Spatial patterns of ecosystem disservices across urban-rural gradients: Municipal wasp extermination data from Kobe, Japan

Spatial patterns of ecosystem disservices across urban-rural gradients: Municipal wasp extermination data from Kobe, Japan | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Spatial patterns of ecosystem disservices across urban-rural gradients: Municipal wasp extermination data from Kobe, Japan Tatsuya Saga, Nakata Taichi, Yuta Uchiyama This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6891890/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 18 Aug, 2025 Read the published version in Urban Ecosystems → Version 1 posted 11 You are reading this latest preprint version Abstract Urban greening initiatives provide numerous ecosystem services but may inadvertently increase human-wildlife conflicts (ecosystem disservices). In Japan, hornets ( Vespa spp.) and paper wasps ( Polistes spp.) represent significant disservices due to their stings, which occasionally result in fatalities. To understand how urban environments shape public encounters with these wasps, we analyzed 7,916 extermination request records (2019–2021) from Kobe City, Japan, a highly urbanized region with steep land-use gradients. Building on previous research in Tokyo and Nagoya, our study presents the first fine-scale analysis of how land-use gradients shape genus-specific wasp extermination patterns, differentiating between Vespa (hornets) and Polistes (paper wasps). We examined the effects of surrounding land-use characteristics and population density on extermination frequencies across 779 neighborhood-level districts. Vespa exterminations peaked in areas with ~ 15–20% developed land and declined sharply in denser urban zones, whereas Polistes exterminations were highest in ~ 40% developed areas and increased linearly with population size. Farmland showed a strong negative effect on Vespa exterminations but no effect on Polistes . The relative share of Polistes increased with urbanization, rising from 6.6% baseline to approximately 30% in highly developed areas, suggesting an urban dominance of this genus. Spatial analysis revealed that Polistes exterminations were concentrated in post-1960s suburban "new towns" where detached housing with gardens provides ideal nesting sites, while new residents may lack familiarity with wasp management. These patterns reflect both ecological habitat preferences and social perceptions of risk, including psychological factors such as fear, disgust, and limited familiarity with insects in urban settings. These findings highlight the importance of integrating disservice considerations into biodiversity-friendly urban planning and provide actionable thresholds for spatially-targeted management strategies. ecosystem disservices human-wildlife conflict spatial analysis wasp management public risk perception urban ecology Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Urban greening initiatives are central to "nature-positive" development (Shaikh and Hamel, 2023 ) and provide numerous ecosystem services including air filtering, micro-climate regulation, and recreational opportunities (Bolund & Hunhammar, 1999 ). However, these initiatives may inadvertently increase human-wildlife conflicts, commonly referred to as ecosystem disservices (Lyytimäki et al., 2008 ; Lyytimäki, 2015 ). Recent large-scale studies in Japan have demonstrated that urban green spaces show positive correlations with wasp abundance and human–wildlife conflicts, with stinging insects accounting for nearly 50% of all pest consultations in Tokyo (Hosaka & Numata, 2016 ). Furthermore, research on 1,030 urban residents revealed that public tolerance toward hornets and wild boars is remarkably low, with over 60% requesting government removal services even when animals cause no damage (Hosaka et al., 2017 ). This low tolerance is particularly pronounced among females and elderly residents, and is significantly influenced by childhood nature experiences and affective attitudes toward wildlife. Although urban greening is expanding worldwide, research addressing the negative impacts caused by wildlife—particularly in urban areas, is still limited (Moesch et al., 2024 ; Shackleton et al., 2016 ). To better understand the trade-offs between urban planning goals and associated risks or costs, it is necessary to quantify the relationship between residential environments and ecological disservices. Ecological studies in Nagoya demonstrated that larger hornet species ( V. mandarinia , V. crabro ) show positive responses to urban greenness at 1–2 km scales, while V. analis , responsible for 92% of nest removals, showed no response to greenness levels (Azmy et al., 2016 ). However, these trap-based studies from parks may not reflect actual human-wildlife conflict patterns, as extermination requests involve both ecological occurrence and human behavioral factors. Wasps are among the primary insect groups associated with ecosystem disservices in urban environments. While previous research demonstrated species-specific responses to urban greenness (Azmy et al., 2016 ), these ecological patterns may not directly translate to human-wildlife conflict patterns, as extermination requests reflect not only wasp abundance but also human risk perception, tolerance levels, and behavioral responses. While they contribute to ecological functions such as predation and pollination (Schmack et al., 2024 ), wasps can pose risks to human health and well-being, especially when people are unaware of how to respond safely to their presence (Sumner et al., 2018 ). Such risks are heightened in areas where wasps were previously uncommon and where local residents may lack experience or knowledge in dealing with them (Schmack et al., 2024 ). In recent years, urban areas around the world have experienced increases in wasp-related problems, sometimes due to the introduction of nonnative species. Invasive Vespidae such as Vespa velutina and Vespula vulgaris have caused substantial ecological, economic, and health impacts in Europe and New Zealand (Robinet et al., 2016 ; Monceau et al., 2014 ). In Japan, Vespa velutina nigrithorax has also been detected on Tsushima Island (Kishi and Goka, 2017 ), suggesting potential for further spread. These cases illustrate how insufficient public preparedness or response capacity, especially in regions with little prior wasp exposure, can exacerbate ecological disservices. As emphasized by Wilson Rankin ( 2021 ), understanding how humans interact with both native and invasive wasps in urban settings is a growing global research priority. In this context, detailed analyses of native wasp and human interactions in Japanese cities can inform both domestic management and future preparedness for biological invasions. In Japan, wasps belonging to the genera Vespa (hornet) and Polistes (paper wasp) are among the most prominent stinging insects in urban areas. Reports to pest control agencies regarding wasp-related complaints have been steadily increasing, reaching 26,463 cases in 2022—1.5 times the figure in 2014 (Osanai, 2024 ). A large-scale, ten-year study in Korea found that nests of wasps were concentrated in low-rise residential areas within 170 meters of parks or green spaces, and that urban heat islands may facilitate nest establishment (Song et al., 2025 ). Our study uses municipal extermination records to examine how human-wasp conflicts manifest across urban-rural gradients, providing a socioecological perspective complementary to previous ecological studies. Although urban greening continues under the banner of nature-positive development, it will likely increase wasp nesting, raising potential costs for residents and local governments. While some individuals may remove nests themselves, most rely on municipal or private pest control services. In addition to ecological and demographic variables, human perceptions and emotional responses toward insects—especially wasps—can strongly mediate the frequency of extermination requests. Recent studies have emphasized that negative attitudes toward social wasps are often rooted in affective biases such as fear and disgust, which are themselves reinforced by limited direct interactions with nature and negative media portrayals (Sumner et al., 2018 ; Oi et al., 2024 ; Schmack et al., 2024 ). The “extinction of experience” phenomenon—the progressive loss of direct contact with nature in daily life— has been linked to diminished understanding of ecological roles and greater intolerance of insects (Gaston and Soga, 2020 ). Moreover, the urbanization–disgust hypothesis suggests that people living in urban areas experience more indoor insect encounters, which trigger stronger disgust reactions, while simultaneously possessing lower insect knowledge—factors that amplify extermination behavior even toward ecologically beneficial taxa such as Polistes (Brock et al., 2021 ; Fukano and Soga, 2021 ). To inform urban planning and relevant policy areas such as environmental education, it is essential to understand how local environmental characteristics are associated with ecosystem disservices. Neither ecological data on wasps nor perception surveys alone are sufficient to capture the full picture. Instead, data that reflect actual interactions between humans and wasps are needed. In this respect, extermination request records are a valuable resource, as they are influenced not only by ecological factors such as nest density, but also by human factors including perceived risk, fear, and tolerance. These records likely correlate with local environments that shape both insect populations and resident attitudes. Previous studies have used such records to reveal seasonal trends and habitat preferences in urban areas (e.g., Song et al., 2025 ; Park and Jung, 2020 ), but few have combined them with administratively comprehensive data linked to spatially explicit environmental variables across a major city. In this study, we take advantage of a rare dataset from Kobe City, where extermination costs were fully subsidized by the local government during the study period and nearly all requests were processed through a centralized municipal system. This allows for robust analysis of how land use and demographic factors shape patterns of human–wasp conflict. This study focuses on Kobe City because it is one of the major cities in Japan and among the large cities in the country, it has rich natural lands and socioecological landscapes shaped by diverse ecosystems including forestlands, agricultural lands, and water bodies, which can be potential habitats of wasps. It spans 553 km² with a green cover ratio of 68.5% and a population of approximately 1.5 million. It is geographically divided by the Rokko mountain range, which peaks at 931 meters in elevation. Compared to surrounding municipalities, Kobe contains many suburban housing developments built on former forested land. The city stretches from urbanized southern districts to northern rural zones, resulting in a high diversity of land-use types and landscapes at the neighborhood scale. From 2010 to 2021, Kobe City fully subsidized the cost of wasp nest extermination for private households. During this period, most extermination cases were handled through the city, providing an administratively centralized and near-complete record of pest control requests. Pest control professionals were able to distinguish between Vespa and Polistes nests in these records. In Japan, the genera Vespa and Polistes both comprise eusocial wasps that form annual colonies initiated by a single overwintered queen (Spradbery, 1973 ; Matsuura & Yamane 1990 ). However, they differ markedly in ecology, colony size, and risk to humans. Vespa species typically build large, enclosed nests in underground cavities, tree hollows, or building structures, and colonies can reach several thousand individuals by late summer. These wasps are highly defensive and are responsible for the majority of sting-related injuries and fatalities. Indeed, in Japan, the number of deaths caused by wasp stings annually exceeds those caused by bears or venomous snakes (Matsuura, 1995 ). In contrast, Polistes wasps construct small, open-comb nests with fewer than 100 individuals, usually under eaves or on fences. While they are generally less aggressive, their close association with human dwellings increases the likelihood of contact and extermination. These ecological and behavioral distinctions shape public perception and management responses, and are central to understanding the contrasting extermination patterns analyzed in this study. The aim of this study was to analyze data on wasp extermination requests from 2019 to 2021 in Kobe City, comparing the two genera ( Vespa and Polistes ) with differing risk profiles. By incorporating gradients of urbanization, population size, and agricultural land cover into statistical models, we sought to clarify the relationship between local environmental characteristics and wasp extermination patterns. Specifically, we treated each neighborhood-level district as a spatial analysis unit and integrated nest removal records with data on land use (urban area, farmland), population, and spatial autocorrelation. Using multivariate regression analysis, we evaluated the extent to which these geographic and demographic factors explained variation in wasp extermination frequencies. These findings may help inform targeted pest control strategies and guide sustainable urban landscape planning. While previous studies have examined hornet abundance in relation to urban greenness using standardized trapping methods (Azmy et al., 2016 ), our approach using municipal extermination records provides a unique perspective on actual human-wasp conflicts. Unlike trap-based studies that sample from predetermined locations (typically parks), extermination records reflect the intersection of ecological occurrence and human behavioral responses across the entire urban