Sight, Scent, and Sound: Leveraging the Landscape of Fear for Effective Wildlife Management

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As wildlife populations, including various deer and predator species, increase across the northern hemisphere, conflicts in densely populated areas have also grown, resulting in social and financial costs. Using the “applied ecology of fear” framework, which leverages fear-induced behaviours to influence prey movement, this study systematically reviews deer responses to predator cues. A total of 256 anti-predator behavioural responses were drawn from 39 peer-reviewed studies following standard meta-analysis guidelines and analysed by cue type, with responses clustered into visual, olfactory, and acoustic groups. Results revealed variation in cue efficacy, with predator acoustic cues (predator playbacks) being the most effective to elicit anti-predator responses in deer, followed by olfactory cues (faeces, urine, hair), and visual cues (photographs, body shape). These findings inform sustainable wildlife management strategies that align with ecocentric principles, offering innovative solutions for human-wildlife coexistence challenges. Further research on cue effectiveness and scalability across relevant spatial and temporal scales of conflict will refine these approaches, fostering harmonious interaction between expanding wildlife populations and human communities. Biological sciences/Ecology/Behavioural ecology Biological sciences/Ecology/Conservation Cervid Conservation Human-Wildlife Coexistence Landscape of fear Wildlife Figures Figure 1 Figure 2 Introduction Wildlife populations are rebounding across the northern hemisphere, with various deer species and large carnivores extending their ranges and numbers (Côté et al., 2004 ; Ramirez, 2021 ; Ramirez et al., 2018 ; Weber & Gonzalez, 2003 ). This resurgence positively impacts biodiversity and ecosystems (Bakker & Svenning, 2018 ; Trepel et al., 2024 ), but it also raises challenges in human-wildlife coexistence, especially in densely populated regions where conflicts can result in significant social and financial costs (Linnell et al., 2020 ). As societal perspectives shift toward more ecocentric values emphasizing animal welfare and balanced ecosystems (Herdoiza et al., 2024 , 2025 ), there is growing advocacy for nonlethal wildlife management strategies (van Eeden et al., 2017 ). Although hunting has been a traditional approach, it does not always suffice to mitigate conflicts effectively (Sudharsan et al., 2006 ; Takatsuki, 2009 ). Consequently, interest has risen in managing wildlife behaviour to reduce these conflicts more effectively (Cromsigt et al., 2013 ; Gaynor et al., 2021 ). The “ecology of fear” concept explains how predation risk prompts prey species to adopt anti-predator behaviours, impacting individual fitness and community interactions (Brown et al., 1999 ; Clinchy et al., 2013 ; Zbyryt et al., 2018 ). Expanding on this, Gaynor et al., ( 2021 ) proposed the “applied ecology of fear” framework, which uses fear of real or simulated predators to guide prey species’ behaviour proactively, such as habitat choice and foraging, to reduce human-wildlife conflicts. An example includes using predator cues to discourage deer from entering areas of human interest. Prey detect predator cues visually, olfactory or acoustically, which can trigger diverse anti-predator responses (Chabot et al., 1996 ; Kuijper et al., 2014 ; Li et al., 2011 ; Stankowich & Coss, 2007 ; Widén et al., 2022 ). The effectiveness of these cues varies depending on environmental factors and the prey’s experiences with predators (Berger et al., 2001 ; Gaynor et al., 2019 ). Further, prey face trade-offs between responding to cues and essential activities like foraging, especially when in poor health (Clare et al., 2023 ; Gaynor et al., 2019 ). Prey however can also stop responding to repetitive predator cues to allocate that energy for survival functions, also known as habituation (Apfelbach et al., 2005 ; Blumstein, 2016 ; Rankin et al., 2009 ; Smith et al., 2021 ). To optimize the use of predator cues in wildlife management and address knowledge gaps, this study systematically reviews from published articles behavioural responses within the Cervidae family, which are increasing in numbers and present human-wildlife coexistence concerns (Côté et al., 2004 ; Martin et al., 2020 ; Ramirez, 2019 ; Ramirez et al., 2019 , 2023 ; Ramirez, Jansen, den Ouden, Li, et al., 2021; Ramirez, Jansen, den Ouden, Moktan, et al., 2021). More specifically, this study quantifies deer responses to different types of predator cues clustered by visual, olfactory and acoustic cues. This study will thus, inform researchers and management about the most promising cues -e.g., visual model, body shape, faeces, urine, playbacks- for deterring deer from areas of human interest at relevant spatial and temporal scales of human-wildlife coexistence (Fig. 1 ). Materials and Methods This study employed a systematic approach to conduct the meta-analysis of the peer reviewed literature on the responses of deer to types of predator cues. I retrieved 114 articles by searching on Scopus and Web of Science online data bases by introducing a specific search criteria on the 4th of February of 2023 (see supplementary information S1 for search criteria). The query did not include geographical restriction as this study employs a global approach. This meta-analysis followed the Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA) guidelines, providing standardized procedures for the systematic collection, analysis, and reporting of data in systematic reviews (Page et al., 2021 ). I excluded a total of 84 studies during the screening process by reading the titles and the abstracts (see supplementary informaction S2 for exclution criteria). During the full-text screening phase, I excluded 11 additional articles after closely evaluating their content. However, I also identified 20 new relevant studies through the reference lists of the screened publications. The most frequent reasons for exclusion included articles that were not peer-reviewed, those written in languages other than English, studies focusing on non-Cervidae species, or those addressing topics outside the defined scope of our review, such as intra-specific communication or responses to chemical or natural cues unrelated to predation. Following this process, I finalized a set of 39 peer-reviewed publications (see supplementary information S3 for data source). From each of these studies, I extracted key information including author(s), study location, predator species, cue type (olfactory, visual, or acoustic), cue specific component (e.g, blood, sweath, physical model), deer species, and recorded behavioral responses. Antipredator responses were defined as variation in foraging, crop damage, diel activity, fleeing and patch utilization compared to the base line (Table 1). For studies that examined multiple deer or predator species, I distilled separate data points for each unique predator-prey combination. See Ramirez et al., ( 2024 ) for more information on the methods used for this study. Due to the diversity of statistical methods and response metrics across studies and the fact that effect sizes were often unreported, I did not rely on exact effect sizes. Instead, I adopted a binary classification system, categorizing each response as either the presence or absence of an anti-predator reaction to a predator