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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Ecol. Evol.</journal-id>
<journal-title>Frontiers in Ecology and Evolution</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Ecol. Evol.</abbrev-journal-title>
<issn pub-type="epub">2296-701X</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fevo.2023.1130075</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Ecology and Evolution</subject>
<subj-group>
<subject>Perspective</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>How chronic anthropogenic noise can affect wildlife communities</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Kok</surname> <given-names>Annebelle C. M.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>&#x002A;</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/2039641/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Berkhout</surname> <given-names>Boris W.</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
</contrib>
<contrib contrib-type="author">
<name><surname>Carlson</surname> <given-names>Nora V.</given-names></name>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<xref ref-type="aff" rid="aff6"><sup>6</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/2248068/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Evans</surname> <given-names>Neil P.</given-names></name>
<xref ref-type="aff" rid="aff7"><sup>7</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/285046/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Khan</surname> <given-names>Nicola</given-names></name>
<xref ref-type="aff" rid="aff8"><sup>8</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/619532/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Potvin</surname> <given-names>Dominique A.</given-names></name>
<xref ref-type="aff" rid="aff9"><sup>9</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1050955/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Radford</surname> <given-names>Andrew N.</given-names></name>
<xref ref-type="aff" rid="aff10"><sup>10</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/2109495/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Sebire</surname> <given-names>Marion</given-names></name>
<xref ref-type="aff" rid="aff11"><sup>11</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/1428908/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Shafiei Sabet</surname> <given-names>Saeed</given-names></name>
<xref ref-type="aff" rid="aff12"><sup>12</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/204525/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Shannon</surname> <given-names>Graeme</given-names></name>
<xref ref-type="aff" rid="aff13"><sup>13</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/410168/overview"/>
</contrib>
<contrib contrib-type="author">
<name><surname>Wascher</surname> <given-names>Claudia A. F.</given-names></name>
<xref ref-type="aff" rid="aff14"><sup>14</sup></xref>
<uri xlink:href="http://loop.frontiersin.org/people/46881/overview"/>
</contrib>
</contrib-group>
<aff id="aff1"><sup>1</sup><institution>Marine Biology Lab, Groningen Institute of Evolutionary Life Sciences, Rijksuniversiteit Groningen</institution>, <addr-line>Groningen</addr-line>, <country>Netherlands</country></aff>
<aff id="aff2"><sup>2</sup><institution>Acoustic Ecology Lab, Scripps Institution of Oceanography, University of California</institution>, <addr-line>San Diego, La Jolla, CA</addr-line>, <country>United States</country></aff>
<aff id="aff3"><sup>3</sup><institution>Faculteit der Natuurwetenschappen, Wiskunde en Informatica, University of Amsterdam</institution>, <addr-line>Amsterdam</addr-line>, <country>Netherlands</country></aff>
<aff id="aff4"><sup>4</sup><institution>Juanes Lab, School of Biology, University of Victoria</institution>, <addr-line>Victoria, BC</addr-line>, <country>Canada</country></aff>
<aff id="aff5"><sup>5</sup><institution>Suzuki Lab, Faculty of Science, Graduate School of Science, Kyoto University</institution>, <addr-line>Kyoto</addr-line>, <country>Japan</country></aff>
<aff id="aff6"><sup>6</sup><institution>Canadian Wildlife Service, Environment and Climate Change Canada</institution>, <addr-line>Sidney, BC</addr-line>, <country>Canada</country></aff>
<aff id="aff7"><sup>7</sup><institution>School of Biodiversity, One Health and Veterinary Medicine, University of Glasgow</institution>, <addr-line>Glasgow</addr-line>, <country>United Kingdom</country></aff>
<aff id="aff8"><sup>8</sup><institution>School of Animal, Rural and Environmental Sciences, Nottingham Trent University</institution>, <addr-line>Southwell</addr-line>, <country>United Kingdom</country></aff>
<aff id="aff9"><sup>9</sup><institution>School of Science Technology and Engineering, University of the Sunshine Coast</institution>, <addr-line>Petrie, QLD</addr-line>, <country>Australia</country></aff>
<aff id="aff10"><sup>10</sup><institution>School of Biological Sciences, University of Bristol</institution>, <addr-line>Bristol</addr-line>, <country>United Kingdom</country></aff>
<aff id="aff11"><sup>11</sup><institution>Centre for Environment, Fisheries and Aquaculture Science, Weymouth Laboratory</institution>, <addr-line>Weymouth</addr-line>, <country>United Kingdom</country></aff>
<aff id="aff12"><sup>12</sup><institution>Fisheries Department, Faculty of Natural Resources, University of Guilan</institution>, <addr-line>Someh Sara</addr-line>, <country>Iran</country></aff>
<aff id="aff13"><sup>13</sup><institution>School of Natural Sciences, Bangor University</institution>, <addr-line>Bangor</addr-line>, <country>United Kingdom</country></aff>
<aff id="aff14"><sup>14</sup><institution>Behavioural Ecology Research Group, School of Life Sciences, Anglia Ruskin University</institution>, <addr-line>Cambridge</addr-line>, <country>United Kingdom</country></aff>
<author-notes>
<fn fn-type="edited-by"><p>Edited by: Jarle Tryti Nordeide, Nord University, Norway</p></fn>
<fn fn-type="edited-by"><p>Reviewed by: Maria Clara P. Amorim, University of Lisbon, Portugal; Ann Bowles, Hubbs-SeaWorld Research Institute, United States</p></fn>
<corresp id="c001">&#x002A;Correspondence: Annebelle C. M. Kok, <email>a.c.m.kok@rug.nl</email></corresp>
<fn fn-type="other" id="fn004"><p>This article was submitted to Behavioral and Evolutionary Ecology, a section of the journal Frontiers in Ecology and Evolution</p></fn>
</author-notes>
<pub-date pub-type="epub">
<day>05</day>
<month>04</month>
<year>2023</year>
</pub-date>
<pub-date pub-type="collection">
<year>2023</year>
</pub-date>
<volume>11</volume>
<elocation-id>1130075</elocation-id>
<history>
<date date-type="received">
<day>22</day>
<month>12</month>
<year>2022</year>
</date>
<date date-type="accepted">
<day>15</day>
<month>03</month>
<year>2023</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#x00A9; 2023 Kok, Berkhout, Carlson, Evans, Khan, Potvin, Radford, Sebire, Shafiei Sabet, Shannon and Wascher.</copyright-statement>