landscape, including private properties where most conflicts occur. Materials and Methods Study System Data on extermination requests for wasps ( Vespa spp. and Polistes spp.) in Kobe City, Japan, were obtained from records maintained by the Kobe City Public Health Bureau between 2019 and 2021. The number of requests for each year is 3270 in 2019, 2601 in 2020, and 2045 in 2021. In total, 7,916 extermination request records were collected. Such datasets are typically fragmented across multiple organizations in Japanese cities, as extermination responsibilities are distributed among several entities, and city governments rarely collect comprehensive data. Kobe City represents a rare case where a single organization handled all exterminations, enabling the city government to collect comprehensive data. This study represents the first analysis of the largest municipal wasp extermination dataset in Japan. The individual record includes the information of location (neighborhood-level district names) of the extermination, exterminated species, and dates. These records were aggregated by neighborhood-level districts (n = 779), which served as the basic spatial units of analysis in this study (hereafter referred to as "units"). Land-use metrics for each unit were derived using QGIS version 3.22.8 and the High Resolution Land-Use and Land-Cover Map published by the Japan Aerospace Exploration Agency (JAXA, 2022). Following ecological studies indicating that Vespinae wasps typically forage over distances of at least 1 km (Matsuura and Yamane, 1990 ) and previous research demonstrating that 1–2 km radii effectively predict hornet abundance (Azmy et al., 2016 ), we calculated the area of developed land, farmland, and forest within a 1 km radius from the centroid of each unit. The original dataset includes address-level extermination records and was provided by the Kobe City Public Health Bureau under a data-sharing agreement that prohibits public dissemination of raw data. Statistical Analysis We analyzed how extermination frequency per unit was influenced by surrounding land use and demographic characteristics between 2019 and 2021. The main response variables were the annual number of exterminations of Vespa and Polistes wasps per unit. Due to the large number of units with zero extermination cases and the presence of overdispersion, we applied generalized linear mixed models (GLMMs) with a negative binomial distribution and a logarithmic link function. Explanatory variables included the developed land area, farmland area, and log₁₀-transformed population size for each unit. To allow for potential non-linear effects, we included quadratic terms for these predictors. The variable "year" was modeled as a random intercept to account for temporal variation. In addition, spatial autocorrelation was accounted for by including spatial autocovariates in the analysis, which were calculated for each study site based on latitude and longitude. Forest land area was excluded from the model due to multicollinearity, as indicated by high variance inflation factor (VIF) values: developed land = 32.74, forest = 26.78, farmland = 7.14, log₁₀(population) = 1.41, and spatial autocovariate = 1.17 (Fox et al., 2007). To assess how land-use characteristics influenced the relative frequency of Polistes exterminations among all wasp exterminations, we conducted a complementary analysis using a binomial GLMM with a logit link function. The response variable was the proportion of Polistes exterminations per unit. Explanatory variables included the ratio of developed land to farmland and log₁₀-transformed population, while spatial autocovariates were again included to control for spatial dependency. Quadratic terms were not included in this model. As the logit link function already captures non-linearity, and exploratory analyses indicated that relationships between predictors and the logit-transformed response were largely monotonic, quadratic terms were not expected to improve model fit and could lead to overfitting or implausible effects. For model simplicity and interpretability, we retained only linear terms. All analyses were conducted in R version 4.2.2 (R Core Team, 2021) using the glmmTMB package (Magnusson et al., 2017) and/or the spdep package (Bivand et al., 2017 ). Results Model Adequacy The three models presented in this study ( Vespa extermination count, Polistes extermination count, and Polistes extermination ratio) were all based on negative binomial or binomial GLMMs and accounted appropriately for overdispersion and year effects. The estimated dispersion parameters (θ = 1.54 for Vespa, 2.75 for Polistes) fell within the empirically acceptable range for ecological data (Ver Hoef & Boveng, 2007 ). The variance of the random effect for year was negligible or small across models (σ² = 2.1 × 10⁻⁹, 0.068, and 0.187), indicating limited inter-annual variation (Nakagawa & Schielzeth, 2013 ). These values confirm that all models were well-specified and structurally sound for inference and prediction. Extermination Patterns by Species Vespa Extermination Counts Among explanatory variables, the proportion of developed land exhibited a significant quadratic relationship: the linear term was positive (β = 1.05, P = 0.022), and the quadratic term was negative (β = −3.30, P < 0.001). The number of exterminations peaked when developed land comprised approximately 15–20% of the unit area (IRR = 2.86, 95% CI = 1.17–6.97), then declined sharply beyond this threshold (Fig. 1 a). Farmland proportion showed a significant negative effect (β = −2.03, P = 0.006), with a 20% increase in farmland associated with a 34% decrease in extermination count. The population variable showed a significant quadratic effect (β = 0.248, P < 0.001), indicating an accelerating increase in exterminations with population growth. Spatial autocorrelation also had a significant positive effect (β = 0.333, P < 0.001), suggesting that Vespa exterminations were spatially clustered. Polistes Extermination Counts Extermination counts exhibited a significant quadratic relationship with developed land: the linear term was positive (β = 2.32, P < 0.001) and the quadratic term was negative (β = −2.78, P < 0.001). The peak extermination count occurred at around 42% developed land (Fig. 1 b). Farmland proportion was not significantly associated with Polistes exterminations. Population size showed a significant positive linear effect (β = 1.67, P = 0.006), indicating higher extermination counts in more densely populated areas. Spatial autocorrelation also showed a significant positive effect (β = 0.336, P < 0.001), reflecting spatial clustering of Polistes exterminations. Proportion of Polistes Among Total Exterminations The proportion of Polistes exterminations increased significantly with the proportion of developed land (β = 2.15, P < 0.001). In central urban areas, this proportion was approximately 8.6 times greater than in less urbanized units (Fig. 2 ). Farmland proportion showed a marginally significant negative effect (β = −0.60, P = 0.077), suggesting that Vespa exterminations may be relatively more common in agricultural areas. Population size was positively associated with the proportion of Polistes (β = 0.234, P < 0.001), indicating that every 10-fold increase in population size raised the odds of a Polistes extermination by approximately 1.26 times. In contrast, spatial autocorrelation showed a significant negative effect (β = −0.14, P = 0.003), suggesting that in units with highly clustered exterminations, Vespa wasps tend to dominate. The estimated intercept (β = −2.71) corresponds to a baseline Polistes extermination proportion of 6.6% under average conditions. In densely urbanized areas (urban proportion = 0.8, log₁₀ population ≈ 5), the model predicts that Polistes may account for approximately 30% of all exterminations. To further clarify the species-specific responses to urbanization, we grouped sites into quartiles based on urbanization level and calculated mean extermination counts per genus. As shown in Fig. 3, Vespa exterminations peaked at moderate levels of urbanization (Q2), whereas Polistes showed higher frequencies in highly urbanized areas (Q3–Q4). Table1. Model results for Vespa and Polistes exterminations Vespa (AIC=7076.2) Polistes (AIC=5216) Predictor Estimate SE p-value Estimate SE p-value (Intercept) -1.058 0.326 0.001 -5.786 0.961 <0.001 Developed land 1.045 0.459 0.022 2.318 0.601 <0.001 Developed land 2 -3.301 0.472 <0.001 -2.780 0.582 <0.001 Farmland -2.031 0.735 0.006 -1.530 1.000 0.126 Farmland 2 0.377 1.047 0.719 0.118 1.522 0.938 Log 10 (Population) -0.095 0.232 0.684 1.666 0.603 0.006 Log 10 (Population) 2 0.248 0.041 <0.001 0.011 0.093 0.904 Spatial Autocovariate 0.333 0.022 <0.001 0.336 0.049 <0.001 Table2. Binomial GLMM results for Polistes extermination proportion Predictor Estimate SE z value p-value (Intercept) -2.715 0.330 -8.222 <0.001 Developed land 2.150 0.126 17.033 <0.001 Farmland -0.595 0.337 -1.769 0.077 Log 10 (Population) 0.234 0.066 3.539 <0.001 Spatial autocovariate -0.143 0.048 -2.989 0.002 Discussion Urban gradient effects on genus-specific human-wasp conflicts This study revealed distinct associations between extermination requests for Vespa (hornets) and Polistes (paper wasps) and land-use characteristics in Kobe City. Our findings complement ecological studies from Nagoya, which showed species-specific responses to urban greenness (Azmy et al., 2016 ). However, our conflict data revealed different patterns: Vespa exterminations peaked at intermediate urbanization (15–20% developed land) rather than increasing monotonically with greenness, highlighting how human behavioral factors modify ecological patterns in real-world conflict scenarios. Extermination counts were also negatively associated with farmland cover and positively associated with population size and spatial autocorrelation, suggesting that both ecological conditions and human behavioral factors shape reported disservices (Fig. 1 a, b). In contrast, Polistes extermination counts were highest in areas with approximately 40% developed land, showed no significant association with farmland, increased linearly with population size, and were similarly clustered spatially. Moreover, the proportion of Polistes exterminations relative to all wasp exterminations increased with urbanization, indicating their dominance in highly developed areas. These patterns refine earlier findings, such as those by Song et al. ( 2025 ), who reported more Vespidae nests near green areas in South Korea. While proximity to green space may indeed promote nesting, our results indicate that wasp-human interactions cannot be explained by developed land alone. Instead, extermination requests are shaped by a complex interplay of urbanization, population density, land-use type, and local perceptions of risk. Vespa Extermination Counts The peak in Vespa exterminations at intermediate levels of development may reflect the species’ preference for semi-natural environments and the increased likelihood of human-wasp encounters in moderately populated zones. Although Vespa nests were less frequently reported in highly urbanized areas, some city units with over 85% developed land still recorded high extermination counts, underscoring the persistence of potential hazards even in dense urban centers. Farmland coverage was negatively associated with Vespa exterminations, which may reflect reduced nesting opportunities or differences in local residents’ attitudes and knowledge about wasps. People familiar with rural insects may be more tolerant of wasp presence or capable of managing nests independently, while newer suburban communities might be more likely to request professional extermination. These behavioral differences may be influenced by media portrayals that emphasize the lethal risks of stings (Oi et al., 2024 ). The accelerated increase in Vespa exterminations with population size, despite a flattening at higher urbanization levels, suggests a trade-off: while denser populations increase wasp-human encounters, extreme urbanization suppresses wasp nesting opportunities. The strong spatial autocorrelation may be driven by limited dispersal distances of overwintered queens and similarities in local environmental and social contexts, such as the prevalence of green spaces or public aversion to wasps in tightly packed residential areas. Pesticide use in farmland might further suppress Vespa populations, either directly or indirectly through prey reduction (Tison et al., 2023 ). However, no similar effect was observed for Polistes , possibly due to differences in diet or nesting behavior. Polistes Extermination Counts Polistes were more common in built environments, consistent with their preference for nesting on human-made structures. Their extermination frequency increased linearly with population size, implying both ecological tolerance and increased human-wasp contact in denser areas. In rural areas, nests may be left undisturbed or managed independently, suggesting that local perceptions of risk and response thresholds vary with setting. Compared to Vespa , Polistes exterminations were more strongly associated with developed land and population, but not farmland, highlighting divergent ecological preferences and public responses to each genus. These differences likely reflect not only biological traits—such as nest size and aggression—but also variation in risk perception. Proportion of Polistes Among Total Exterminations The proportion of Polistes among all exterminations was highest in highly urbanized areas, potentially reflecting a loss of suitable nesting habitats for Vespa (Fig. 2 ). Similar shifts in species dominance in urban contexts have been observed elsewhere (e.g., Baker et al., 2020), where smaller wasps thrive on unique urban nesting substrates and prey. Importantly, the number of extermination requests does not directly reflect wasp nest density but rather a combination