Understanding Ixodes ricinus occurrence in private yards: influence of yard and landscape features

Understanding Ixodes ricinus occurrence in private yards: influence of yard and landscape features | 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 Understanding Ixodes ricinus occurrence in private yards: influence of yard and landscape features Anna Mazaleyrat, Jonas Durand, Irene Carravieri, Christophe Caillot, and 5 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-4453852/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 10 Oct, 2024 Read the published version in International Journal of Health Geographics → Version 1 posted 9 You are reading this latest preprint version Abstract Background: Lyme borreliosis is the most frequent zoonotic disease in the northern hemisphere and is transmitted by ticks of the genus Ixodes . Although many people are bitten by ticks in private yards, our understanding of the factors associated with their presence in these areas remains limited. To address this gap, we used a citizen science approach to identify the local and landscape features associated with tick presence in yards. Methods: This study was conducted near Nancy, a city in northeastern France, from 2020 to 2022. Citizen scientists collected ticks in their yard on a single event (n = 185) and measured 13 yard features. Additionally, we computed 11 features related to the landscape composition and spatial configuration surrounding these yards. Using mixed-effect generalized linear models, we determined the yard and landscape features associated with the presence of ticks and nymphal Ixodes ricinus (hereafter nymphs), the life stage, and species that mostly bite humans. Results: Despite a low density, ticks were found in 32% of the yards, including yards in urbanized areas. At the transect level, the likelihood of finding a nymph was nearly three times higher in transects shaded by vegetation compared to those in open areas, with no relationship between nymph occurrence and transect location or grass height. At the yard level, the occurrence of ticks and nymphs was related to both yard and landscape characteristics. Nymph and tick occurrence were more than twice as high in yards with signs of deer and a wood/brush pile compared to those without these characteristics, and increased with the connectivity of vegetation areas and the percentage of forest areas in the landscape. Conclusions: Our study reveals that private yards across an urbanization gradient are locations of tick exposure with tick presence linked to both yard and landscape factors. These findings emphasize the importance of public awareness regarding tick exposure in yards and provide crucial insights for future public health prevention campaigns. Ixodes ricinus tick occurrence yard landscape citizen science tick-borne disease prevention Figures Figure 1 Figure 2 Background Ticks are hematophagous ectoparasites and can transmit many bacteria, viruses, and protozoa causing human and animal diseases [ 1 , 2 ]. In the northern hemisphere, Lyme borreliosis is the most frequently reported zoonotic disease and an important economic burden for countries [ 3 , 4 ]. The primary vectors of Lyme borreliosis are ticks from the Ixodes ricinus species complex (hereafter Ixodes ), namely I. ricinus in Europe, I. pacificus in western North America, I. scapularis in eastern and mid-western USA and southern Canada, and I. persulcatus in Asia [ 5 ]. These ticks have three active life stages, larva, nymph, and adult, each requiring a blood meal to molt into the next stage (larva and nymph) or produce eggs (adult female) [ 6 ]. Although they can feed on a broad range of vertebrate hosts, including birds, small mammals, carnivores, and ungulates [ 7 – 10 ], in forests subadults mainly feed on birds and small- and medium-sized mammals, while adults are mainly fed by medium- and large-sized mammals (e.g. ungulates) [ 8 , 9 , 11 ]. Identifying the drivers of Ixodes tick distribution and abundance is necessary to predict and mitigate tick exposure, thereby potentially reducing the risk of tick-borne diseases [ 12 ]. The distribution and abundance of ticks are linked to habitat suitability for both ticks and their hosts, which depend on complex interactions between abiotic (e.g. climatic conditions) and biotic conditions (host community and vegetation) playing differently at various spatial scales [ 13 – 15 ]. At the local scale, tick populations are also influenced by both abiotic and biotic conditions. Microclimatic conditions, impact the survival, development, and activity of Ixodes ticks [ 14 , 16 – 22 ]. For example, low humidity and high temperatures usually have a detrimental effect on the survival of Ixodes ticks [ 16 , 18 , 19 ]. Microclimatic conditions are directly influenced by biotic habitat and soil characteristics such as vegetation structure (e.g. cover of near-ground vegetation) and composition (e.g. diversity of plant species), as well as soil texture. Besides buffering climatic extremes detrimental to tick survival [ 23 , 24 ], trees and shrubs also create a litter layer with a humid microhabitat protecting ticks from desiccation during their off-host period [ 25 – 27 ]. Tick abundance also depends on the presence and abundance of their hosts [ 14 ], which are influenced by vegetation structure and composition (e.g. small mammals and birds: [ 28 , 29 ] through changes in the availability of food resources, shelters, breeding, or resting sites for hosts. Overall, several studies demonstrated a strong association between Ixodes ticks and woodland habitats (reviewed in: [ 25 , 30 ]), likely reflecting both the higher availability of some propagation hosts (e.g. deer) and the more suitable microclimatic conditions in forest habitats compared to non-forest habitats. At a broader scale, landscape composition (e.g. proportion of habitat types) and configuration (i.e. spatial arrangement of these habitats) can influence the host availability through changes in the host community composition, host movement, and habitat use [ 13 ]. Landscape fragmentation, common in urban areas, results in a mosaic of patches of various sizes and land-use types, potentially impeding tick host movement [ 13 ] and isolating suitable habitats for ticks and their hosts from one another and from the source of large propagation hosts (e.g. deer in forests). In this context, ticks form a metapopulation, with subpopulations connected and reliant on each other for persistence. The degree of connectivity between subpopulations, enhanced by the tick host movement, determines whether tick subpopulations persist. Consistent with this explanation, previous studies in both urban and rural environments have demonstrated that connectivity between suitable patches for ticks and their hosts, as well as connectivity to populations of deer, are important for tick persistence (urban: [ 31 – 33 ]; rural: [ 34 – 36 ]). Several studies have reported an increased abundance or occurrence of Ixodes ticks with an increasing proportion of forest/tree canopy or the number of forest patches in the landscape [ 33 , 36 – 40 ] and a decreasing proportion of agricultural areas and built-up and paved areas [ 32 ]. Considering the complex interplay between local and landscape habitat characteristics on ticks and their hosts, studies should examine both spatial scales concurrently (e.g. [ 37 ]). Until now, most studies on tick ecology in Europe have primarily focused on forests, where ticks are most abundant [ 30 ]. Our understanding of the local and landscape features associated with tick occurrence in private yards thus remains limited (but see: Richter et al. in Germany [ 40 ] and Gregory et al. in the USA [ 31 ]). Nevertheless, this knowledge is fundamental to the prevention of tick-borne diseases as yards are important places for tick encounters and tick bites. Studies conducted in France, Belgium, and the Netherlands indicated that approximately 30% of reported tick bites occurred in yards [ 41 – 44 ]. The scarcity of research on tick populations in private yards can be primarily attributed to the challenge associated with tick sampling, as these private areas are not easily accessible to scientists. Citizen science, when members of the public collaborate with scientists to answer research questions, helps to overcome this challenge while likely increasing citizens’ tick-borne disease literacy (as seen in a researcher – community partnership to promote Lyme disease prevention [ 45 ]). Citizen science has emerged as a powerful means of advancing tick research (reviewed in [ 46 ]) and is a valuable approach to gain insight into the complex ecology of ticks in yards. TIQUoJARDIN, which translates to TICKinYARD in English, is a citizen science project conducted in northeastern France. This project aims to characterize the risk associated with the presence of ticks in private yards, encompassing the occurrence of ticks, their pathogens, and human exposure. The project also seeks to identify the local and landscape features related to this risk. To reach these objectives, citizen scientists followed a standardized protocol to collect ticks in their yard and completed surveys about the attributes of their yard and their tick bites. In this paper, we aim to characterize the presence of ticks in private yards and determine whether habitat characteristics at various scales, ranging from the local scale to the landscape scale, were associated with the occurrence of ticks. This study mainly focuses on Ixodes ricinus , i.e. the most widely distributed tick species in Europe, the primary vector of Lyme borreliosis, and a vector of many other human and animal diseases (e.g. tick-borne encephalitis, Babesiosis, piroplasmosis, and anaplasmosis) [ 1 , 47 ]. Methods Study site This study took place within a 35 km radius of Nancy (48°41′ 31.3′′ N, 6°11′ 3.9′′ E), a city located in northeastern France. Nancy (104 403 inhabitants) is part of a metropolitan community that covers 142 km 2 and has ~ 255 000 inhabitants [ 48 ]. The climate is semi-continental with hot summers (mean daily maximum temperature during July is 25.8°C) and cold winters (mean daily temperature during January is 2.6°C) (1991 − 2020). The mean annual temperature is 11.0°C and mean annual precipitation is 746 mm (1991 − 2020) [ 49 ]. The landscape is composed of agricultural areas (35% of the territory), non-forest vegetated areas (e.g. grassland and herbaceous areas, permeable surfaces in urban areas, and meadows), and forest areas (both representing 29% of the territory). Artificial surfaces (i.e. impervious surfaces only), and water bodies and wetlands cover 4% and 2% of the territory, respectively. In the metropolitan community, artificial surfaces cover 27% of the territory (based on data provided by DataGrand Est [ 50 ]). Tick sampling A massive local media coverage of the project has been organized each year to recruit participating citizens. Ticks were initially sampled in 213 private yards by the residents in 2020 (n = 28), 2021 (n = 73), and 2022 (n = 112). However, due to missing yard features or key information to carry our analysis, we only kept data from 185 yards (n = 26 in 2020, 59 in 2021, and 100 in 2022) (Fig. 1 ). A yard was defined as a collective or an individual vegetated surface of at least 100 m 2 adjacent or away from the dwelling maintained by the residents. Each yard was sampled once from May to mid-July, i.e. the period of peak activity of I. ricinus nymphs in the forest in northeastern France [ 51 , 52 ]. Citizen scientists conducted the sampling following a standardized protocol using a “tick collection kit”, containing all the material to sample, identify, handle, and store ticks (see supplementary material S1 for details). Participants dragged a 1 m 2 white cloth across