<copyright-year>2023</copyright-year>
<copyright-holder>Kok, Berkhout, Carlson, Evans, Khan, Potvin, Radford, Sebire, Shafiei Sabet, Shannon and Wascher</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/"><p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p></license>
</permissions>
<abstract>
<p>Anthropogenic noise is a major pollutant in terrestrial and aquatic ecosystems. Since the industrial revolution, human activities have become increasingly noisy, leading to both acute and chronic disturbance of a wide variety of animals. Chronic noise exposure can affect animals over their lifespan, leading to changes in species interactions and likely altering communities. However, the community-level impacts of chronic noise are not well-understood, which impairs our ability for effective mitigation. In this review, we address the effects of chronic noise exposure on communities and explore possible mechanisms underlying these effects. The limited studies on this topic suggest that noise can affect communities by changing the behavior and/or physiology of species in a community, which results in direct or knock-on consequences for other species in the ecosystem. Major knowledge gaps remain due to the logistically complex and financially expensive nature of the long-term studies needed to address these questions. By identifying these gaps and suggesting approaches to answer them, we provide a road map toward mitigating the effects of a noisy world.</p>
</abstract>
<kwd-group>
<kwd>anthropogenic noise</kwd>
<kwd>chronic</kwd>
<kwd>community</kwd>
<kwd>behavior</kwd>
<kwd>ecology</kwd>
</kwd-group>
<counts>
<fig-count count="1"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="112"/>
<page-count count="10"/>
<word-count count="9132"/>
</counts>
</article-meta>
</front>
<body>
<sec id="S1">
<title>1. Chronic noise can change communities</title>
<sec id="S1.SS1">
<title>1.1. Introduction</title>
<p>Anthropogenic noise is recognized as a major global pollutant that has considerable implications for human health (<xref ref-type="bibr" rid="B56">Matheson et al., 2003</xref>; <xref ref-type="bibr" rid="B39">Hammer et al., 2014</xref>; <xref ref-type="bibr" rid="B63">Mohamed et al., 2021</xref>) and the behavior, physiology and fitness of wildlife (<xref ref-type="bibr" rid="B5">Barber et al., 2010</xref>). Indeed, a substantial body of research has been published over the past two decades that has explored the effects of noise pollution on animals across terrestrial and aquatic ecosystems (<xref ref-type="bibr" rid="B88">Shannon et al., 2016b</xref>; <xref ref-type="bibr" rid="B98">Sordello et al., 2020</xref>; <xref ref-type="bibr" rid="B23">Duarte et al., 2021</xref>; <xref ref-type="bibr" rid="B44">Jerem and Mathews, 2021</xref>). Anthropogenic noise that significantly alters the acoustic environment has increased markedly since the Industrial Revolution both in terms of the level of sound exposure and geographical extent. The rise in noise pollution has been driven by population growth, mechanization, infrastructure development and increasing demand for resources (<xref ref-type="bibr" rid="B5">Barber et al., 2010</xref>). For example, in the US continent-wide changes in the soundscape extend well beyond the boundaries of urban environments (<xref ref-type="bibr" rid="B62">Mennitt and Fristrup, 2016</xref>), with 63% of protected areas experiencing sound exposure double that of the ambient level (<xref ref-type="bibr" rid="B11">Buxton et al., 2017</xref>). As such, noise has the potential to be a major selective force that can restructure wildlife communities (<xref ref-type="bibr" rid="B101">Swaddle et al., 2015</xref>). At the same time, noise does not persist long-term in the environment like many other pollutants&#x2014;as was evidenced during the COVID-19 pandemic when lockdown measures led to sustained global quieting (<xref ref-type="bibr" rid="B53">Lecocq et al., 2020</xref>)&#x2014;which can facilitate mitigation measures and the reestablishment of natural soundscapes. The latency of recovery following noise removal will depend on the specific behavioral and/or physiological mechanisms that restructure wildlife communities, as well as the flexibility of species to adapt to the return of a more natural acoustic environment.</p>
<p>Field-based studies and laboratory experiments have provided considerable evidence that exposure to noise can cause a wide range of ecological impacts to wildlife (<xref ref-type="bibr" rid="B88">Shannon et al., 2016b</xref>; <xref ref-type="bibr" rid="B50">Kunc and Schmidt, 2019</xref>; <xref ref-type="bibr" rid="B44">Jerem and Mathews, 2021</xref>). These include changing spatial distribution and deterring wildlife from important feeding and breeding areas, or interfering with crucial biological functions such as foraging performance (more food handling errors and discrimination errors), predator avoidance, prey detection and conspecific communication. Furthermore, there are direct physiological costs associated with exposure to noise from reduced sleep (<xref ref-type="bibr" rid="B34">Grunst et al., 2021</xref>) to increasing stress hormone levels (<xref ref-type="bibr" rid="B104">Tro&#x00EF;anowski et al., 2017</xref>). These varied impacts may lead to negative consequences for individual fitness, population levels and community structure (<xref ref-type="bibr" rid="B96">Slabbekoorn et al., 2010</xref>; <xref ref-type="bibr" rid="B26">Francis and Barber, 2013</xref>). Similarly, human communities experience physiological and behavioral effects from noise exposure that include sleep deprivation, cognitive impairment, elevated stress and annoyance, and a higher incidence of cardiovascular disease (<xref ref-type="bibr" rid="B99">Stansfeld and Matheson, 2003</xref>; <xref ref-type="bibr" rid="B102">Szalma and Hancock, 2011</xref>; <xref ref-type="bibr" rid="B39">Hammer et al., 2014</xref>).</p>
<p>The behavioral effects of noise exposure are driven by four key mechanisms that are not mutually exclusive: (1) Noise masks critical sounds including communication of conspecifics and other relevant cues such as approaching danger; (2) noise distracts animals from attending to pertinent information in the environment; (3) noise is perceived as a direct threat, thereby altering behavioral responses of the animal; and/or (4) noise initiates chronic stress, leading to long-term behavioral and physiological changes in the animal. It is important to note that each of these mechanisms is capable of driving a change in animal distribution and habitat use, which can alter the composition and interaction of species at the community-level.</p>
</sec>
<sec id="S1.SS2">
<title>1.2. Chronic noise exposure</title>