cue. This approach enabled consistent comparisons across studies with highly varied methodologies. A p-value threshold of < 0.05 was used to distinguish statistically significant anti-predator responses from non-responses. I then clustered the behavioural responses of deer as visual, olfactory and acoustic for the statistical analysis. I constructed contingency tables for three chi-square tests (including 2000 simulations) by setting anti-predator behavior by deer (as yes vs. no) as response and cues from predators as the predictor, clustered by type of visual, olfactory and acoustic cues. Variables with < 3 samples were omitted from the analysis to control for bias results. I treated behavioral responses independently and not as nested because of three reasons. Accounting for the nested designs would first lead to insufficient number of replicates for an advanced analysis. Second, the studies screened significantly vary in their methodological approach and thus, it is unrealistic to statistically account also for this variation. Third, the opted statistical analysis informs researchers and practitioners in a straightforward way about the effectiveness of different type of predator cues to promote human-wildlife coexistence in areas of human interest. Results In terms of geographical distribution, responses were registered in 12 different countries located in the northern hemisphere (13 territories when including Greenland, Table 2). A total of 17 predator species (subspecies counted individually) were simulates and the responses of 11 deer species were recoreded (subspecies counted individually, Table 3–4). A total of 256 antipredator behavioural responses were recorded from the 39 articles, from which 34 were visual, 133 olfactory and 89 acoustic (supplemenaty information S4 for individual responses). There was a large variation in type of predator cues with olfactory cues being the richest (including blood, faeces, urine, hair, dead body, cologne and sweat, Fig. 2 ) followed by visual (including body shape, photogram and human object) and acoustic (including animal playbacks). The effectiveness of the cues varied according to the type of component used in each of the sense categories: visual, olfactory and acoustic. There was a difference in anti-predator response between type of component in visual cues (x 2 = 5.23, p = 0.036) and the strongest components were body shape (in 100%, n = 5, of all cases employing predator ‘s body shape) and real size photograph (44%, n = 27, Fig. 2 ). There was a difference in anti-predator response between type of component in olfactory cues (x 2 = 12.16, p = 0.003) and the strongest component ranked as faeces (in 72%, n = 47, of all cases employing faeces from predators), urine (58%, n = 62), hair (57%, n = 14) and cologne (0%, n = 6). All acoustic experiments (n = 89) used animal playbacks as cues, with an overall effectiveness rate of 85%. This effect was statistically significant (x 2 = 44.60, p < 0.001). Please refer to Ramirez et al., ( 2024 ) for an overview of results not included in this publication but that are relevant for this study; for example, effectivenes of predator cues by deer foraging strategy and predator hunting strategy, pen vs field experiments, and habituation. Discussion There were important variations in the effectiveness of predator cues, suggesting that certain types may be more suitable for deterring deer from areas of human interest at spatial and temporal scales relevant to human–wildlife coexistence. Understanding these differences is crucial for developing targeted, context-specific strategies to mitigate negative human-deer interactions effectively. In the case of visual cues, the visual presence (body shape) of a predator is extremely useful for managing wildlife behaviour. Human presence for example triggered anti-predator responses in deer, particularly in fawns. Yet the relationship weakens when deer were in large groups (i.e., safety in numbers) and with increasing distance between deer and human (Espmark & Langvatn, 1985 ; LaGory, 1987 ; Padié et al., 2015 ). The practicality of introducing to a particular environment the body of a non-human predator for wildlife management is not entirely feasible because of the logistical challenges and the ethical concerns. A second option is to use models that replicate the visual presence of a predator, yet this had limited effectiveness in stimulating antipredator responses in deer. The main factor that modulates anti-predator responses of deer is whether the visual model represents a naïve or a co-evolutionary predator (Li et al., 2011 ; Stankowich & Coss, 2007 ). Deer are generally expected to exhibit stronger anti-predator responses to co-evolutionary predators—those they have encountered over evolutionary time—than to naïve or novel predators. This pattern can be explained by the multi-predator hypothesis (Blumstein, 2006), which suggests that prey species may retain a generalized recognition of predator traits, such as body shape or movement, allowing them to respond to a range of predatory threats. However, the degree to which deer perceive naïve predators as risky can vary. For instance, if a novel predator’s appearance, particularly its camouflage pattern or coloration, does not resemble that of historical predators, the deer may fail to recognize it as a threat (Stankowich & Coss, 2007 ).This highlights the importance of both evolutionary history and visual similarity in shaping prey responses to predator cues. It also suggests that predator recognition is not solely based on form, but also on specific visual patterns associated with threat. In the case of olfactory cues, faeces were the most effective, followed closely by urine and hair, and these cues have been widely tested in the northern hemisphere (Berger et al., 2001 ; Chabot et al., 1996 ; Kuijper et al., 2014 ; Osada et al., 2014 ; Sullivan et al., 1985 ). From an applied management perspective, faeces and hair are among the most practical predator cues to deploy at larger spatial scales aimed at promoting human–wildlife coexistence. These materials can be readily and ethically collected from areas where predators naturally occur, avoiding the significant animal welfare concerns associated with obtaining urine or blood, which typically require invasive procedures. Their accessibility makes them promising candidates for use in deterrence strategies across broad landscapes. However, several challenges remain. Critically, the effective dose needed to elicit a reliable anti-predator response in prey species is still poorly understood. Furthermore, the longevity of these cues is uncertain, as organic materials such as faeces and hair degrade over time due to environmental exposure, including decomposition, desiccation, and microbial activity (Ramirez et al., 2024 ; Sullivan et al., 1985 ). These limitations highlight the need for further research to determine not only optimal deployment strategies but also how long such cues remain effective in real-world conditions. Acoustic cues in the form of predator playbacks showed strong potential to manage deer behaviour, especially since predator playbacks have been extensively tested in wide variety of environments and are easily scalable to manage wildlife-human interactions at relevant temporal and spatial scales of conflict (Berger, 2007 ; Berger et al., 2001 ). However, it remains unclear to what extent does repetitive predator playbacks lead to diminished cue efficacy and eventually