the vegetation surface or over the leaf litter (e.g. [ 31 ]). To maximize tick detection probability, participants had to avoid tick sampling when vegetation was wet (ticks attach less well to a wet drag cloth) and sampling between 11 a.m. and 4 p.m. ( I. ricinus activity decreases when temperature increases and/or relative humidity decreases [ 21 , 22 ]). Participants had to sample 12 transects of 10 m long (~ 10 steps), four along property edges, and eight in the core of the yard. After each transect, they inspected the drag cloth (and their clothing to prevent tick bites and missing ticks) and transferred any tick (or what they thought might be a tick) into a labeled ethanol tube. Date and time of tick sampling were recorded. An expert later counted and morphologically identified the ticks to the species level and developmental stage according to Estrada-Peña et al. [ 53 ] and Pérez-Eid [ 54 ]. This study mainly focuses on I. ricinus nymphs due to the higher number of individuals collected compared to other stages and species (see result section) and that humans are predominantly bitten by I. ricinus nymphs [ 55 , 56 ]. Given the low variability in the abundance of I. ricinus nymphs per yard (see result section), count data were converted into occurrence data. To enhance tick presence assessment and to account for the single sampling event per yard, participants noted if they had been bitten (or likely bitten) by ticks in their yard during the last three years. Habitat characteristics To assess the influence of habitat characteristics on tick occurrence, we focused on three spatial scales: the transect scale, the yard scale, and the landscape scale. Transect features For each transect, citizen scientists recorded the transect location (edge vs core of the yard), grass height ( 10 cm, and mixed, such as litter or various grass heights), shading conditions (partly under trees/bushes vs no tree branches/bushes above the transect), and approximate length (m), since it was not always 10 m long. Yard features We computed 13 yard features potentially influencing habitat suitability for I. ricinus or its hosts based mainly on the responses of participants to a questionnaire (Table 1 ). These included the presence/absence of a wood/brush pile, a stone wall or a pile of stone, a vegetable garden, a compost, an unmanaged long grass area, a bird feeder, and nut-producing trees (i.e. oaks, beech, hazel, and/or walnut). Participants also reported the presence of six fruit-producing group species and we computed the “number of fruit-producing species groups” per yard. Groups were based on the type of vegetation (i.e. trees vs. brushes and shrubs), and the period of harvesting (Table S1 ). Citizen scientists also indicated whether their yard was fully closed (preventing medium and large-sized mammals from entering the yard) or not. The mowing intensity was computed as the cumulative number of mowing events in spring, summer, and autumn based on participants’ reported lawn mowing frequency per season. Using the location of the yard, satellite images from Google Earth (including CNES/Airbus, GeoContent, and Maxar technologies images), and cadastral information [ 57 ], we computed the “vegetated surface of the yard” as the property surface excluding buildings (e.g. house) and unvegetated areas (e.g. pool and paved alleys). To characterize host presence, participants reported deer sightings or signs of their presence (e.g. feces and footprints) near their property (i.e. inside or within 5 m from the edges of their property). Participants also reported if they had a dog and if cats were present in their yard. Only the variable “presence of a dog” was considered due to the high prevalence of cats (96% of yards in 2021 and 2022, n = 159 yards) and missing information for 2020. The presence of free-ranging chicken was not considered due to the rarity of its occurrence (7% of the yards). Table 1 Expected relationships between yard features computed and the occurrence of I. ricinus in yards. Yard features Expected relationship with I. ricinus occurrence in yards Explanation Habitat suitability for tick hosts Presence of • a wood/brush pile • a stone wall or a pile of stone • a vegetable garden • a compost • a bird feeder + These attributes can provide a shelter (i.e. the presence of a wood/brush pile, a stone wall or pile of stone) or food for tick hosts (i.e. the presence of a vegetable garden, a compost, and a bird feeder) leading to an increase in host richness or abundance [ 89 , 90 , 99 ]. Previous studies have thus shown a positive relationship between the presence of these features and tick occurrence in yards or recreational sites [ 31 , 86 , 87 ]. No. of fruit-producing species groups + Fruit or nut-producing trees can provide food for tick host species (mainly small mammals and birds) influencing their abundance (e.g. [ 100 – 102 ]). Host species abundance may in turn influence tick abundance [ 100 ]. The abundance of I. ricinus nymphs in pasture has been shown to be positively related to the presence of fleshy fruit trees at pasture edges [ 79 ]. Presence of nut-producing trees Vegetated surface of the yard (m 2 ) + The relative abundance of tick host species can increase with yard size [ 89 ] and tick abundance has been shown to increase with the vegetated surface of the yard [ 103 ]. Yard accessibility and presence of tick hosts Yard closure - Yard fencing can reduce the diversity and abundance of host species by preventing medium and large-sized mammals from entering a yard [ 90 , 104 , 105 ]. Full fencing around the yards has been shown to decrease the odds of finding I. scapularis [ 31 ] (but see: [ 87 ]). Signs of deer + Deer play a key role in the persistence of I. ricinus populations as adult ticks mainly feed and copulate on deer [ 8 ]. Previous studies showed that the density of I. ricinus nymphs in forests was positively associated with deer presence [ 106 ]. In green spaces (mainly parks), I. ricinus density increased with the connectivity to a known population of roe deer ( Capreolus capreolus ) [ 32 ]. Presence of a dog +/- Dogs can bring back ticks into the yard after a walk outside the yard (e.g. in park [ 107 ]) or serve as hosts for feeding ticks in yards. Alternatively, a free dog in a yard can prevent some hosts from entering the yard [ 104 ]. Habitat suitability for ticks Mowing intensity - Increasing mowing intensity and the presence of an unmanaged long grass area are expected to decrease and increase the availability of moist microhabitats for ticks, respectively. As low humidity and high temperatures usually have a detrimental effect on the survival of Ixodes ticks [ 16 , 18 , 19 ], these features are expected to affect tick occurrence or abundance. However, the influence of mowing intensity and the presence of an unmanaged long grass area on tick abundance or occurrence has been seldom tested appropriately. A study showed that a single mowing event did not affect the abundance of I. scapularis and Dermacentor variabilis on recreational hiking trails [ 85 ], while another demonstrated that tick density was not associated with grass height in pasture edges [ 79 ]. Presence of an unmanaged long grass area + Landscape features We used land cover data obtained from high-resolution orthophotos [ 50 ] to classify land cover into five classes: 1) artificial surfaces (i.e. impervious surfaces only), 2) non-forest vegetated areas (e.g. grassland and herbaceous areas, permeable surfaces in developed areas, and abandoned agricultural areas), 3) forest areas (e.g. deciduous forest, clearcuts, and young plantations), 4) agricultural areas (e.g. annual and perennial crops), and 5) open water and wetlands. Subsequently, we derived eleven features describing the landscape composition and spatial configuration that can influence host availability and the availability of suitable habitats for ticks (Table 2 ). The six spatial configuration variables were the density of forest patches, the shortest distance to a forest patch, the effective mesh size of vegetation areas (i.e. combining forest and non-forest vegetated areas), the effective mesh size of vegetation and agricultural areas combined, the total edge density, and the edge density of vegetation areas. The effective mesh size (MESH) is a fragmentation index and represents the average size (ha) of areas (i.e. vegetation areas or vegetation and agricultural areas combined) that an organism is connected to in a landscape starting from a randomly chosen point [ 58 ]. The more barriers in the landscape (e.g. artificial surfaces), the less vegetation areas organisms will have access to, and the lower the effective mesh size. Except for the shortest distance to a forest patch, we computed these variables within buffer sizes of 300, 400, and 500 m to maximize variability and avoid missing values. Indeed, when the landscape consists of only one vegetation patch, the MESH cannot be computed. To describe the landscape composition, we computed five variables, the percentage of artificial surfaces, non-forest vegetated areas, forest areas, and agricultural areas within the three buffer sizes, and the percentage of vegetation areas within a 50-m buffer, to characterize the immediate environment surrounding the yard (Table 2 ). Table 2 Expected relationships between landscape features and the occurrence of I. ricinus in yards. Landscape features Expected relationship with I. ricinus occurrence in yards Explanation Landscape composition Percentage of • artificial surfaces - Due to the lack of a litter layer and unsuitable microclimatic conditions, artificial surfaces are unsuitable habitats for ticks. Moreover, they can impede the movement of hosts, especially large-sized mammals which are key hosts of adults I. ricinus [ 8 ], and isolate the yard from putative sources of ticks and hosts such as forests. Consistently, densities of I. ricinus (all stages) in green spaces in Belgium decreased with increasing urbanization (i.e. the proportion of built-up and paved areas [ 32 ]). • agricultural areas +/- On one hand, crops are less suitable habitats for ticks (compared to forests) due to the lack of a litter layer and less suitable microclimatic conditions [ 108 ]. On the other hand, agricultural practices can alter host movements, and habitat use, with some mammals being more (or less) abundant in crops compared to mature forests [ 109 , 110 ]. Previous studies showed that the densities of I. ricinus larvae and adults (but not nymphs) in green spaces in Belgium decreased with the percentage of agricultural areas [ 32 ]. • Non-forest vegetated areas + Non-forest vegetated areas (e.g. parks and pasture) can be suitable habitats for ticks [ 32 , 79 ] and are used by some tick hosts [ 109 , 111 ]. In urban green spaces (e.g. parks, natural and amenity green spaces), the density of I. ricinus nymph was positively associated with the proportion of vegetation areas in the landscape (i.e. urban forests, open green spaces, and green corridors such as wildflower verges and hedgerows [ 73 ]), while in forests it decreased with the percentage of pastures in the landscape [ 37 ]. • Vegetation areas (forest and non-forest vegetated areas combined) • forest areas + Given that the abundance of some host species increases with the percentage of forest areas in the landscape [ 38 ] and that I. ricinus is most abundant in forests than in other habitat types [ 30 ], many studies reported an increase in I. ricinus abundance or occurrence in forests or green spaces with the proportion of forests or the number of forest patches [ 36 – 40 ]. The density and occurrence of nymphal I. ricinus in green spaces were negatively related to the distance to woodland [ 73 ]. Landscape configuration Density of forest patch (number.100 ha − 1 ) Min. distance