Research Paper

INTRODUCTION

Noise pollution effects

Noise pollution is defined as the diffusion of noise in natural or human-made environments occurring in ways detrimental to the physiological or psychological health of humans and other creatures. Anthropogenic noise comes from many sources including power operations, transportation (land, air, and water vehicles), construction and machinery works, and human activities (Schomer et al. 2001, Haviland-Howell et al. 2007). Like other forms of pollution, noise sources can be point (e.g., airports, power plants, construction sites) or non-point (traffic noises, ground, and air), and noise can be chronic (constant; freeways, power lines) or episodic and acute (fireworks, sonic booms, mechanical tools). Because of human infrastructure expansion, anthropogenic noise is increasingly extensive and its intensity in urban and even natural/rural lands frequently exceeds government standards limiting noise pollution (Buxton et al. 2017, Yuan et al. 2019, Farooqi et al. 2020). Some effects of noise pollution on people include interference with communication, insomnia followed by decreased work efficiency, and even deafness and mental breakdown can occur (Singh and Davar 2004). Known physiological and mental health declines occur when humans live and work in noisy environments (Cohen and Weinstein 1981, Ahmadi Kanrash et al. 2019). Similarly, detections of negative effects of noise on wildlife activity, community integrity, population viability, individual health, and communication, in both urban and natural areas, are rising fast (McLure et al. 2013, Basner et al. 2015, Halfwerk et al. 2016, Shannon et al. 2016, Buxton et al. 2017, Ciach and Fröhlich 2017, Madadi et al. 2017, Injaian et al. 2018, Aulsebrook 2020). Understanding how animals survive and operate in noise polluted environments is important to continue improving wildlife conservation and education about noise effects (Miller 1974, Fardell et al. 2022).

Consequences of anthropogenic noise on ecosystems can include altered forest soundscape patterns (Munro et al. 2018), changes in the structure and function of wildlife communities (Chen and Koprowski 2015, Brischoux et al. 2017, Blom et al. 2019), interference with animal communication (Rossi-Santos 2015, Grade and Sieving 2016), as well as reductions in reproductive success (Lagardère 1982, Siemers and Schaub 2011, Luo et al. 2015, Senzaki et al. 2016, Ru et al. 2018, Blom et al. 2019, Elmer et al. 2021). Proximate effects of noise pollution include hormonal signatures of stress in adults and juveniles (Kleist et al. 2018), reduced clutch size and juvenile growth rates, and shortened telomeres in offspring (Salmon et al. 2016, Dorado-Correa et al. 2018, Injaian et al. 2018, Zollinger et al. 2019). The latter suggests chronic noise exposure may shorten attainable lifespan (Monaghan and Haussmann 2006). Grade and Sieving (2016) identified that 500 m on either side of interstate highways in natural areas show significant negative effects of noise on communication effectiveness. Along with urbanized areas and additional episodic noise from oil and gas development, military training (aircraft sonic booms, explosions, gunfire: Wakstein 2019, Kuehne and Olden 2020) and recreational activities outside of urban areas (concerts, off-road vehicles, fireworks: Taylor 2006, Bernat-Ponce et al. 2021) North American land areas affected by unhealthful noise levels are extensive (Hammer et al. 2014). Given mounting evidence of noise-caused reproductive declines in wildlife, and failure by some species to avoid these areas (e.g., Plummer et al. 2021), this raises the possibility that large areas exposed to diverse human noise may create population sinks (Schlaepfer et al. 2002, Rodewald et al. 2011, Francis and Barber 2013, Robertson et al. 2013, Swaddle et al. 2015).

Most studies addressing wildlife responses to noise at community, population, and individual levels concern chronic or continuously produced noises (e.g., Bayne et al. 2008, McClure et al. 2017) but increasingly the focus is on episodic and intermittent noises (de Jong et al. 2020, Hahn and Yosef 2021, Rosa and Koper 2022). Presumably, mobile species such as birds can choose to settle on quieter ends of noise gradients and community structure does indeed vary with distance to noise sources (McClure et al. 2017). Whereas some species like bluebirds (Sialia spp) appear indiscriminate regarding nest site choice along chronic noise gradients (Ferraro et al. 2020, Plummer et al. 2021), other species prefer quieter sites (Injaian et al. 2018). In species choosing to nest in chronically noisy areas, different traits (e.g., personality) may be more common in noisy habitats because of assortative habitat selection, adaptation, and/or transgenerational effects (Caizergues et al. 2022, Caspi et al. 2022). Episodic noise impacts on wildlife responses are less well understood than chronic highway noise (Francis and Barber 2013, Kok et al. 2023) but are also abundant in human landscapes, and less predictable in time and space. For humans, intermittent noises are highly disruptive of sleep, concentration, emotional well-being, and physiological stress responses, typically more so than constant noise of similar loudness (Kawada 2011). In general, chronic, and episodic noise disturbances appear to have different effects on animals, including some observable behaviors (Shannon et al. 2016, Berkhout et al. 2023). For example, chronic noise increases vigilance (Meillère et al. 2015) but may be accepted during a nesting cycle, whereas intermittent loud noises may cause serious disruptions to nesting (i.e., nest abandonment: Ortega 2012; reduced incubation time: Williams et al. 2021).

Is female incubation behavior linked to nest failure?

A common mechanism underlying reproductive failure in birds is disruption of incubation behaviors, but this has generally been overlooked when explaining noise-related failures (e.g., Schroeder et al. 2012). In most songbirds, females are the primary incubating parent, alternating between sitting on the eggs all night and most of the day with frequent brief foraging forays during daylight hours (Matysioková and Remeš 2014). Males can significantly reduce the burden of maintaining high incubation temperatures by trading off incubation bouts with females or by feeding the female and reducing the foraging opportunity costs of incubation (Reid et al. 2002). Indeed, females incubating alone can achieve 60–80% constancy (% of time eggs are being warmed) whereas when males of a species also feed the female frequently or solely (e.g., Kemp 1969) or share incubation duties, constancy can be higher (Skutch 1962). Maintaining optimal egg temperatures for sufficient time is key to successful hatching in birds (French 2009). Embryos of most passerine species stop developing at or below physiological zero (approx. = 26 °C: Webb 1987, Haftorn 1988, Cooper et al. 2005) and develop best between 35 °C and 39 °C. Extended time periods at or below physiological zero or outside of the optimal zone can halt or slow development, cause egg mortality, and reduce hatching success (Wang and Beissinger 2009). Even in species where females incubate alone, effective behavioral responses can buffer eggs from moderate and predictable environmental fluctuations, such as nightly cooling and inclement weather, with fine adjustments in time spent sitting quietly on eggs (Williams and DeLeon 2020). However, the physiological limits of females, which must balance food intake with sitting on eggs (Jones 1987) can be exceeded. For example, in the earliest and coldest days of temperate breeding seasons, optimal egg temperatures are regularly unattainable by lone female incubators (Ardia et al. 2009) and unseasonable cold snaps can significantly exceed females’ foraging-incubation trade-offs, and mass nest failures ensue (Moreno et al. 2015, Glądalski et al. 2020). Thus, incubation behaviors within the nest are key to hatching and reproductive success. Parents must continually adjust to changing environmental conditions (Conway and Martin 2000, Coe et al. 2015) to maintain egg development given constraints of the incubator (age, condition, whether mate helps or not; Skutch 1962, Jones 1987). If noise disrupts typical behavioral states of incubators, then additional challenges are likely to arise.