deer habituation; for example, if individuals learn that the cue is not followed by actual predation risk (Cromsigt et al., 2013 ). Since habituation is a dynamic, context-dependent process (Blumstein, 2016 ; Rankin et al., 2009 ), future studies should examine deer responses across a variety of ecological conditions, including different times of day and seasons, fluctuations in food availability and quality, levels of sociality among prey, and variations in playback schemes (e.g., frequency, volume, and duration). For current management efforts, strategies should consider rotating cue locations, adjusting playback parameters, incorporating multiple cue types, or occasionally reinforcing cues with real threats to minimize the risk of desensitization. From an applied perspective, wildlife managers can enhance the effectiveness of predator cues by rotating cue locations, incorporating multiple cue types, adjusting cue dosage, and occasionally reinforcing cues with real threats to minimize the risk of desensitization (Ramirez et al., 2024 ). These strategies are particularly advantageous in areas prone to frequent conflicts, as they allow for adaptive management tailored to the specific dynamics of each situation. Additionally, inducing a landscape of fear can be more cost-effective than other forms of wildlife management, such as hunting, relocation, or the use of barriers, as predator cues are inexpensive and scalable. These approaches align with the growing preference for ethical, ecocentric strategies in modern wildlife conservation. Conclusion The application of these results is meant for researchers and wildlife managers to identify the most promising type of predator cue for their system and to subsequently test the cues in future experiments, scale the distribution of cues to real spatiotemporal scenarios and steer deer away from areas of human interest. Visual and olfactory cues require further testing given their prelaminar potential to stimulate antipredator responses in prey and the large variation of visual and scent option to experiment with. Overall, it is expected that wildlife will present stronger antipredator responses to predator playbacks (acoustic) in general. Future research should experiment with the process of habituation to predator cues and the effectiveness of presenting visual, olfactory and acoustic cues simultaneously for promoting human-wildlife coexistence. Declarations Acknowledgements JIR’s salary during this project was kindly provided by The Arctic University of Norway. Funding: not applicable Conflict of interest: The authors declare no conflict of interest associated to this publication. Ethics approval: not applicable Consent to participate: not applicable Available of data and material: Data associated to this publication is included as supplementary information. Code availability: not applicable Authors’ contributions: J.I.R. conducted and led all aspects of the research. N.H. extracted and curated data from literature, wrote part of the discussion and provided cfeedback. All authors reviewed and approved the manuscript. References Apfelbach, R., Blanchard, C. D., Blanchard, R. J., Hayes, R. A., & McGregor, I. S. (2005). The effects of predator odors in mammalian prey species: a review of field and laboratory studies. Neuroscience & Biobehavioral Reviews , 29 (8), 1123–1144. https://doi.org/10.1016/j.neubiorev.2005.05.005 Bakker, E. S., & Svenning, J. C. (2018). Trophic rewilding: impact on ecosystems under global change. 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Do wild ungulates experience higher stress with humans than with large carnivores? Behavioral Ecology , 29 (1), 19–30. https://doi.org/10.1093/beheco/arx142 Tables Tables 1. List and description of anti-predator behavioural responses of deer to predator cues. The table is a partially modified version of table 1 in Ramirez et al., (2024) Response name Response description Presence of anti-predator response Evading behaviour Variation in behaviour such as vigilance, snorting, tail flagging & stamping Increase in behaviour incidence Crop damage Variation in crop damage Decrease in crop damage Diel activity Variation in daily activity pattern compared to baseline activity Increase or decrease in activity Fleeing behaviour Variation in incidence of flee Increase flee incidence Foraging behaviour Variation in foraging time, food consumption or food selection. Some studies reported crop damage as a proxy for foraging. Decrease foraging time & quality Heart rate Variation in heart rate compared to baseline Increase or decrease in heart rate Patch use Variation in time spend at a specific patch compared to baseline Increase or decrease utilization Visitation Variation in visitation to a specific site or visitation span Decrease in visitation Tables 2. List of countries where experiments were conducted from the collection of reviewed studies. Countries or Territories with number of studies Canada - 4 China - 1 Denmark - 1 France - 1 Gernmany - 1 Greenland - 1 Japan - 1 Norway - 3 Poland - 3 Russia - 1 Sweden - 4 The Netherlands - 2 USA - 23 Tables 3 . List of predator species simulated in the behavioural experiments from the collection of reviewed studies. Number of studies per predator species or subspecies Canis latrans - 13 Canis lupus - 18 Canis lupus familiaris -7 Gulo gulo -1 Homo sapiens - 15 Lynx lynx - 3 Lynx rufus - 5 Panthera leo - 2 Panthera onca - 2 Panthera pardus -2 Panthera tigirs - 5 Panthera uncia - 1 Puma concolor - 7 Ursus americanus - 3 Ursus arctos - 2 Ursus arctos horribilis -2 Vulpes vulpes - 3 Tables 4. List of deer species included in the behavioural experiments from the collection of reviewed studies. Number of studies per deer species or subspecies Alces alces - 7 Capreolus capreolus - 7 Cervus canadensis - 3 Cervus elaphus - 8 Cervus nippon - 1 Dama dama - 3 Elaphurus davidianus -1 Odocoileus hemionus - 2 Odocoileus h. columbianus - 7 Odocoileus virginianus -13 Rangifer tarandus -1 Additional Declarations No competing interests reported. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6955116","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":502177443,"identity":"26e63f86-afb2-42e3-93ec-c17c4325c8c5","order_by":0,"name":"Juan Ignacio Ramirez","email":"data:image/png;base64,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","orcid":"","institution":"UiT The Arctic University of Norway","correspondingAuthor":true,"prefix":"","firstName":"Juan","middleName":"Ignacio","lastName":"Ramirez","suffix":""},{"id":502177444,"identity":"5abaed81-a958-44b6-9030-12b0b5893f55","order_by":1,"name":"Natalie Herdoiza","email":"","orcid":"","institution":"Utrecht University","correspondingAuthor":false,"prefix":"","firstName":"Natalie","middleName":"","lastName":"Herdoiza","suffix":""}],"badges":[],"createdAt":"2025-06-23 09:23:36","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6955116/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6955116/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":89602632,"identity":"fe080db1-c218-4798-87d1-ed9c5f6f7e99","added_by":"auto","created_at":"2025-08-21 18:44:30","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":539307,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eImage illustrating the application of the landscape of fear for human-wildlife coexistence. This AI-generated image was created using ChatGPT with the following promt: wolf inducing fear in a forest inhabited by deer drawn with pencil.