to the forest (m) - Effective mesh size (MESH, ha) • of vegetation areas • of combined vegetation and agricultural areas + Fragmentation can alter host availability through changes in the host community composition, host movement, and habitat use. Highly fragmented habitats may hold populations of smaller and/or more mobile tick host species (e.g. rodents and birds), but not of larger animals (e.g. deer) [ 13 ]. Previous studies in both urban environments [ 32 ] and rural environments [ 34 – 36 ] have demonstrated that connectivity between suitable patches for ticks and their hosts, as well as connectivity to source populations of large propagation hosts (i.e. forest), are important for I. ricinus persistence. Edge density (m. ha − 1 ) • of landscape • of vegetation areas +/- On one hand, some mammal species can be more abundant at habitat edges, i.e. zones of transition between adjacent ecological systems, than in the habitat interior (e.g. forest edges) [ 112 , 113 ]. On the other hand, increasing edge density implies increased habitat fragmentation, which could be detrimental to the movement of tick host species. The abundance of adult I. ricinus has been shown to increase with the forest edge density, while that of nymph was not related to the edge density of forest [ 36 ]. Meteorological data To account for variation in meteorological conditions during sampling, we computed the average of daily saturation deficit per yard during the five days preceding tick sampling. Indeed, the saturation deficit affects the questing activity of I. ricinus [ 21 , 22 , 59 ] and could thus affect the likelihood of finding ticks. For each yard and each of the five days preceding tick sampling, we computed the daily saturation deficit (mmHg) using the daily average of relative humidity (%), the daily average of temperature (°C), and the formula provided by Randolph and Storey [ 59 ]. We then averaged those five values. For each yard, we used hourly temperature and relative humidity data from the nearest weather station (mean distance between yards and the weather stations: 7770 m, range: 724 − 26835 m, n = 185) [ 49 ]. Statistical analysis In this paper, we investigated the relationships between the occurrence of I. ricinus nymphs and habitat characteristics at transect, yard, and landscape levels. Moreover, we examined how incorporating tick bites in the assessment of tick occurrence could influence the relationships between tick occurrence and environmental variables. We thus also explored the relationships between the occurrence of ticks, and the corrected occurrence of ticks (0 when no ticks were found, and the household reported no tick bites; otherwise, 1) and yard and landscape features. For all the following analyses, we used mixed-effect generalized linear regression models (GLMMs) with a binomial error distribution (logit link). Analysis of the occurrence of nymphal I. ricinus at the transect-level We assessed the relationships between transect characteristics and the presence of I. ricinus nymphs at the transect-level. In this analysis, we only considered yards where nymphs were found, to only include suitable conditions for tick presence and avoid background noise due to unsuitable landscape characteristics. Fixed effects included transect location (edge vs core), grass height ( 10 cm, and other), and shading (shaded vs not shaded). The length of the transect, log-transformed, was included as an offset variable to adjust models for differences in sampling effort among transects. The offset variable makes model adjustments with its regression coefficient being fixed at 1 [ 60 ]. To account for variations in sampling conditions between years and repeated measures within yards, we included the yard ID nested in the sampling year as a random effect. For this analysis, we only kept data from yards for which information was available for all transects (i.e. length, location, grass height, and shading), i.e. 434 transects (n = 48 in 2020, 226 in 2021, and 160 in 2022) across 37 yards (4 in 2020, 19 in 2021, and 14 in 2022). Analysis of the occurrence of nymphal I. ricinus and ticks, and the corrected occurrence of ticks at the yard level To assess the relationships between yard and landscape features, and the occurrence of I. ricinus nymphs, ticks, and the corrected occurrence of ticks, we used occurrence data at the yard level. The total length of transects sampled per yard (log-transformed) was included as an offset variable, to account for variations in sampling effort among yards, and the sampling year as a random effect. Fixed effects included 13 yard features (Table 1 ), 11 landscape features (Table 2 ) (see below for the selected buffer size), and the 5-day average of daily saturation deficit to account for temporal variability of meteorological conditions. These analyses rely on data from 185 yards (n = 26 in 2020, 59 in 2021, and 100 in 2022). For each of the three response variables, we built models that included different combinations of yard and landscape features. To avoid collinearity issues, we did not build candidate models that included variables with an absolute value of the Pearson correlation coefficient greater than 0.6. Model selection was based on the Akaike information criterion (AIC). We selected models within ΔAICc < 2, as there is substantial evidence to support them as the best models to explain the observed patterns in the data [ 61 ]. To gain insight into the respective influence of yard and landscape features, the same model selection procedure was applied using combinations of either yard or landscape features. Model accuracy was evaluated using the AIC, AUC, sensitivity, and specificity. For each best model, the ROC curve was used to compute the AUC and determine the threshold that optimizes both sensitivity and specificity (Youden’s Index, [ 62 ]). A predicted occurrence greater than the cut-off threshold was assigned as tick (corrected or not) or nymphs were present. The sensitivity and specificity were computed using the optimal cut-off. Before exploring the relationships between environmental factors and the three response variables, we identified the optimal buffer size at which the relationships between landscape features and the occurrence of ticks were the strongest. Indeed, variables computed with different buffer sizes were highly correlated (Fig. S1 ) which can cause multicollinearity. For each of the three buffer sizes, we built candidate models that included as fixed effects different combinations of the nine variables computed at various buffer sizes. We included the total length of transects sampled per yard (log-transformed) as an offset variable and the sampling year as a random effect. The best model for each buffer size was selected based on the AIC and the parsimony principle, as previously described. We then compared the AIC of the three best models and chose the buffer size at which the AIC was the lowest. A buffer size of 500 m was the best fit for tick occurrence (AIC = 205.71 compared to AIC = 206.71 and AIC = 207.50 for buffers of 300 and 400 m, respectively). As a result, we kept landscape features computed within a 500 m buffer. All analyses were carried out in R v 4.3.0 [ 63 ] using the packages g lmmTMB [ 64 ] and MuMIN [ 65 ] for GLMMs and model selection. Collinearity was assessed using the variance inflation factor (VIF) (package performance , [ 66 ]. The DHARMA package [ 67 ] was used to check for spatial autocorrelation (Moran’s I test) and for patterns in the residuals using simulation-based standardized residuals. The package cutpointr [ 68 ] was used to find the optimal cutpoint and compute the accuracy metrics (i.e. AUC, sensitivity, and specificity) using these thresholds. The 95% CI of the accuracy metrics was computed using the package pROC [ 69 ]. Predicted odd ratios and their confidence intervals were obtained with the sjPlot package [ 70 ]. Results Description of sampled yards Following the classification of a territory according to its degree of urbanization [ 71 , 72 ], 33% of sampled private yards with complete data (n = 185) were in densely populated urban areas (cities), 29% in urban areas of intermediate density (towns and suburbs), and 38% in thinly populated rural areas. These yards were almost exclusively private individual yards adjacent to or surrounding to dwelling (96.2%). Others were individual yards away from the dwelling (2.2%), or collective yards (1.6%). Yards differed in their yard and landscape features (Table 3 ). On average, yards had a vegetated surface of 953 m 2 (SD = 1246) and were located 310 m from a forest patch (SD = 318). The three most common yard features were the presence of a compost, the presence of a vegetable garden, and the presence of a bird feeder (83%,76%, and 71% of the yards, respectively). Additionally, over half of the yards had a wood/brush pile, a stone wall or pile of stone, or nut-producing trees while having a dog, a fully closed yard, and signs of deer were less common. On average participants mowed their lawn approximately twice a month in spring, summer, and autumn. Quickly, within a 500 m buffer surrounding the yard, the mean percentages of artificial surfaces and non-forest vegetated areas were 28% (SD = 17) and 50% (SD = 12), respectively, while the mean density of forest patches was 4.0 patches.100 ha − 1 (SD = 3.7). Table 3 Variation of the yard and landscape features in private yards (n = 185). Freq. of occurrence of I. ricinus nymphs in yards when the feature is Continuous variables Mean ± SD Range Below average Above average Mowing intensity 18.3 ± 9.5 2 − 36 22% 26% Vegetated surface of the yard (m 2 ) 953 ± 1246 91 − 8846 19% 35% No. of fruit-producing species groups 3.4 ± 1.8 0 − 6 13% 34% 5-day avg. daily saturation deficit (mmHg) 4.7 ± 2.0 1.3 − 11.7 26% 22% Percentage of artificial surfaces 500 28 ± 17 2 − 70 32% 12% Percentage of agricultural area 500 9 ± 13 0 − 51 28% 16% Percentage of forest areas 500 10 ± 13 0 − 57 14% 43% Percentage of non-forest vegetated areas 500 50 ± 12 21 − 80 27% 20% Percentage of vegetation areas 50 59 ± 16 11 − 100 14% 33% Edge density of landscape 500 (m. ha − 1 ) 354 ± 118 82 − 652 29% 18% Edge density of vegetation areas 500 (m.ha − 1 ) 346 ± 131 65–651 30% 17% Density of forest patch 500 (number. 100 ha − 1 ) 4.0 ± 3.7 0.0–18.3 20% 30% MESH of vegetation areas 500 (ha) 9.6 ± 10.2 0.2–60.6 15% 44% MESH of combined vegetation and agricultural areas 500 (ha) 14.6 ± 13.7 0.2 − 74.5 16% 35% Min. distance to the forest (m) 310 ± 318 0 − 1856 31% 11% Dichotomous variables Percentage of yards with the feature (number) Freq. of occurrence of I. ricinus nymphs in yards With the feature Without the feature Compost 83% (153) 27% 9% Vegetable garden 76% (140) 25% 20% Bird feeder 71% (132) 22% 28% Nut-producing trees 61% (112) 31% 12% Stone wall or pile of stone 57% (105) 22% 26% Wood/brush pile 55% (102) 33% 12% Unmanaged long grass area 41% (75) 29% 20% Dog 22% (41) 20% 25% Signs of deer 20% (37) 57% 16% Yard closure (fully closed) 18% (34) 18% 25% These variables were used to explore the relationships between environmental factors and the occurrence of nymphal I. ricinus , ticks, and the corrected occurrence of ticks. Variables were computed within a 500-m ( 500 ) or a 50-m buffer ( 50 ) from the edges of the property. MESH: effective mesh size. Vegetation areas are composed of forest areas and non-forest vegetated areas. Tick collection and tick bites Citizen scientists sampled a total of 2030 transects in 185 yards. As indicated in the protocol, tick sampling took place at the appropriate time of day (i.e. before 11 a.m. or after 4 p.m.) 93% of the time. On average, citizen scientists sampled 11 transects per yard (SD = 2.0, range: 4 − 12), covering an average area of 109 m 2 (SD = 22.7, range: 36 − 200 m 2 ). 