Research design

Given that noise disrupts the sleep of incubating females in nest boxes (Grunst et al. 2021) and that incubation behavior of females can influence reproductive success (Nilsson and Smith 1988) and other key responses to noise (e.g., telomere length; Stier et al. 2020, Hope et al. 2023), we propose that noise-related hatching failure may be closely related to disruption of female incubation behavior. Indeed, Williams et al. (2021) used experimental addition of noises at quiet nest boxes and found a connection between the addition of noise and reduced incubation time and hatching success. Greater incubation constancy is known to produce higher incubation temperatures and hatching success (Deeming 2002, Bertin et al. 2018, Wada et al. 2018), but constancy has several dimensions that can vary with disturbances at the nest. An incubation bout is the total time it takes a female to first settle on eggs and incubate, then leave to forage, and then return to settle on the eggs again, and includes both the on and off egg periods. Females can alter the length of each on and off period, the entire incubation bout length, and the number of bouts per day (Ward 1990). When females are distracted from incubation when they should be deeply settled on eggs, they may stand up on the nest rim without leaving the nest, stand and look outside (called head-poking in cavity-nesting birds; Hatakeyama 2017), or they may preen excessively or snap at insects (Maurelli 2022). And if they rise up and down frequently, they will also turn eggs more frequently (Coe et al. 2015). More frequent egg-turning enhances embryo development when temperatures are controlled within the optimal zone (as in incubators; Oliveira et al. 2020), but in natural nests with restless females unable to maintain optimal egg temperatures, frequent egg-turning could contribute to faster cooling.

In this study we tested for the effects of chronic and acute noise on both female incubation behavior and reproductive success in Eastern Bluebirds (Sialia sialis). We utilized a gradient of chronic traffic noise (from approximately < 60 to > 75 dBA) and experimentally added intermittent noise disturbances to nest boxes where females were incubating completed clutches of eggs in a combined comparative-experimental design. We assessed whether background traffic noise and added, intermittent construction noise affected (a) hatching success and (b) female incubation behaviors (constancy and others, see below). Over one breeding season, we monitored bluebird nest success in an urban population using nest boxes, and quantified incubation behaviors of female bluebirds using temperature signatures obtained from iButton data loggers placed in nests.

METHODS

Study system

The Eastern bluebird (EABL) breeds throughout the Eastern U.S. and prefers nesting in open woodland with sparse ground vegetation (Berger et al. 2001). EABL nests in natural cavities and artificial nesting boxes. After nest building in early spring females typically lay one egg per day (clutch size ranges from three to seven eggs; four or five is most common) and begin incubating the eggs just before or after the final egg is laid (Cooper et al. 2005). Incubation lasts 11–19 days (14 days is typical; Buxton et al. 2017). In north-central Florida, where this study occurred, females are the principal incubators and mean incubation bout lengths (on bouts) are 15.6 min (± 8.7 SD) interspersed with mean off bouts lasting 10.1 min (± 9.7 SD; Malone et al. 2017).

The University of Florida campus (in Gainesville, Alachua County, Florida USA; 29.6465° N, 82.3533° W) and its surrounding areas are becoming increasingly urbanized. As an agricultural land grant campus (est. 1862; LaCharite 2016), large areas suitable for EABL nesting still occurred in the year of the study and included a wide range of land uses including livestock and crop fields, open greenspace, high-rise buildings, hardscape, heavily trafficked areas (foot and vehicular), and a range of noise levels. Average annual temperatures range from 26.7 °C (high) to 14.4 °C (low) with annual rainfall at 119.4 cm.

Nest box placement, monitoring, and hatching success

In January and February 2019, we placed 98 Gilbertson style bluebird boxes in selected areas to attract breeding pairs. This nest box is made of a cylindrical PVC chamber with a drain in a wood bottom and covered by a wood roof that is attached to a stainless-steel pipe (1.27 cm diameter, 1.2 m tall) supported by a 1.5 m length of steel rebar driven halfway into the ground. The Gilbertson nest box was previously tested in this area and performs well for EABL (see Perreau and Sieving 2015 for photos of the box type; Malone et al. 2017). We systematically visited every area of campus with suitable space for EABL nesting with at least a 100 m diameter grassy area at or within 100 m of a suitable nest location. Moreover, all nest boxes were placed in the open without overhanging canopy or vegetation touching the nest pole (Kight and Swaddle 2015; see Plummer et al. 2021 for a map of nest boxes and their occupancy). Such sites were available close to and far from the noisiest traffic arteries surrounding campus (Florida State Routes 441 [E], 24 [S], 121 [W] and 26 [N]). Boxes ended up in a variety of sites, near roads, buildings, on a golf course, in pastures, and open manicured lawns. The spacing between any two nests was at least 100 m to minimize inter-pair aggression.

Student volunteers checked and recorded the status of nest boxes at least once a week from February through early July. If volunteers found any trace of nesting material (dried grass or pine needles), they would increase the frequency of monitoring and identify the nest as potentially active. After the first eggs were laid in a box, a nest was monitored daily until the egg number did not change for 2 days, at which point the nest was randomly assigned as a playback treatment or control nest, and the necessary equipment was set up near the box. Nearby each nest (approx. 3–5 m away from the nest) we placed a second metal post with a time lapse camera (Wingscapes Birdcam Pro) aimed at the nest entrance to sample nest visits from dawn to dusk. At playback treatment nests, a speaker (OontZ Angle 3rd generation) plus an mp3 player (RUIZU X02) wrapped in plastic were also mounted on the pole using duct tape. After a female laid her last egg and before we implemented any playbacks, we placed an iButton (temperature logger; DS1921G-F5) in the bottom of each nest cup set to record nest temperature every 10 minutes. Loggers were secured using a piece of fishing line (glued to the logger and threaded outside the box and tied to the pole).

We retrieved mean ambient temperatures for each day of observation (for 10 AM and 6 PM) from the Florida Automated Weather Network (FAWN, https://fawn.ifas.ufl.edu/), using data from the closest station to campus (Gainesville Regional Airport). Ambient temperatures are typically strongly correlated with female incubation behaviors and hatching success in birds (Coe et al. 2015). Given that optimal incubation temperatures for avian embryos fall between 35.5 °C and 38.5 °C, females will experience a wide range of ambient temperatures well below this range early in the season and closer to this range later in the breeding season, and ambient temperatures can influence incubation behavior (Webb 1987, Simmonds et al. 2017). Therefore, a range of activity levels in incubating females was expected over the course of the study in response to widely varying ambient temperatures. For all nests that successfully reached hatching stage, we recorded the number of eggs at clutch completion as well as the number of chicks that hatched at the end of the incubation stage.

Ambient noise and application of noise treatments

To classify nests into noisier and quieter classes, volunteers measured the ambient noise levels at all nest boxes using mobile phone software (the app, Decibel X). Four different ambient noise measurements were taken on different days throughout the season. For each measurement taken, the phone was placed on top of the nest box and the app was opened to record for at least 3 minutes to derive the average ambient noise levels (decibels, A scale) per minute. We summarized the mean dBA per nest box taking the mean of 3 minutes per day averaged over 4 days. The grand mean ambient traffic noise was 70 dBA, and we used this value to identify two classes of nest boxes: quieter (< 70 dBA) versus noisier (≥ 70 dBA). Range of noise overall was 55–88 dBA. We note that smartphone sound apps do not allow precise estimation of sound levels (Cheng 2020). However, they do produce values highly correlated with high quality sound meters (Crossley et al. 2021), typically overestimating true sound pressure levels. Therefore, although we note that the actual sound levels reported here are likely to be overestimated, we simply needed them to identify two classes of nests: noisier and quieter (Table 1).