of ecological occurrence and human behavior. Thus, extermination data serve as a proxy for ecosystem disservices from a socioecological perspective. Future research should integrate field nest surveys with social surveys on risk tolerance to better assess the true societal cost of urban greening under “nature-positive” agendas. Quantifying disservices in monetary terms—including extermination costs, medical expenses, and labor losses—may also help optimize policy decisions balancing ecosystem services and disservices. Spatial patterns and housing development characteristics The detailed spatial distribution of extermination requests reveals important relationships between urban development patterns and human-wasp conflicts (Fig. 4 ). Vespa exterminations were primarily observed in relatively rural areas on the northern side of Kobe City and at the boundaries between urban and rural areas, while Polistes exterminations were concentrated in northern urban areas and urban-rural boundary zones. These spatial patterns are consistent with our statistical model results and provide additional insights into the socioecological drivers of extermination behavior. The northern urban areas, developed primarily after the 1960s-1970s, represent what are commonly called "new towns" in Japan—suburban housing developments popular among families seeking new homes. Unlike dense urban cores dominated by apartments, these new towns predominantly feature detached houses with gardens and eaves, providing ideal nesting sites for both genera. Both Polistes and Vespa commonly nest in garden hedges and under eaves, while some Vespa species additionally utilize enclosed spaces such as attics and building foundations (Spradbery, 1973 ; Matsuura 1995 ). This housing typology creates abundant nesting opportunities while increasing human-wasp encounter rates through gardening and outdoor activities around residences. The combination of suitable nesting habitat and resident characteristics may explain the elevated Polistes extermination rates in these areas. New residents may lack familiarity with Polistes behavior and management techniques, leading to increased reliance on professional extermination services. In contrast, residents in surrounding rural areas, who typically grew up locally, may possess greater tolerance and experience cohabiting with these wasps. For Vespa , this residential experience difference appears less pronounced, likely reflecting the more universal perception of hornets as a significant threat regardless of residents' background. These findings highlight how urban development patterns inadvertently create conditions that promote both wasp abundance and human-wildlife conflicts, exemplifying the ecosystem disservices associated with suburban expansion. Sociocultural drivers of extermination behavior Beyond ecological distribution, human emotional and cognitive responses play a crucial role in shaping wasp extermination patterns. In particular, urban residents are more likely to encounter insects indoors, a context that tends to evoke heightened feelings of disgust (Fukano and Soga, 2021 ). This tendency is reinforced by the general lack of direct nature experience in urban environments—a phenomenon described as the extinction of experience—which contributes to a lower tolerance for insects, even when they are harmless or beneficial (Gaston and Soga, 2020 ). Media portrayals that emphasize the danger or aggressiveness of wasps, in contrast to the more favorable framing of bees, further amplify public aversion and may distort risk perception (Oi et al., 2024 ; Sumner et al., 2018 ). Importantly, recent findings indicate that even individuals with some knowledge about wasps, such as urban gardeners, often express strong emotional aversion. Schmack et al. ( 2024 ) showed that knowledge alone may not suffice to overcome affective biases, underscoring the importance of addressing both cognitive and emotional dimensions in any intervention strategy. Environmental education implications The spatial distribution of extermination requests showed different patterns between the two genera. This result suggests that contents of education to deal with hornets and paper wasps should be tailored based on degree of urbanization in each neighborhood-level district. For example, urban centers and areas in between urban and rural areas might need more knowledge on Polistes . For rural areas, dissemination of knowledge on Vespa should be prioritized. Such contextualized approach can effectively reduce the negative influence of the disservices from these species and to raise awareness of their importance in local ecosystems based on hands-on experience (Fig. 3). In addition to species-specific knowledge, emotional and cultural biases must also be considered when designing educational programs. For instance, the extinction of experience has reduced familiarity with insects among urban residents, amplifying feelings of disgust and fear (Gaston and Soga, 2020 ; Fukano and Soga, 2021 ). Moreover, negative portrayals of wasps in the media may further reinforce aversion (Oi et al., 2024 ; Sumner et al., 2018 ). Therefore, effective education should incorporate both factual knowledge and indirect experiential learning, such as observing wasps safely in the environment or learning through case studies, to foster tolerance and reduce unnecessary extermination. These findings complement ecological research showing species-specific habitat preferences (Azmy et al., 2016 ) by revealing how human behavioral responses vary across the urban-rural gradient. While ecological studies inform us about where wasps are likely to be abundant, extermination data reveals where conflicts actually occur, providing crucial information for targeted education and management interventions. Implications for invasive species risk assessment While the introduction highlighted the growing global concern surrounding Vespidae invasions, our findings also contribute to this field by offering actionable insights into how urban wasp disservices are mediated by local ecological and sociocultural contexts. Around the world, invasive species in the Vespidae family—such as Vespa velutina , Vespula vulgaris , and Vespula germanica —have caused severe ecological, economic, and public health impacts, particularly in Europe and New Zealand (Lester et al., 2013 ; Monceau et al., 2014 ). However, as noted in recent syntheses, the severity of such conflicts is not solely determined by wasp behavior or population dynamics, but also by how humans perceive, tolerate, and respond to these insects (Wilson Rankin, 2021 ; Lester et al., 2025 ). Our spatially explicit data on extermination requests—capturing variation in urbanization, land use, and perceived risk—may provide a valuable empirical basis for modeling future wasp-human interactions under scenarios of biological invasion. These findings underscore the need to incorporate socioecological perspectives into early-warning frameworks for invasive wasp management, especially in urban landscapes where human responses can critically shape outcomes. Methodological Contributions and Limitations Our study demonstrates the value of municipal extermination records as a complementary approach to standardized ecological sampling. While trap-based studies like Azmy et al. ( 2016 ) provide controlled measures of hornet abundance, they are limited to accessible public spaces and may not reflect private property conflicts where most human-wasp interactions occur. Conversely, extermination records capture actual conflict patterns but are influenced by socioeconomic factors, risk perception, and reporting behaviors that vary across communities. The species-specific patterns we observed with Vespa conflicts peaking at intermediate urbanization and Polistes increasing with urban density, suggest that extermination data provides unique insights into the socioecological dimensions of ecosystem disservices that cannot be captured through ecological sampling alone. Future research integrating both approaches would provide a more complete understanding of urban wasp dynamics. Conclusion This study demonstrates how spatial patterns of extermination requests for social wasps in Kobe City are shaped by land use, population density, and species-specific ecological and behavioral traits. Vespa and Polistes differ not only in nesting preferences and colony size but also in how the public perceives and responds to them across urban gradients. These findings emphasize that urban ecosystem disservices are socioecological in nature and should be addressed through context-specific strategies. Our results provide actionable insights for designing pest management and biodiversity education programs that reflect both local ecological conditions and human risk perception. Furthermore, by establishing spatial baselines of human–wasp interaction, this study supports the development of early-warning frameworks and response policies for invasive wasp species. Such approaches are especially relevant in rapidly urbanizing regions where environmental change and public sensitivity converge to shape future biodiversity challenges. By integrating ecological and conflict pattern analyses, urban planners can develop strategies that account for both biological distributions and human behavioral responses to urban wildlife. Declarations Acknowledgments We are grateful to Dr. Atsushi Ushimaru for his valuable advice on data analysis. We also thank the Kobe City Public Health Bureau and the Hyogo Pest Control Association for their cooperation in data collection. Funding Statement This study was supported by a research grant from the Graduate School of Human Development and Environment, Kobe University. Data availability statement The extermination request data analyzed in this study were obtained from the Kobe City Public Health Bureau (Kobe City Kenkōkyoku). These data contain address-level information and are not publicly shareable due to privacy regulations. To protect individual confidentiality, only aggregated statistics and model results are available. Summary tables and regression outputs can be provided by the corresponding author upon reasonable request and with permission from the Kobe City Public Health Bureau. Author contribution T.S. led the project and was responsible for conceptualization, methodology, investigation, formal analysis, funding acquisition, and original draft writing. T.N. contributed to data analysis and visualization. Y.U. contributed to conceptual development, data processing, visualization, and discussions on methodology. All authors reviewed and edited the manuscript. Declaration of Competing Interests The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. 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Biological Reviews, 96(4), 1645–1675. https://doi.org/10.1111/brv.12720 Fox, J., & Weisberg, S. (2007). An R companion to applied regression. SAGE Publications. Fukano, Y., & Soga, M. (2021). Why do so many modern people hate insects? The urbanization–disgust hypothesis. Science of the Total Environment, 777, 146229. https://doi.org/10.1016/j.scitotenv.2021.146229 Gaston, K. J., & Soga, M. (2020). Extinction of experience: The need to be more specific. People and Nature, 2(3), 575–581. https://doi.org/10.1002/pan3.10118 Hosaka, T., & Numata, S. (2016). Spatiotemporal dynamics of urban green spaces and human–wildlife conflicts in Tokyo. Scientific Reports, 6, 30911. https://doi.org/10.1038/srep30911 Hosaka, T., Sugimoto, K., & Numata, S. (2017). Effects of childhood experience with nature on tolerance of urban residents toward hornets and wild boars in Japan. PLOS ONE, 12(4), e0175243. https://doi.org/10.1371/journal.pone.0175243 Kishi, S., & Goka, K. (2017). Review of the invasion and expansion of Vespa velutina nigrithorax in Japan. Applied Entomology and Zoology, 52(4), 541-548. Lester, P.J., Beggs, J.R., Brown, B.A., Edwards, E.D., Gorenteman, R., Toft, R.J., Twidle, A.M., & Ward, D.F. (2013). The outlook for control of New Zealand’s most abundant, widespread and damaging invertebrate pests: social wasps. New Zealand Science Review , 70(4), 56–62. Lester, P. J., & Beggs, J. R. (2019). Invasion success and management strategies for social Vespula wasps. Annual Review of Entomology , 64, 51–71. https://doi.org/10.1146/annurev-ento-011118-111812 Lester, P. J., Shortall, C. R., & Beggs, J. R. (2025). Recent advances and avenues for the pest management of invasive social wasps and hornets. Current Opinion in Insect Science , 68, 101336. https://doi.org/10.1016/j.cois.2025.101336 Lyytimäki, J. (2015). Ecosystem disservices: Embrace the catchword. Ecosystem Services, 12, 136. https://doi.org/10.1016/j.ecoser.2014.11.008 Lyytimäki, J., Petersen, L. K., Normander, B., & Bezák, P. (2008). Nature as a nuisance? Ecosystem services and disservices to urban lifestyle. Environmental Sciences, 5(3), 161–172. https://doi.org/10.1080/15693430802055524 Matsuura, M. (1995). Social wasps of Japan in color. Hokkaido University Press. Matsuura, M., & Yamane, S. (1990). Biology of the vespine wasps. Springer. Moesch, S. S., Wellmann, T., Haase, D., & Bhardwaj, M. (2024). Mammal Mia: A review on how ecological and human dimension research on urban wild mammals can benefit future biophilic cities. Basic and Applied Ecology, 79, 90-101. https://doi.org/10.1016/j.baae.2024.05.004 Monceau, K., Bonnard, O., & Thiéry, D. (2014). Vespa velutina : A new invasive predator of honeybees in Europe. Journal of Pest Science , 87(1), 1–16. https://doi.org/10.1007/s10340-013-0537-3 Nakagawa, S., & Schielzeth, H. (2013). A general and simple method for obtaining R² from generalized linear mixed-effects models. Methods in Ecology and Evolution, 4(2), 133–142. https://doi.org/10.1111/j.2041-210x.2012.00261.x Oi, C. A., Brown, R. L., & Sumner, S. (2024). Bee-ing positive about wasp-negative media reporting: The opinions of scientists and their influence on the media. Insectes