84% of citizen scientists sampled at least 10 transects. Yards with fewer than ten transects had a mean vegetated surface of 339 m 2 (SD = 295), compared to a mean of 1072 m 2 (SD = 1323) for yards with at least ten transects. Citizen scientists, collected 501 ticks (112 larvae, 365 nymphs, and 24 adults) belonging to four species: Ixodes ricinus , Ixodes frontalis , Dermacentor marginatus , and Dermacentor reticulatus . Ixodes ricinus was by far the most collected species (96% of the ticks collected) and I. ricinus nymphs (hereafter “ nymphs ”) represented 71% of the collected ticks (Table 4 ). Ticks occurred in 8.6% of the transects and 32% of sampled yards (Table S2). At least one nymph was found in 24% of yards (Table S2), with an occurrence of 18% in yards in densely populated urban areas, 30% in urban areas of intermediate density, and 24% in thinly populated rural areas. Other species and life stages occurred in less than 6% of the yards, except for I. frontalis larvae in 2020 (15% of the yards, 4 yards) and I. ricinus male in 2021 (8% of yards, 5 yards) (Table S2). The average density of nymphs was 1.6 individuals per 100 m 2 (SD = 6.3, range: 0 − 62.9). In yards with nymphs (n = 44), the median density was 2.5 nymphs per 100 m 2 (mean: 6.7, SD = 11.7, range: 0.8 − 62.9). Among households (n = 185), 29% indicated that at least one family member had been (very likely) bitten by ticks in their yard in the last three years. In the last three years, 48% of family members reported being bitten by ticks and 18% reported being bitten by ticks in the yard (n = 485, data only available for 2021 and 2022). When corrected by tick bites in the yard, ticks potentially occur in 45% of the yards (35% in 2020, 56% in 2021, and 41% in 2022). Table 4 Frequency of each tick species, stage, and sex (number collected) per sampling year. Species Stage and sex 2020 2021 2022 Ixodes ricinus larvae 52% (96) 2% (4) 5% (4) Ixodes ricinus nymph 41% (75) 91% (224) 81% (59) Ixodes ricinus adult female 1% (2) 1% (3) 3% (2) Ixodes ricinus adult male 1% (2) 3% (7) 3% (2) Ixodes frontalis larvae 4% (8) 0% (0) 0% (0) Ixodes frontalis nymph 0% (0) 1% (3) 5% (4) Dermacentor marginatus adult female 0% (0) 0% (0) 1% (1) Dermacentor reticulatus adult female 0% (0) 1% (2) 0% (0) Dermacentor reticulatus adult male 0% (0) 1% (2) 1% (1) Total 100% (183) 100% (245) 100% (73) Tick collection took place in 185 private yards between from May to mid-and July 2020 (n = 26), 2021 (n = 59), and 2022 (n = 100) within a 35 km radius of Nancy, a city located in the northeastern of France. Relationships between transect features and the occurrence of nymphal I. ricinus at the transect-level In yards with nymphal I. ricinus and for which information was available for all transects (n = 434 transects in 37 yards), nymphs occurred in 27% of the transects (SD = 24). In these yards, nymphs occurred in 34% of shaded transects compared to 17% of unshaded transects (Table S3). The likelihood of finding at least one nymph increased by 2.75 (95% CI: 1.52–4.97) when the transect was shaded by vegetation compared to not shaded transect. Grass height and the location of the transect (edge vs core) were not related to the likelihood of observing nymphs. Relationships between yard and landscape features, and the occurrence of I. ricinus nymphs, ticks, and the corrected occurrence of ticks Nine, twelve, and ten models explaining variations in the occurrence of nymphs, ticks, and corrected occurrence of ticks, respectively, were within ΔAIC < 2 (Tables S4, S5, S6). Among these models, one model best explained the occurrence of I. ricinus nymphs (Table S4), while two models best explained the occurrence of ticks and the corrected occurrence of ticks (Tables S5, S6). These models always included three explanatory variables. Overall, the presence of a wood/brush pile and signs of deer in/near the yard were included in all best models (except for a model for signs of deer) and were positively associated with all three occurrence variables (Fig. 2 ). In yards with a wood/brush pile, the odds of finding ticks or nymphs was 2.37 (95% CI: 1.18–4.75) to 3.57 (95% CI: 1.45–8.77) greater compared to yards without this feature. Nymphs were observed in 33% of yards with a wood/brush pile compared to 12% of yards without it (Table 3 ). Similarly, the presence of deer signs in/near the yard increased the odds of finding ticks or nymphs by 2.67 (95% CI: 1.02–6.97) to 3.33 (95% CI: 1.36–8.14) compared to yards without signs of deer (Fig. 2 ). Nymphs were observed in 16% of the yard without signs of deer, while in contrast, they were found in 57% of the yard with signs of deer nearby (Table 3 ). In addition to being positively associated with signs of deer and the presence of a wood/brush pile, the odds of finding a nymph also increased by 1.09 (95% CI: 1.04–1.15) with a one-unit increase in MESH of vegetation areas (Table S4, Fig. S2). Nymphs were observed in 15% of the yard with a MESH of vegetation areas less than 9.6 ha (average) compared to 44% when the MESH of vegetation areas was higher. One of the best models explaining the occurrence of ticks was similar to the best model explaining the occurrence of nymphs as it also included signs of deer, the presence of a wood/brush pile, and the MESH of vegetation areas. The second best model explaining the occurrence of ticks included the percentage of forest areas instead of the MESH of vegetation areas (Table S5, Fig. 2 ). This model, which included signs of deer, the presence of a wood/brush pile, and the percentage of forest areas, was identical to one of the best models explaining the corrected occurrence of ticks (Table S6, Fig. 2 ). In these models, the odds of finding ticks or ticks corrected by tick bites increased by 1.05 (95% CI: 1.02–1.08) and 1.07 (95% CI: 1.04–1.11) with a one-unit increase in the percentage of forest areas (see Fig. S3 for the occurrence of ticks), respectively. In particular, ticks were observed in 20% of yards with less than 10% of the landscape covered by forest areas (i.e. a below-average percentage), while in contrast, they were found in 54% of the yard with an above-average percentage of forest. Finally, the second best model explaining the corrected occurrence of ticks included the presence of a compost, in addition to the presence of a wood/brush pile, and the percentage of forest areas (Fig. 2 , Table S6), but its association with the corrected occurrence of ticks was not significant (OR = 2.53, 95% CI: 0.98–6.55; Fig. 2 ). For the three response variables, the AIC was the lowest for the models including yard and landscape features, intermediate for models with only landscape features, and the highest for models with only yard features. AUC values (0.72–0.84) were similar irrespective of yard and/or landscape features inclusion. Overall, the sensitivity and specificity of the best models was around 0.67 (range: 0.49 − 0.89) and 0.78 (range: 0.60 − 0.88), respectively (Fig. S5). For the occurrence of nymphs, the model with yard and landscape features had a similar specificity and sensibility (~ 0.8), while these values differed in models with only yard or landscape features. Differences in specificity and sensitivity were also observed for all models explaining the variation in the occurrence of ticks or the corrected occurrence of ticks with yard features (Fig. S5). Discussion In Europe, there has been limited research on the factors associated with ticks presence or abundance in vegetated areas other than forests (e.g. green spaces including parks [ 32 , 73 ]), with even fewer studies focusing on private yards [ 40 ]. Using citizen science, we assessed, for the first time in France, the occurrence of ticks in private yards within and around a metropolitan community, and investigated the yard and landscape features related to tick presence. Our study reveals that ticks are frequent in private yards, irrespective of whether they are located in densely populated urban areas, urban areas of intermediate density, or thinly populated rural areas, with ticks detected in 32% of the sampled yards and potentially present in 45% of yards when considering collected ticks and reported tick bites. We also demonstrated that their occurrence was linked to both yard and landscape features. Consistent with previous studies conducted in yards in Germany [ 40 ] and in urban green spaces in Europe (reviewed in [ 74 ]), nymph density in yards was low (mean of 1.6 individuals per 100 m 2 ). This suggests that tick presence is primarily due to introductions by hosts that acquired ticks elsewhere (sink populations), and could be sporadic. Nonetheless, 29% of households and 18% of family members reported tick bites in their yard over the last three years, aligning with previous studies highlighting the risk of tick bites in yards [ 41 – 44 ]. Although the participating households were not randomly selected across the study area, which can lead to some biases that limit the representativeness of our study, there was no apparent massive participation of citizens who were previously bitten by ticks in our study. Taken together our findings emphasize the importance of informing the public about the risk of tick exposure and tick bites in yards. Tick distribution depends on habitat suitability for ticks and their hosts [ 14 , 15 ]. At the transect level, nymphs were found across all types of transects, regardless of vegetation shading, location, or grass height, indicating their ability to occur in habitats typically deemed less suitable due to less favorable microclimatic conditions. Moreover, in yards with nymphs, the likelihood of finding a nymph was similar in long and short grass areas, as well as in the core or the edge of the yard, whereas it was nearly three times higher in transects beneath trees, hedges, and shrubs compared to transects in open areas without vegetation shading. Compared to open areas, the cover provided by trees and shrubs can buffer temperature extremes and maintain a higher humidity [ 23 ], thereby enhancing tick survival [ 17 , 18 ]. Moreover, trees and shrubs may provide food resources, shelters, breeding, or resting sites for hosts. The abundance or cover of trees and shrubs is an important predictor of I. ricinus nymphs density in various environments, including woodlands [ 37 , 75 – 78 ], agricultural areas [ 79 ], and urban parks and green spaces [ 80 – 82 ]. However, contrary to our results, Richter et al. [ 40 ] found no relationship between I. ricinus occurrence in transects and transect shading in yards, suggesting that the effect of vegetation shading might be variable in yards according to other features (type of vegetation, etc.). The lower shading effectiveness of herb layer vegetation compared to trees and shrubs [ 83 ] may explain the lack of a relationship between nymph occurrence and grass height (see also: [ 84 ]). However, the influence of the herb layer on ticks may be context-dependent (e.g. depending on yard size and availability of suitable habitats nearby), explaining why some studies found an association between tick density and the herb layer (e.g. [ 80 ] in urban parks), while others have not (e.g. [ 79 ] in pasture edges, [ 82 , 85 ] in sub-urban and rural parks). At the yard level, models for the three occurrence variables (i.e. occurrence of nymphs and ticks, and the corrected occurrence of ticks) were similar, given that most ticks were nymphal I. ricinus . In all models, the likelihood of finding ticks or nymphs in a yard increased with the presence of a wood/brush pile and signs of deer in/near the yard (Fig. 2 ). This aligns with previous studies in the US showing a higher occurrence or abundance of three tick species (including I. scapularis) in residential properties with a log or brush pile [ 31 , 86 ] (but see: [ 87 ]). Despite the expectation that wood/brush piles attract birds and small mammals, primary hosts for immature I. ricinus ticks, studies investigating its influence on tick hosts in yards remain scarce and have shown weak or no relationship with mammal species abundance or richness [ 88 – 90 ]. Therefore, the causal relationships between the presence of a wood/brush pile and tick occurrence should be further investigated. Cervids are vital for the persistence of I. ricinus populations, as adult ticks mainly feed and copulate on deer [ 8 ]. While cervids may contribute to tick presence in some yards, signs of deer more likely indicate yard proximity to forests. Indeed, the likelihood of observing signs of deer decreased with the minimal distance to the forest (OR = 0.99, 95% CI: 0.99–1.0; mean minimal distance to the forest: 121 m [SD = 157 m] for yards with signs of deer vs 357 m [SD = 331 m] for yards without signs of deer). Mowing intensity, the presence of an unmanaged long grass area, a bird feeder, a vegetable garden, and yard closure were yard features absent from the top models (within ΔAIC < 2 from the lowest scoring model). The lack of relationship between the occurrence variables and mowing intensity and the presence of an unmanaged long grass area aligns with our finding at the transect level, where nymph occurrence was not related to grass height. Surprisingly, yard closure was also not related to the occurrence variables (but see: [ 31 ] in the USA). This might be explained by the fact that we relied on citizen’s perception and did not accurately estimate fence permeability based on explicit criteria (e.g. fence type, height, and structural integrity). Alternatively, this finding might indicate that ticks are primarily introduced by hosts little or not affected by the most common types of fences in our yards (i.e. hedge and chain link), such as small mammals and birds. At the landscape scale, we found evidence that the odds of at least one of the three tick occurrence variables were positively linked to the percentage of forest areas (tick occurrence and corrected occurrence of ticks) and the MESH of vegetation areas (occurrence of nymphs and corrected occurrence of ticks). This finding aligns with previous literature indicating that, in Europe, forests are more suitable habitats for ticks compared to non-forest land-use types [ 30 ]. As previously shown for tick persistence in meadows, forests can also be considered as a source of ticks for yards, while more artificialized or open landscape types (e.g. crops, artificial surfaces, or even yard lawns) act as sinks [ 91 ]. The positive association between the MESH of vegetation areas (i.e. a decrease in the fragmentation) and the occurrence variables is unsurprising considering the positive correlation between the MESH of vegetation areas and the percentage of forest areas (r = 0.66, Fig. S4). The importance of landscape connectivity for tick populations has already been identified in rural areas in Spain [ 34 , 35 ], as well as in green spaces in urban areas in Belgium [ 32 ], and the US [ 31 , 33 ]. Given favorable conditions for tick survival in yards, enhanced connectivity between suitable habitats for ticks, their hosts, and potential source host populations (e.g. forest) could promote tick presence in yards. Citizen science is a valuable approach for advancing ecological knowledge, yet concerns persist regarding data quality [ 46 , 92 ]. We implemented several recommended measures to ensure data quality and reliability [ 93 ], including a standardized protocol for tick collection, available in written and video format, as well as a support for tick recognition. Additionally, we asked participants to record easily identifiable yard characteristics. Although we did not quantify it, the participation of citizens in this study likely increased their tick risk awareness and tick-associated literacy, based on their feedbacks. This may in turn increase the acceptance of methods to prevent tick bites (but see: [ 94 ]). One of the objectives of our project was to determine whether certain features of the yard or landscape which can be modified to reduce the risk of tick occurrence. However, our results should be interpreted with caution regarding public health. For example, tick occurrence was related to the percentage of forest areas in the landscape and the connectivity of vegetated areas. This suggests that initiatives aimed at enhancing landscape connectivity, and/or creating vegetated areas near yards may increase tick exposure risk, but final decisions must be balanced against other benefits, such as an increase in biodiversity or human well-being [ 95 ]. Additionally, tick occurrence was linked to vegetation shading, the presence of a wood/brush pile, and signs of deer in/near the yard. However, we caution against providing clear recommendations to yard owners based solely on these results. Indeed, further research is needed to establish causal relationships between these features and tick occurrence, as well as to assess the potential impact of yard management measures on fauna and the associated financial implications for yard owners. Nonetheless, yard owners should be particularly vigilant when using shaded areas (e.g. while trimming hedges or lying down a tree), as nymph occurrence was three times higher than in unshaded areas. In our study the risk of encountering ticks did not appear to be associated with the presence of a stone wall, a vegetable garden, a compost, a bird feeder, a dog, and nut-producing trees, the vegetated surface of the yard, yard closure, mowing intensity, grass height, and the number of fruit-producing trees. Moreover, it was similar in the core and the edge of the yard. Health authorities (e.g. USA and Canada) have recommended various property management measures to control tick populations (e.g. [ 96 ]), not supported by the results of this study, but often cited in scientific or popular science publications (e.g. [ 97 , 98 ]). As previously stated [ 87 ], we recommend that public health officials provide clear information on the scientific support and uncertainties surrounding yard management measures, as most of them seem to lack scientific support. European health authorities should also be careful when using results from North American studies, as yard features (especially the size), tick species, and host ecology, might differ greatly from European yards. Further studies are necessary to identify features associated with tick occurrence in yards and to implement management strategies adapted to the private yards to mitigate the risk of tick exposure. Conclusions Ticks are frequent in private yards across an urbanization gradient in our study area in northeastern France and their presence is shaped by both yard and landscape features. Despite official recommendations on some yard measures expected to control ticks abundance or occurrence, we found no evidence linking I. ricinus occurrence in yards with the mowing intensity and other relevant yard features. However, we found that vegetation shading, the presence of a wood/brush pile, and signs of deer in/near the yard were related to the occurrence of ticks. At the landscape level, tick occurrence were positively related to the percentage of forest areas, and the connectivity of vegetated areas. We suggest further studies are needed to implement adapted management strategies against tick risk in yards. To better understand the threat posed by ticks in yards and enhance management measures effectiveness, future research should investigate the prevalence of pathogens in the ticks present in yards, as well as identify the hosts responsible for tick presence in yards. Declarations Ethics approval and consent to participate Study participation was voluntary and all participants provided signed informed consent before entering the study. This study was declared to the National Commission on Informatics and Liberty (Commission Nationale de l'Informatique et des Libertés [CNIL]), the independent French administrative regulatory body responsible for data protection, which ensures that all information is kept confidential and anonymous, in compliance with the Declaration of Helsinki. Consent for publication Not applicable Availability of data and materials The data used in this paper includes personal residential information. Therefore, to protect study participant privacy the data are not publicly available. The anonymized data collected are, however, available from the corresponding author upon reasonable request. Competing interests The authors declare that they have no competing interests. Funding This study was funded by the French Agency for Food, Environmental and Occupational Health Safety (ANSES), grant no.: ANSES-21-EST-050, as well as the French National Research Agency (ANR) as part of the “Investissements d'Avenir” program (ANR-11-LABX-0002-01, Lab of Excellence ARBRE), the project “Des Hommes et Des Arbres” (DHDA) supported by the French government through its program “Plan d'Investissement d'Avenir - Territoire d'Innovation”, and the Metropole du Grand Nancy (grant no. 7434). Authors’ contributions Project conceptualization (L.B, J.D, P.FK, F.B, C.G, I.C). Design of the work (L.B, J.D, I.C, S.C); data acquisition (Citizen scientists, A.M, L.B, J.D, C.C, and S.C); data modelling (A.M with valuable support from J.D and L.B); interpretation of results (A.M, L.B, J.D) and manuscript preparation (A.M, J.D, and L.B). All authors read, revised, and approved the final manuscript. Acknowledgements We would like to acknowledge the contribution of staff from the Centre Permanent d’Initiatives pour l’Environnement Nancy Champenoux (particularly Marcel Klein), the Nancy Laboratory for Rabies and Wildlife of The French Agency for Food, Environmental and Occupational Health and Safety (ANSES), and the Tous Chercheurs Laboratory (French National Research Institute for Agriculture, Food and Environment, INRAE), as well as the Jean-Marie Pelt botanical garden (Villers-lès-Nancy), for their invaluable assistance in distributing the tick collection kit. Thanks are extended to all colleagues at the ANSES Nancy Laboratory for Rabies and Wildlife, especially Olivier Musquar and Jean-Marc Boucher, as well as Sandrine Warion and Lucero Petrowski from INRAE Tous Chercheurs laboratory for their help in preparing the tick collection kit. Special thanks to Gerald Umhang (ANSES Nancy Laboratory for Rabies and Wildlife) for his assistance in tick sampling and valuable feedback on the sampling protocol. We are grateful to the departments of communication and Institutional relations of INRAE and ANSES for their support in recruiting citizen scientists through media campaigns. Last, but not least, we sincerely thank all of the citizen scientists whose participation made this project a true success. References Fuente JDL. Overview: Ticks as vectors of pathogens that cause disease in humans and animals. Front Biosci. 2008;13:6938–46. Jongejan F, Uilenberg G. The global importance of ticks. Parasitology. 2004;129:S3–14. Mac S, Da Silva SR, Sander B. The economic burden of Lyme disease and the cost-effectiveness of Lyme disease interventions: a scoping review. PLOS ONE. 2019;14:1–17. Rosenberg R, Lindsey NP, Fischer M, Gregory CJ, Hinckley AF, Mead PS, et al. Vital signs : Trends in reported vectorborne disease cases — United States and territories, 2004–2016. MMWR Morb Mortal Wkly Rep. 2018;67:496–501. Steere AC, Strle F, Wormser GP, Hu LT, Branda JA, Hovius JWR, et al. Lyme borreliosis. Nat Rev Dis Primer. 