As active nests were identified, we attempted to assign playback to a comparable number of clutches in each category of background noise (noisy and quiet), but this was not entirely under our control. Different numbers of playback days were dictated by the total days of active incubation we could monitor at each nest. This varied depending on how soon we could begin playbacks after a clutch was complete and on how many days total a nest was incubated. The former was influenced by when volunteers spotted an active nest and when equipment could be freed from another nest to begin the playback period. Actual playback treatments varied from 0 (control) up to 4 (1–4) treatment days of construction noise playback (Table 1). Each clutch receiving playback treatment was exposed to at least one full day of playback (up to four days maximum) during the incubation period. If a nest received more than one day of playback, nests received noise on one day, then the speakers were silenced for two days and then noise was broadcast again for a day (etc.). In this way, background noise (noisy or quiet) was a categorical variable, but days of playback (0–4) was treated as a quantitative variable because the number of nests receiving 0 or 1 or 2 (etc.) playbacks varied opportunistically.

On playback days, construction noise was broadcast from the speaker and mp3 player mounted on the poles set near the nest boxes. Construction noises included jackhammers, drills, hammering, excavators, and other noises associated with building construction. Recordings were constructed to have noisy and quiet sections to mimic the tempo of a construction site with bursts of noise either every few minutes (5–10) or every 30 minutes. All recordings included a quiet period in the middle of the day (2 hours) to represent a lunch break. Sections with noise included a sequence of all five types of construction noises but on different recording copies the sequence of each type of sound was randomized. We made five different recordings with the same timing (noisy and quiet bouts) but with different sequences of the sound types (sound effects obtained from https://www.zapsplat.com). Sound levels of all playbacks were normalized to 90% of the maximum pressure levels, and playback volumes at each nest were standardized to ~60 dBA at the nest hole (using a professional sound meter; Extech 407730), and the total length of each playback file was 10 hours. On treatment days, the recordings were started manually between 8 and 10 AM, and ended between 6 and 8 PM such that each nest received 8 hours of exposure.

Characterizing female incubation behavior

iButtons recorded the temperature inside the nest cup every 10 minutes, and we downloaded all data for each nest recorded during daylight hours, and for each 10-minute period we retrieved the average nest temperature. Using the detailed iButton data, we created nest temperature plots for the hours between 10 AM to 6 PM (all nests shared this time span: Fig. 1). Using the iButton temperature plots created for each active nest for each day of incubation, we counted the number of (1) incubation bouts, the (2) mean bout length over each day, the number of (3) small drops in temperature (< 2 °C; fluctuations), and the (4) total minutes during each day that nests were increasing in temperature (warming minutes) as measures expressing the range of female incubation behaviors. Normal incubation bouts (attentive periods) are characterized by regular cycles of significant box temperature increases and decreases as females first settle on eggs, and then leave the nest to forage. We defined a bout as a warming temperature rise of more than 2 °C, and bout length was equal to the total amount of time between the start of a bout and when the temperature peaked. If a nest temperature rise did not exceed 2 °C before falling again, we identified the pattern as a small fluctuation that we interpreted as restless or agitated incubation behavior (blue line in Fig. 1). Total warming minutes per day was calculated as the total time, summarized over the 8 hours between 10 AM and 6 PM, over which the nest temperature was rising because of female warming efforts during regular bouts. The latter represents incubation constancy (Skutch 1962).

We confirmed that the large and small temperature changes (bouts and fluctuations) recorded by iButtons represented female on- and off-nest behaviors by using the time lapse photographs taken at nest boxes (Audubon BirdCam Pro). We counted the number of observed exits and entrances made by females as well as the number of head-pokes (when a female stands on the nest and pushes only her head out of the nest entrance) captured in photos for the hours between 10 AM and 6 PM for a few of the nests with the highest and lowest number of temperature changes (bouts and fluctuations). We confirmed that on days with high numbers of bouts and/or fluctuations, we also observed the highest number of exits, entrances, and head-pokes in the photo data (authors’ unpublished data). Time lapse cannot capture every entry, exit, and head-poke by females, but the numbers were highly correlated with the number of daily up and down changes in nest temperatures.

Data analysis

All analyses (five in total) were run in STATA (version 17, BE) as generalized linear mixed models with four terms, one random effect (Table 1), maximum likelihood estimation, and identity link function. Predictor variables included metrics representing noise and daily mean temperatures. We used two separate variables to represent the combination of construction noise playback with ambient noise at the nest. Ambient noise was implemented as a categorical variable identifying either quiet or noisy boxes, depending on whether the mean noise measured at the boxes was less or greater than 70 dBA. Additions of construction noise playbacks were coded as a count variable indicating the number of total days that a single nest received construction noise playback (0–4, 8-hr days) during incubation. All ambient temperature measures obtained from FAWN (hourly and daily mean lows, means and maximums) were strongly correlated with day of the year (counting 1 Jan as the first day). Therefore, in analyses, we used the day of the year when each nest was first identified as a treatment or control nest for playback treatment to represent the seasonal progression in temperature (days of the year marking the beginning of nest observations varied from day 68 to 191). In models, all three of these factors were included as main effects along with one 2-way interaction between ambient noise (categorical) and number of days of construction noise playback for a total of four terms. Because some nest boxes were re-used a maximum of two times (probably by the same pairs) we used the box ID number as a random effect. If a model term produced a p-value < 0.10, we also ran marginal contrasts to determine if any were significant over the predicted range of outcomes. We present significant effects contrasts in graphs.

Response variables included hatching success, and four measures representing female incubation behaviors extracted from iButton data: mean number of bouts, mean bout length, number of fluctuations, and mean total number of warming minutes per day (assessed between the hours of 10 AM to 6 PM in all nests). For hatching success, we divided the total number of chicks that hatched and lived for at least a week by the number of eggs laid to get the proportion of eggs that led to viable nestling. Using an arcsin(sqrt(# hatchlings/# eggs laid)) angular transformation to normalize the data, the proportion of eggs hatched was extended from 0–1.0 to 0–1.57 and was normally distributed (Shapiro-Wilk W test for normality was ns). Unlike hatching success, behavioral measures inferred from iButton data were measured multiple times per day and over 1–11 days during incubation periods. To construct a very conservative analysis that is unlikely to produce a false positive (finding an effect of noise when it does not occur) with our relatively small data set we calculated mean values for fluctuations, bouts, warming minutes, and bout lengths across all days of observation at a nest and obtained grand means of these variables (one line of data for each nest). For example, given a nest that received 3 days of construction noise playback, we also had at least 4 days of data when no playback occurred but that came after playback days. If the effects of playback accumulate, we thought it best to average the behaviors on playback and non-playback days to get the most balanced picture of incubation constancy over the course of each female’s incubation efforts. Each measure was then tested for normality using the Shapiro-Wilk W test. Mean bout and fluctuation number were both normally distributed according to this test but mean daily warming minutes and bout lengths were skewed in minor ways. To overcome the skew, we applied a Box-Cox transformation, which solved the issues with both. We did not utilize information theoretic approaches and apply model comparisons in analysis because every term included expressed hypothesized causal factors that we designed this study to address. The data set used in these analyses is permanently archived here (https://original-ufdc.uflib.ufl.edu/IR00012055/00001). Our research protocol was (1) reviewed and approved by University of Florida’s Institutional Animal Care and Use Committee under protocol # 201910544, and (2) further permitted by Bird Banding master permit # 022541 (issued to KES) under USGS Federal Regulation number 16 U.S.C. 703-712 (Migratory Bird Treaty Act).