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-6955116/v1/17ccae4906e162c5dc0060d1.png"},{"id":89602630,"identity":"171923dd-6c3b-4ba9-8fd8-3ae88a406ff0","added_by":"auto","created_at":"2025-08-21 18:44:30","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":51064,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cem\u003eBar graph for the percentage of anti-predator responses by deer to predator cues clustered by type of cue. The number between parenthesis on the x-axis indicates the total number of responses. Species with n\u0026lt;3 were not included in the analysis. Level of significance is presented in the x-axis title: p value= *** \u0026lt;0.001, ** \u0026lt;0.01, * \u0026lt;0.05.\u003c/em\u003e\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-6955116/v1/01c0cb2ef0b78fad2b2033aa.png"},{"id":100358243,"identity":"207ccd22-13e4-478d-800a-49ac5f6064af","added_by":"auto","created_at":"2026-01-16 07:20:47","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1144103,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6955116/v1/f9b7bc8e-dc4a-43d5-baea-d3d1e419f325.pdf"},{"id":89602848,"identity":"48ee7228-e30f-4c21-a8a6-5405f5d4ef73","added_by":"auto","created_at":"2025-08-21 18:52:30","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":136221,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryFear2.docx","url":"https://assets-eu.researchsquare.com/files/rs-6955116/v1/207c5d165763bf5414441856.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Sight, Scent, and Sound: Leveraging the Landscape of Fear for Effective Wildlife Management","fulltext":[{"header":"Introduction","content":"\u003cp\u003eWildlife populations are rebounding across the northern hemisphere, with various deer species and large carnivores extending their ranges and numbers (C\u0026ocirc;t\u0026eacute; et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Ramirez, \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Ramirez et al., \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Weber \u0026amp; Gonzalez, \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). This resurgence positively impacts biodiversity and ecosystems (Bakker \u0026amp; Svenning, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Trepel et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), but it also raises challenges in human-wildlife coexistence, especially in densely populated regions where conflicts can result in significant social and financial costs (Linnell et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). As societal perspectives shift toward more ecocentric values emphasizing animal welfare and balanced ecosystems (Herdoiza et al., \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2024\u003c/span\u003e, \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), there is growing advocacy for nonlethal wildlife management strategies (van Eeden et al., \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). Although hunting has been a traditional approach, it does not always suffice to mitigate conflicts effectively (Sudharsan et al., \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Takatsuki, \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2009\u003c/span\u003e). Consequently, interest has risen in managing wildlife behaviour to reduce these conflicts more effectively (Cromsigt et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Gaynor et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eThe \u0026ldquo;ecology of fear\u0026rdquo; concept explains how predation risk prompts prey species to adopt anti-predator behaviours, impacting individual fitness and community interactions (Brown et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e1999\u003c/span\u003e; Clinchy et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Zbyryt et al., \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Expanding on this, Gaynor et al., (\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) proposed the \u0026ldquo;applied ecology of fear\u0026rdquo; framework, which uses fear of real or simulated predators to guide prey species\u0026rsquo; behaviour proactively, such as habitat choice and foraging, to reduce human-wildlife conflicts. An example includes using predator cues to discourage deer from entering areas of human interest.\u003c/p\u003e\u003cp\u003ePrey detect predator cues visually, olfactory or acoustically, which can trigger diverse anti-predator responses (Chabot et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Kuijper et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Li et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Stankowich \u0026amp; Coss, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Wid\u0026eacute;n et al., \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). The effectiveness of these cues varies depending on environmental factors and the prey\u0026rsquo;s experiences with predators (Berger et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Gaynor et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Further, prey face trade-offs between responding to cues and essential activities like foraging, especially when in poor health (Clare et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Gaynor et al., \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Prey however can also stop responding to repetitive predator cues to allocate that energy for survival functions, also known as habituation (Apfelbach et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Blumstein, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Rankin et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Smith et al., \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eTo optimize the use of predator cues in wildlife management and address knowledge gaps, this study systematically reviews from published articles behavioural responses within the Cervidae family, which are increasing in numbers and present human-wildlife coexistence concerns (C\u0026ocirc;t\u0026eacute; et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Martin et al., \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Ramirez, \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Ramirez et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2019\u003c/span\u003e, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2023\u003c/span\u003e; Ramirez, Jansen, den Ouden, Li, et al., 2021; Ramirez, Jansen, den Ouden, Moktan, et al., 2021). More specifically, this study quantifies deer responses to different types of predator cues clustered by visual, olfactory and acoustic cues. This study will thus, inform researchers and management about the most promising cues -e.g., visual model, body shape, faeces, urine, playbacks- for deterring deer from areas of human interest at relevant spatial and temporal scales of human-wildlife coexistence (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003eThis study employed a systematic approach to conduct the meta-analysis of the peer reviewed literature on the responses of deer to types of predator cues. I retrieved 114 articles by searching on Scopus and Web of Science online data bases by introducing a specific search criteria on the 4th of February of 2023 (see supplementary information S1 for search criteria). The query did not include geographical restriction as this study employs a global approach. This meta-analysis followed the Preferred Reporting Items for Systematic Reviews and Meta-Analyses (PRISMA) guidelines, providing standardized procedures for the systematic