2016;2:1–18. Johnson N. Ticks: biology, ecology, and diseases. London, United Kingdom: Elsevier, Academic Press, an Imprint of Elsevier; 2023. Castro MB, Wright SA. Vertebrate hosts of Ixodes pacificus (Acari: Ixodidae) in California. J Vector Ecol. 2007;32:140–9. Hofmeester TR, Coipan EC, Van Wieren SE, Prins HHT, Takken W, Sprong H. Few vertebrate species dominate the Borrelia burgdorferi s.l. life cycle. Environ Res Lett. 2016;11:1–16. Keirans JE, Hutcheson HJ, Durden LA, Klompen JSH. Ixodes (Ixodes) scapularis (Acari: Ixodidae) : redescription of all active stages, distribution, hosts, geographical variation, and medical and veterinary importance. J Med Entomol. 1996;33:297–318. Wang S-S, Liu J-Y, Wang B-Y, Wang W-J, Cui X-M, Jiang J-F, et al. Geographical distribution of Ixodes persulcatus and associated pathogens: analysis of integrated data from a China field survey and global published data. One Health. 2023;16:1–9. Uspensky I. The Taiga Tick Ixodes Persulcatus (Acari: Ixodidae), the main vector of Borrelia Burgdorferi Sensu Lato in Eurasia. In: Capinera JL, editor. Encycl Entomol. 2nd ed. 2016. Fischhoff IR, Keesing F, Ostfeld RS. Risk factors for bites and diseases associated with black-legged ticks: a meta-analysis. Am J Epidemiol. 2019;188:1742–50. Diuk-Wasser MA, VanAcker MC, Fernandez MP. Impact of land use changes and habitat fragmentation on the eco-epidemiology of tick-borne diseases. J Med Entomol. 2021;58:1546–64. Pfäffle M, Littwin N, Muders SV, Petney TN. The ecology of tick-borne diseases. Int J Parasitol. 2013;43:1059–77. Estrada-Peña A, De La Fuente J. The ecology of ticks and epidemiology of tick-borne viral diseases. Antiviral Res. 2014;108:104–28. Eisen RJ, Eisen L, Ogden NH, Beard CB. Linkages of weather and climate with Ixodes scapularis and Ixodes pacificus (Acari: Ixodidae), enzootic transmission of Borrelia burgdorferi , and Lyme disease in North America. J Med Entomol. 2016;53:250–61. Grigoryeva LA. Influence of air humidity on the survival rate, lifetime, and development of Ixodes ricinus (L., 1758) and Ixodes persulcatus Schulze, 1930 (Acari: Ixodidae). Syst Appl Acarol. 2022;27:2241–8. Herrmann C, Gern L. Survival of Ixodes ricinus (Acari: Ixodidae) under challenging conditions of temperature and humidity is influenced by Borrelia burgdorferi sensu lato infection. J Med Entomol. 2010;47:1196–204. Leal B, Zamora E, Fuentes A, Thomas DB, Dearth RK. Questing by tick larvae (Acari: Ixodidae): a review of the influences that affect off-host survival. Reddy GVP, editor. Ann Entomol Soc Am. 2020;113:425–38. Ogden NH, Lindsay LR, Beauchamp G, Charron D, Maarouf A, O’Callaghan CJ, et al. Investigation of relationships between temperature and developmental rates of tick Ixodes scapularis (Acari: Ixodidae) in the laboratory and field. J Med Entomol. 2004;41:622–33. Perret JL, Rais O, Gern L. Influence of climate on the proportion of Ixodes ricinus nymphs and adults questing in a tick population. J Med Entomol. 2004;41:361–5. Perret J-L, Guerin PM, Diehl PA, Vlimant M, Gern L. Darkness induces mobility, and saturation deficit limits questing duration, in the tick Ixodes ricinus . J Exp Biol. 2003;206:1809–15. De Frenne P, Lenoir J, Luoto M, Scheffers BR, Zellweger F, Aalto J, et al. Forest microclimates and climate change: importance, drivers and future research agenda. Glob Change Biol. 2021;27:2279–97. Milling CR, Rachlow JL, Olsoy PJ, Chappell MA, Johnson TR, Forbey JS, et al. Habitat structure modifies microclimate: an approach for mapping fine‐scale thermal refuge. Mahon A, editor. Methods Ecol Evol. 2018;9:1648–57. Mathisson DC, Kross SM, Palmer MI, Diuk-Wasser MA. Effect of vegetation on the abundance of tick vectors in the northeastern United States: a review of the literature. Ginsberg H, editor. J Med Entomol. 2021;58:2030–7. Medlock JM, Shuttleworth H, Copley V, Hansford KM, Leach S. Woodland biodiversity management as a tool for reducing human exposure to Ixodes ricinus ticks: a preliminary study in an English woodland. J Vector Ecol. 2012;37:307–15. Schulze TL, Jordan RA. Influence of meso- and microscale habitat structure on focal distribution of sympatric Ixodes scapularis and Amblyomma americanum (Acari: Ixodidae). J Med Entomol. 2005;42:285–94. Ecke F, Löfgren O, Sörlin D. Population dynamics of small mammals in relation to forest age and structural habitat factors in northern Sweden. J Appl Ecol. 2002;39:781–92. Ferger SW, Schleuning M, Hemp A, Howell KM, Böhning‐Gaese K. Food resources and vegetation structure mediate climatic effects on species richness of birds. Glob Ecol Biogeogr. 2014;23:541–9. Bourdin A, Dokhelar T, Bord S, Van Halder I, Stemmelen A, Scherer-Lorenzen M, et al. Forests harbor more ticks than other habitats: a meta-analysis. For Ecol Manag. 2023;541:1–9. Gregory N, Fernandez MP, Diuk-Wasser M. Risk of tick-borne pathogen spillover into urban yards in New York City. Parasit Vectors. 2022;15:1–14. Heylen D, Lasters R, Adriaensen F, Fonville M, Sprong H, Matthysen E. Ticks and tick-borne diseases in the city: role of landscape connectivity and green space characteristics in a metropolitan area. Sci Total Environ. 2019;670:941–9. VanAcker MC, Little EAH, Molaei G, Bajwa WI, Diuk-Wasser MA. Enhancement of risk for Lyme disease by landscape connectivity, New York, New York, USA. Emerg Infect Dis. 2019;25:1136–43. Estrada-Peña A. The relationships between habitat topology, critical scales of connectivity and tick abundance Ixodes ricinus in a heterogeneous landscape in northern Spain. Ecography. 2003;26:661–71. Estrada-Peña A. Effects of habitat suitability and landscape patterns on tick (Acarina) metapopulation processes. Landsc Ecol. 2005;20:529–41. Li S, Heyman P, Cochez C, Simons L, Vanwambeke SO. A multi-level analysis of the relationship between environmental factors and questing Ixodes ricinus dynamics in Belgium. Parasit Vectors. 2012;5:1–11. Ehrmann S, Liira J, Gärtner S, Hansen K, Brunet J, Cousins SAO, et al. Environmental drivers of Ixodes ricinus abundance in forest fragments of rural European landscapes. BMC Ecol. 2017;17:1–14. Perez G, Bastian S, Agoulon A, Bouju A, Durand A, Faille F, et al. Effect of landscape features on the relationship between Ixodes ricinus ticks and their small mammal hosts. Parasit Vectors. 2016;9:1–18. Talbot B, Slatculescu A, Thickstun CR, Koffi JK, Leighton PA, McKay R, et al. Landscape determinants of density of blacklegged ticks, vectors of Lyme disease, at the northern edge of their distribution in Canada. Sci Rep. 2019;9:1–12. Richter D, Schneider A-K, Schibalski A, Dahlkamp A, Schröder B. Features in and around residential gardens affecting the presence and abundance of questing Ixodes ricinus ticks. Infect Ecol Epidemiol. 2023;13:1–12. Hassler L, Durand J, Frey-Klett P. CiTIQUE-Tracker [Internet]. INRAE, SK8; 2024 [cited 2024 May 17]. Available from: https://ci-tique-tracker.sk8.inrae.fr/ Mulder S, Van Vliet AJH, Bron WA, Gassner F, Takken W. High risk of tick bites in Dutch gardens. Vector-Borne Zoonotic Dis. 2013;13:865–71. Raguet S, Couturier E. Étude ALSA(CE)TIQUE 2014-2015. Principaux résultats descriptifs. Saint-Maurice: Santé Publique France; 2016 p. 1–9. Sciensano. TIQUESNET 2020, surveillance des morsures de tiques en Belgique. Épidémiologie des maladies infectieuses. Bruxelles; 2021 p. 1–12. Seifert VA, Wilson S, Toivonen S, Clarke B, Prunuske A. Community partnership designed to promote Lyme disease prevention and engagement in citizen science. J Microbiol Biol Educ. 2016;17:63–9. Eisen L, Eisen RJ. Benefits and drawbacks of citizen science to complement traditional data gathering approaches for medically important hard ticks (Acari: Ixodidae) in the United States. J Med Entomol. 2021;58:1–9. Heyman P, Cochez C, Hofhuis A, Van Der Giessen J, Sprong H, Porter SR, et al. A clear and present danger: tick-borne diseases in Europe. Expert Rev Anti Infect Ther. 2010;8:33–50. INSEE. Growth and structure of the population in 2020 inter-communality metropolis of Métropole du Grand Nancy (245400676) [Internet]. 2023 Jun. Available from: https://www.insee.fr/fr/statistiques/1405599?geo=EPCI-245400676 Météo France. Fiche climatologique; indicatif : 54526001 [Internet]. 2023 [cited 2024 May 6]. Available from: https://donneespubliques.meteofrance.fr/FichesClim/FICHECLIM_54526001.pdf DataGrand Est. Occupation du sol à grande échelle en région Grand Est [Internet]. 2022 [cited 2024 May 6]. Available from: https://www.datagrandest.fr/portail/fr/projets/occupation-du-sol Wongnak P, Bord S, Jacquot M, Agoulon A, Beugnet F, Bournez L, et al. Meteorological and climatic variables predict the phenology of Ixodes ricinus nymph activity in France, accounting for habitat heterogeneity. Sci Rep. 2022;12:1–16. Perez G, Bournez L, Boulanger N, Fite J, Livoreil B, McCoy KD, et al. The distribution, phenology, host range and pathogen prevalence of Ixodes ricinus in France: a systematic map and narrative review. Peer Community J. 2023;3:1–40. Estrada-Peña A, Mihalca AD, Petney T, Domşa C, Gragera Artal J, editors. Ticks of Europe and North Africa: a guide to species identification. Cham: Springer Switzerland; 2017. Pérez-Eid C. Les tiques : identification, biologie, importance medicale et veterinaire. Lavoisier, Editions médicales internationales; 2007. Durand J, Bournez L, Marchand J, Schmid C, Carravieri I, Palin B, et al. Are orienteers protected enough against tick bites? Estimating human exposure to tick bites through a participative science survey during an orienteering competition. Int J Environ Res Public Health. 2021;18:1–19. Wilhelmsson P, Lindblom P, Fryland L, Nyman D, Jaenson TG, Forsberg P, et al. Ixodes ricinus ticks removed from humans in Northern Europe: seasonal pattern of infestation, attachment sites and duration of feeding. Parasit Vectors. 2013;6:1–11. ETALAB. Cadastre data of France [Internet]. 2022 [cited 2023 Aug 2]. Available from: https://cadastre.data.gouv.fr/data/etalab-cadastre/2020-10-01/shp/departements/ Jaeger JAG. Landscape division, splitting index, and effective mesh size: new measures of landscape fragmentation. Landsc Ecol. 2000;15:115–30. Randolph SE, Storey K. Impact of microclimate on immature tick-rodent host interactions (Acari: Ixodidae): implications for parasite transmission. J Med Entomol. 1999;36:741–8. Hilbe JM. Negative binomial regression. 2nd ed. Cambridge, UK ; New York: Cambridge University Press; 2011. Burnham KP, Anderson DR, editors. Model selection and multimodel inference. New York, NY: Springer New York; 2004. Youden WJ. Index for rating diagnostic tests. Cancer. 1950;3:32–5. R Core Team. R: a language and environment for statistical computing [Internet]. Vienna, Austria: R Foundation for Statistical Computing; 2023. Available from: https://www.R-project.org/ Brooks ME, Kristensen K, van Benthem KJ, Magnusson A, Berg CW, Nielsen A, et al. glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. 2017; Available from: http://hdl.handle.net/20.500.11850/242692 Bartoń K. MuMIn: Multi-Model Inference. 2023; Available from: https://CRAN.R-project.org/package=MuMIn Lüdecke D, Ben-Shachar MS, Patil I, Waggoner P, Makowski D. An R package for assessment, comparison and testing of statistical models. J Open Source Softw. 2021;6:1–8. Hartig F. DHARMa: Residual Diagnostics for Hierarchical (Multi-Level / Mixed) Regression Models. 2022; Available from: https://CRAN.R-project.org/package=DHARMa Thiele C, Hirschfeld G. cutpointr: improved estimation and validation of optimal cutpoints in R . J Stat Softw. 2021;98:1–27. Robin X, Turck N, Hainard A, Tiberti N, Lisacek F, Sanchez J-C, et al. pROC: an open-source package for R and S+ to analyze and compare ROC curves. BMC Bioinformatics. 2011;12:1–8. Lüdecke D. sjPlot: data visualization for statistics in social science. 2023; Available from: https://CRAN.R-project.org/package=sjPlot Dijkstra L, Poelman H. A harmonised definition of cities and rural areas: the new degree of urbanisation. 2021;European Commission Working Paper WP 01/2014. INSEE. La grille communale de densité à 7 niveaux [Internet]. 