<p>Although there have been considerable advances in our understanding of the biological responses associated with anthropogenic noise exposure over the past two decades, most are based on comparatively short-term experiments and observations (<xref ref-type="bibr" rid="B88">Shannon et al., 2016b</xref>; <xref ref-type="bibr" rid="B44">Jerem and Mathews, 2021</xref>). In addition, these studies have predominantly focused on a single species. However, transport networks, industry and urban environments are major sources of chronic anthropogenic noise that permeate natural and human transformed environments over the long-term. These noise sources are also characterized by acoustic energy mainly being concentrated in the low-frequency spectrum (&#x003C;2 kHz), which travel further than high-frequency sounds and therefore potentially impact a wide range of species simultaneously.</p>
<p>Sound in nature is a complex and challenging physical phenomenon to measure, as it varies in terms of duration, frequency and amplitude. The acoustic characteristics of a given noise source and the perceptual capabilities (e.g., hearing sensitivity) of the species exposed to it will be instrumental in determining whether it is likely to significantly impact animal behavior and physiology (<xref ref-type="bibr" rid="B26">Francis and Barber, 2013</xref>). Indeed, the duration, spectral and temporal characteristics of chronic noise may ultimately be as important as the amplitude when it comes to mediating long-term effects on animal communities. It is therefore crucial that scientists and wildlife practitioners accurately measure and report the specific sound characteristics of the noise source, as this information is critical for comparing research findings and understanding the specific levels of exposure that can drive a biological response (<xref ref-type="bibr" rid="B60">McKenna et al., 2016</xref>). Currently, many of the noise metrics used in research and monitoring are focused on human perceptual abilities, rather than considering the varied hearing thresholds of animal communities.</p>
<p>To fully understand the impacts of chronic anthropogenic noise exposure on animal communities it is necessary to conduct long-term studies (<xref ref-type="bibr" rid="B44">Jerem and Mathews, 2021</xref>). These need to consider the effects of chronic exposure and lasting impacts after exposure has ended. However, a definition of chronic exposure is rarely given (<xref ref-type="bibr" rid="B44">Jerem and Mathews, 2021</xref>) and short-term and chronic exposure are on a continuum, making it difficult to classify exposure events in the field. For example, repeated short bursts of exposure (e.g., sonar) over longer periods (a few times a year for years) differ from medium-term exposure (continuous for weeks, e.g., construction site) in their effects. Additionally, what is considered chronic will also depend on the lifespan of the organism, which ranges from weeks to decades (<xref ref-type="bibr" rid="B4">Austad, 2010</xref>). Construction of a gas pipeline that takes a year would thus be multi-generation exposure for some animals, whilst only a brief exposure for others. Finally, the effects of chronic exposure are expected to impact multiple species in the community, and should therefore be long enough to affect species with varying lifespans. Chronic noise exposure can thus have a low or high duty cycle, as long as it occurs regularly or long enough to impact species with varying lifespans. Therefore, we suggest a rather broad, but practical chronic exposure definition: &#x201C;Exposure throughout a significant part of the lifespan of an animal, at regular enough intervals to have the potential of lasting impacts from the individual- to community-scale.&#x201D;</p>
</sec>
<sec id="S1.SS3">
<title>1.3. State of the knowledge about chronic noise effects on communities</title>
<p>We conducted a search of peer-reviewed journal articles that have been published since 1970 and have addressed the effects of noise on animal communities. Our article search used the datasets provided in four previous systematic review papers (<xref ref-type="bibr" rid="B88">Shannon et al., 2016b</xref>; <xref ref-type="bibr" rid="B98">Sordello et al., 2020</xref>; <xref ref-type="bibr" rid="B23">Duarte et al., 2021</xref>; <xref ref-type="bibr" rid="B44">Jerem and Mathews, 2021</xref>), as well as using the Web of Science and Google Scholar search engines to identify any further papers that were omitted from the reviews or that have been published in the last 2 years. The search terms we used were &#x201C;anthropogenic,&#x201D; &#x201C;noise,&#x201D; &#x201C;wildlife,&#x201D; &#x201C;animal,&#x201D; and &#x201C;community.&#x201D; We specifically focused on studies that explored whether noise exposure affected community-level composition (e.g., changes in abundance and diversity of multiple species) over time. The dataset comprised 48 papers published between 1995 and 2021 (<xref ref-type="table" rid="T1">Table 1</xref> and <xref ref-type="supplementary-material" rid="FS1">Supplementary Material 1</xref>), of which 16 were not included in the previous review papers. Overall, the most common method to explore the community effects of noise was through direct observation (<italic>n</italic> = 29) followed by playback (<italic>n</italic> = 10) and natural experiment (<italic>n</italic> = 9). Seventy-nine percent of the studies (<italic>n</italic> = 38) reported negative effects of noise exposure in one taxonomic group (decreased abundance/species richness, decreased nesting, and decreased offspring survival/hatching success) while only one study documented a positive effect (increased abundance/species richness). The remainder revealed either no effect (<italic>n</italic> = 7) or mixed effects within the same taxonomic group (<italic>n</italic> = 7), with the direction depending upon the species. Birds featured in 38 (79%) of the studies; mammals, amphibians, reptiles and invertebrates were the least represented, featuring in one study each. Seventy-nine percent of the studies were conducted in either North America or Europe (<italic>n</italic> = 38). Seventy-one percent of the studies (<italic>n</italic> = 34) explored whether noise influenced abundance and/or species diversity.</p>
<table-wrap position="float" id="T1">
<label>TABLE 1</label>
<caption><p>Number of studies in the literature review (S2) that investigated the effects of noise on communities separated by taxonomic group.</p></caption>
<table cellspacing="5" cellpadding="5" frame="box" rules="all">
<thead>
<tr>