collection, analysis, and reporting of data in systematic reviews (Page et al., \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e\u003cp\u003eI excluded a total of 84 studies during the screening process by reading the titles and the abstracts (see supplementary informaction S2 for exclution criteria). During the full-text screening phase, I excluded 11 additional articles after closely evaluating their content. However, I also identified 20 new relevant studies through the reference lists of the screened publications. The most frequent reasons for exclusion included articles that were not peer-reviewed, those written in languages other than English, studies focusing on non-Cervidae species, or those addressing topics outside the defined scope of our review, such as intra-specific communication or responses to chemical or natural cues unrelated to predation. Following this process, I finalized a set of 39 peer-reviewed publications (see supplementary information S3 for data source). From each of these studies, I extracted key information including author(s), study location, predator species, cue type (olfactory, visual, or acoustic), cue specific component (e.g, blood, sweath, physical model), deer species, and recorded behavioral responses. Antipredator responses were defined as variation in foraging, crop damage, diel activity, fleeing and patch utilization compared to the base line (Table\u0026nbsp;1). For studies that examined multiple deer or predator species, I distilled separate data points for each unique predator-prey combination. See Ramirez et al., (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) for more information on the methods used for this study.\u003c/p\u003e\u003cp\u003eDue to the diversity of statistical methods and response metrics across studies and the fact that effect sizes were often unreported, I did not rely on exact effect sizes. Instead, I adopted a binary classification system, categorizing each response as either the presence or absence of an anti-predator reaction to a predator cue. This approach enabled consistent comparisons across studies with highly varied methodologies. A p-value threshold of \u0026lt;\u0026thinsp;0.05 was used to distinguish statistically significant anti-predator responses from non-responses. I then clustered the behavioural responses of deer as visual, olfactory and acoustic for the statistical analysis. I constructed contingency tables for three chi-square tests (including 2000 simulations) by setting anti-predator behavior by deer (as yes vs. no) as response and cues from predators as the predictor, clustered by type of visual, olfactory and acoustic cues. Variables with \u0026lt;\u0026thinsp;3 samples were omitted from the analysis to control for bias results. I treated behavioral responses independently and not as nested because of three reasons. Accounting for the nested designs would first lead to insufficient number of replicates for an advanced analysis. Second, the studies screened significantly vary in their methodological approach and thus, it is unrealistic to statistically account also for this variation. Third, the opted statistical analysis informs researchers and practitioners in a straightforward way about the effectiveness of different type of predator cues to promote human-wildlife coexistence in areas of human interest.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003eIn terms of geographical distribution, responses were registered in 12 different countries located in the northern hemisphere (13 territories when including Greenland, Table\u0026nbsp;2). A total of 17 predator species (subspecies counted individually) were simulates and the responses of 11 deer species were recoreded (subspecies counted individually, Table\u0026nbsp;3\u0026ndash;4). A total of 256 antipredator behavioural responses were recorded from the 39 articles, from which 34 were visual, 133 olfactory and 89 acoustic (supplemenaty information S4 for individual responses). There was a large variation in type of predator cues with olfactory cues being the richest (including blood, faeces, urine, hair, dead body, cologne and sweat, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) followed by visual (including body shape, photogram and human object) and acoustic (including animal playbacks).\u003c/p\u003e\u003cp\u003e\u003c/p\u003e\u003cp\u003e The effectiveness of the cues varied according to the type of component used in each of the sense categories: visual, olfactory and acoustic. There was a difference in anti-predator response between type of component in visual cues (x\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;5.23, p\u0026thinsp;=\u0026thinsp;0.036) and the strongest components were body shape (in 100%, n\u0026thinsp;=\u0026thinsp;5, of all cases employing predator \u0026lsquo;s body shape) and real size photograph (44%, n\u0026thinsp;=\u0026thinsp;27, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). There was a difference in anti-predator response between type of component in olfactory cues (x\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;12.16, p\u0026thinsp;=\u0026thinsp;0.003) and the strongest component ranked as faeces (in 72%, n\u0026thinsp;=\u0026thinsp;47, of all cases employing faeces from predators), urine (58%, n\u0026thinsp;=\u0026thinsp;62), hair (57%, n\u0026thinsp;=\u0026thinsp;14) and cologne (0%, n\u0026thinsp;=\u0026thinsp;6). All acoustic experiments (n\u0026thinsp;=\u0026thinsp;89) used animal playbacks as cues, with an overall effectiveness rate of 85%. This effect was statistically significant (x\u003csup\u003e2\u003c/sup\u003e\u0026thinsp;=\u0026thinsp;44.60, p\u0026thinsp;\u0026lt;\u0026thinsp;0.001). Please refer to Ramirez et al., (\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) for an overview of results not included in this publication but that are relevant for this study; for example, effectivenes of predator cues by deer foraging strategy and predator hunting strategy, pen vs field experiments, and habituation.\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThere were important variations in the effectiveness of predator cues, suggesting that certain types may be more suitable for deterring deer from areas of human interest at spatial and temporal scales relevant to human\u0026ndash;wildlife coexistence. Understanding these differences is crucial for developing targeted, context-specific strategies to mitigate negative human-deer interactions effectively.