RESULTS

Among the 98 nest boxes placed, 37 were occupied by nesting birds and those boxes hosted 66 total nesting attempts, including primarily EABL, but also other species. Considering only Eastern Bluebird nests, the first-egg date was 26 February 2019, and 23 nests were established in a first round of reproduction (all of these fledged by late May). The first re-nest in a box that had already fledged chicks from the first nesting was started 18 April, a date we considered to mark the beginning of the second nesting period more generally. Twenty nests were established in the second round of reproduction, and five nest boxes hosted third clutches initiated in late July. We could not include third clutches in analysis because two failed early and we could not fully apply noise treatments with the remainder. Of the 43 EABL nests in first and second nestings that were completed by June, two were parasitized by a Brown-headed Cowbird (Molothrus ater) and another failed for unknown reasons prior to hatching. Figure 2A displays the raw hatching success according to whether the nests were in high or low ambient noise areas and by how many days of construction noise playback they received. N = 40 nests total used in analyses.

Generalized linear models (all results are in Table 2) suggest that hatching success of nests placed in quieter areas (< 70 dBA ambient noise) was significantly greater than for noisy nest boxes (Fig. 2B) and the addition of construction noise playback to quieter nests caused significant reductions in hatching success, but not for already noisy nests (Fig. 3A). Day of the year was not a significant factor in determining hatching success. Regarding the effects of noise on behavioral measures, only fluctuation number appeared to be responsive: mean fluctuation number peaked in nests placed in quiet areas that also received 3 or 4 days of construction noise playback (Fig. 3B). As day of the year increased and temperatures warmed, both fluctuations and mean bout lengths increased significantly and bout number per day decreased significantly (Table 2). Total warming minutes did not vary by sound treatments or by day of the year.

DISCUSSION

Both chronic and intermittent noise reduced hatching success

Reduced hatching success was associated with both types of noise, chronic daytime traffic noise from major state roads that mark the perimeter of the University of Florida campus (441, 20, 21, and 26; Plummer et al. 2021) and recorded construction noises broadcast near nests. There was no marked indication that birds preferred quieter nest sites in this study (Plummer et al. 2021), though bluebirds did choose nest sites with more open grass, as is typical for this species (e.g., Munro and Rounds 1985). More data are needed to see if lack of preference for quiet habitat is a mark of urban adapting species that either already have noise-adapted traits (e.g., parids can adjust social behavior in noisy environs; Owens et al. 2012) or that will eventually develop them (Bowles 1995). Current evidence suggests that negative effects of chronic noise on birds occur beginning around 50 dBA (communication loss; Grade and Sieving 2016) and where reproductive success declines, levels are typically higher (Shannon et al. 2016). Studies that compare incubation versus nestling stage effects of chronic noise find that hatching failure or reduced clutch size drives the reduction in reproductive output, but post-hatching effects on fledgling growth also occur (lab settings: Halfwerk et al. 2011, Potvin and MacDougall-Shackleton 2015; natural settings: Injaian et al. 2018, Kleist et al. 2018).

Intermittent noise altered female behavior and hatching success

One robust response we detected was that increases in the number of days of construction noise playback significantly reduced hatching success but only in the quietest of nests (Figs. 2A, 3A). We had less data to assess construction noise additions to noisy nests (> 70 dBA), but patterns suggest little or even an opposite effect of playback in this group (Figs. 2A, 3B). In our experience, the best explanation for playback effects is that intermittent noise causes frequent, periodic arousal of females when resting and/or sleeping. Nocturnal video data (Maurelli 2022) suggest that when females are disturbed, they may stand up or shift and turn eggs more frequently than when undisturbed. Others note that awakened incubating adults are more likely to move, engaging in preening, egg-turning, or standing on the nest cup off the eggs (Eberhardt 1988, Winkler 2016, Grunst et al. 2021). All such activities are natural, but if artificially increased in frequency, then this could cause more numerous small drops in nest cup temperature (e.g., fluctuations; Fig. 3B) and hatching success (Fig. 3A) that we detected with more construction playback days per quiet nest. We note that additions of construction noise here were only during daylight hours, when traffic volume and noise levels are highest. Yet the behavioral effects we detected in daytime were correlated with hatching success suggesting that birds respond to the highest levels of noise stress experienced during 24-hour incubation cycles, and that reactions to daytime noise might carry over into nighttime incubation behaviors, even considering that traffic drops off in the wee morning hours on most roadways. Daytime winter temperatures in central Florida can be quite mild, but night-time temperatures can be within 5–10 degrees of freezing, especially during first nestings; deviations from steady incubation during the night hours likely also occurred in order to depress hatching success. As others have found, adding intermittent to chronic noise did not decrease reproductive success the most (e.g., Blom et al. 2019). Some have detected a down-regulation of corticosteroid secretions in response to chronic high traffic noise (Rich and Romero 2005, Kleist et al. 2018). Therefore, we can speculate that addition of acute noise stressors might not elicit additional behavioral stress responses (e.g., Katz et al. 1981) connected to hatching failure, but we cannot explain this pattern without additional data (e.g., time synched video, temperature, and noise measures taken continuously day and night inside the nest day).

CONCLUSIONS

Based on our experiences with low-tech nest box monitoring procedures, we suggest more varied and precise measures will be needed to disentangle the links between noise and decreased reproduction. For example, what specific female behaviors caused the small fluctuations? Video recordings time-synched with temperature readings would have helped understand how temperature fluctuations in the daytime inside the nest were related to hatching success. Nest box technologies are rapidly developing, however, and we suggest future studies consider smart nest boxes with remote sensing tech to obtain and analyze multiple data streams (including sensors for audio, video, temperature, heart rate, etc.; e.g., Caorsi et al. 2019, Kubizňák et al. 2019, Grunst et al. 2020, Williams and DeLeon 2020, Grunst et al. 2021). Remote sensing would eliminate disturbance from nest checks that can be a problem during sensitive incubation periods, and cause abandonment. Smart tech can assess individual level response variation within and between nesting families, another critical factor to address in this type of research (yet often ignored; Harding et al. 2019). Individual-specific measures of personality, condition, age, and stress response would also allow distinctions between direct and indirect effects of noise and other disturbances on both parents and offspring (Heathcote 2019, Injaian et al. 2019).

We found that (a) increasing exposure (days) to intermittent construction noises at the nest boxes caused restlessness in female incubating bluebirds and this, in turn, may have (b) caused reductions in hatching success and that (c) chronic high traffic noise greatly reduced hatching success. Intermittent noise can disrupt rest, concentration, coordination, and task completion and causes increases activity in vertebrates (Brackbill 1970, Szalma and Hancock 2011, Nassiri et al. 2021). We note again that our analysis was conservative, taking the mean number of fluctuations / per day averaged over all days of incubation observed at a nest. For those nests receiving playback, this includes non-playback and playback days, suggesting that behavioral effects of added days of playback are cumulative, and likely to be affecting restlessness during night-time incubation as well as days without playback. Chronic traffic noise was associated with depressed hatching success in our study, possibly due to direct effects on embryos (Potvin and MacDougall-Shackleton 2015) because we did not register any altered female behaviors in chronic noise. Addition of acute noise (playbacks) to chronically noisy nests had little to no detectable effect on either hatching or female incubation behavior; this aligns with other tests for combined effects of chronic and intermittent stressors in vertebrates (Katz et al. 1981, Rich and Romero 2005).

We detected one clear sign that adult incubation behavior can mediate the impacts of noise on hatching success; female restlessness increased, and hatching success declined, with increasing incidence of intermittent sounds broadcast near quiet nests. Females exposed to high levels of chronic noise did not respond to playback in the same way. Implications for conservation are two-fold. (1) Bluebird boxes should be placed as far from busy roads as possible because they do not necessarily choose available quiet nest boxes (Plummer et al. 2021) despite responding poorly to traffic noise. (2) Once placed, nest boxes should be protected from noisy incursions near the nest.