<td valign="top" align="left" colspan="25" style="color:#ffffff;background-color: #7f8080;">Taxonomic group</td>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left" style="color:#ffffff;background-color: #7f8080;"></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;" colspan="3"><bold>Plants</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;" colspan="3"><bold>Worms</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;" colspan="3"><bold>Arthropods</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;" colspan="3"><bold>Fish</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;" colspan="3"><bold>Amphibians</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;" colspan="3"><bold>Reptiles</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;" colspan="3"><bold>Birds</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;" colspan="3"><bold>Mammals</bold></td>
</tr>
<tr>
<td valign="top" align="left" style="color:#ffffff;background-color: #7f8080;"><bold>Effect</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>N</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>0</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>P</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>N</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>0</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>P</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>N</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>0</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>P</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>N</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>0</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>P</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>N</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>0</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>P</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>N</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>0</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>P</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>N</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>0</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>P</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>N</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>0</bold></td>
<td valign="top" align="center" style="color:#ffffff;background-color: #7f8080;"><bold>P</bold></td>
</tr>
<tr>
<td valign="top" align="left">Negative</td>
<td/>
<td valign="top" align="center">1</td>
<td valign="top" align="center">1<xref ref-type="table-fn" rid="t1fns1">&#x002A;</xref></td>
<td valign="top" align="center">1</td>
<td/>
<td/>
<td valign="top" align="center">1</td>
<td/>
<td valign="top" align="center"><bold>1</bold>&#x2020;</td>
<td/>
<td valign="top" align="center">2</td>
<td/>
<td/>
<td/>
<td valign="top" align="center">1</td>
<td valign="top" align="center">1&#x00B0;</td>
<td/>
<td/>
<td valign="top" align="center">3&#x00B0;</td>
<td valign="top" align="center">18&#x2021;</td>
<td valign="top" align="center">7&#x2020;</td>
<td valign="top" align="center">1&#x00B0;</td>
<td/>
<td/>
</tr>
<tr>
<td valign="top" align="left">Positive</td>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="center">1</td>
<td/>
<td/>
<td/>
<td/>
</tr>
<tr>
<td valign="top" align="left">Neutral</td>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="center">1</td>
<td/>
<td/>
<td/>
<td valign="top" align="center">1&#x2021;</td>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="center">1</td>
<td valign="top" align="center">4</td>
<td/>
<td/>
<td/>
<td/>
</tr>
<tr>
<td valign="top" align="left">Mixed</td>
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<td valign="top" align="center">1</td>
<td valign="top" align="center">1<xref ref-type="table-fn" rid="t1fns1">&#x002A;</xref></td>
<td/>
<td valign="top" align="center">1</td>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td/>
<td valign="top" align="center">2</td>
<td valign="top" align="center">1</td>
<td valign="top" align="center">1</td>
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</tr>
</tbody>
</table>
<table-wrap-foot>
<fn id="t1fns1"><p>N, natural experiment; O, observational; P, playback experiment. Mixed effects = effects differed between or within species within the same taxonomic group. Note that studies focusing on more than one taxonomic group are featured multiple times, once per taxonomic group. Those studies have been indicated with matching symbols (i.e. the study &#x002A; features plants and arthropods).</p></fn>
</table-wrap-foot>
</table-wrap>
<p>The early research on community-level effects of noise was conducted by observing the abundance and diversity of bird species as a function of distance from a chronic noise source, such as a busy roadway. This proved an effective method for understanding the effects of noise with clear evidence that species composition, density and abundance were more negatively impacted the higher the noise levels (<xref ref-type="bibr" rid="B78">Reijnen and Foppen, 1995</xref>; <xref ref-type="bibr" rid="B80">Reijnen et al., 1995</xref>, <xref ref-type="bibr" rid="B79">1996</xref>). Later studies highlighted how species with low frequency calls that overlapped considerably with traffic noise were likely to be impacted to a much greater extent than species with higher frequency calls [(e.g., lower occupancy (<xref ref-type="bibr" rid="B32">Goodwin and Shriver, 2011</xref>)]. However, there were challenges associated with this observational approach such as accounting for confounding variables, including habitat fragmentation, chemical pollution, elevated human activity, and increased mortality (e.g., vehicle strike), that also occur to a greater extent in close proximity to roads (<xref ref-type="bibr" rid="B100">Summers et al., 2011</xref>).</p>
<p>The expansion of gas extraction across North America led to the development of natural experiments where areas with noise-generating compressor stations could be compared to areas structurally equivalent to compressor stations but with quiet well pads. These natural experiments largely controlled for the confounding variables associated with other forms of disturbance &#x2014; such as habitat transformation and human activity &#x2014; and explicitly investigated the effects of noise on the abundance and diversity of bird species in an otherwise natural environment (<xref ref-type="bibr" rid="B6">Bayne et al., 2008</xref>). These studies demonstrated that occupancy, diversity and abundance of avian species were negatively impacted by anthropogenic noise (<xref ref-type="bibr" rid="B6">Bayne et al., 2008</xref>), while also revealing that species exhibited varying levels of sensitivity to noise exposure depending on the extent of vocal masking that they experienced (<xref ref-type="bibr" rid="B28">Francis et al., 2009</xref>, <xref ref-type="bibr" rid="B29">2011a</xref>). Larger bodied birds with lower frequency calls were found to use noisy areas considerably less than smaller bodied species with higher frequency vocalizations, which presents a strong selective force shaping avian community structure and species interactions such as predator&#x2014;prey relationships (<xref ref-type="bibr" rid="B28">Francis et al., 2009</xref>, <xref ref-type="bibr" rid="B29">2011a</xref>). However, researchers working in grassland prairies documented an effect of the gas extraction infrastructure, rather than the noise that was associated with it (<xref ref-type="bibr" rid="B69">Nenninger and Koper, 2018</xref>), showing the complexity of the mechanisms underlying this disturbance. The physiological costs of noise have also been documented using this natural study system demonstrating impacts to glucocorticoid-signaling and reduced fitness across bird species (<xref ref-type="bibr" rid="B48">Kleist et al., 2018</xref>), while truly long-term cascading impacts of noise exposure were recently found in a study that revealed chronic noise exposure (15 years) impacted seedling recruitment and woody plant community structure (<xref ref-type="bibr" rid="B74">Phillips et al., 2021</xref>). These effects were still in evidence 2&#x2013;4 years after the removal of noise.</p>
