting birds common to the Colorado Plateau. We found different effects of noise and landscape features depending on species, with strong effects of noise observed in breeding habitat selection of Myiarchus cinerascens, the Ash-throated Flycatcher, and Sialia currucoides, the Mountain Bluebird. In contrast, the amount of forest cover less frequently explained habitat selection for those species or had a smaller standardized effect than the acoustic environment. Although forest cover characterization and management is commonly employed by natural resource managers, our results show that characterizing and managing the acoustic environment should be an important tool in protected area management.

Acoustic environments matter: Synergistic benefits to humans and ecological communities.

Protected areas are critical locations worldwide for biodiversity preservation and offer important opportunities for increasingly urbanized humans to experience nature. However, biodiversity preservation and visitor access are often at odds and creative solutions are needed to safeguard protected area natural resources in the face of high visitor use. Managing human impacts to natural soundscapes could serve as a powerful tool for resolving these conflicting objectives. Here, we review emerging research that demonstrates that the acoustic environment is critical to wildlife and that sounds shape the quality of nature-based experiences for humans. Human-made noise is known to affect animal behavior, distributions and reproductive success, and the organization of ecological communities. Additionally, new research suggests that interactions with nature, including natural sounds, confer benefits to human welfare termed psychological ecosystem services. In areas influenced by noise, elevated human-made noise not only limits the variety and abundance of organisms accessible to outdoor recreationists, but also impairs their capacity to perceive the wildlife that remains. Thus soundscape changes can degrade, and potentially limit the benefits derived from experiences with nature via indirect and direct mechanisms. We discuss the effects of noise on wildlife and visitors through the concept of listening area and demonstrate how the perceptual worlds of both birds and humans are reduced by noise. Finally, we discuss how management of soundscapes in protected areas may be an innovative solution to safeguarding both and recommend several key questions and research directions to stimulate new research.

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.

Understanding park visitors' soundscape perception using subjective and objective measurement.

Environmental noise knows no boundaries, affecting even protected areas. Noise pollution, originating from both external and internal sources, imposes costs on these areas. It is associated with adverse health effects, while natural sounds contribute to cognitive and emotional improvements as ecosystem services. When it comes to parks, individual visitors hold unique perceptions of soundscapes, which can be shaped by various factors such as their motivations for visiting, personal norms, attitudes towards specific sounds, and expectations. In this study, we utilized linear models and geospatial data to evaluate how visitors' personal norms and attitudes, the park's acoustic environment, visitor counts, and the acoustic environment of visitors' neighborhoods influenced their perception of soundscapes at Muir Woods National Monument. Our findings indicate that visitors' subjective experiences had a greater impact on their perception of the park's soundscape compared to purely acoustic factors like sound level of the park itself. Specifically, we found that motivations to hear natural sounds, interference caused by noise, sensitivity to noise, and the sound levels of visitors' home neighborhoods influenced visitors' perception of the park's soundscape. Understanding how personal factors shape visitors' soundscape perception can assist urban and non-urban park planners in effectively managing visitor experiences and expectations.

Unravelling the determinants of insular body size shifts.

The island rule, a pattern of size shifts on islands, is an oft-cited but little understood phenomenon of evolutionary biology. Here, we explore the evolutionary mechanisms behind the rule in 184 mammal species, testing climatic, ecological and phylogenetic hypotheses in a robust quantitative framework. Our findings confirm the importance of species' ecological traits in determining both the strength and the direction of body size changes on islands. Although the island rule pattern appears relatively weak overall, we find strongest support for models incorporating trait, climatic and geographical factors in a phylogenetic context, lending support to the idea that the island rule is a complex phenomenon driven by interacting intrinsic and extrinsic mechanisms. Overall, we find that different clades may be evolutionarily predisposed to dwarfism or gigantism, but the magnitude of size changes depends more on adaptation to the novel island environment.

Noise pollution alters ecological services: enhanced pollination and disrupted seed dispersal.

Noise pollution is a novel, widespread environmental force that has recently been shown to alter the behaviour and distribution of birds and other vertebrates, yet whether noise has cumulative, community-level consequences by changing critical ecological services is unknown. Herein, we examined the effects of noise pollution on pollination and seed dispersal and seedling establishment within a study system that isolated the effects of noise from confounding stimuli common to human-altered landscapes. Using observations, vegetation surveys and pollen transfer and seed removal experiments, we found that effects of noise pollution can reverberate through communities by disrupting or enhancing these ecological services. Specifically, noise pollution indirectly increased artificial flower pollination by hummingbirds, but altered the community of animals that prey upon and disperse Pinus edulis seeds, potentially explaining reduced P. edulis seedling recruitment in noisy areas. Despite evidence that some ecological services, such as pollination, may benefit indirectly owing to noise, declines in seedling recruitment for key-dominant species such as P. edulis may have dramatic long-term effects on ecosystem structure and diversity. Because the extent of noise pollution is growing, this study emphasizes that investigators should evaluate the ecological consequences of noise alongside other human-induced environmental changes that are reshaping human-altered landscapes worldwide.