LITERATURE CITED

Ahmadi Kanrash, F., I. Alimohammad, J. Abolaghasemi, and K. Rahmani. 2019. A study of mental and physiological effects of chronic exposure to noise in an automotive industry. Iranian Journal of Ergonomics 7(1):54-62. https://doi.org/10.30699/jergon.7.1.54

Ardia, D. R., J. H. Pérez, E. K. Chad, M. A. Voss, and E. D. Clotfelter. 2009. Temperature and life history: experimental heating leads female Tree Swallows to modulate egg temperature and incubation behaviour. Journal of Animal Ecology 78:4-13. https://doi.org/10.1111/j.1365-2656.2008.01453.x

Aulsebrook, L. C., M. G. Bertram, J. M. Martin, A. E. Aulsebrook, T. Brodin, J. P. Evans, M. D. Hall, M. K. O’Bryan, A. J. Pask, C. R. Tyler, and B. B. Wong. 2020. Reproduction in a polluted world: implications for wildlife. Reproduction 160(2):R13-R23. https://doi.org/10.1530/REP-20-0154

Basner, M., M. Brink, A. Bristow, Y. De Kluizenaar, L. Finegold, J. Hong, S. A. Janssen, R. Klaeboe, T., Leroux, A. Liebl, and T. Matsui. 2015. ICBEN review of research on the biological effects of noise 2011-2014. Noise & Health 17(75):57. https://doi.org/10.4103/1463-1741.153373

Bayne, E. M., L. Habib, and S. Boutin. 2008. Impacts of chronic anthropogenic noise from energy‐sector activity on abundance of songbirds in the boreal forest. Conservation Biology 22(5):1186-1193. https://doi.org/10.1111/j.1523-1739.2008.00973.x

Berger, C., K. Kridler, and J. L. Griggs. 2001. The bluebird monitor’s guide. HarperCollins, New York, New York, USA.

Berkhout, B. W., A. Budria, D. W. Thieltges, and H. Slabbekoorn. 2023. Anthropogenic noise pollution and wildlife diseases. Trends in Parasitology 39:181-190. https://doi.org/10.1016/j.pt.2022.12.002

Bernat-Ponce, E., J. A. Gil-Delgado, and G. M. López-Iborra. 2021. Recreational noise pollution of traditional festivals reduces the juvenile productivity of an avian urban bioindicator. Environmental Pollution 286:117247. https://doi.org/10.1016/j.envpol.2021.117247

Bertin, A., L. Calandreau, M. Meurisse, M. Georgelin, R. Palme, S. Lumineau, C. Houdelier, A. S. Darmaillacq, L. Dickel, V. Colson, F. Cornilleau, C. Rat, J. Delaveau, and C. Arnould. 2018. Incubation temperature affects the expression of young precocial birds’ fear-related behaviours and neuroendocrine correlates. Scientific Reports 8:1857. https://doi.org/10.1038/s41598-018-20319-y

Blom, E. L., C. Kvarnemo, I. Dekhla, S. Schöld, M. H. Andersson, O. Svensson, and M. C. P. Amorim. 2019. Continuous but not intermittent noise has a negative impact on mating success in a marine fish with paternal care. Scientific Reports 9:5494. https://doi.org/10.1038/s41598-019-41786-x

Bowles, A. E. 1995. Responses of wildlife to noise. Pages 109-156 in R. L. Knight and K. J. Gutzwiller, editors. Wildlife and recreationists: coexistence through management and research. Island, Washington, D.C., USA.

Brackbill, Y. 1970. Acoustic variation and arousal level in infants. Psychophysiology 6(5):517-526. https://doi.org/10.1111/j.1469-8986.1970.tb02241.x

Brischoux, F., A. Meillère, A. Dupoué, O. Lourdais, and F. Angelier. 2017. Traffic noise decreases nestlings’ metabolic rates in an urban exploiter. Journal of Avian Biology 48(7):905-909. https://doi.org/10.1111/jav.01139

Buxton, R. T., M. F. McKenna, D. Mennitt, K. Fristrup, K. Crooks, L. Angeloni, and G. Wittemyer. 2017. Noise pollution is pervasive in U.S. protected areas. Science 356(6337):531-533. https://doi.org/10.1126/science.aah4783

Caizergues, A. E., A. Grégoire, R. Choquet, S. Perret, and A. Charmantier. 2022. Are behaviour and stress‐related phenotypes in urban birds adaptive? Journal of Animal Ecology 91(8):1627-1641. https://doi.org/10.1111/1365-2656.13740

Caorsi, V., P. Sprau, S. A. Zollinger, and H. Brumm. 2019. Nocturnal resting behaviour in urban great tits and its relation to anthropogenic disturbance and microclimate. Behavioral Ecology and Sociobiology 73:19. https://doi.org/10.1007/s00265-018-2624-1

Caspi, T., J. R. Johnson, M. R. Lambert, C. J. Schell, and A. Sih. 2022. Behavioral plasticity can facilitate evolution in urban environments. Trends in Ecology & Evolution 37(12):1092-1103. https://doi.org/10.1016/j.tree.2022.08.002

Chen, H. L., and J. L. Koprowski. 2015. Animal occurrence and space use change in the landscape of anthropogenic noise. Biological Conservation 192:315-322. https://doi.org/10.1016/j.biocon.2015.10.003

Cheng, T. 2020. Evaluation of smartphone-based sound level meters. Pages 1-8 in 2020 IEEE Integrated STEM Education Conference. Institute of Electrical and Electronics Engineers, Piscataway, New Jersey, USA. https://doi.org/10.1109/ISEC49744.2020.9280746

Ciach, M., and A. Fröhlich. 2017. Habitat type, food resources, noise and light pollution explain the species composition, abundance and stability of a winter bird assemblage in an urban environment. Urban Ecosystems 20:547-559. https://doi.org/10.1007/s11252-016-0613-6

Coe, B. H., M. L. Beck, S. Y. Chin, C. M. Jachowski, and W. A. Hopkins. 2015. Local variation in weather conditions influences incubation behavior and temperature in a passerine bird. Journal of Avian Biology 46(4):385-394. https://doi.org/10.1111/jav.00581

Cohen, S., and N. Weinstein. 1981. Nonauditory effects of noise on behavior and health. Journal of Social Issues 37(1):36-70. https://doi.org/10.1111/j.1540-4560.1981.tb01057.x

Conway, C. J., and T. E. Martin. 2000. Effects of ambient temperature on avian incubation behavior. Behavioral Ecology 11(2):178-188. https://doi.org/10.1093/beheco/11.2.178

Cooper, C. B., W. M. Hochachka, G. Butcher, and A. A. Dhondt. 2005. Seasonal and latitudinal trends in clutch size: thermal constraints during laying and incubation. Ecology 86(8):2018-2031. https://doi.org/10.1890/03-8028

Crossley, E., T. Biggs, P. Brown, and T. Singh. 2021. The accuracy of iPhone applications to monitor environmental noise levels. Laryngoscope 131(1):E59-E62. https://doi.org/10.1002/lary.28590

de Jong, K., T. N. Forland, M. C. P. Amorim, G. Rieucau, H. Slabbekoorn, and L. D. Sivle. 2020. Predicting the effects of anthropogenic noise on fish reproduction. Reviews in Fish Biology and Fisheries 30:245-268. https://doi.org/10.1007/s11160-020-09598-9

Deeming, D. C. 2002. Avian incubation: behaviour, environment and evolution. Oxford University Press, Oxford, UK. https://doi.org/10.1093/oso/9780198508106.001.0001