<p>Scientists have also employed playbacks across the landscape to assess experimentally the effects of introduced noise on wildlife communities. The advantages of the playback approach include the ability to control the specific location and duration of noise exposure, as well as the noise source sound level. Furthermore, playbacks allow for the effects of noise to be investigated in isolation of confounding variables that are generally associated with noise disturbance. The first of these landscape-level studies broadcast traffic noise along a 500 m &#x201C;phantom road&#x201D; in habitat favored by migratory birds, with the researchers documenting a 25% reduction in bird abundance during playback periods (<xref ref-type="bibr" rid="B58">McClure et al., 2013</xref>). Further work revealed that 31% of the bird community avoided the phantom road, while those individuals that remained experienced a reduction in body condition that was associated with an altered trade-off in foraging and vigilance (<xref ref-type="bibr" rid="B109">Ware et al., 2015</xref>). There were also age effects with younger birds being impacted by noise to a greater extent than adults (<xref ref-type="bibr" rid="B57">McClure et al., 2017</xref>). Subsequent research has demonstrated similar effects of introduced traffic noise on invertebrates, particularly species that rely on acoustic signals in the environment (<xref ref-type="bibr" rid="B83">Senzaki et al., 2020</xref>), while breeding birds in North American grassland prairies declined in abundance when exposed to playbacks of noise associated with energy extraction at the landscape scale (<xref ref-type="bibr" rid="B15">Cinto Mejia et al., 2019</xref>; <xref ref-type="bibr" rid="B81">Rosa and Koper, 2022</xref>). There appear to be complex interactions between infrastructure, noise and species-specific physiological and behavioral responses that can potentially mediate negative effects, and in some cases noise effects may be dwarfed by the effects of physical structures in natural areas [i.e., oil pumps: (<xref ref-type="bibr" rid="B69">Nenninger and Koper, 2018</xref>)]. However, we believe these complexities further highlight gaps in noise research on community ecology where interactions between different ecosystem components, noise, and the health of the community network still exist (<xref ref-type="bibr" rid="B16">Curry et al., 2018</xref>).</p>
</sec>
</sec>
<sec id="S2">
<title>2. Why do we see differences in community composition due to noise?</title>
<p>Noise can have a profound effect on community composition through a variety of mechanisms (<xref ref-type="fig" rid="F1">Figure 1</xref> and <xref ref-type="supplementary-material" rid="FS1">Supplementary Figure 1</xref>). It can directly impact local abundance of different species due to avoidance, increased mortality and decreased recruitment, while indirectly impacting predator&#x2014;prey and parasite&#x2014;host interactions, competitor interactions, and species-driven ecosystem structure. Noise can have such a broad impact because species within a community respond differently (e.g., declines, altered predator&#x2013;prey relationships, etc.). This combination of direct and indirect effects and species variability in response alter composition and structure of the community</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption><p>Theoretical knock-on consequences from direct noise impacts on the population of one species. This example is based on decline in recruitment as a result of lower fecundity in female great tits and lower nestling survival due to noise. The resulting decrease in the local population could affect a number of other species though a variety of mechanisms: <bold>(A)</bold> Removing hosts for parasites (e.g., blowfly larvae), <bold>(B)</bold> reducing prey availability resulting in prey switching, causing declines in sympatric species due to increased predation pressure, <bold>(C)</bold> removing anti-predator information resulting in higher mortality due to predation, or altered spatial use within a mixed-species community by <bold>(D)</bold> opening a niche previously used by the declining species (<xref ref-type="bibr" rid="B14">Cimprich and Grubb Jnr, 1994</xref>), <bold>(E)</bold> removing information/presence that allowed species to broaden their own microhabitat use when the declining species were present (<xref ref-type="bibr" rid="B22">Dolby and Grubb, 1998</xref>), or <bold>(F)</bold> removing information/perceived safety that allowed species to cross barriers such as open spaces (e.g., heterospecific using chick-a-dee calls to initiate movement across roads, man-made fields, etc.) (<xref ref-type="bibr" rid="B91">Sieving et al., 2004</xref>).</p></caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fevo-11-1130075-g001.tif"/>
</fig>
<sec id="S2.SS1">
<title>2.1. Direct effects</title>
<p>One of the direct influences of noise is avoidance behavior; many species will temporarily (<xref ref-type="bibr" rid="B97">Slotte et al., 2004</xref>; <xref ref-type="bibr" rid="B58">McClure et al., 2013</xref>; <xref ref-type="bibr" rid="B109">Ware et al., 2015</xref>; <xref ref-type="bibr" rid="B10">Bunkley et al., 2017</xref>; <xref ref-type="bibr" rid="B12">Carral-Murrieta et al., 2020</xref>) or permanently (<xref ref-type="bibr" rid="B71">Nicholson et al., 1992</xref>; <xref ref-type="bibr" rid="B72">Pearson et al., 1992</xref>; <xref ref-type="bibr" rid="B65">Morton and Symonds, 2002</xref>; <xref ref-type="bibr" rid="B28">Francis et al., 2009</xref>; <xref ref-type="bibr" rid="B103">Thompson et al., 2010</xref>; <xref ref-type="bibr" rid="B41">Herrera-Montes and Aide, 2011</xref>; <xref ref-type="bibr" rid="B13">Ciach and Fr&#x00F6;hlich, 2017</xref>) avoid noisy areas. A variety of migrating bird species, for example, avoided stopping over in noisy areas, even though they used the same areas under quieter conditions (<xref ref-type="bibr" rid="B58">McClure et al., 2013</xref>; <xref ref-type="bibr" rid="B109">Ware et al., 2015</xref>), while many insects demonstrate reduced abundance in noisy areas (<xref ref-type="bibr" rid="B10">Bunkley et al., 2017</xref>). Species respond differently to noise (<xref ref-type="bibr" rid="B86">Shafiei Sabet et al., 2016</xref>; <xref ref-type="bibr" rid="B108">Voellmy et al., 2016</xref>; <xref ref-type="bibr" rid="B10">Bunkley et al., 2017</xref>). While some species may disappear entirely from a noisy area (<xref ref-type="bibr" rid="B108">Voellmy et al., 2016</xref>), others may not exhibit such a drastic decline, or, in some cases, may even increase (<xref ref-type="bibr" rid="B28">Francis et al., 2009</xref>; <xref ref-type="bibr" rid="B108">Voellmy et al., 2016</xref>; <xref ref-type="bibr" rid="B10">Bunkley et al., 2017</xref>), altering the composition and interactions of the species remaining.</p>