Sociaux, 71(1), 29–42. https://doi.org/10.1007/s00040-024-00952-9 Osanai, R. (2024). Ecology and control of hornets. Pest Control [in Japanese], 207, 10-14. Park, J., & Jung, C. (2020). Analysis of reports on wasp nest removal in Seoul, Korea: Spatial and temporal patterns of human–insect conflict. Proceedings of the National Institute of Ecology of the Republic of Korea, 2 (2), 101–107. https://doi.org/10.22920/PNIE.2020.2.2.101 Robinet, C., Suppo, C., & Darrouzet, E. (2016). Rapid spread of the invasive yellow-legged hornet in France: The role of human-mediated dispersal and the effects of control measures. Journal of Applied Ecology , 54(1), 205–215. https://doi.org/10.1111/1365-2664.12724 Schmack, J. M., Egerer, M. H., Karlebowski, S., Neumann, A. E., & Sturm, U. (2024). Overlooked and misunderstood: How urban community gardeners perceive social wasps and their ecosystem functions. Journal of Insect Conservation, 28(2), 283–289. https://doi.org/10.1007/s10841-024-00548-5 Shackleton, C. M., Ruwanza, S., Sinasson Sanni, G. K., Bennett, S., De Lacy, P., Modipa, R., ... & Thondhlana, G. (2016). Unpacking Pandora's box: understanding and categorising ecosystem disservices for environmental management and human wellbeing. Ecosystems, 19(4), 587-600. Shaikh, S. F. E. A., & Hamel, P. (2023). Identifying nature-positive futures in new cities: An application of the Urban Nature Futures Framework. Sustainability Science, 18(1), 123–134. https://doi.org/10.1007/s11625-023-01411-3 Song, W., Kim, H., & Kim, W. (2025). Modeling urban wasp nest occurrences using 119 fire service reports, LiDAR, and hyperspectral imagery: The role of green spaces and structural factors. Journal of Environmental Management, 379, 124776. Spradbery, J. P. (1973). Wasps: An account of the biology and natural history of solitary and social wasps. University of Washington Press. Sumner, S., Law, G., & Cini, A. (2018). Why we love bees and hate wasps. Ecological Entomology, 43(6), 836–845. https://doi.org/10.1111/een.12676 Tison, L., Franc, C., Burkart, L., Jactel, H., Monceau, K., de Revel, G., & Thiéry, D. (2023). Pesticide contamination in an intensive insect predator of honey bees. Environment International, 176, 107975. https://doi.org/10.1016/j.envint.2023.107975 Ver Hoef, J. M., & Boveng, P. L. (2007). Quasi-Poisson vs. negative binomial regression: How should we model overdispersed count data? Ecology, 88(11), 2766–2772. https://doi.org/10.1890/07-0043.1 Wilson Rankin, E. E. (2021). Emerging patterns in social wasp invasions. Current Opinion in Insect Science , 46, 72–77. https://doi.org/10.1016/j.cois.2021.02.014 Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 18 Aug, 2025 Read the published version in Urban Ecosystems → Version 1 posted Editorial decision: Revision requested 08 Jul, 2025 Reviews received at journal 08 Jul, 2025 Reviews received at journal 07 Jul, 2025 Reviews received at journal 25 Jun, 2025 Reviewers agreed at journal 17 Jun, 2025 Reviewers agreed at journal 17 Jun, 2025 Reviewers agreed at journal 17 Jun, 2025 Reviewers invited by journal 16 Jun, 2025 Editor assigned by journal 16 Jun, 2025 Submission checks completed at journal 16 Jun, 2025 First submitted to journal 14 Jun, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. 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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-6891890","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":472762778,"identity":"ca0a192c-952c-4b9d-81bd-f710dee9099b","order_by":0,"name":"Tatsuya Saga","email":"data:image/png;base64,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","orcid":"","institution":"Kobe University","correspondingAuthor":true,"prefix":"","firstName":"Tatsuya","middleName":"","lastName":"Saga","suffix":""},{"id":472762779,"identity":"086b6cc1-4915-4af9-af86-1606c9fa967b","order_by":1,"name":"Nakata Taichi","email":"","orcid":"","institution":"Kyushu University","correspondingAuthor":false,"prefix":"","firstName":"Nakata","middleName":"","lastName":"Taichi","suffix":""},{"id":472762780,"identity":"7d1e7ddb-7069-480a-8103-1030b814efff","order_by":2,"name":"Yuta Uchiyama","email":"","orcid":"","institution":"Kobe University","correspondingAuthor":false,"prefix":"","firstName":"Yuta","middleName":"","lastName":"Uchiyama","suffix":""}],"badges":[],"createdAt":"2025-06-14 05:23:12","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6891890/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6891890/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s11252-025-01788-2","type":"published","date":"2025-08-18T16:12:59+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":84923345,"identity":"6f484dad-ce40-433e-ac64-1d7b2ef84f77","added_by":"auto","created_at":"2025-06-18 20:46:34","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":917622,"visible":true,"origin":"","legend":"\u003cp\u003eRelationships between developed land area and the number of extermination: (a) \u003cem\u003eVespa\u003c/em\u003e; (b) \u003cem\u003ePolistes\u003c/em\u003e. Regression lines were drawn using estimated coefficients from the GLMMs.\u003c/p\u003e","description":"","filename":"floatimage1.png","url":"https://assets-eu.researchsquare.com/files/rs-6891890/v1/7c8c1884a42f863073d34a73.png"},{"id":84923349,"identity":"1f4cdf15-6c6e-4628-8331-94ae41599562","added_by":"auto","created_at":"2025-06-18 20:46:35","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":600033,"visible":true,"origin":"","legend":"\u003cp\u003eRelationship between developed land area and the proportion of \u003cem\u003ePolistes\u003c/em\u003eexterminations. Circle size indicates the number of \u003cem\u003ePolistes\u003c/em\u003e extermination. Regression lines were drawn using estimated coefficients from the GLMMs.\u003c/p\u003e","description":"","filename":"floatimage2.png","url":"https://assets-eu.researchsquare.com/files/rs-6891890/v1/4bdb0ca2f833face6e33ac51.png"},{"id":84923347,"identity":"43274a62-dba5-4797-9ad3-e81e081a124e","added_by":"auto","created_at":"2025-06-18 20:46:35","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":157413,"visible":true,"origin":"","legend":"\u003cp\u003eMean annual extermination counts (± SE) of \u003cem\u003eVespa \u003c/em\u003eand \u003cem\u003ePolistes\u003c/em\u003e wasps across urbanization quartiles in Kobe City. Extermination counts were averaged across three years (2019–2021) within each urbanization quartile and genus to represent yearly means.\u003c/p\u003e\n\u003cp\u003eUrbanization quartiles were calculated from the proportion of developed land in each neighborhood-level district.\u003c/p\u003e","description":"","filename":"floatimage3.png","url":"https://assets-eu.researchsquare.com/files/rs-6891890/v1/9a8df681b165f403f1c43478.png"},{"id":84923348,"identity":"ebe1b5b4-271e-427a-ac52-4185d035bc4d","added_by":"auto","created_at":"2025-06-18 20:46:35","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":954418,"visible":true,"origin":"","legend":"\u003cp\u003eNumber of extermination request in three years (2019-2021) per person and urban area of Kobe City. Left: \u003cem\u003eVepa \u003c/em\u003eextermination; Right: \u003cem\u003ePolistes \u003c/em\u003eextermination.\u003c/p\u003e","description":"","filename":"floatimage4.png","url":"https://assets-eu.researchsquare.com/files/rs-6891890/v1/ef51ea9fb95c010431b64186.png"},{"id":89847113,"identity":"2f6d839a-bd75-4939-b00d-7c68bc6a388c","added_by":"auto","created_at":"2025-08-25 16:40:25","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":3010099,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6891890/v1/3a2d4630-c729-4886-9f09-b0e6115027fa.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"Spatial patterns of ecosystem disservices across urban-rural gradients: Municipal wasp extermination data from Kobe, Japan","fulltext":[{"header":"Introduction","content":"\u003cp\u003eUrban greening initiatives are central to \"nature-positive\" development (Shaikh and Hamel, \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) and provide numerous ecosystem services including air filtering, micro-climate regulation, and recreational opportunities (Bolund \u0026amp; Hunhammar, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e1999\u003c/span\u003e). However, these initiatives may inadvertently increase human-wildlife conflicts, commonly referred to as ecosystem disservices (Lyytim\u0026auml;ki et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Lyytim\u0026auml;ki, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Recent large-scale studies in Japan have demonstrated that urban green spaces show positive correlations with wasp abundance and human\u0026ndash;wildlife conflicts, with stinging insects accounting for nearly 50% of all pest consultations in Tokyo (Hosaka \u0026amp; Numata, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Furthermore, research on 1,030 urban residents revealed that public tolerance toward hornets and wild boars is remarkably low, with over 60% requesting government removal services even when animals cause no damage (Hosaka et al., \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). This low tolerance is particularly pronounced among females and elderly residents, and is significantly influenced by childhood nature experiences and affective attitudes toward wildlife.\u003c/p\u003e \u003cp\u003eAlthough urban greening is expanding worldwide, research addressing the negative impacts caused by wildlife\u0026mdash;particularly in urban areas, is still limited (Moesch et al., \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Shackleton et al., \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). To better understand the trade-offs between urban planning goals and associated risks or costs, it is necessary to quantify the relationship between residential environments and ecological disservices. Ecological studies in Nagoya demonstrated that larger hornet species (\u003cem\u003eV. mandarinia\u003c/em\u003e, \u003cem\u003eV. crabro\u003c/em\u003e) show positive responses to urban greenness at 1\u0026ndash;2 km scales, while \u003cem\u003eV. analis\u003c/em\u003e, responsible for 92% of nest removals, showed no response to greenness levels (Azmy et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). However, these trap-based studies from parks may not reflect actual human-wildlife conflict patterns, as extermination requests involve both ecological occurrence and human behavioral factors.\u003c/p\u003e \u003cp\u003eWasps are among the primary insect groups associated with ecosystem disservices in urban environments. While previous research demonstrated species-specific responses to urban greenness (Azmy et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), these ecological patterns may not directly translate to human-wildlife conflict patterns, as extermination requests reflect not only wasp abundance but also human risk perception, tolerance levels, and behavioral responses. While they contribute to ecological functions such as predation and pollination (Schmack et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), wasps can pose risks to human health and well-being, especially when people are unaware of how to respond safely to their presence (Sumner et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Such risks are heightened in areas where wasps were previously uncommon and where local residents may lack experience or knowledge in dealing with them (Schmack et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). In recent years, urban areas around the world have experienced increases in wasp-related problems, sometimes due to the introduction of nonnative species. Invasive Vespidae such as \u003cem\u003eVespa velutina\u003c/em\u003e and \u003cem\u003eVespula vulgaris\u003c/em\u003e have caused substantial ecological, economic, and health impacts in Europe and New Zealand (Robinet et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Monceau et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). In Japan, \u003cem\u003eVespa velutina nigrithorax\u003c/em\u003e has also been detected on Tsushima Island (Kishi and Goka, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2017\u003c/span\u003e), suggesting potential for further spread. These cases illustrate how insufficient public preparedness or response capacity, especially in regions with little prior wasp exposure, can exacerbate ecological disservices. As emphasized by Wilson Rankin (\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), understanding how humans interact with both native and invasive wasps in urban settings is a growing global research priority. In this context, detailed analyses of native wasp and human interactions in Japanese cities can inform both domestic management and future preparedness for biological invasions.\u003c/p\u003e \u003cp\u003eIn Japan, wasps belonging to the genera \u003cem\u003eVespa\u003c/em\u003e (hornet) and \u003cem\u003ePolistes\u003c/em\u003e (paper wasp) are among the most prominent stinging insects in urban areas. Reports to pest control agencies regarding wasp-related complaints have been steadily increasing, reaching 26,463 cases in 2022\u0026mdash;1.5 times the figure in 2014 (Osanai, \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). A large-scale, ten-year study in Korea found that nests of wasps were concentrated in low-rise residential areas within 170 meters of parks or green spaces, and that urban heat islands may facilitate nest establishment (Song et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Our study uses municipal extermination records to examine how human-wasp conflicts manifest across urban-rural gradients, providing a socioecological perspective complementary to previous ecological studies.