\u003c/p\u003e\u003cp\u003eIn the case of visual cues, the visual presence (body shape) of a predator is extremely useful for managing wildlife behaviour. Human presence for example triggered anti-predator responses in deer, particularly in fawns. Yet the relationship weakens when deer were in large groups (i.e., safety in numbers) and with increasing distance between deer and human (Espmark \u0026amp; Langvatn, \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e1985\u003c/span\u003e; LaGory, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1987\u003c/span\u003e; Padi\u0026eacute; et al., \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). The practicality of introducing to a particular environment the body of a non-human predator for wildlife management is not entirely feasible because of the logistical challenges and the ethical concerns. A second option is to use models that replicate the visual presence of a predator, yet this had limited effectiveness in stimulating antipredator responses in deer. The main factor that modulates anti-predator responses of deer is whether the visual model represents a na\u0026iuml;ve or a co-evolutionary predator (Li et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Stankowich \u0026amp; Coss, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). Deer are generally expected to exhibit stronger anti-predator responses to co-evolutionary predators\u0026mdash;those they have encountered over evolutionary time\u0026mdash;than to na\u0026iuml;ve or novel predators. This pattern can be explained by the multi-predator hypothesis (Blumstein, 2006), which suggests that prey species may retain a generalized recognition of predator traits, such as body shape or movement, allowing them to respond to a range of predatory threats. However, the degree to which deer perceive na\u0026iuml;ve predators as risky can vary. For instance, if a novel predator\u0026rsquo;s appearance, particularly its camouflage pattern or coloration, does not resemble that of historical predators, the deer may fail to recognize it as a threat (Stankowich \u0026amp; Coss, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2007\u003c/span\u003e).This highlights the importance of both evolutionary history and visual similarity in shaping prey responses to predator cues. It also suggests that predator recognition is not solely based on form, but also on specific visual patterns associated with threat.\u003c/p\u003e\u003cp\u003eIn the case of olfactory cues, faeces were the most effective, followed closely by urine and hair, and these cues have been widely tested in the northern hemisphere (Berger et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Chabot et al., \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1996\u003c/span\u003e; Kuijper et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Osada et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Sullivan et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1985\u003c/span\u003e). From an applied management perspective, faeces and hair are among the most practical predator cues to deploy at larger spatial scales aimed at promoting human\u0026ndash;wildlife coexistence. These materials can be readily and ethically collected from areas where predators naturally occur, avoiding the significant animal welfare concerns associated with obtaining urine or blood, which typically require invasive procedures. Their accessibility makes them promising candidates for use in deterrence strategies across broad landscapes. However, several challenges remain. Critically, the effective dose needed to elicit a reliable anti-predator response in prey species is still poorly understood. Furthermore, the longevity of these cues is uncertain, as organic materials such as faeces and hair degrade over time due to environmental exposure, including decomposition, desiccation, and microbial activity (Ramirez et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Sullivan et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1985\u003c/span\u003e). These limitations highlight the need for further research to determine not only optimal deployment strategies but also how long such cues remain effective in real-world conditions.\u003c/p\u003e\u003cp\u003eAcoustic cues in the form of predator playbacks showed strong potential to manage deer behaviour, especially since predator playbacks have been extensively tested in wide variety of environments and are easily scalable to manage wildlife-human interactions at relevant temporal and spatial scales of conflict (Berger, \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Berger et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). However, it remains unclear to what extent does repetitive predator playbacks lead to diminished cue efficacy and eventually deer habituation; for example, if individuals learn that the cue is not followed by actual predation risk (Cromsigt et al., \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). Since habituation is a dynamic, context-dependent process (Blumstein, \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Rankin et al., \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2009\u003c/span\u003e), future studies should examine deer responses across a variety of ecological conditions, including different times of day and seasons, fluctuations in food availability and quality, levels of sociality among prey, and variations in playback schemes (e.g., frequency, volume, and duration). For current management efforts, strategies should consider rotating cue locations, adjusting playback parameters, incorporating multiple cue types, or occasionally reinforcing cues with real threats to minimize the risk of desensitization.\u003c/p\u003e\u003cp\u003eFrom an applied perspective, wildlife managers can enhance the effectiveness of predator cues by rotating cue locations, incorporating multiple cue types, adjusting cue dosage, and occasionally reinforcing cues with real threats to minimize the risk of desensitization (Ramirez et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). These strategies are particularly advantageous in areas prone to frequent conflicts, as they allow for adaptive management tailored to the specific dynamics of each situation. Additionally, inducing a landscape of fear can be more cost-effective than other forms of wildlife management, such as hunting, relocation, or the use of barriers, as predator cues are inexpensive and scalable. These approaches align with the growing preference for ethical, ecocentric strategies in modern wildlife conservation.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThe application of these results is meant for researchers and wildlife managers to identify the most promising type of predator cue for their system and to subsequently test the cues in future experiments, scale the distribution of cues to real spatiotemporal scenarios and steer deer away from areas of human interest. Visual and olfactory cues require further testing given their prelaminar potential to stimulate antipredator responses in prey and the large variation of visual and scent option to experiment with. Overall, it is expected that wildlife will present stronger antipredator responses to predator playbacks (acoustic) in general. Future research should experiment with the process of habituation to predator cues and the effectiveness of presenting visual, olfactory and acoustic cues simultaneously for promoting human-wildlife coexistence.\u003c/p\u003e"},{"header":"Declarations","content":"\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eJIR\u0026rsquo;s salary during this project was kindly provided by The Arctic University of Norway.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding:\u0026nbsp;\u003c/strong\u003enot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflict of interest:\u0026nbsp;\u003c/strong\u003eThe authors declare no conflict of interest associated to this publication.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval:\u0026nbsp;\u003c/strong\u003enot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to participate:\u0026nbsp;\u003c/strong\u003enot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailable of data and material:\u0026nbsp;\u003c/strong\u003eData associated to this publication is included as supplementary information.