Dorado-Correa, A. M., S. A. Zollinger, B. Heidinger, and H. Brumm. 2018. Timing matters: traffic noise accelerates telomere loss rate differently across developmental stages. Frontiers in Zoology 15:29. https://doi.org/10.1186/s12983-018-0275-8

Eberhardt, J. L. 1988. The influence of road traffic noise on sleep. Journal of Sound and Vibration 127(3):449-455. https://doi.org/10.1016/0022-460X(88)90369-0

Elmer, L. K., C. L. Madliger, D. T. Blumstein, C. K. Elvidge, E. Fernández-Juricic, A. Z.Horodysky, N. S. Johnson, L. P. McGuire, R. R. Swaisgood, and S. J. Cooke. 2021. Exploiting common senses: sensory ecology meets wildlife conservation and management. Conservation Physiology 9(1):coab002. https://doi.org/10.1093/conphys/coab002

Fardell, L. L., C. R. Pavey, and C. R. Dickman. 2022. Backyard biomes: is anyone there? Improving public awareness of urban wildlife activity. Diversity 14(4):263. https://doi.org/10.3390/d14040263

Farooqi, Z. U. R., M. Sabir, J. Latif, Z. Aslam, H. R. Ahmad, I. Ahmad, M. Imran, and P. Ilić. 2020. Assessment of noise pollution and its effects on human health in industrial hub of Pakistan. Environmental Science and Pollution Research 27:2819-2828. https://doi.org/10.1007/s11356-019-07105-7

Ferraro, D. M., M. L. T. Le, and C. D. Francis. 2020. Combined effect of anthropogenic noise and artificial night lighting negatively affect Western Bluebird chick development. Condor 122(4):duaa037. https://doi.org/10.1093/condor/duaa037

Francis, C. D., and J. R. Barber. 2013. A framework for understanding noise impacts on wildlife: an urgent conservation priority. Frontiers in Ecology and the Environment 11(6):305-313. https://doi.org/10.1890/120183

French, N. A. 2009. The critical importance of incubation temperature. Avian Biology Research 2(1-2):55-59. https://doi.org/10.3184/175815509X431812

Glądalski, M., M. Bańbura, A. Kaliński, M. Markowski, J. Skwarska, J. Wawrzyniak, P. Zieliński, and J. Bańbura. 2020. Extreme temperature drop alters hatching delay, reproductive success, and physiological condition in great tits. International Journal of Biometeorology 64:623-629. https://doi.org/10.1007/s00484-019-01851-6

Grade, A. M., and K. E. Sieving. 2016. When the birds go unheard: highway noise disrupts information transfer between bird species. Biology Letters 12(4):20160113. https://doi.org/10.1098/rsbl.2016.0113

Grunst, A. S., M. L. Grunst, R. Pinxten, and M. Eens. 2020. Sources of individual variation in problem-solving performance in urban Great Tits (Parus major): exploring effects of metal pollution, urban disturbance and personality. Science of the Total Environment 749:141436. https://doi.org/10.1016/j.scitotenv.2020.141436

Grunst, M. L., A. S. Grunst, R. Pinxten, and M. Eens. 2021. Variable and consistent traffic noise negatively affect the sleep behavior of a free-living songbird. Science of the Total Environment 778:146338. https://doi.org/10.1016/j.scitotenv.2021.146338

Haftorn, S. 1988. Incubating female passerines do not let the egg temperature fall below the ‘physiological zero temperature’ during their absences from the nest. Ornis Scandinavica 19(2):97-110. https://doi.org/10.2307/3676458

Hahn, A., and R. Yosef. 2021. Reactions of breeding Common Swifts (Apus apus) to explosions. Journal of Vertebrate Biology 71(21060):21060-21061. https://doi.org/10.25225/JVB.21060

Halfwerk, W., L. J. Holleman, C. K. M. Lessells, and H. Slabbekoorn. 2011. Negative impact of traffic noise on avian reproductive success. Journal of Applied Ecology 48(1):210-219. https://doi.org/10.1111/j.1365-2664.2010.01914.x

Halfwerk, W., A. M. Lea, M. A. Guerra, R. A. Page, and M. J. Ryan. 2016. Vocal responses to noise reveal the presence of the Lombard effect in a frog. Behavioral Ecology 27(2):669-676. https://doi.org/10.1093/beheco/arv204

Hammer, M. S., T. K. Swinburn, and R. L. Neitzel. 2014. Environmental noise pollution in the United States: developing an effective public health response. Environmental Health Perspectives 122(2):115-119. https://doi.org/10.1289/ehp.1307272

Harding, H. R., T. A. Gordon, E. Eastcott, S. D. Simpson, and A. N. Radford. 2019. Causes and consequences of intraspecific variation in animal responses to anthropogenic noise. Behavioral Ecology 30(6):1501-1511. https://doi.org/10.1093/beheco/arz114

Hatakeyama, Y. 2017. Behavior during nesting of Varied Tit. Binos 24:1-14.

Haviland-Howell, G., A. S. Frankel, C. M. Powell, A. Bocconcelli, R. L. Herman, and L. S. Sayigh. 2007. Recreational boating traffic: a chronic source of anthropogenic noise in the Wilmington, North Carolina Intracoastal Waterway. Journal of the Acoustical Society of America 122(1):151-160. https://doi.org/10.1121/1.2717766

Heathcote, A. L. 2019. Effects of oil infrastructure and associated noise on the stress physiology, growth and development of an altricial grassland songbird nestling. Thesis. University of Manitoba, Winnipeg, Manitoba, Canada.

Hope, S. F., F. Angelier, C. Ribout, J. Groffen, R. A. Kennamer, and W. A. Hopkins. 2023. Warmer incubation temperatures and later lay‐orders lead to shorter telomere lengths in Wood Duck (Aix sponsa) ducklings. Journal of Experimental Zoology Part A: Ecological and Integrative Physiology 339(1):101-111. https://doi.org/10.1002/jez.2659

Injaian, A. S., L. Y. Poon, and G. L. Patricelli. 2018. Effects of experimental anthropogenic noise on avian settlement patterns and reproductive success. Behavioral Ecology 29(5):1181-1189. https://doi.org/10.1093/beheco/ary097

Injaian, A. S., P. L. Gonzalez-Gomez, C. C. Taff, A. K. Bird, A. D. Ziur, G. L. Patricelli, M. F. Haussmann, and J. C. Wingfield. 2019. Traffic noise exposure alters nestling physiology and telomere attrition through direct, but not maternal, effects in a free-living bird. General and Comparative Endocrinology 276:14-21. https://doi.org/10.1016/j.ygcen.2019.02.017

Jones, G. 1987. Time and energy constraints during incubation in free-living swallows (Hirundo rustica): an experimental study using precision electronic balances. Journal of Animal Ecology 56(1)229-245. https://doi.org/10.2307/4812

Katz, R. J., K. A. Roth, and B. J. Carroll. 1981. Acute and chronic stress effects on open field activity in the rat: implications for a model of depression. Neuroscience & Biobehavioral Reviews 5(2):247-251. https://doi.org/10.1016/0149-7634(81)90005-1

Kawada, T. 2011. Noise and health—sleep disturbance in adults. Journal of Occupational Health 53(6):413-416. https://doi.org/10.1539/joh.11-0071-RA

Kemp, A. C. 1969. Some observations on the sealed-in nesting method of hornbills (Family: Bucerotidae). Ostrich 40(S1):149-155. https://doi.org/10.1080/00306525.1969.9639117