\u003c/p\u003e \u003cp\u003eAlthough urban greening continues under the banner of nature-positive development, it will likely increase wasp nesting, raising potential costs for residents and local governments. While some individuals may remove nests themselves, most rely on municipal or private pest control services. In addition to ecological and demographic variables, human perceptions and emotional responses toward insects\u0026mdash;especially wasps\u0026mdash;can strongly mediate the frequency of extermination requests. Recent studies have emphasized that negative attitudes toward social wasps are often rooted in affective biases such as fear and disgust, which are themselves reinforced by limited direct interactions with nature and negative media portrayals (Sumner et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Oi et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Schmack et al., \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). The \u0026ldquo;extinction of experience\u0026rdquo; phenomenon\u0026mdash;the progressive loss of direct contact with nature in daily life\u0026mdash; has been linked to diminished understanding of ecological roles and greater intolerance of insects (Gaston and Soga, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Moreover, the urbanization\u0026ndash;disgust hypothesis suggests that people living in urban areas experience more indoor insect encounters, which trigger stronger disgust reactions, while simultaneously possessing lower insect knowledge\u0026mdash;factors that amplify extermination behavior even toward ecologically beneficial taxa such as \u003cem\u003ePolistes\u003c/em\u003e (Brock et al., \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Fukano and Soga, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo inform urban planning and relevant policy areas such as environmental education, it is essential to understand how local environmental characteristics are associated with ecosystem disservices. Neither ecological data on wasps nor perception surveys alone are sufficient to capture the full picture. Instead, data that reflect actual interactions between humans and wasps are needed. In this respect, extermination request records are a valuable resource, as they are influenced not only by ecological factors such as nest density, but also by human factors including perceived risk, fear, and tolerance. These records likely correlate with local environments that shape both insect populations and resident attitudes. Previous studies have used such records to reveal seasonal trends and habitat preferences in urban areas (e.g., Song et al., \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2025\u003c/span\u003e; Park and Jung, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), but few have combined them with administratively comprehensive data linked to spatially explicit environmental variables across a major city. In this study, we take advantage of a rare dataset from Kobe City, where extermination costs were fully subsidized by the local government during the study period and nearly all requests were processed through a centralized municipal system. This allows for robust analysis of how land use and demographic factors shape patterns of human\u0026ndash;wasp conflict.\u003c/p\u003e \u003cp\u003eThis study focuses on Kobe City because it is one of the major cities in Japan and among the large cities in the country, it has rich natural lands and socioecological landscapes shaped by diverse ecosystems including forestlands, agricultural lands, and water bodies, which can be potential habitats of wasps. It spans 553 km\u0026sup2; with a green cover ratio of 68.5% and a population of approximately 1.5\u0026nbsp;million. It is geographically divided by the Rokko mountain range, which peaks at 931 meters in elevation. Compared to surrounding municipalities, Kobe contains many suburban housing developments built on former forested land. The city stretches from urbanized southern districts to northern rural zones, resulting in a high diversity of land-use types and landscapes at the neighborhood scale. From 2010 to 2021, Kobe City fully subsidized the cost of wasp nest extermination for private households. During this period, most extermination cases were handled through the city, providing an administratively centralized and near-complete record of pest control requests. Pest control professionals were able to distinguish between \u003cem\u003eVespa\u003c/em\u003e and \u003cem\u003ePolistes\u003c/em\u003e nests in these records.\u003c/p\u003e \u003cp\u003eIn Japan, the genera \u003cem\u003eVespa\u003c/em\u003e and \u003cem\u003ePolistes\u003c/em\u003e both comprise eusocial wasps that form annual colonies initiated by a single overwintered queen (Spradbery, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1973\u003c/span\u003e; Matsuura \u0026amp; Yamane \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1990\u003c/span\u003e). However, they differ markedly in ecology, colony size, and risk to humans. Vespa species typically build large, enclosed nests in underground cavities, tree hollows, or building structures, and colonies can reach several thousand individuals by late summer. These wasps are highly defensive and are responsible for the majority of sting-related injuries and fatalities. Indeed, in Japan, the number of deaths caused by wasp stings annually exceeds those caused by bears or venomous snakes (Matsuura, \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). In contrast, \u003cem\u003ePolistes\u003c/em\u003e wasps construct small, open-comb nests with fewer than 100 individuals, usually under eaves or on fences. While they are generally less aggressive, their close association with human dwellings increases the likelihood of contact and extermination. These ecological and behavioral distinctions shape public perception and management responses, and are central to understanding the contrasting extermination patterns analyzed in this study.\u003c/p\u003e \u003cp\u003eThe aim of this study was to analyze data on wasp extermination requests from 2019 to 2021 in Kobe City, comparing the two genera (\u003cem\u003eVespa\u003c/em\u003e and \u003cem\u003ePolistes\u003c/em\u003e) with differing risk profiles. By incorporating gradients of urbanization, population size, and agricultural land cover into statistical models, we sought to clarify the relationship between local environmental characteristics and wasp extermination patterns. Specifically, we treated each neighborhood-level district as a spatial analysis unit and integrated nest removal records with data on land use (urban area, farmland), population, and spatial autocorrelation. Using multivariate regression analysis, we evaluated the extent to which these geographic and demographic factors explained variation in wasp extermination frequencies. These findings may help inform targeted pest control strategies and guide sustainable urban landscape planning. While previous studies have examined hornet abundance in relation to urban greenness using standardized trapping methods (Azmy et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), our approach using municipal extermination records provides a unique perspective on actual human-wasp conflicts. Unlike trap-based studies that sample from predetermined locations (typically parks), extermination records reflect the intersection of ecological occurrence and human behavioral responses across the entire urban landscape, including private properties where most conflicts occur.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003eStudy System\u003c/p\u003e \u003cp\u003eData on extermination requests for wasps (\u003cem\u003eVespa\u003c/em\u003e spp. and \u003cem\u003ePolistes\u003c/em\u003e spp.) in Kobe City, Japan, were obtained from records maintained by the Kobe City Public Health Bureau between 2019 and 2021. The number of requests for each year is 3270 in 2019, 2601 in 2020, and 2045 in 2021. In total, 7,916 extermination request records were collected. Such datasets are typically fragmented across multiple organizations in Japanese cities, as extermination responsibilities are distributed among several entities, and city governments rarely collect comprehensive data. Kobe City represents a rare case where a single organization handled all exterminations, enabling the city government to collect comprehensive data. This study represents the first analysis of the largest municipal wasp extermination dataset in Japan. The individual record includes the information of location (neighborhood-level district names) of the extermination, exterminated species, and dates. These records were aggregated by neighborhood-level districts (n\u0026thinsp;=\u0026thinsp;779), which served as the basic spatial units of analysis in this study (hereafter referred to as \"units\").\u003c/p\u003e \u003cp\u003eLand-use metrics for each unit were derived using QGIS version 3.22.8 and the High Resolution Land-Use and Land-Cover Map published by the Japan Aerospace Exploration Agency (JAXA, 2022). Following ecological studies indicating that Vespinae wasps typically forage over distances of at least 1 km (Matsuura and Yamane, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1990\u003c/span\u003e) and previous research demonstrating that 1\u0026ndash;2 km radii effectively predict hornet abundance (Azmy et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e), we calculated the area of developed land, farmland, and forest within a 1 km radius from the centroid of each unit. The original dataset includes address-level extermination records and was provided by the Kobe City Public Health Bureau under a data-sharing agreement that prohibits public dissemination of raw data.\u003c/p\u003e \u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStatistical Analysis\u003c/h2\u003e \u003cp\u003eWe analyzed how extermination frequency per unit was influenced by surrounding land use and demographic characteristics between 2019 and 2021. The main response variables were the annual number of exterminations of Vespa and Polistes wasps per unit. Due to the large number of units with zero extermination cases and the presence of overdispersion, we applied generalized linear mixed models (GLMMs) with a negative binomial distribution and a logarithmic link function. Explanatory variables included the developed land area, farmland area, and log₁₀-transformed population size for each unit. To allow for potential non-linear effects, we included quadratic terms for these predictors. The variable \"year\" was modeled as a random intercept to account for temporal variation. In addition, spatial autocorrelation was accounted for by including spatial autocovariates in the analysis, which were calculated for each study site based on latitude and longitude. Forest land area was excluded from the model due to multicollinearity, as indicated by high variance inflation factor (VIF) values: developed land\u0026thinsp;=\u0026thinsp;32.74, forest\u0026thinsp;=\u0026thinsp;26.78, farmland\u0026thinsp;=\u0026thinsp;7.14, log₁₀(population)\u0026thinsp;=\u0026thinsp;1.41, and spatial autocovariate\u0026thinsp;=\u0026thinsp;1.17 (Fox et al., 2007).\u003c/p\u003e \u003cp\u003eTo assess how land-use characteristics influenced the relative frequency of \u003cem\u003ePolistes\u003c/em\u003e exterminations among all wasp exterminations, we conducted a complementary analysis using a binomial GLMM with a logit link function. The response variable was the proportion of \u003cem\u003ePolistes\u003c/em\u003e exterminations per unit. Explanatory variables included the ratio of developed land to farmland and log₁₀-transformed population, while spatial autocovariates were again included to control for spatial dependency. Quadratic terms were not included in this model. As the logit link function already captures non-linearity, and exploratory analyses indicated that relationships between predictors and the logit-transformed response were largely monotonic, quadratic terms were not expected to improve model fit and could lead to overfitting or implausible effects. For model simplicity and interpretability, we retained only linear terms. All analyses were conducted in R version 4.2.2 (R Core Team, 2021) using the \u003cem\u003eglmmTMB\u003c/em\u003e package (Magnusson et al., 2017) and/or the \u003cem\u003espdep\u003c/em\u003e package (Bivand et al., \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cp\u003eModel Adequacy\u003c/p\u003e \u003cp\u003eThe three models presented in this study (\u003cem\u003eVespa\u003c/em\u003e extermination count, \u003cem\u003ePolistes\u003c/em\u003e extermination count, and \u003cem\u003ePolistes\u003c/em\u003e extermination ratio) were all based on negative binomial or binomial GLMMs and accounted appropriately for overdispersion and year effects. The estimated dispersion parameters (θ\u0026thinsp;=\u0026thinsp;1.54 for Vespa, 2.75 for Polistes) fell within the empirically acceptable range for ecological data (Ver Hoef \u0026amp; Boveng, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2007\u003c/span\u003e). The variance of the random effect for year was negligible or small across models (σ\u0026sup2; = 2.1 \u0026times; 10⁻⁹, 0.068, and 0.187), indicating limited inter-annual variation (Nakagawa \u0026amp; Schielzeth, \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2013\u003c/span\u003e). These values confirm that all models were well-specified and structurally sound for inference and prediction.