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCode availability:\u0026nbsp;\u003c/strong\u003enot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors\u0026rsquo; contributions:\u0026nbsp;\u003c/strong\u003eJ.I.R. conducted and led all aspects of the research. N.H. \u0026nbsp;extracted and curated data from literature, wrote part of the discussion and provided cfeedback. All authors reviewed and approved the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eApfelbach, R., Blanchard, C. D., Blanchard, R. J., Hayes, R. A., \u0026amp; McGregor, I. S. (2005). The effects of predator odors in mammalian prey species: a review of field and laboratory studies. \u003cem\u003eNeuroscience \u0026amp; Biobehavioral Reviews\u003c/em\u003e, \u003cem\u003e29\u003c/em\u003e(8), 1123\u0026ndash;1144. https://doi.org/10.1016/j.neubiorev.2005.05.005\u003c/li\u003e\n\u003cli\u003eBakker, E. S., \u0026amp; Svenning, J. C. (2018). Trophic rewilding: impact on ecosystems under global change. \u003cem\u003ePhilosophical Transactions of the Royal Society B: Biological Sciences\u003c/em\u003e, \u003cem\u003e373\u003c/em\u003e(1761), 20170432. https://doi.org/10.1098/rstb.2017.0432\u003c/li\u003e\n\u003cli\u003eBerger, J. 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R., Ritchie, E. G., \u0026amp; Newsome, T. M. (2017). Shifting public values and what they mean for increasing democracy in wildlife management decisions. \u003cem\u003eBiodiversity and Conservation\u003c/em\u003e, \u003cem\u003e26\u003c/em\u003e, 2759\u0026ndash;2763. https://doi.org/10.1007/s10531-017-1378-9\u003c/li\u003e\n\u003cli\u003eWeber, M., \u0026amp; Gonzalez, S. (2003). Latin American deer diversity and conservation: A review of status and distribution. \u003cem\u003eEcoscience\u003c/em\u003e, \u003cem\u003e10\u003c/em\u003e(4), 443\u0026ndash;454. https://doi.org/10.1080/11956860.2003.11682792\u003c/li\u003e\n\u003cli\u003eWid\u0026eacute;n, A., Clinchy, M., Felton, A. M., Hofmeester, T. R., Kuijper, D. P. J., Singh, N. J., Widemo, F., Zanette, L. Y., \u0026amp; Cromsigt, J. P. G. M. (2022). Playbacks of predator vocalizations reduce crop damage by ungulates. \u003cem\u003eAgriculture, Ecosystems \u0026amp; Environment\u003c/em\u003e, \u003cem\u003e328\u003c/em\u003e, 107853. https://doi.org/10.1016/j.agee.2022.107853\u003c/li\u003e\n\u003cli\u003eZbyryt, A., Bubnicki, J. W., Kuijper, D. P. J., Dehnhard, M., Churski, M., \u0026amp; Schmidt, K. (2018). Do wild ungulates experience higher stress with humans than with large carnivores? \u003cem\u003eBehavioral Ecology\u003c/em\u003e, \u003cem\u003e29\u003c/em\u003e(1), 19\u0026ndash;30. https://doi.org/10.1093/beheco/arx142\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cem\u003eTables 1. List and description of anti-predator behavioural responses of deer to predator cues.\u0026nbsp;The table is a partially modified version of table 1 in Ramirez et al., (2024)\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"598\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 18.7605%;\"\u003e\n \u003cp\u003eResponse name\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 43.7186%;\"\u003e\n \u003cp\u003eResponse description\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 37.5209%;\"\u003e\n \u003cp\u003ePresence of anti-predator response\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 18.7605%;\"\u003e\n \u003cp\u003eEvading behaviour\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 43.7186%;\"\u003e\n \u003cp\u003eVariation in behaviour such as vigilance, snorting, tail flagging \u0026amp; stamping\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 37.5209%;\"\u003e\n \u003cp\u003eIncrease in behaviour incidence\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 18.7605%;\"\u003e\n \u003cp\u003eCrop damage\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 43.7186%;\"\u003e\n \u003cp\u003eVariation in crop damage\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 37.5209%;\"\u003e\n \u003cp\u003eDecrease in crop damage\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 18.7605%;\"\u003e\n \u003cp\u003eDiel activity\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 43.7186%;\"\u003e\n \u003cp\u003eVariation in daily activity pattern compared to baseline activity\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 37.5209%;\"\u003e\n \u003cp\u003eIncrease or decrease in activity\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 18.7605%;\"\u003e\n \u003cp\u003eFleeing behaviour\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 43.7186%;\"\u003e\n \u003cp\u003eVariation in incidence of flee\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 37.5209%;\"\u003e\n \u003cp\u003eIncrease flee incidence\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 18.7605%;\"\u003e\n \u003cp\u003eForaging behaviour\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 43.7186%;\"\u003e\n \u003cp\u003eVariation in foraging time, food consumption or food selection. Some studies reported crop damage as a proxy for foraging.\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 37.5209%;\"\u003e\n \u003cp\u003eDecrease foraging time \u0026amp; quality\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 18.7605%;\"\u003e\n \u003cp\u003eHeart rate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 43.7186%;\"\u003e\n \u003cp\u003eVariation in heart rate compared to baseline\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 37.5209%;\"\u003e\n \u003cp\u003eIncrease or decrease in heart rate\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 18.7605%;\"\u003e\n \u003cp\u003ePatch use\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 43.7186%;\"\u003e\n \u003cp\u003eVariation in time spend at a specific patch compared to baseline\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 37.5209%;\"\u003e\n \u003cp\u003eIncrease or decrease utilization\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 18.7605%;\"\u003e\n \u003cp\u003eVisitation\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 43.7186%;\"\u003e\n \u003cp\u003eVariation in visitation to a specific site or visitation span\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 37.5209%;\"\u003e\n \u003cp\u003eDecrease in visitation\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cem\u003eTables 2. List of countries where experiments were conducted from the collection of reviewed studies.\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"580\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"4\" style=\"width: 580px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eCountries or Territories with number of studies\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 130px;\"\u003e\n \u003cp\u003e\u003cem\u003eCanada - 4\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 131px;\"\u003e\n \u003cp\u003e\u003cem\u003eChina - 1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp\u003e\u003cem\u003eDenmark - 1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 202px;\"\u003e\n \u003cp\u003e\u003cem\u003eFrance - 1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 130px;\"\u003e\n \u003cp\u003e\u003cem\u003eGernmany - 1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 131px;\"\u003e\n \u003cp\u003e\u003cem\u003eGreenland - 1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp\u003e\u003cem\u003eJapan - 1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 202px;\"\u003e\n \u003cp\u003e\u003cem\u003eNorway - 3\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 130px;\"\u003e\n \u003cp\u003e\u003cem\u003ePoland - 3\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 131px;\"\u003e\n \u003cp\u003e\u003cem\u003eRussia - 1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp\u003e\u003cem\u003eSweden - 4\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 202px;\"\u003e\n \u003cp\u003e\u003cem\u003eThe Netherlands - 2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 130px;\"\u003e\n \u003cp\u003e\u003cem\u003eUSA - 23\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 131px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 116px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 202px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cem\u003eTables 3\u003c/em\u003e\u003cem\u003e. List of predator species simulated in the behavioural experiments from the collection of reviewed studies.