<p>While avoidance is a common strategy employed by many species, chronic noise can also alter species abundance through increased mortality. Noise can directly lead to individual mortality through noise-induced permanent injury (excluding hearing loss) or reduction of predator detection. To the best of our knowledge, no examples of noise-induced injury have been demonstrated in terrestrial systems; however, they are unfortunately common in marine and other aquatic systems (<xref ref-type="bibr" rid="B23">Duarte et al., 2021</xref>). Noise can kill by causing injuries such as swim bladder and kidney rupture in fish (<xref ref-type="bibr" rid="B66">National Academies of Sciences, Engineering and Medicine, 2011</xref>; <xref ref-type="bibr" rid="B38">Halvorsen et al., 2012</xref>; <xref ref-type="bibr" rid="B43">Jenkins et al., 2022</xref>), or through alterations in behavior that can lead to death [e.g., diving behavior (<xref ref-type="bibr" rid="B25">Fern&#x00E1;ndez et al., 2005</xref>), strandings (<xref ref-type="bibr" rid="B8">Bernaldo de Quir&#x00F3;s et al., 2019</xref>)]. This is likely in part due to the physical properties of the two environments. Sound pressure in water is relatively much higher than in air, due to the density difference between the two media (<xref ref-type="bibr" rid="B75">Popper and Hawkins, 2019</xref>). Additionally, sound travels further and faster underwater than in air, increasing the potential area of effect of noise. These factors have been exploited by many marine and freshwater species that have evolved sensitive specialized anatomical structures for hearing and sensing pressure (<xref ref-type="bibr" rid="B75">Popper and Hawkins, 2019</xref>). However, noisy activities such as pile driving, sonar, dredging and seismic airguns for oil and gas exploration are common in underwater environments and can overwhelm these highly adapted systems.</p>
<p>Though not as immediately fatal, animals living in noisy conditions often show altered levels of stress hormones, which are linked to increased stress-responsiveness (<xref ref-type="bibr" rid="B54">Leshyk et al., 2013</xref>). These can lead to poorer body condition (<xref ref-type="bibr" rid="B2">Anderson et al., 2011</xref>) and reduced reproductive success, such as lower hatching success (<xref ref-type="bibr" rid="B48">Kleist et al., 2018</xref>) and poorer growth and development in those offspring that survive (<xref ref-type="bibr" rid="B42">Injaian et al., 2018</xref>). Poorer body condition can be a result of increased glucocorticoid concentrations, decreased foraging success, and changes in social relationships and behavior. Noise can result in activation of the hypothalamic&#x2013;pituitary&#x2013;adrenocortical (HPA) axis. If the animal is unable to escape the source of stress, the HPA system remains activated (<xref ref-type="bibr" rid="B76">Pravosudov et al., 2001</xref>), and the short-term physiological or behavioral changes that ameliorate stress and promote survival can become deleterious. These effects have the potential to be far-reaching: for example, in birds, short-term corticosterone manipulation has been shown to reduce survivorship 2 years after the manipulation (<xref ref-type="bibr" rid="B33">Goutte et al., 2010</xref>). Extended bouts of stress do not only deplete energy reserves, but also impair growth and immune function, accelerate aging, negatively impact cognitive ability and increase atrophy of nerve cells in the brain (<xref ref-type="bibr" rid="B111">Wingfield et al., 1998</xref>; <xref ref-type="bibr" rid="B95">Sims and Holberton, 2000</xref>; <xref ref-type="bibr" rid="B76">Pravosudov et al., 2001</xref>; <xref ref-type="bibr" rid="B110">Wingfield and Kitaysky, 2002</xref>; <xref ref-type="bibr" rid="B24">Ellenberg et al., 2007</xref>). However, it is important to note that measuring stress response is inherently complex because stress is experienced on multiple levels&#x2014;social, psychological, and physiological&#x2014;and varies with life stage and life history, circadian rhythm, hormone interactions, and the extent and longevity of environmental disturbance [e.g., (<xref ref-type="bibr" rid="B3">Angelier et al., 2009</xref>; <xref ref-type="bibr" rid="B21">Dickens and Romero, 2013</xref>; <xref ref-type="bibr" rid="B54">Leshyk et al., 2013</xref>; <xref ref-type="bibr" rid="B82">Schultner et al., 2013</xref>)].</p>
<p>Aside from causing avoidance and increased mortality, noise can alter local populations by impacting reproductive success. This can occur through several mechanisms including decreased breeding rates and decreased juvenile recruitment, as well as altered resource allocation. For example, noise can reduce breeding rates through disrupting detection of potential mates [e.g., greater sage grouse (<xref ref-type="bibr" rid="B9">Blickley et al., 2012</xref>), ovenbirds, <italic>Seiurus aurocapilla</italic> (<xref ref-type="bibr" rid="B36">Habib et al., 2007</xref>)] and alter or reduce mating behavior [e.g., decreasing display, <xref ref-type="bibr" rid="B17">de Jong et al. (2018b)</xref>]. In a similar fashion to causing direct mortality and poor condition, noise can result in lower clutch size (<xref ref-type="bibr" rid="B37">Halfwerk et al., 2011</xref>), hatching success (<xref ref-type="bibr" rid="B48">Kleist et al., 2018</xref>), juvenile growth/condition (<xref ref-type="bibr" rid="B52">Lagard&#x00E8;re, 1982</xref>; <xref ref-type="bibr" rid="B47">Kight et al., 2012</xref>), and juvenile survival (<xref ref-type="bibr" rid="B19">De Soto et al., 2013</xref>; <xref ref-type="bibr" rid="B68">Nedelec et al., 2017</xref>; <xref ref-type="bibr" rid="B18">de Jong et al., 2018a</xref>). Many of these effects are thought to be either a result of changes in parental behavior (e.g., increased aggression and decreased attention to young, <xref ref-type="bibr" rid="B68">Nedelec et al. (2017)</xref>, increases in larval/juvenile mutation (<xref ref-type="bibr" rid="B52">Lagard&#x00E8;re, 1982</xref>), and/or <italic>via</italic> direct or indirect interactions with elevated stress hormones resulting in higher mortality or poorer condition e.g., <xref ref-type="bibr" rid="B92">Silverin (1986)</xref>; <xref ref-type="bibr" rid="B42">Injaian et al. (2018)</xref>. In combination, this is especially damaging as populations under chronic stress are less likely to recover, due to increased mortality and decreased fitness of remaining adults. Fitness costs can fundamentally influence population dynamics; for instance, changes in breeding success can drastically alter population size and range by decreasing the numbers of new individuals coming into the population, and in dispersing species, potentially alter population range as a result of fewer individuals to disperse or lower survival whilst dispersing.</p>