\u003c/p\u003e \u003cp\u003eExtermination Patterns by Species\u003c/p\u003e \u003cp\u003e \u003cem\u003eVespa\u003c/em\u003e Extermination Counts\u003c/p\u003e \u003cp\u003eAmong explanatory variables, the proportion of developed land exhibited a significant quadratic relationship: the linear term was positive (β\u0026thinsp;=\u0026thinsp;1.05, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.022), and the quadratic term was negative (β = \u0026minus;3.30, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). The number of exterminations peaked when developed land comprised approximately 15\u0026ndash;20% of the unit area (IRR\u0026thinsp;=\u0026thinsp;2.86, 95% CI\u0026thinsp;=\u0026thinsp;1.17\u0026ndash;6.97), then declined sharply beyond this threshold (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eFarmland proportion showed a significant negative effect (β = \u0026minus;2.03, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.006), with a 20% increase in farmland associated with a 34% decrease in extermination count. The population variable showed a significant quadratic effect (β\u0026thinsp;=\u0026thinsp;0.248, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), indicating an accelerating increase in exterminations with population growth. Spatial autocorrelation also had a significant positive effect (β\u0026thinsp;=\u0026thinsp;0.333, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), suggesting that Vespa exterminations were spatially clustered.\u003c/p\u003e \u003cp\u003e \u003cem\u003ePolistes\u003c/em\u003e Extermination Counts\u003c/p\u003e \u003cp\u003eExtermination counts exhibited a significant quadratic relationship with developed land: the linear term was positive (β\u0026thinsp;=\u0026thinsp;2.32, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001) and the quadratic term was negative (β = \u0026minus;2.78, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). The peak extermination count occurred at around 42% developed land (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003eb). Farmland proportion was not significantly associated with Polistes exterminations.\u003c/p\u003e \u003cp\u003ePopulation size showed a significant positive linear effect (β\u0026thinsp;=\u0026thinsp;1.67, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.006), indicating higher extermination counts in more densely populated areas. Spatial autocorrelation also showed a significant positive effect (β\u0026thinsp;=\u0026thinsp;0.336, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), reflecting spatial clustering of Polistes exterminations.\u003c/p\u003e \u003cp\u003eProportion of \u003cem\u003ePolistes\u003c/em\u003e Among Total Exterminations\u003c/p\u003e \u003cp\u003eThe proportion of Polistes exterminations increased significantly with the proportion of developed land (β\u0026thinsp;=\u0026thinsp;2.15, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001). In central urban areas, this proportion was approximately 8.6 times greater than in less urbanized units (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Farmland proportion showed a marginally significant negative effect (β = \u0026minus;0.60, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.077), suggesting that Vespa exterminations may be relatively more common in agricultural areas. Population size was positively associated with the proportion of Polistes (β\u0026thinsp;=\u0026thinsp;0.234, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;\u0026lt;\u0026thinsp;0.001), indicating that every 10-fold increase in population size raised the odds of a Polistes extermination by approximately 1.26 times. In contrast, spatial autocorrelation showed a significant negative effect (β = \u0026minus;0.14, \u003cem\u003eP\u003c/em\u003e\u0026thinsp;=\u0026thinsp;0.003), suggesting that in units with highly clustered exterminations, \u003cem\u003eVespa\u003c/em\u003e wasps tend to dominate. The estimated intercept (β = \u0026minus;2.71) corresponds to a baseline \u003cem\u003ePolistes\u003c/em\u003e extermination proportion of 6.6% under average conditions. In densely urbanized areas (urban proportion\u0026thinsp;=\u0026thinsp;0.8, log₁₀ population\u0026thinsp;\u0026asymp;\u0026thinsp;5), the model predicts that Polistes may account for approximately 30% of all exterminations. To further clarify the species-specific responses to urbanization, we grouped sites into quartiles based on urbanization level and calculated mean extermination counts per genus. As shown in Fig.\u0026nbsp;3, \u003cem\u003eVespa\u003c/em\u003e exterminations peaked at moderate levels of urbanization (Q2), whereas \u003cem\u003ePolistes\u003c/em\u003e showed higher frequencies in highly urbanized areas (Q3\u0026ndash;Q4).\u003c/p\u003e\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" style=\"width: 78px;\"\u003e\n \u003cp\u003eTable1. Model results for \u003cem\u003eVespa\u003c/em\u003e and \u003cem\u003ePolistes\u003c/em\u003e exterminations\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003e \u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e \u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 24px;\"\u003e\n \u003cp\u003e\u003cem\u003eVespa\u003c/em\u003e (AIC=7076.2)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 25px;\"\u003e\n \u003cp\u003e\u003cem\u003ePolistes\u003c/em\u003e (AIC=5216)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\u003cbr\u003e\u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\n \u003cp\u003e\u003cem\u003ePredictor\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003eEstimate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 9px;\"\u003e\n \u003cp\u003eSE\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003ep-value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003eEstimate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003eSE\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003ep-value\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\n \u003cp\u003e(Intercept)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e-1.058\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 9px;\"\u003e\n \u003cp\u003e0.326\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e-5.786\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003e0.961\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\n \u003cp\u003eDeveloped land\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e1.045\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 9px;\"\u003e\n \u003cp\u003e0.459\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e0.022\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e2.318\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003e0.601\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\n \u003cp\u003eDeveloped land\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e-3.301\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 9px;\"\u003e\n \u003cp\u003e0.472\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e-2.780\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003e0.582\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\n \u003cp\u003eFarmland\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e-2.031\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 9px;\"\u003e\n \u003cp\u003e0.735\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e0.006\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e-1.530\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003e1.000\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e0.126\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\n \u003cp\u003eFarmland\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e0.377\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 9px;\"\u003e\n \u003cp\u003e1.047\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e0.719\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e0.118\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003e1.522\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e0.938\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\n \u003cp\u003eLog\u003csub\u003e10\u003c/sub\u003e(Population)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e-0.095\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 9px;\"\u003e\n \u003cp\u003e0.232\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e0.684\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e1.666\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003e0.603\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e0.006\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\n \u003cp\u003eLog\u003csub\u003e10\u003c/sub\u003e(Population)\u003csup\u003e2\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e0.248\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 9px;\"\u003e\n \u003cp\u003e0.041\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e0.011\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003e0.093\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e0.904\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 27px;\"\u003e\n \u003cp\u003eSpatial Autocovariate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e0.333\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 9px;\"\u003e\n \u003cp\u003e0.022\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 14px;\"\u003e\n \u003cp\u003e0.336\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 10px;\"\u003e\n \u003cp\u003e0.049\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 11px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cbr\u003e\n\u003ctable border=\"0\" cellspacing=\"0\" cellpadding=\"0\" width=\"100%\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd colspan=\"5\" style=\"width: 100px;\"\u003e\n \u003cp\u003eTable2. Binomial GLMM results for Polistes extermination proportion\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 33px;\"\u003e\n \u003cp\u003e\u003cem\u003ePredictor\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003eEstimate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15px;\"\u003e\n \u003cp\u003eSE\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 17px;\"\u003e\n \u003cp\u003ez value\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003ep-value\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 33px;\"\u003e\n \u003cp\u003e(Intercept)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e-2.715\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15px;\"\u003e\n \u003cp\u003e0.330\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 17px;\"\u003e\n \u003cp\u003e-8.222\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 33px;\"\u003e\n \u003cp\u003eDeveloped land\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e2.150\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15px;\"\u003e\n \u003cp\u003e0.126\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 17px;\"\u003e\n \u003cp\u003e17.033\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 33px;\"\u003e\n \u003cp\u003eFarmland\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e-0.595\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15px;\"\u003e\n \u003cp\u003e0.337\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 17px;\"\u003e\n \u003cp\u003e-1.769\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e0.077\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 33px;\"\u003e\n \u003cp\u003eLog\u003csub\u003e10\u003c/sub\u003e(Population)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e0.234\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15px;\"\u003e\n \u003cp\u003e0.066\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 17px;\"\u003e\n \u003cp\u003e3.539\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e\u0026lt;0.001\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 33px;\"\u003e\n \u003cp\u003eSpatial autocovariate\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e-0.143\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 15px;\"\u003e\n \u003cp\u003e0.048\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 17px;\"\u003e\n \u003cp\u003e-2.989\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 16px;\"\u003e\n \u003cp\u003e0.002\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e"},{"header":"Discussion","content":"\u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eUrban gradient effects on genus-specific human-wasp conflicts\u003c/h2\u003e \u003cp\u003eThis study revealed distinct associations between extermination requests for \u003cem\u003eVespa\u003c/em\u003e (hornets) and \u003cem\u003ePolistes\u003c/em\u003e (paper wasps) and land-use characteristics in Kobe City. Our findings complement ecological studies from Nagoya, which showed species-specific responses to urban greenness (Azmy et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). However, our conflict data revealed different patterns: \u003cem\u003eVespa\u003c/em\u003e exterminations peaked at intermediate urbanization (15\u0026ndash;20% developed land) rather than increasing monotonically with greenness, highlighting how human behavioral factors modify ecological patterns in real-world conflict scenarios.\u003c/p\u003e \u003cp\u003eExtermination counts were also negatively associated with farmland cover and positively associated with population size and spatial autocorrelation, suggesting that both ecological conditions and human behavioral factors shape reported disservices (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003ea, b). In contrast, \u003cem\u003ePolistes\u003c/em\u003e extermination counts were highest in areas with approximately 40% developed land, showed no significant association with farmland, increased linearly with population size, and were similarly clustered spatially. Moreover, the proportion of \u003cem\u003ePolistes\u003c/em\u003e exterminations relative to all wasp exterminations increased with urbanization, indicating their dominance in highly developed areas. These patterns refine earlier findings, such as those by Song et al. (\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2025\u003c/span\u003e), who reported more \u003cem\u003eVespidae\u003c/em\u003e nests near green areas in South Korea. While proximity to green space may indeed promote nesting, our results indicate that wasp-human interactions cannot be explained by developed land alone. Instead, extermination requests are shaped by a complex interplay of urbanization, population density, land-use type, and local perceptions of risk.