\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"561\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"3\" style=\"width: 561px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNumber of studies per predator species or subspecies\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 199px;\"\u003e\n \u003cp\u003e\u003cem\u003eCanis latrans - 13\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 168px;\"\u003e\n \u003cp\u003e\u003cem\u003eCanis lupus - 18\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 194px;\"\u003e\n \u003cp\u003e\u003cem\u003eCanis lupus familiaris -7\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 199px;\"\u003e\n \u003cp\u003e\u003cem\u003eGulo gulo -1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 168px;\"\u003e\n \u003cp\u003e\u003cem\u003eHomo sapiens - 15\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 194px;\"\u003e\n \u003cp\u003e\u003cem\u003eLynx lynx - 3\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 199px;\"\u003e\n \u003cp\u003e\u003cem\u003eLynx rufus - 5\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 168px;\"\u003e\n \u003cp\u003e\u003cem\u003ePanthera leo - 2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 194px;\"\u003e\n \u003cp\u003e\u003cem\u003ePanthera onca - 2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 199px;\"\u003e\n \u003cp\u003e\u003cem\u003ePanthera pardus -2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 168px;\"\u003e\n \u003cp\u003e\u003cem\u003ePanthera tigirs - 5\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 194px;\"\u003e\n \u003cp\u003e\u003cem\u003ePanthera uncia - 1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 199px;\"\u003e\n \u003cp\u003e\u003cem\u003ePuma concolor - 7\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 168px;\"\u003e\n \u003cp\u003e\u003cem\u003eUrsus americanus - 3\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 194px;\"\u003e\n \u003cp\u003e\u003cem\u003eUrsus arctos - 2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 199px;\"\u003e\n \u003cp\u003e\u003cem\u003eUrsus arctos horribilis -2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 168px;\"\u003e\n \u003cp\u003e\u003cem\u003eVulpes vulpes - 3\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 194px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cem\u003eTables 4. List of deer species included in the behavioural experiments from the collection of reviewed studies.\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"561\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"3\" style=\"width: 561px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eNumber of studies per deer species or subspecies\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 183px;\"\u003e\n \u003cp\u003e\u003cem\u003eAlces alces - 7\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 174px;\"\u003e\n \u003cp\u003e\u003cem\u003eCapreolus capreolus - 7\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 204px;\"\u003e\n \u003cp\u003e\u003cem\u003eCervus canadensis - 3\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 183px;\"\u003e\n \u003cp\u003e\u003cem\u003eCervus elaphus - 8\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 174px;\"\u003e\n \u003cp\u003e\u003cem\u003eCervus nippon - 1\u0026nbsp;\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 204px;\"\u003e\n \u003cp\u003e\u003cem\u003eDama dama - 3\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 183px;\"\u003e\n \u003cp\u003e\u003cem\u003eElaphurus davidianus -1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 174px;\"\u003e\n \u003cp\u003e\u003cem\u003eOdocoileus hemionus - 2\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 204px;\"\u003e\n \u003cp\u003e\u003cem\u003eOdocoileus h. columbianus - 7\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 183px;\"\u003e\n \u003cp\u003e\u003cem\u003eOdocoileus virginianus -13\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 174px;\"\u003e\n \u003cp\u003e\u003cem\u003eRangifer tarandus -1\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"bottom\" style=\"width: 204px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Cervid, Conservation, Human-Wildlife Coexistence, Landscape of fear, Wildlife","lastPublishedDoi":"10.21203/rs.3.rs-6955116/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6955116/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThis study examines the effectiveness of different types of predator cues\u0026mdash;visual, olfactory, and acoustic\u0026mdash;in eliciting anti-predator responses in deer, with the aim of supporting non-lethal, ecocentric management strategies for human-wildlife coexistence. As wildlife populations, including various deer and predator species, increase across the northern hemisphere, conflicts in densely populated areas have also grown, resulting in social and financial costs. Using the \u0026ldquo;applied ecology of fear\u0026rdquo; framework, which leverages fear-induced behaviours to influence prey movement, this study systematically reviews deer responses to predator cues. A total of 256 anti-predator behavioural responses were drawn from 39 peer-reviewed studies following standard meta-analysis guidelines and analysed by cue type, with responses clustered into visual, olfactory, and acoustic groups. Results revealed variation in cue efficacy, with predator acoustic cues (predator playbacks) being the most effective to elicit anti-predator responses in deer, followed by olfactory cues (faeces, urine, hair), and visual cues (photographs, body shape). These findings inform sustainable wildlife management strategies that align with ecocentric principles, offering innovative solutions for human-wildlife coexistence challenges. Further research on cue effectiveness and scalability across relevant spatial and temporal scales of conflict will refine these approaches, fostering harmonious interaction between expanding wildlife populations and human communities.\u003c/p\u003e","manuscriptTitle":"Sight, Scent, and Sound: Leveraging the Landscape of Fear for Effective Wildlife Management","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-08-21 18:44:25","doi":"10.21203/rs.3.rs-6955116/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"c565dde8-b918-43e4-84ea-8c333915d645","owner":[],"postedDate":"August 21st, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[{"id":53339833,"name":"Biological sciences/Ecology/Behavioural ecology"},{"id":53339834,"name":"Biological sciences/Ecology/Conservation"}],"tags":[],"updatedAt":"2026-01-09T08:55:07+00:00","versionOfRecord":[],"versionCreatedAt":"2025-08-21 18:44:25","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6955116","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6955116","identity":"rs-6955116","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}