Kight, C. R., and J. P. Swaddle. 2015. Eastern bluebirds alter their song in response to anthropogenic changes in the acoustic environment. Integrative and Comparative Biology 55(3):418-431. https://doi.org/10.1093/icb/icv070

Kleist, N. J., R. P. Guralnick, A. Cruz, C. A. Lowry, and C. D. Francis. 2018. Chronic anthropogenic noise disrupts glucocorticoid signaling and has multiple effects on fitness in an avian community. Proceedings of the National Academy of Sciences 115(4):E648-E657. https://doi.org/10.1073/pnas.1709200115

Kok, A., B. W. Berkhout, N. V. Carlson, N. P. Evans, N. Khan, D. A. Potvin, A. N. Radford, M. Sebire, Shafiei S. Sabet, G. Shannon, and C. A. Wascher. 2023. How chronic anthropogenic noise can affect wildlife communities. Frontiers in Ecology and Evolution 11:289. https://doi.org/10.3389/fevo.2023.1130075

Kubizňák, P., W. M. Hochachka, V. Osoba, T. Kotek, J. Kuchař, V. Klapetek, K. Hradcová, J. Růžička, and M. Zárybnická. 2019. Designing network-connected systems for ecological research and education. Ecosphere 10(6):e02761. https://doi.org/10.1002/ecs2.2761

Kuehne, L. M., and J. D. Olden. 2020. Military flights threaten the wilderness soundscapes of the Olympic Peninsula, Washington. Northwest Science 94(2):188-202. https://doi.org/10.3955/046.094.0208

LaCharite, K. 2016. Re-visioning agriculture in higher education: the role of campus agriculture initiatives in sustainability education. Agriculture and Human Values 33(3):521-535. https://doi.org/10.1007/s10460-015-9619-6

Lagardère, J. P. 1982. Effects of noise on growth and reproduction of Crangon crangon in rearing tanks. Marine Biology 71:177-185. https://doi.org/10.1007/BF00394627

Luo, J., B. M. Siemers, and K. Koselj. 2015. How anthropogenic noise affects foraging. Global Change Biology 21(9):3278-3289. https://doi.org/10.1111/gcb.12997

Madadi, H., H. Moradi, A. Soffianian, A. Salmanmahiny, J. Senn, and D. Geneletti. 2017. Degradation of natural habitats by roads: comparing land-take and noise effect zone. Environmental Impact Assessment Review 65:147-155. https://doi.org/10.1016/j.eiar.2017.05.003

Malone, K. M., A. C. Powell, F. Hua, and K. E. Sieving. 2017. Bluebirds perceive prey switching by Cooper’s Hawks across an urban gradient and adjust reproductive effort. Écoscience 24(1-2):21-31. https://doi.org/10.1080/11956860.2017.1346449

Matysioková, B. and V. Remeš. 2014. The importance of having a partner: male help releases females from time limitation during incubation in birds. Frontiers in Zoology 11:24. https://doi.org/10.1186/1742-9994-11-24

Maurelli, O. V. J. 2022. Sleepless and restless: effects of traffic noise and mosquito intrusions on Eastern Bluebird (Sialia sialis) incubation behavior. Thesis, University of Florida, Gainesville, Florida, USA. https://ufdc.ufl.edu/UFE0059505/00001/pdf

McClure, C. J., H. E. Ware, J. D. Carlisle, and J. R. Barber. 2017. Noise from a phantom road experiment alters the age structure of a community of migrating birds. Animal Conservation 20(2):164-172. https://doi.org/10.1111/acv.12302

McClure, C. J., H. E. Ware, J. Carlisle, G. Kaltenecker, and J. R. Barber. 2013. An experimental investigation into the effects of traffic noise on distributions of birds: avoiding the phantom road. Proceedings of the Royal Society B: Biological Sciences 280(1773):20132290. https://doi.org/10.1098/rspb.2013.2290

Meillère, A., F. Brischoux, and F. Angelier. 2015. Impact of chronic noise exposure on antipredator behavior: an experiment in breeding House Sparrows. Behavioral Ecology 26(2):569-577. https://doi.org/10.1093/beheco/aru232

Miller, J. D. 1974. Effects of noise on people. Journal of the Acoustical Society of America 56(3):729-764. https://doi.org/10.1121/1.1903322

Monaghan, P., and M. F. Haussmann. 2006. Do telomere dynamics link lifestyle and lifespan? Trends in Ecology & Evolution 21(1):47-53. https://doi.org/10.1016/j.tree.2005.11.007

Moreno, J., S. González-Braojos, and R. Ruiz-de-Castañeda. 2015. A spring cold snap is followed by an extreme reproductive failure event in a mountain population of Pied Flycatchers Ficedula hypoleuca. Bird Study 62(4):466-473. https://doi.org/10.1080/00063657.2015.1073680

Munro, H. L., and R. C. Rounds. 1985. Selection of artificial nest sites by five sympatric passerines. Journal of Wildlife Management 49:264-276. https://doi.org/10.2307/3801882

Munro, J., I. Williamson, and S. Fuller. 2018. Traffic noise impacts on urban forest soundscapes in south‐eastern Australia. Austral Ecology 43(2):180-190. https://doi.org/10.1111/aec.12555

Nassiri, P., M. R. Monazzam, S. F. Dehghan, G. Teimori, S. A. Zakerian, K. Azam, and M. Asghari. 2021. The combined effect of industrial noise type, level and frequency characteristics on hand motor skills: a lab trial study. Work 68(3):711-719. https://doi.org/10.3233/WOR-203405

Nilsson, J. Å. and H. G. Smith. 1988. Incubation feeding as a male tactic for early hatching. Animal Behaviour 36(3):641-647. https://doi.org/10.1016/S0003-3472(88)80145-3

Oliveira, G. D. S., V. M. Dos Santos, J. C. Rodrigues, and S. T. Nascimento. 2020. Effects of different egg turning frequencies on incubation efficiency parameters. Poultry Science 99(9):4417-4420. https://doi.org/10.1016/j.psj.2020.05.045

Ortega, C. P. 2012. Chapter 2: Effects of noise pollution on birds: a brief review of our knowledge. Ornithological Monographs 74(1):6-22. https://doi.org/10.1525/om.2012.74.1.6

Owens, J. L., C. L. Stec, and A. O’Hatnick. 2012. The effects of extended exposure to traffic noise on parid social and risk-taking behavior. Behavioural Processes 91(1):61-69. https://doi.org/10.1016/j.beproc.2012.05.010

Perreau, J., and K. E. Sieving. 2015. Choice of bluebird nest box: a potential for ecological traps? University of Florida Journal of Undergraduate Research 16(3):1-4.