\u003c/p\u003e \u003cp\u003e \u003cb\u003eVespa\u003c/b\u003e \u003cb\u003eExtermination Counts\u003c/b\u003e\u003c/p\u003e \u003cp\u003eThe peak in \u003cem\u003eVespa\u003c/em\u003e exterminations at intermediate levels of development may reflect the species\u0026rsquo; preference for semi-natural environments and the increased likelihood of human-wasp encounters in moderately populated zones. Although \u003cem\u003eVespa\u003c/em\u003e nests were less frequently reported in highly urbanized areas, some city units with over 85% developed land still recorded high extermination counts, underscoring the persistence of potential hazards even in dense urban centers. Farmland coverage was negatively associated with \u003cem\u003eVespa\u003c/em\u003e exterminations, which may reflect reduced nesting opportunities or differences in local residents\u0026rsquo; attitudes and knowledge about wasps. People familiar with rural insects may be more tolerant of wasp presence or capable of managing nests independently, while newer suburban communities might be more likely to request professional extermination. These behavioral differences may be influenced by media portrayals that emphasize the lethal risks of stings (Oi et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2024\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe accelerated increase in \u003cem\u003eVespa\u003c/em\u003e exterminations with population size, despite a flattening at higher urbanization levels, suggests a trade-off: while denser populations increase wasp-human encounters, extreme urbanization suppresses wasp nesting opportunities. The strong spatial autocorrelation may be driven by limited dispersal distances of overwintered queens and similarities in local environmental and social contexts, such as the prevalence of green spaces or public aversion to wasps in tightly packed residential areas. Pesticide use in farmland might further suppress \u003cem\u003eVespa\u003c/em\u003e populations, either directly or indirectly through prey reduction (Tison et al., \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, no similar effect was observed for \u003cem\u003ePolistes\u003c/em\u003e, possibly due to differences in diet or nesting behavior.\u003c/p\u003e \u003cp\u003e \u003cb\u003ePolistes\u003c/b\u003e \u003cb\u003eExtermination Counts\u003c/b\u003e\u003c/p\u003e \u003cp\u003e \u003cem\u003ePolistes\u003c/em\u003e were more common in built environments, consistent with their preference for nesting on human-made structures. Their extermination frequency increased linearly with population size, implying both ecological tolerance and increased human-wasp contact in denser areas. In rural areas, nests may be left undisturbed or managed independently, suggesting that local perceptions of risk and response thresholds vary with setting. Compared to \u003cem\u003eVespa\u003c/em\u003e, \u003cem\u003ePolistes\u003c/em\u003e exterminations were more strongly associated with developed land and population, but not farmland, highlighting divergent ecological preferences and public responses to each genus. These differences likely reflect not only biological traits\u0026mdash;such as nest size and aggression\u0026mdash;but also variation in risk perception.\u003c/p\u003e \u003cp\u003e \u003cb\u003eProportion of\u003c/b\u003e \u003cb\u003ePolistes\u003c/b\u003e \u003cb\u003eAmong Total Exterminations\u003c/b\u003e\u003c/p\u003e \u003cp\u003eThe proportion of \u003cem\u003ePolistes\u003c/em\u003e among all exterminations was highest in highly urbanized areas, potentially reflecting a loss of suitable nesting habitats for \u003cem\u003eVespa\u003c/em\u003e (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Similar shifts in species dominance in urban contexts have been observed elsewhere (e.g., Baker et al., 2020), where smaller wasps thrive on unique urban nesting substrates and prey.\u003c/p\u003e \u003cp\u003eImportantly, the number of extermination requests does not directly reflect wasp nest density but rather a combination of ecological occurrence and human behavior. Thus, extermination data serve as a proxy for ecosystem disservices from a socioecological perspective. Future research should integrate field nest surveys with social surveys on risk tolerance to better assess the true societal cost of urban greening under \u0026ldquo;nature-positive\u0026rdquo; agendas. Quantifying disservices in monetary terms\u0026mdash;including extermination costs, medical expenses, and labor losses\u0026mdash;may also help optimize policy decisions balancing ecosystem services and disservices.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eSpatial patterns and housing development characteristics\u003c/h3\u003e\n\u003cp\u003eThe detailed spatial distribution of extermination requests reveals important relationships between urban development patterns and human-wasp conflicts (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e4\u003c/span\u003e). \u003cem\u003eVespa\u003c/em\u003e exterminations were primarily observed in relatively rural areas on the northern side of Kobe City and at the boundaries between urban and rural areas, while \u003cem\u003ePolistes\u003c/em\u003e exterminations were concentrated in northern urban areas and urban-rural boundary zones. These spatial patterns are consistent with our statistical model results and provide additional insights into the socioecological drivers of extermination behavior.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eThe northern urban areas, developed primarily after the 1960s-1970s, represent what are commonly called \"new towns\" in Japan\u0026mdash;suburban housing developments popular among families seeking new homes. Unlike dense urban cores dominated by apartments, these new towns predominantly feature detached houses with gardens and eaves, providing ideal nesting sites for both genera. Both \u003cem\u003ePolistes\u003c/em\u003e and \u003cem\u003eVespa\u003c/em\u003e commonly nest in garden hedges and under eaves, while some \u003cem\u003eVespa\u003c/em\u003e species additionally utilize enclosed spaces such as attics and building foundations (Spradbery, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e1973\u003c/span\u003e; Matsuura \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e1995\u003c/span\u003e). This housing typology creates abundant nesting opportunities while increasing human-wasp encounter rates through gardening and outdoor activities around residences.\u003c/p\u003e \u003cp\u003eThe combination of suitable nesting habitat and resident characteristics may explain the elevated \u003cem\u003ePolistes\u003c/em\u003e extermination rates in these areas. New residents may lack familiarity with \u003cem\u003ePolistes\u003c/em\u003e behavior and management techniques, leading to increased reliance on professional extermination services. In contrast, residents in surrounding rural areas, who typically grew up locally, may possess greater tolerance and experience cohabiting with these wasps. For \u003cem\u003eVespa\u003c/em\u003e, this residential experience difference appears less pronounced, likely reflecting the more universal perception of hornets as a significant threat regardless of residents' background.\u003c/p\u003e \u003cp\u003eThese findings highlight how urban development patterns inadvertently create conditions that promote both wasp abundance and human-wildlife conflicts, exemplifying the ecosystem disservices associated with suburban expansion.\u003c/p\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eSociocultural drivers of extermination behavior\u003c/h2\u003e \u003cp\u003eBeyond ecological distribution, human emotional and cognitive responses play a crucial role in shaping wasp extermination patterns. In particular, urban residents are more likely to encounter insects indoors, a context that tends to evoke heightened feelings of disgust (Fukano and Soga, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). This tendency is reinforced by the general lack of direct nature experience in urban environments\u0026mdash;a phenomenon described as the extinction of experience\u0026mdash;which contributes to a lower tolerance for insects, even when they are harmless or beneficial (Gaston and Soga, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Media portrayals that emphasize the danger or aggressiveness of wasps, in contrast to the more favorable framing of bees, further amplify public aversion and may distort risk perception (Oi et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Sumner et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eImportantly, recent findings indicate that even individuals with some knowledge about wasps, such as urban gardeners, often express strong emotional aversion. Schmack et al. (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2024\u003c/span\u003e) showed that knowledge alone may not suffice to overcome affective biases, underscoring the importance of addressing both cognitive and emotional dimensions in any intervention strategy.\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eEnvironmental education implications\u003c/h3\u003e\n\u003cp\u003eThe spatial distribution of extermination requests showed different patterns between the two genera. This result suggests that contents of education to deal with hornets and paper wasps should be tailored based on degree of urbanization in each neighborhood-level district. For example, urban centers and areas in between urban and rural areas might need more knowledge on \u003cem\u003ePolistes\u003c/em\u003e. For rural areas, dissemination of knowledge on \u003cem\u003eVespa\u003c/em\u003e should be prioritized. Such contextualized approach can effectively reduce the negative influence of the disservices from these species and to raise awareness of their importance in local ecosystems based on hands-on experience (Fig.\u0026nbsp;3).\u003c/p\u003e \u003cp\u003eIn addition to species-specific knowledge, emotional and cultural biases must also be considered when designing educational programs. For instance, the extinction of experience has reduced familiarity with insects among urban residents, amplifying feelings of disgust and fear (Gaston and Soga, \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Fukano and Soga, \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Moreover, negative portrayals of wasps in the media may further reinforce aversion (Oi et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2024\u003c/span\u003e; Sumner et al., \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). Therefore, effective education should incorporate both factual knowledge and indirect experiential learning, such as observing wasps safely in the environment or learning through case studies, to foster tolerance and reduce unnecessary extermination. These findings complement ecological research showing species-specific habitat preferences (Azmy et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) by revealing how human behavioral responses vary across the urban-rural gradient. While ecological studies inform us about where wasps are likely to be abundant, extermination data reveals where conflicts actually occur, providing crucial information for targeted education and management interventions.\u003c/p\u003e\n\u003ch3\u003eImplications for invasive species risk assessment\u003c/h3\u003e\n\u003cp\u003eWhile the introduction highlighted the growing global concern surrounding Vespidae invasions, our findings also contribute to this field by offering actionable insights into how urban wasp disservices are mediated by local ecological and sociocultural contexts. Around the world, invasive species in the Vespidae family\u0026mdash;such as \u003cem\u003eVespa velutina\u003c/em\u003e, \u003cem\u003eVespula vulgaris\u003c/em\u003e, and \u003cem\u003eVespula germanica\u003c/em\u003e\u0026mdash;have caused severe ecological, economic, and public health impacts, particularly in Europe and New Zealand (Lester et al., \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Monceau et al., \u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). However, as noted in recent syntheses, the severity of such conflicts is not solely determined by wasp behavior or population dynamics, but also by how humans perceive, tolerate, and respond to these insects (Wilson Rankin, \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2021\u003c/span\u003e; Lester et al., \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2025\u003c/span\u003e). Our spatially explicit data on extermination requests\u0026mdash;capturing variation in urbanization, land use, and perceived risk\u0026mdash;may provide a valuable empirical basis for modeling future wasp-human interactions under scenarios of biological invasion. These findings underscore the need to incorporate socioecological perspectives into early-warning frameworks for invasive wasp management, especially in urban landscapes where human responses can critically shape outcomes.\u003c/p\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eMethodological Contributions and Limitations\u003c/h2\u003e \u003cp\u003eOur study demonstrates the value of municipal extermination records as a complementary approach to standardized ecological sampling. While trap-based studies like Azmy et al. (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) provide controlled measures of hornet abundance, they are limited to accessible public spaces and may not reflect private property conflicts where most human-wasp interactions occur. Conversely, extermination records capture actual conflict patterns but are influenced by socioeconomic factors, risk perception, and reporting behaviors that vary across communities.\u003c/p\u003e \u003cp\u003eThe species-specific patterns we observed with \u003cem\u003eVespa\u003c/em\u003e conflicts peaking at intermediate urbanization and Polistes increasing with urban density, suggest that extermination data provides unique insights into the socioecological dimensions of ecosystem disservices that cannot be captured through ecological sampling alone. Future research integrating both approaches would provide a more complete understanding of urban wasp dynamics.