</sec>
<sec id="S2.SS2">
<title>2.2. Indirect effects</title>
<p>Changes in one species&#x2019; local presence can have indirect consequences across trophic and taxonomic lines, especially if those relocating are predators or parasites. For example, while bird abundance and diversity decreased near chronic playback of traffic noise, grasshoppers and odonates decreased in areas far from the traffic noise, likely because their predators relocated to those places (<xref ref-type="bibr" rid="B83">Senzaki et al., 2020</xref>). Changes in the predator assemblage can alter both the communities they move to (increased/different predation pressure) and those they move from (lower/different predation pressure). They may alter the predator&#x2013;prey relationships with third parties as predators may be forced to switch to uncommonly eaten prey or start eating prey they have never hunted before. Indirect effects of noise can also fundamentally alter the habitat entire species communities live in, for example, if important species like seed dispersers are impacted. By altering the presence of both pollinators and seed predators/dispersers, noise has been shown to alter both the tree and flower communities that are at the foundation of most terrestrial ecosystems (<xref ref-type="bibr" rid="B27">Francis et al., 2012</xref>; <xref ref-type="bibr" rid="B74">Phillips et al., 2021</xref>).</p>
<p>Noise can also change the interaction with other species in the community. It can alter predator&#x2014;prey interactions, both through the increase of mortality by predation and through changes in foraging success. Similarly, it can change parasite&#x2013;host interactions by affecting parasite species&#x2019; ability to find their hosts (<xref ref-type="bibr" rid="B7">Berkhout et al., 2023</xref>). By masking important cues (<xref ref-type="bibr" rid="B90">Siemers and Schaub, 2011</xref>; <xref ref-type="bibr" rid="B55">Mason et al., 2016</xref>; <xref ref-type="bibr" rid="B84">Senzaki et al., 2016</xref>; <xref ref-type="bibr" rid="B45">Jung et al., 2020</xref>) or reducing hearing abilities e.g., <xref ref-type="bibr" rid="B112">Wysocki and Ladich (2005)</xref>; <xref ref-type="bibr" rid="B51">Ladich (2013)</xref>; <xref ref-type="bibr" rid="B46">Kastelein et al. (2016)</xref>, noise can reduce the distance before which a predator or prey is perceived, and likely impedes localization of a host (<xref ref-type="bibr" rid="B61">McMahon et al., 2017</xref>; <xref ref-type="bibr" rid="B73">Phillips et al., 2019</xref>). Noise can distract from responses to predator or prey, e.g., by increasing food handling and food discrimination errors (<xref ref-type="bibr" rid="B77">Purser and Radford, 2011</xref>; <xref ref-type="bibr" rid="B85">Shafiei Sabet et al., 2015</xref>), or by distracting from alarm signals in another sensory modality (<xref ref-type="bibr" rid="B40">Hasan et al., 2018</xref>).</p>
<p>Noise has both been shown to reduce responses to predation in some species (<xref ref-type="bibr" rid="B93">Simpson et al., 2015</xref>, <xref ref-type="bibr" rid="B94">2016</xref>) but also to increase anti-predator behavior in others (<xref ref-type="bibr" rid="B70">Neo et al., 2014</xref>; <xref ref-type="bibr" rid="B87">Shannon et al., 2014</xref>, <xref ref-type="bibr" rid="B89">2016a</xref>; <xref ref-type="bibr" rid="B107">Voellmy et al., 2014</xref>; <xref ref-type="bibr" rid="B105">van der Knaap et al., 2022</xref>). These changes in the effectiveness in which predators hunt their prey can have multiple outcomes. If predators in a noisy area preferentially switch to new or less commonly hunted prey as they become easier prey, this might exacerbate that prey&#x2019;s local decline, while less drastically affected species may then be temporarily released from predation pressure. Therefore, the outcome of the changes in predator&#x2014;prey interactions can only be predicted on a case-by-case basis. Furthermore, many of the studies that showed these effects only investigated short-term noise exposure, and might therefore have studied behavioral changes that fade over time.</p>
</sec>
</sec>
<sec id="S3">
<title>3. Future directions</title>
<p>There is growing evidence that the impact of chronic noise on animals affects the composition of communities. Direct effects of chronic noise exposure on one species, such as population declines and long-term alterations in behavior, can lead to knock-on consequences for other species in the community. By differentially affecting species, chronic noise has the ability to drastically change community structure and function. Although the growing body of work on this topic is commendable, given the expensive and logistically complex nature of long-term studies, some key knowledge gaps remain. These gaps need to be addressed to ensure effective mitigation of chronic noise exposure. Below, we discuss the main gaps that we see, and suggest possible approaches to fill these.</p>
<p>There is a discrepancy in our understanding of how chronic noise affects different types of communities. The majority of the literature that we reviewed (90%, <xref ref-type="table" rid="T1">Table 1</xref>) investigated terrestrial ecosystems and diurnal species. While studying freshwater and marine ecosystems is more challenging than studying terrestrial systems, the effects of sound are likely to be different for aquatic communities. Sound propagates further in water than on land, so the same noise source will affect a larger area in the water. Furthermore, terrestrial species can often compensate for acoustic information loss by visual signals and perception. This option is less readily available for aquatic species, as light attenuates rapidly in water. Similarly, nocturnal communities might suffer larger impacts from chronic noise disturbance because of their dependence on acoustic signals.</p>