2023 [cited 2024 May 6]. Available from: https://www.insee.fr/fr/information/6439600 Hansford KM, Gillingham EL, Vaux AGC, Cull B, McGinley L, Catton M, et al. Impact of green space connectivity on urban tick presence, density and Borrelia infected ticks in different habitats and seasons in three cities in southern England. Ticks Tick-Borne Dis. 2023;14:1–11. Hansford KM, Wheeler BW, Tschirren B, Medlock JM. Questing Ixodes ricinus ticks and Borrelia spp. in urban green space across Europe: A review. Zoonoses Public Health. 2022;69:153–66. Bourdin A, Bord S, Durand J, Galon C, Moutailler S, Scherer-Lorenzen M, et al. Forest diversity reduces the prevalence of pathogens transmitted by the tick Ixodes ricinus . Front Ecol Evol. 2022;10:1–13. Kiewra D, Stefańska-Krzaczek E, Szymanowski M, Szczepańska A. Local-scale spatio-temporal distribution of questing Ixodes ricinus L. (Acari: Ixodidae)-A case study from a riparian urban forest in Wrocław, SW Poland. Ticks Tick-Borne Dis. 2017;8:362–9. Tack W, Madder M, Baeten L, Vanhellemont M, Gruwez R, Verheyen K. Local habitat and landscape affect Ixodes ricinus tick abundances in forests on poor, sandy soils. For Ecol Manag. 2012;265:30–6. Tack W, Madder M, Baeten L, Verheyen K. The abundance of Ixodes ricinus ticks depends on tree species composition and shrub cover. Parasitology. 2012;139:1273–81. Boyard C, Barnouin J, Bord S, Gasqui P, Vourc’h G. Reproducibility of local environmental factors for the abundance of questing Ixodes ricinus nymphs on pastures. Ticks Tick-Borne Dis. 2011;2:104–10. Hansford KM, McGinley L, Wilkinson S, Gillingham EL, Cull B, Gandy S, et al. Ixodes ricinus and Borrelia burgdorferi sensu lato in the Royal Parks of London, UK. Exp Appl Acarol. 2021;84:593–606. Mathews-Martin L, Namèche M, Vourc’h G, Gasser S, Lebert I, Poux V, et al. Questing tick abundance in urban and peri-urban parks in the French city of Lyon. Parasit Vectors. 2020;13:1–9. Van Gestel M, Verheyen K, Matthysen E, Heylen D. Danger on the track? Tick densities near recreation infrastructures in forests. Urban For Urban Green. 2021;59:1–7. Sodoudi S, Zhang H, Chi X, Müller F, Li H. The influence of spatial configuration of green areas on microclimate and thermal comfort. Urban For Urban Green. 2018;34:85–96. Van Gestel M, Matthysen E, Heylen D, Verheyen K. Survival in the understorey: testing direct and indirect effects of microclimatological changes on Ixodes ricinus . Ticks Tick-Borne Dis. 2022;13:1–7. Lee X, Maxson G-A, Paskewitz S. Single mowing event does not reduce abundance of Ixodes scapularis (Acari: Ixodidae) and Dermacentor variabilis (Acari: Ixodidae) on recreational hiking trails. J Med Entomol. 2023;60:228–34. Hahn MB, Bjork JKH, Neitzel DF, Dorr FM, Whitemarsh T, Boegler KA, et al. Evaluating acarological risk for exposure to Ixodes scapularis and Ixodes scapularis -borne pathogens in recreational and residential settings in Washington County, Minnesota. Ticks Tick-Borne Dis. 2018;9:340–8. Fischhoff IR, Keesing F, Pendleton J, DePietro D, Teator M, Duerr STK, et al. Assessing effectiveness of recommended residential yard management measures against ticks. J Med Entomol. 2019;56:1420–7. Grade AM, Warren PS, Lerman SB. Managing yards for mammals: mammal species richness peaks in the suburbs. Landsc Urban Plan. 2022;220:1–11. Hansen CP, Parsons AW, Kays R, Millspaugh JJ. Does Use of backyard resources explain the abundance of urban wildlife? Front Ecol Evol. 2020;8:1–12. Johansson EP, DeGregorio BA. The effects of landscape and yard features on mammal diversity in residential yards within Northwest Arkansas, USA. Urban Ecosyst. 2024;27:275–87. Hoch T, Monnet Y, Agoulon A. Influence of host migration between woodland and pasture on the population dynamics of the tick Ixodes ricinus : a modelling approach. Ecol Model. 2010;221:1798–806. Fraisl D, Hager G, Bedessem B, Gold M, Hsing P-Y, Danielsen F, et al. Citizen science in environmental and ecological sciences. Nat Rev Methods Primer. 2022;2:1–20. Kosmala M, Wiggins A, Swanson A, Simmons B. Assessing data quality in citizen science. Front Ecol Environ. 2016;14:551–60. Potes L, Bouchard C, Rocheleau J-P, Richard L, Leighton P, Pelletier J, et al. Evaluation of a community-based One Health intervention to reduce the risk of Lyme disease in a high-incidence municipality. CABI One Health. 2023;2:1–12. Ecosystems and human well-being: wetlands and water synthesis: a report of the Millennium Ecosystem Assessment. Washington, DC: World Resources Institute; 2005. Stafford KC. Tick management handbook; an integrated guide for homeowners, pest control operators, and public health officials for the prevention of tick-associated disease. New Haven, CT: The Connecticut Agricultural Experiment Station; 2007. Černý J, Lynn G, Hrnková J, Golovchenko M, Rudenko N, Grubhoffer L. Management options for Ixodes ricinus -associated pathogens: a review of prevention strategies. Int J Environ Res Public Health. 2020;17:1–18. Hansen J. How to kill and prevent ticks in your lawn [Internet]. [cited 2024 May 6]. Available from: https://www.gardentech.com/blog/pest-id-and-prevention/protecting-your-lawn-and-garden-against-ticks Reed JH, Bonter DN. Supplementing non-target taxa: bird feeding alters the local distribution of mammals. Ecol Appl. 2018;28:761–70. Clotfelter ED, Pedersen AB, Cranford JA, Ram N, Snajdr EA, Nolan V, et al. Acorn mast drives long-term dynamics of rodent and songbird populations. Oecologia. 2007;154:493–503. Ostfeld RS, Levi T, Keesing F, Oggenfuss K, Canham CD. Tick‐borne disease risk in a forest food web. Ecology. 2018;99:1562–73. Schnurr JL, Ostfeld RS, Canham CD. Direct and indirect effects of masting on rodent populations and tree seed survival. Oikos. 2002;96:402–10. Maupin GO, Fish D, Zultowsky J, Campos EG, Piesman J. Landscape ecology of Lyme disease in a residential area of Westchester County, New York. Am J Epidemiol. 1991;133:1105–13. Kays R, Parsons AW. Mammals in and around suburban yards, and the attraction of chicken coops. Urban Ecosyst. 2014;17:691–705. Vercauteren KC, Vandeelen TR, Lavelle MJ, Hall WH. Assessment of abilities of white‐tailed deer to jump fences. J Wildl Manag. 2010;74:1378–81. Hofmeester TR, Sprong H, Jansen PA, Prins HHT, van Wieren SE. Deer presence rather than abundance determines the population density of the sheep tick, Ixodes ricinus , in Dutch forests. Parasit Vectors. 2017;10:1–8. Jennett AL, Smith FD, Wall R. Tick infestation risk for dogs in a peri-urban park. Parasit Vectors. 2013;6:1–10. Medlock JM, Vaux AGC, Hansford KM, Pietzsch ME, Gillingham EL. Ticks in the ecotone: the impact of agri‐environment field margins on the presence and intensity of Ixodes ricinus ticks (Acari: Ixodidae) in farmland in southern England. Med Vet Entomol. 2020;34:175–83. Gehring TM, Swihart RK. Body size, niche breadth, and ecologically scaled responses to habitat fragmentation: mammalian predators in an agricultural landscape. Biol Conserv. 2003;109:283–95. Panzacchi M, Linnell JDC, Melis C, Odden M, Odden J, Gorini L, et al. Effect of land-use on small mammal abundance and diversity in a forest–farmland mosaic landscape in south-eastern Norway. For Ecol Manag. 2010;259:1536–45. Baker PJ, Harris S. Urban mammals: what does the future hold? An analysis of the factors affecting patterns of use of residential gardens in Great Britain. Mammal Rev. 2007;37:297–315. Anderson CS, Cady AB, Meikle DB. Effects of vegetation structure and edge habitat on the density and distribution of white-footed mice ( Peromyscus leucopus ) in small and large forest patches. Can J Zool. 2003;81:897–904. Ries L, Fletcher RJ, Battin J, Sisk TD. Ecological responses to habitat edges: mechanisms, models, and variability explained. Annu Rev Ecol Evol Syst. 2004;35:491–522. Additional Declarations No competing interests reported. Supplementary Files supplementarymaterialfinal.docx Cite Share Download PDF Status: Published Journal Publication published 10 Oct, 2024 Read the published version in International Journal of Health Geographics → Version 1 posted Editorial decision: Revision requested 05 Jul, 2024 Reviews received at journal 26 Jun, 2024 Reviews received at journal 04 Jun, 2024 Reviewers agreed at journal 29 May, 2024 Reviewers agreed at journal 28 May, 2024 Reviewers invited by journal 28 May, 2024 Submission checks completed at journal 21 May, 2024 Editor assigned by journal 21 May, 2024 First submitted to journal 21 May, 2024 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. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. 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-4453852","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":308070998,"identity":"2e859619-fcdb-4142-bf99-b26eb550b221","order_by":0,"name":"Anna Mazaleyrat","email":"","orcid":"","institution":"ANSES, Nancy Laboratory for Rabies and Wildlife","correspondingAuthor":false,"prefix":"","firstName":"Anna","middleName":"","lastName":"Mazaleyrat","suffix":""},{"id":308070999,"identity":"c1f0d904-314d-4397-95d3-4f422e425c4c","order_by":1,"name":"Jonas Durand","email":"","orcid":"","institution":"UMR 1136 ‘Interactions Arbres Micro-Organismes’, Université de Lorraine, INRAE, Centre INRAE Grand Est-Nancy","correspondingAuthor":false,"prefix":"","firstName":"Jonas","middleName":"","lastName":"Durand","suffix":""},{"id":308071001,"identity":"3d4e5103-f205-41ec-9ccd-e5dd374b87f3","order_by":2,"name":"Irene Carravieri","email":"","orcid":"","institution":"Centre Permanent d’Initiatives pour l’Environnement Nancy Champenoux","correspondingAuthor":false,"prefix":"","firstName":"Irene","middleName":"","lastName":"Carravieri","suffix":""},{"id":308071003,"identity":"ea3d8689-8386-467c-9251-26507cf95973","order_by":3,"name":"Christophe Caillot","email":"","orcid":"","institution":"ANSES, Nancy Laboratory for Rabies and Wildlife","correspondingAuthor":false,"prefix":"","firstName":"Christophe","middleName":"","lastName":"Caillot","suffix":""},{"id":308071005,"identity":"4525bc53-e982-41d4-a3ee-be0ca819a044","order_by":4,"name":"Cyril Galley","email":"","orcid":"","institution":"Centre Permanent d’Initiatives pour l’Environnement Nancy Champenoux","correspondingAuthor":false,"prefix":"","firstName":"Cyril","middleName":"","lastName":"Galley","suffix":""},{"id":308071007,"identity":"aaa61def-c6d9-416a-9153-625bdb08f1c3","order_by":5,"name":"Sandrine Capizzi","email":"","orcid":"","institution":"UMR 1136 ‘Interactions Arbres Micro-Organismes’, Université de Lorraine, INRAE, Centre INRAE Grand Est-Nancy","correspondingAuthor":false,"prefix":"","firstName":"Sandrine","middleName":"","lastName":"Capizzi","suffix":""},{"id":308071014,"identity":"c311241a-e385-4381-aa63-52c72eebd87a","order_by":6,"name":"Franck Boué","email":"","orcid":"","institution":"ANSES, Nancy Laboratory for Rabies and Wildlife","correspondingAuthor":false,"prefix":"","firstName":"Franck","middleName":"","lastName":"Boué","suffix":""},{"id":308071015,"identity":"9e5ed1f6-9018-4df4-9498-8974958c4198","order_by":7,"name":"Pascale Frey-Klett","email":"","orcid":"","institution":"UMR 1136 ‘Interactions Arbres Micro-Organismes’, Université de Lorraine, INRAE, Centre INRAE Grand Est-Nancy","correspondingAuthor":false,"prefix":"","firstName":"Pascale","middleName":"","lastName":"Frey-Klett","suffix":""},{"id":308071016,"identity":"f4f6d8aa-2c74-4350-a14b-e451807e1920","order_by":8,"name":"Laure Bournez","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA4klEQVRIiWNgGAWjYBACPgh1AMqtQKOxATZULWfQaMJaGNuI0cJ+xuzDB4Y7cga3zx58+HOeXZ58e/sDhoN78GjhyTGeOYPhmbHBubxkA8ltycUGZ84YMBx4hs9hOcbMPAyHEzec4TGTMNzGnLhBIoeB+cMBPFr43xgz/4FoMf+ROKc+cf785w8YDuDTIgG0hQFqC8PBhsOJDTcYDAhoeVbM2GPwzFjyDI+xZMOx40C9OQYH8Gnh50/ezPCj4o4c3xkew48/aqoT57cff/gAnxYIMEDjE9QwCkbBKBgFowA/AADgwlRmUhPmvQAAAABJRU5ErkJggg==","orcid":"","institution":"ANSES, Nancy Laboratory for Rabies and Wildlife","correspondingAuthor":true,"prefix":"","firstName":"Laure","middleName":"","lastName":"Bournez","suffix":""}],"badges":[],"createdAt":"2024-05-21 09:49:03","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-4453852/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-4453852/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1186/s12942-024-00380-9","type":"published","date":"2024-10-10T15:57:23+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":57614802,"identity":"f98d3c21-e9c7-43d1-9b49-ea561504fa44","added_by":"auto","created_at":"2024-06-03 11:14:06","extension":"jpg","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":4320633,"visible":true,"origin":"","legend":"\u003cp\u003eLocation of the 185 sampled yards near Nancy, a city in the northeast of France. Tick sampling took place from May to mid-July 2020 (n = 26), 2021 (n = 59), and 2022 (n = 100) within a 35 km radius of Nancy. Land cover classes were modified from DataGrand Est [50].