\u003c/p\u003e \u003c/div\u003e"},{"header":"Conclusion","content":"\u003cp\u003eThis study demonstrates how spatial patterns of extermination requests for social wasps in Kobe City are shaped by land use, population density, and species-specific ecological and behavioral traits. \u003cem\u003eVespa\u003c/em\u003e and \u003cem\u003ePolistes\u003c/em\u003e differ not only in nesting preferences and colony size but also in how the public perceives and responds to them across urban gradients. These findings emphasize that urban ecosystem disservices are socioecological in nature and should be addressed through context-specific strategies. Our results provide actionable insights for designing pest management and biodiversity education programs that reflect both local ecological conditions and human risk perception. Furthermore, by establishing spatial baselines of human\u0026ndash;wasp interaction, this study supports the development of early-warning frameworks and response policies for invasive wasp species. Such approaches are especially relevant in rapidly urbanizing regions where environmental change and public sensitivity converge to shape future biodiversity challenges. By integrating ecological and conflict pattern analyses, urban planners can develop strategies that account for both biological distributions and human behavioral responses to urban wildlife.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgments\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe are grateful to Dr. Atsushi Ushimaru for his valuable advice on data analysis. We also thank the Kobe City Public Health Bureau and the Hyogo Pest Control Association for their cooperation in data collection.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;Funding Statement\u003c/p\u003e\n\u003cp\u003eThis study was supported by a research grant from the Graduate School of Human Development and Environment, Kobe University.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;Data availability statement\u003c/p\u003e\n\u003cp\u003eThe extermination request data analyzed in this study were obtained from the Kobe City Public Health Bureau (Kobe City Kenkōkyoku). These data contain address-level information and are not publicly shareable due to privacy regulations. To protect individual confidentiality, only aggregated statistics and model results are available. Summary tables and regression outputs can be provided by the corresponding author upon reasonable request and with permission from the Kobe City Public Health Bureau.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;Author contribution\u003c/p\u003e\n\u003cp\u003eT.S. led the project and was responsible for conceptualization, methodology, investigation, formal analysis, funding acquisition, and original draft writing. T.N. contributed to data analysis and visualization. Y.U. contributed to conceptual development, data processing, visualization, and discussions on methodology. All authors reviewed and edited the manuscript.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;Declaration of Competing Interests\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAzmy, M.M., Hosaka, T., Numata, S., 2016. Responses of four hornet species to levels of urban greenness in Nagoya city, Japan: Implications for ecosystem disservices of urban green spaces. Urban Forestry \u0026amp; Urban Greening 18, 117-125. https://doi.org/10.1016/j.ufug.2016.05.014\u003c/li\u003e\n\u003cli\u003eBaker, A. M., \u0026amp; Potter, D. A. (2020). Invasive paper wasp turns urban pollinator gardens into ecological traps for monarch butterfly larvae. 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Recent advances and avenues for the pest management of invasive social wasps and hornets. \u003cem\u003eCurrent Opinion in Insect Science\u003c/em\u003e, 68, 101336. https://doi.org/10.1016/j.cois.2025.101336\u003c/li\u003e\n\u003cli\u003eLyytim\u0026auml;ki, J. (2015). Ecosystem disservices: Embrace the catchword. Ecosystem Services, 12, 136. https://doi.org/10.1016/j.ecoser.2014.11.008\u003c/li\u003e\n\u003cli\u003eLyytim\u0026auml;ki, J., Petersen, L. K., Normander, B., \u0026amp; Bez\u0026aacute;k, P. (2008). Nature as a nuisance? Ecosystem services and disservices to urban lifestyle. Environmental Sciences, 5(3), 161\u0026ndash;172. https://doi.org/10.1080/15693430802055524\u003c/li\u003e\n\u003cli\u003eMatsuura, M. (1995). Social wasps of Japan in color. Hokkaido University Press.\u003c/li\u003e\n\u003cli\u003eMatsuura, M., \u0026amp; Yamane, S. (1990). Biology of the vespine wasps. Springer.\u003c/li\u003e\n\u003cli\u003eMoesch, S. S., Wellmann, T., Haase, D., \u0026amp; Bhardwaj, M. (2024). Mammal Mia: A review on how ecological and human dimension research on urban wild mammals can benefit future biophilic cities. Basic and Applied Ecology, 79, 90-101. https://doi.org/10.1016/j.baae.2024.05.004\u003c/li\u003e\n\u003cli\u003eMonceau, K., Bonnard, O., \u0026amp; Thi\u0026eacute;ry, D. (2014). \u003cem\u003eVespa velutina\u003c/em\u003e: A new invasive predator of honeybees in Europe. \u003cem\u003eJournal of Pest Science\u003c/em\u003e, 87(1), 1\u0026ndash;16. https://doi.org/10.1007/s10340-013-0537-3\u003c/li\u003e\n\u003cli\u003eNakagawa, S., \u0026amp; Schielzeth, H. (2013). A general and simple method for obtaining R\u0026sup2; from generalized linear mixed-effects models. Methods in Ecology and Evolution, 4(2), 133\u0026ndash;142. https://doi.org/10.1111/j.2041-210x.2012.00261.x\u003c/li\u003e\n\u003cli\u003eOi, C. A., Brown, R. L., \u0026amp; Sumner, S. (2024). Bee-ing positive about wasp-negative media reporting: The opinions of scientists and their influence on the media. Insectes Sociaux, 71(1), 29\u0026ndash;42. https://doi.org/10.1007/s00040-024-00952-9\u003c/li\u003e\n\u003cli\u003eOsanai, R. (2024). Ecology and control of hornets. Pest Control [in Japanese], 207, 10-14.\u003c/li\u003e\n\u003cli\u003ePark, J., \u0026amp; Jung, C. (2020). Analysis of reports on wasp nest removal in Seoul, Korea: Spatial and temporal patterns of human\u0026ndash;insect conflict. \u003cem\u003eProceedings of the National Institute of Ecology of the Republic of Korea, 2\u003c/em\u003e(2), 101\u0026ndash;107. https://doi.org/10.22920/PNIE.2020.2.2.101\u003c/li\u003e\n\u003cli\u003eRobinet, C., Suppo, C., \u0026amp; Darrouzet, E. (2016). Rapid spread of the invasive yellow-legged hornet in France: The role of human-mediated dispersal and the effects of control measures. \u003cem\u003eJournal of Applied Ecology\u003c/em\u003e, 54(1), 205\u0026ndash;215. https://doi.org/10.1111/1365-2664.12724\u003c/li\u003e\n\u003cli\u003eSchmack, J. M., Egerer, M. H., Karlebowski, S., Neumann, A. E., \u0026amp; Sturm, U. (2024). Overlooked and misunderstood: How urban community gardeners perceive social wasps and their ecosystem functions. Journal of Insect Conservation, 28(2), 283\u0026ndash;289. https://doi.org/10.1007/s10841-024-00548-5\u003c/li\u003e\n\u003cli\u003eShackleton, C. M., Ruwanza, S., Sinasson Sanni, G. K., Bennett, S., De Lacy, P., Modipa, R., ... \u0026amp; Thondhlana, G. (2016). Unpacking Pandora\u0026apos;s box: understanding and categorising ecosystem disservices for environmental management and human wellbeing. Ecosystems, 19(4), 587-600.\u003c/li\u003e\n\u003cli\u003eShaikh, S. F. E. A., \u0026amp; Hamel, P. (2023). Identifying nature-positive futures in new cities: An application of the Urban Nature Futures Framework. Sustainability Science, 18(1), 123\u0026ndash;134. https://doi.org/10.1007/s11625-023-01411-3\u003c/li\u003e\n\u003cli\u003eSong, W., Kim, H., \u0026amp; Kim, W. (2025). Modeling urban wasp nest occurrences using 119 fire service reports, LiDAR, and hyperspectral imagery: The role of green spaces and structural factors. Journal of Environmental Management, 379, 124776.\u003c/li\u003e\n\u003cli\u003eSpradbery, J. P. (1973). Wasps: An account of the biology and natural history of solitary and social wasps. University of Washington Press.\u003c/li\u003e\n\u003cli\u003eSumner, S., Law, G., \u0026amp; Cini, A. (2018). Why we love bees and hate wasps. Ecological Entomology, 43(6), 836\u0026ndash;845. https://doi.org/10.1111/een.12676\u003c/li\u003e\n\u003cli\u003eTison, L., Franc, C., Burkart, L., Jactel, H., Monceau, K., de Revel, G., \u0026amp; Thi\u0026eacute;ry, D. (2023). Pesticide contamination in an intensive insect predator of honey bees. Environment International, 176, 107975. https://doi.org/10.1016/j.envint.2023.107975\u003c/li\u003e\n\u003cli\u003eVer Hoef, J. M., \u0026amp; Boveng, P. L. (2007). Quasi-Poisson vs. negative binomial regression: How should we model overdispersed count data? Ecology, 88(11), 2766\u0026ndash;2772. https://doi.org/10.1890/07-0043.1\u003c/li\u003e\n\u003cli\u003eWilson Rankin, E. E. (2021). Emerging patterns in social wasp invasions. \u003cem\u003eCurrent Opinion in Insect Science\u003c/em\u003e, 46, 72\u0026ndash;77. https://doi.org/10.1016/j.cois.2021.02.014\u003c/li\u003e\n\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"ecosystem disservices, human-wildlife conflict, spatial analysis, wasp management, public risk perception, urban ecology","lastPublishedDoi":"10.21203/rs.3.rs-6891890/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6891890/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eUrban greening initiatives provide numerous ecosystem services but may inadvertently increase human-wildlife conflicts (ecosystem disservices). In Japan, hornets (\u003cem\u003eVespa\u003c/em\u003e spp.) and paper wasps (\u003cem\u003ePolistes\u003c/em\u003e spp.) represent significant disservices due to their stings, which occasionally result in fatalities. To understand how urban environments shape public encounters with these wasps, we analyzed 7,916 extermination request records (2019\u0026ndash;2021) from Kobe City, Japan, a highly urbanized region with steep land-use gradients. Building on previous research in Tokyo and Nagoya, our study presents the first fine-scale analysis of how land-use gradients shape genus-specific wasp extermination patterns, differentiating between \u003cem\u003eVespa\u003c/em\u003e (hornets) and \u003cem\u003ePolistes\u003c/em\u003e (paper wasps). We examined the effects of surrounding land-use characteristics and population density on extermination frequencies across 779 neighborhood-level districts. \u003cem\u003eVespa\u003c/em\u003e exterminations peaked in areas with ~\u0026thinsp;15\u0026ndash;20% developed land and declined sharply in denser urban zones, whereas \u003cem\u003ePolistes\u003c/em\u003e exterminations were highest in ~\u0026thinsp;40% developed areas and increased linearly with population size. Farmland showed a strong negative effect on \u003cem\u003eVespa\u003c/em\u003e exterminations but no effect on \u003cem\u003ePolistes\u003c/em\u003e. The relative share of \u003cem\u003ePolistes\u003c/em\u003e increased with urbanization, rising from 6.6% baseline to approximately 30% in highly developed areas, suggesting an urban dominance of this genus. Spatial analysis revealed that \u003cem\u003ePolistes\u003c/em\u003e exterminations were concentrated in post-1960s suburban \"new towns\" where detached housing with gardens provides ideal nesting sites, while new residents may lack familiarity with wasp management. These patterns reflect both ecological habitat preferences and social perceptions of risk, including psychological factors such as fear, disgust, and limited familiarity with insects in urban settings. These findings highlight the importance of integrating disservice considerations into biodiversity-friendly urban planning and provide actionable thresholds for spatially-targeted management strategies.\u003c/p\u003e","manuscriptTitle":"Spatial patterns of ecosystem disservices across urban-rural gradients: Municipal wasp extermination data from Kobe, Japan","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-18 20:46:30","doi":"10.21203/rs.3.rs-6891890/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-07-09T02:59:26+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-08T23:13:13+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-07-07T13:04:06+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-25T17:23:34+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"123893870423444001174671667912762171421","date":"2025-06-17T22:21:00+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"195803462679436423198307844171738804665","date":"2025-06-17T12:16:37+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"124228193410049617591766005134750445068","date":"2025-06-17T07:01:05+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-17T02:53:08+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-06-17T01:57:22+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-06-16T06:28:06+00:00","index":"","fulltext":""},{"type":"submitted","content":"Urban Ecosystems","date":"2025-06-14T05:13:29+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"urban-ecosystems","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ueco","sideBox":"Learn more about [Urban Ecosystems](https://www.springer.com/journal/11252)","snPcode":"11252","submissionUrl":"https://submission.nature.com/new-submission/11252/3","title":"Urban Ecosystems","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"6d09f0e1-a384-4f65-aa22-caff28ab7690","owner":[],"postedDate":"June 18th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2025-08-25T16:31:01+00:00","versionOfRecord":{"articleIdentity":"rs-6891890","link":"https://doi.org/10.1007/s11252-025-01788-2","journal":{"identity":"urban-ecosystems","isVorOnly":false,"title":"Urban Ecosystems"},"publishedOn":"2025-08-18 16:12:59","publishedOnDateReadable":"August 18th, 2025"},"versionCreatedAt":"2025-06-18 20:46:30","video":"","vorDoi":"10.1007/s11252-025-01788-2","vorDoiUrl":"https://doi.org/10.1007/s11252-025-01788-2","workflowStages":[]},"version":"v1","identity":"rs-6891890","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6891890","identity":"rs-6891890","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}