Environmental variability and acoustic signals: a multi-level approach in songbirds.

Among songbirds, growing evidence suggests that acoustic adaptation of song traits occurs in response to habitat features. Despite extensive study, most research supporting acoustic adaptation has only considered acoustic traits averaged for species or populations, overlooking intraindividual variation of song traits, which may facilitate effective communication in heterogeneous and variable environments. Fewer studies have explicitly incorporated sexual selection, which, if strong, may favour variation across environments. Here, we evaluate the prevalence of acoustic adaptation among 44 species of songbirds by determining how environmental variability and sexual selection intensity are associated with song variability (intraindividual and intraspecific) and short-term song complexity. We show that variability in precipitation can explain short-term song complexity among taxonomically diverse songbirds, and that precipitation seasonality and the intensity of sexual selection are related to intraindividual song variation. Our results link song complexity to environmental variability, something previously found for mockingbirds (Family Mimidae). Perhaps more importantly, our results illustrate that individual variation in song traits may be shaped by both environmental variability and strength of sexual selection.

Artificial light at night and anthropogenic noise alter the foraging activity and structure of vertebrate communities.

Light and noise pollution from human activity are increasing at a dramatic rate. These sensory stimuli can have a wide range of effects on animal behavior, reproductive success, and physiology. However, less is known about the functional and community-level consequences of these sensory pollutants, especially when they co-occur. Using camera traps in a manipulative field experiment, we studied the effects of anthropogenic light and noise, singularly and in tandem, on richness and community turnover at both the taxa and functional group level as well as foraging activity. We showed that both light and noise pollution did alter taxonomic richness and that these effects can differ depending on the scale of observation. Increases in light levels had a negative effect on richness at the camera-level scale, but light-treated sites had the highest pooled (i.e., cumulative) richness of all treatment types. In contrast, noise was found to have a negative effect on cumulative richness; however, when both stimuli were present, the addition of night-lighting mitigated the effects of noise. Artificial light and moonlight had the strongest influence on community turnover, and results remained consistent at both the taxa and functional group level. Additionally, increases in ambient noise and moonlight, but not artificial light, reduced foraging activity. Our study provides evidence that alterations to the sensory environment can alter the richness and composition of communities and that effects can be scale-dependent and also alter foraging behavior. Unexpectedly, the addition of artificial light may have mitigated the negative effects of noise on cumulative taxonomic richness. This highlights the importance of researching the consequences of co-exposure to these globally common pollutants.

Experimentally broadcast ocean surf and river noise alters birdsong.

Anthropogenic noise and its effects on acoustic communication have received considerable attention in recent decades. Yet, the natural acoustic environment's influence on communication and its role in shaping acoustic signals remains unclear. We used large-scale playbacks of ocean surf in coastal areas and whitewater river noise in riparian areas to investigate how natural sounds influences song structure in six songbird species. We recorded individuals defending territories in a variety of acoustic conditions across 19 study sites in California and 18 sites in Idaho. Acoustic characteristics across the sites included naturally quiet 'control' sites, 'positive control' sites that were adjacent to the ocean or a whitewater river and thus were naturally noisy, 'phantom' playback sites that were exposed to continuous broadcast of low-frequency ocean surf or whitewater noise, and 'shifted' playback sites with continuous broadcast of ocean surf or whitewater noise shifted up in frequency. We predicted that spectral and temporal song structure would generally correlate with background sound amplitude and that signal features would differ across site types based on the spectral profile of the acoustic environment. We found that the ways in which song structure varied with background acoustics were quite variable from species to species. For instance, in Idaho both the frequency bandwidth and duration of lazuli bunting (Passerina amoena) and song sparrow (Melospiza melodia) songs decreased with elevated background noise, but these song features were unrelated to background noise in the warbling vireo (Vireo gilvus), which tended to increase both the minimum and maximum frequency of songs with background noise amplitude. In California, the bandwidth of the trill of white-crowned sparrow (Zonotrichia leucophrys) song decreased with background noise amplitude, matching results of previous studies involving both natural and anthropogenic noise. In contrast, wrentit (Chamaea fasciata) song bandwidth was positively related to the amplitude of background noise. Although responses were quite heterogeneous, song features of all six species varied with amplitude and/or frequency of background noise. Collectively, these results provide strong evidence that natural soundscapes have long influenced vocal behavior. More broadly, the evolved behavioral responses to the long-standing challenges presented by natural sources of noise likely explain the many responses observed for species communicating in difficult signal conditions presented by human-made noise.