<p>To understand community impacts of noise properly, we must gain a better understanding of the mechanisms underlying species differences in response to the same noise source (<xref ref-type="bibr" rid="B35">Gunn et al., 2022</xref>). While in some instances there is a clear link between species traits and the impact of noise, in other cases this link has not yet been established. For instance, while the effect of chronic compressor noise on birds could be linked to the overlap between their vocalizations and the noise (<xref ref-type="bibr" rid="B28">Francis et al., 2009</xref>, <xref ref-type="bibr" rid="B29">2011a</xref>), it is unknown why changes in fish anti-predator behavior in response to noise exposure vary from diminished responses to faster responses (<xref ref-type="bibr" rid="B107">Voellmy et al., 2014</xref>; <xref ref-type="bibr" rid="B93">Simpson et al., 2015</xref>, <xref ref-type="bibr" rid="B94">2016</xref>; <xref ref-type="bibr" rid="B59">McCormick et al., 2018</xref>; <xref ref-type="bibr" rid="B49">Kok et al., 2021</xref>).</p>
<p>Besides species differences in response, changing species interactions are likely to be an important influence on community change. Parasites that cannot locate their hosts (<xref ref-type="bibr" rid="B61">McMahon et al., 2017</xref>; <xref ref-type="bibr" rid="B73">Phillips et al., 2019</xref>), predators that leave an area (<xref ref-type="bibr" rid="B31">Fr&#x00F6;hlich and Ciach, 2019</xref>), seed dispersers that decline in abundance (<xref ref-type="bibr" rid="B27">Francis et al., 2012</xref>), all will directly impact the species that they are interacting with, whether or not that species is sensitive to noise itself. Because these interactions lead to indirect effects of noise on the community, the outcome may be difficult to predict. When a predator leaves, does that always lead to a release of predation pressure for the prey, or do other predators fill up the gap? Conversely, when the prey leaves, does the predator switch prey type, or does it follow the prey to the new area?</p>
<p>While a lot of the effects of chronic noise on communities have been uncovered with observational studies, understanding the mechanisms behind these effects will require long-term experimental studies. Current experimental studies focus mostly on short-term changes due to noise exposure, while long-term changes in species interactions are likely to have a more profound effect on the community. These long-term changes may be quite different from the changes observed on a short time scale (<xref ref-type="bibr" rid="B49">Kok et al., 2021</xref>). A decrease in foraging behavior, for example, might not persist over time, but might be compensated for when the noise stops. While long-term experimental exposure studies are expensive and logistically challenging, the recent developments in citizen science and technology have opened up possibilities that were previously unheard of, such as continental-scale analysis of acoustic niche partitioning in frogs (<xref ref-type="bibr" rid="B1">Allen-Ankins and Schwarzkopf, 2022</xref>) or biodiversity monitoring of insect communities for which many species are undescribed (<xref ref-type="bibr" rid="B106">van Klink et al., 2022</xref>). Additionally, existing differences in chronic noise exposure in natural settings can be exploited to study community effects, as has been done for the gas compressor stations in North America (<xref ref-type="bibr" rid="B28">Francis et al., 2009</xref>, <xref ref-type="bibr" rid="B29">2011a</xref>,<xref ref-type="bibr" rid="B30">2011b</xref>,<xref ref-type="bibr" rid="B27">2012</xref>; <xref ref-type="bibr" rid="B10">Bunkley et al., 2017</xref>; <xref ref-type="bibr" rid="B74">Phillips et al., 2021</xref>).</p>
<p>Finally, to mitigate effects of chronic noise pollution effectively, we need to know what happens when the noise stops. In contrast to other forms of pollution, noise pollution does not leave long-term traces in the environment once removed. However, does the community immediately change back to pre-noise conditions once noise is removed, or is the changes that occurred due to the noise exposure permanent? The few studies that investigated this have shown mixed results. While white-crowned sparrows (<italic>Zonotrichia leucophrys</italic>) reverted back to low frequency songs (<xref ref-type="bibr" rid="B20">Derryberry et al., 2020</xref>) and spiny chromis (<italic>Acanthochromis polyacanthus</italic>) larvae had increased survival rates (<xref ref-type="bibr" rid="B67">Nedelec et al., 2022</xref>) in quieted conditions, juniper and pinyon seedling recruitment did not recover following the removal of noise (<xref ref-type="bibr" rid="B74">Phillips et al., 2021</xref>). The persistence of effects might depend on the time it takes for the effect to reverse, as well as the behavioral plasticity of the affected species. While the COVID-19 pandemic inadvertently created a natural experiment to test the effects of global quieting (<xref ref-type="bibr" rid="B64">Montgomery et al., 2021</xref>), we should also start experimentally removing noise from communities, to see if the effects of noise can truly be removed by noise mitigation. Those data will be vital in developing effective conservation strategies for the future.</p>
</sec>
<sec id="S4" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in this study are included in the article/<xref ref-type="supplementary-material" rid="FS1">Supplementary material</xref>, further inquiries can be directed to the corresponding author.</p>
</sec>
<sec id="S5" sec-type="author-contributions">
<title>Author contributions</title>
<p>AK, GS, MS, SS, NK, NC, and CW wrote the manuscript. GS, MS, and SS performed the literature research. NC made the figures. AK led the collaboration and guided the writing process. All authors critically reviewed the manuscript, conceived the idea behind the manuscript, approved the submitted version, and revised the manuscript.</p>
</sec>
</body>
<back>
<sec id="S6" sec-type="funding-information">
<title>Funding</title>
<p>This manuscript was conceived during an interdisciplinary workshop entitled &#x201C;Allying Conservation &#x0026; Welfare Research on the Impacts of Human-Generated Sounds on Wild &#x0026; Captive Animals&#x201D; funded by the Association for the Study of Animal Behaviour. The funding for the publication of this manuscript was provided by the Canadian Wildlife Service, Environment and Climate Change Canada. During the writing of this manuscript NC was supported by two grants: Japan Society for the Promotion of Science no. PE21015 and Mitacs Accelerate IT29508.</p>
</sec>
<ack><p>We thank the organizers of the workshop for bringing us together and providing a platform for new ideas. We also thank Patrick O&#x2019;Hara and the Canadian Wildlife Service, Environment and Climate Change Canada for the funds to publish this manuscript.</p>
</ack>
<sec id="S7" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="S8" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
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