Plummer, I., Y. Liu, and K. E. Sieving. 2021. Urban greenspace is for the bluebirds: nest-box selection across a noise gradient on an urbanizing university campus. Southeastern Naturalist 20(1):152-161. https://doi.org/10.1656/058.020.0115

Potvin, D. A., and S. A. MacDougall‐Shackleton. 2015. Traffic noise affects embryo mortality and nestling growth rates in captive zebra finches. Journal of Experimental Zoology Part A: Ecological Genetics and Physiology 323(10):722-730. https://doi.org/10.1002/jez.1965

Reid, J. M., P. Monaghan, and R. G. Nager. 2002. Incubation and the costs of reproduction. Pages 314-325 in D. C. Deeming, editor. Avian incubation: behaviour, environment and evolution. Oxford University Press, Oxford, UK. https://doi.org/10.1093/oso/9780198508106.003.0021

Rich, E. L., and L. M. Romero. 2005. Exposure to chronic stress down regulates corticosterone responses to acute stressors. American Journal of Physiology: Regulatory, Integrative and Comparative Physiology 288(6):R1628-R1636. https://doi.org/10.1152/ajpregu.00484.2004

Robertson, B. A., J. S. Rehage, and A. Sih. 2013. Ecological novelty and the emergence of evolutionary traps. Trends in Ecology & Evolution 28(9):552-560. https://doi.org/10.1016/j.tree.2013.04.004

Rodewald, A. D., D. P. Shustack, and T. M. Jones. 2011. Dynamic selective environments and evolutionary traps in human‐dominated landscapes. Ecology 92(9):1781-1788. https://doi.org/10.1890/11-0022.1

Rosa, P., and N. Koper. 2022. Impacts of oil well drilling and operating noise on abundance and productivity of grassland songbirds. Journal of Applied Ecology 59(2):574-584. https://doi.org/10.1111/1365-2664.14075

Rossi-Santos, M. R. 2015. Oil industry and noise pollution in the humpback whale (Megaptera novaeangliae) soundscape ecology of the southwestern Atlantic breeding ground. Journal of Coastal Research 31(1):184-195. https://doi.org/10.2112/JCOASTRES-D-13-00195.1

Ru, H., J. Xu, M. Li, Z. Duan, and Z. Li. 2018. Impact of traffic noise on Tibetan antelopes: a preliminary experiment on the Qinghai-Tibet highway in China. Applied Ecology and Environmental Research 16:2923-2932. https://doi.org/10.15666/aeer/1603_29232932

Salmon, P., J. F. Nilsson, A. Nord, S. Bensch, and C. Isaksson. 2016. Urban environment shortens telomere length in nestling Great Tits, Parus major. Biology Letters 12(6):20160155. https://doi.org/10.1098/rsbl.2016.0155

Schlaepfer, M. A., M. C. Runge, and P. W. Sherman. 2002. Ecological and evolutionary traps. Trends in Ecology & Evolution 17(10):474-480. https://doi.org/10.1016/S0169-5347(02)02580-6

Schomer, P. D., Y. Suzuki, and F. Saito. 2001. Evaluation of loudness-level weightings for assessing the annoyance of environmental noise. Journal of the Acoustical Society of America 110(5):2390-2397. https://doi.org/10.1121/1.1402116

Schroeder, J., S. Nakagawa, I. R. Cleasby, and T. Burke. 2012. Passerine birds breeding under chronic noise experience reduced fitness. PLoS ONE 7(7):e39200. https://doi.org/10.1371/journal.pone.0039200

Senzaki, M., Y. Yamaura, C. D. Francis, and F. Nakamura. 2016. Traffic noise reduces foraging efficiency in wild owls. Scientific Reports 6:30602. https://doi.org/10.1038/srep30602

Shannon, G., M. F. McKenna, L. M. Angeloni, K. R. Crooks, K. M. Fristrup, E. Brown, K. A. Warner, M. D. Nelson, C. White, J. Briggs, and S. McFarland. 2016. A synthesis of two decades of research documenting the effects of noise on wildlife. Biological Reviews 91(4):982-1005. https://doi.org/10.1111/brv.12207

Siemers, B. M., and A. Schaub. 2011. Hunting at the highway: traffic noise reduces foraging efficiency in acoustic predators. Proceedings of the Royal Society B: Biological Sciences 278(1712):1646-1652. https://doi.org/10.1098/rspb.2010.2262

Simmonds, E. G., B. C. Sheldon, T. Coulson, and E. F. Cole. 2017. Incubation behavior adjustments, driven by ambient temperature variation, improve synchrony between hatch dates and caterpillar peak in a wild bird population. Ecology and Evolution 7(22):9415-9425. https://doi.org/10.1002/ece3.3446

Singh, N., and S. C. Davar. 2004. Noise pollution-sources, effects and control. Journal of Human Ecology 16(3):181-187. https://doi.org/10.1080/09709274.2004.11905735

Skutch, A. F. 1962. The constancy of incubation. Wilson Bulletin 74(2):115-152.

Stier, A., N. B. Metcalfe, and P. Monaghan. 2020. Pace and stability of embryonic development affect telomere dynamics: an experimental study in a precocial bird model. Proceedings of the Royal Society B 287(1933):20201378. https://doi.org/10.1098/rspb.2020.1378

Swaddle, J. P., C. D. Francis, J. R. Barber, C. B. Cooper, C. C. Kyba, D. M. Dominoni, G. Shannon, E. Aschehoug, S. E. Goodwin, A. Y. Kawahara, et al. 2015. A framework to assess evolutionary responses to anthropogenic light and sound. Trends in Ecology & Evolution 30(9):550-560. https://doi.org/10.1016/j.tree.2015.06.009

Szalma, J. L. and P. A. Hancock. 2011. Noise effects on human performance: a meta-analytic synthesis. Psychological Bulletin 137(4):682-707. https://doi.org/10.1037/a0023987

Taylor, R. B. 2006. The effects of off-road vehicles on ecosystems. Texas Parks and Wildlife, Uvalde, USA. https://tpwd.texas.gov/publications/pwdpubs/media/pwd_rp_t3200_1081.pdf

Wada, H., B. P. Kriengwatana, T. D. Steury, and S. A. MacDougall-Shackleton. 2018. Incubation temperature influences sex ratio and offspring’s body composition in Zebra Finches (Taeniopygia guttata). Canadian Journal of Zoology 96(9):1010-1015. https://doi.org/10.1139/cjz-2017-0099

Wakstein, C. 2019. Pollution by noise: social and technical aspects. Pages 209-270 in A. D. McKnight, P. K. Marstrand, T. C. Sinclair, editors. Environmental pollution control. Routledge, London, UK. https://doi.org/10.4324/9780429345036-13

Wang, J. M., and S. R. Beissinger. 2009. Variation in the onset of incubation and its influence on avian hatching success and asynchrony. Animal Behaviour 78(3):601-613. https://doi.org/10.1016/j.anbehav.2009.05.022

Ward, D. 1990. Incubation temperatures and behavior of crowned, black-winged, and lesser black-winged plovers. Auk 107(1):10-17.

Webb, D. R. 1987. Thermal tolerance of avian embryos: a review. Condor 89(4):874-898. https://doi.org/10.2307/1368537

Williams, D. P., J. D. Avery, T. B. Gabrielson, and M. C. Brittingham. 2021. Experimental playback of natural gas compressor noise reduces incubation time and hatching success in two secondary cavity-nesting bird species. Condor 123(1):duaa066. https://doi.org/10.1093/ornithapp/duaa066

Williams, H. M. and R. L. DeLeon. 2020. Deep learning analysis of nest camera video recordings reveals temperature-sensitive incubation behavior in the Purple Martin (Progne subis). Behavioral Ecology and Sociobiology 74:7. https://doi.org/10.1007/s00265-019-2789-2

Winkler, D. W. 2016. Breeding biology of birds. Pages 407-450 in I. J. Lovette and J. W. Fitzpatrick, editors. Handbook of bird biology. Wiley, Hoboken, New Jersey, USA.

Yuan, M., C. Yin, Y. Sun, and W. Chen. 2019. Examining the associations between urban built environment and noise pollution in high-density high-rise urban areas: a case study in Wuhan, China. Sustainable Cities and Society 50:101678. https://doi.org/10.1016/j.scs.2019.101678

Zollinger, S. A., A. Dorado-Correa, W. Goymann, W. Forstmeier, U. Knief, A. M. BastidasUrrutia, and H. Brumm. 2019. Traffic noise exposure depresses plasma corticosterone and delays offspring growth in breeding zebra finches. Conservation Physiology 7(1):coz056. https://doi.org/10.1093/conphys/coz056