\u003c/p\u003e","description":"","filename":"Figure1.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4453852/v1/5f829e761c25524c25bd1ae8.jpg"},{"id":57614509,"identity":"0ba5c11e-e8c2-4c5d-9271-d11f727a7613","added_by":"auto","created_at":"2024-06-03 11:06:06","extension":"jpg","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":1007000,"visible":true,"origin":"","legend":"\u003cp\u003eSummary of the best models for risk factors associated with the three occurrence variables. Odd ratios with 95% confidence interval are shown for each explanatory variable. Bold indicates confidence interval excluding one. Non-significant relationships are shown as dotted lines. In model summaries, reference levels are no signs of deer and no wood/brush pile. Variables were computed within a 500-m (\u003csup\u003e500\u003c/sup\u003e) or a 50-m buffer (\u003csup\u003e50\u003c/sup\u003e) from property edges. MESH: Effective mesh size.\u003c/p\u003e","description":"","filename":"Figure2.jpg","url":"https://assets-eu.researchsquare.com/files/rs-4453852/v1/687d660462eb4d6af75d62f0.jpg"},{"id":66597127,"identity":"262cdb95-da74-46b6-86bd-cdee138ba940","added_by":"auto","created_at":"2024-10-14 16:07:29","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":6309145,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-4453852/v1/c4a5f388-1886-41ea-871e-66c4e3059f8a.pdf"},{"id":57614511,"identity":"27843d0a-10e0-48a1-b8c0-611df8a27b47","added_by":"auto","created_at":"2024-06-03 11:06:06","extension":"docx","order_by":8,"title":"","display":"","copyAsset":false,"role":"supplement","size":1323949,"visible":true,"origin":"","legend":"","description":"","filename":"supplementarymaterialfinal.docx","url":"https://assets-eu.researchsquare.com/files/rs-4453852/v1/cb9fcf6237e4501525fcc773.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Understanding Ixodes ricinus occurrence in private yards: influence of yard and landscape features","fulltext":[{"header":"Background","content":"\u003cp\u003eTicks are hematophagous ectoparasites and can transmit many bacteria, viruses, and protozoa causing human and animal diseases [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2\u003c/span\u003e]. In the northern hemisphere, Lyme borreliosis is the most frequently reported zoonotic disease and an important economic burden for countries [\u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e3\u003c/span\u003e, \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e4\u003c/span\u003e]. The primary vectors of Lyme borreliosis are ticks from the \u003cem\u003eIxodes ricinus\u003c/em\u003e species complex (hereafter \u003cem\u003eIxodes\u003c/em\u003e), namely \u003cem\u003eI. ricinus\u003c/em\u003e in Europe, \u003cem\u003eI. pacificus\u003c/em\u003e in western North America, \u003cem\u003eI. scapularis\u003c/em\u003e in eastern and mid-western USA and southern Canada, and \u003cem\u003eI. persulcatus\u003c/em\u003e in Asia [\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e5\u003c/span\u003e]. These ticks have three active life stages, larva, nymph, and adult, each requiring a blood meal to molt into the next stage (larva and nymph) or produce eggs (adult female) [\u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e6\u003c/span\u003e]. Although they can feed on a broad range of vertebrate hosts, including birds, small mammals, carnivores, and ungulates [\u003cspan additionalcitationids=\"CR8 CR9\" citationid=\"CR7\" class=\"CitationRef\"\u003e7\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e10\u003c/span\u003e], in forests subadults mainly feed on birds and small- and medium-sized mammals, while adults are mainly fed by medium- and large-sized mammals (e.g. ungulates) [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e, \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e9\u003c/span\u003e, \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e11\u003c/span\u003e]. Identifying the drivers of \u003cem\u003eIxodes\u003c/em\u003e tick distribution and abundance is necessary to predict and mitigate tick exposure, thereby potentially reducing the risk of tick-borne diseases [\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e12\u003c/span\u003e].\u003c/p\u003e \u003cp\u003eThe distribution and abundance of ticks are linked to habitat suitability for both ticks and their hosts, which depend on complex interactions between abiotic (e.g. climatic conditions) and biotic conditions (host community and vegetation) playing differently at various spatial scales [\u003cspan additionalcitationids=\"CR14\" citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e]. At the local scale, tick populations are also influenced by both abiotic and biotic conditions. Microclimatic conditions, impact the survival, development, and activity of \u003cem\u003eIxodes\u003c/em\u003e ticks [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan additionalcitationids=\"CR17 CR18 CR19 CR20 CR21\" citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]. For example, low humidity and high temperatures usually have a detrimental effect on the survival of \u003cem\u003eIxodes\u003c/em\u003e ticks [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e]. Microclimatic conditions are directly influenced by biotic habitat and soil characteristics such as vegetation structure (e.g. cover of near-ground vegetation) and composition (e.g. diversity of plant species), as well as soil texture. Besides buffering climatic extremes detrimental to tick survival [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e24\u003c/span\u003e], trees and shrubs also create a litter layer with a humid microhabitat protecting ticks from desiccation during their off-host period [\u003cspan additionalcitationids=\"CR26\" citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e27\u003c/span\u003e]. Tick abundance also depends on the presence and abundance of their hosts [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e], which are influenced by vegetation structure and composition (e.g. small mammals and birds: [\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e28\u003c/span\u003e, \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e29\u003c/span\u003e] through changes in the availability of food resources, shelters, breeding, or resting sites for hosts. Overall, several studies demonstrated a strong association between \u003cem\u003eIxodes\u003c/em\u003e ticks and woodland habitats (reviewed in: [\u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e25\u003c/span\u003e, \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]), likely reflecting both the higher availability of some propagation hosts (e.g. deer) and the more suitable microclimatic conditions in forest habitats compared to non-forest habitats.\u003c/p\u003e \u003cp\u003eAt a broader scale, landscape composition (e.g. proportion of habitat types) and configuration (i.e. spatial arrangement of these habitats) can influence the host availability through changes in the host community composition, host movement, and habitat use [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]. Landscape fragmentation, common in urban areas, results in a mosaic of patches of various sizes and land-use types, potentially impeding tick host movement [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e] and isolating suitable habitats for ticks and their hosts from one another and from the source of large propagation hosts (e.g. deer in forests). In this context, ticks form a metapopulation, with subpopulations connected and reliant on each other for persistence. The degree of connectivity between subpopulations, enhanced by the tick host movement, determines whether tick subpopulations persist. Consistent with this explanation, previous studies in both urban and rural environments have demonstrated that connectivity between suitable patches for ticks and their hosts, as well as connectivity to populations of deer, are important for tick persistence (urban: [\u003cspan additionalcitationids=\"CR32\" citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e]; rural: [\u003cspan additionalcitationids=\"CR35\" citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e]). Several studies have reported an increased abundance or occurrence of \u003cem\u003eIxodes\u003c/em\u003e ticks with an increasing proportion of forest/tree canopy or the number of forest patches in the landscape [\u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e, \u003cspan additionalcitationids=\"CR37 CR38 CR39\" citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e] and a decreasing proportion of agricultural areas and built-up and paved areas [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]. Considering the complex interplay between local and landscape habitat characteristics on ticks and their hosts, studies should examine both spatial scales concurrently (e.g. [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e]).\u003c/p\u003e \u003cp\u003eUntil now, most studies on tick ecology in Europe have primarily focused on forests, where ticks are most abundant [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]. Our understanding of the local and landscape features associated with tick occurrence in private yards thus remains limited (but see: Richter et al. in Germany [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e] and Gregory et al. in the USA [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]). Nevertheless, this knowledge is fundamental to the prevention of tick-borne diseases as yards are important places for tick encounters and tick bites. Studies conducted in France, Belgium, and the Netherlands indicated that approximately 30% of reported tick bites occurred in yards [\u003cspan additionalcitationids=\"CR42 CR43\" citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e]. The scarcity of research on tick populations in private yards can be primarily attributed to the challenge associated with tick sampling, as these private areas are not easily accessible to scientists. Citizen science, when members of the public collaborate with scientists to answer research questions, helps to overcome this challenge while likely increasing citizens\u0026rsquo; tick-borne disease literacy (as seen in a researcher \u0026ndash; community partnership to promote Lyme disease prevention [\u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e45\u003c/span\u003e]). Citizen science has emerged as a powerful means of advancing tick research (reviewed in [\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e]) and is a valuable approach to gain insight into the complex ecology of ticks in yards.\u003c/p\u003e \u003cp\u003eTIQUoJARDIN, which translates to TICKinYARD in English, is a citizen science project conducted in northeastern France. This project aims to characterize the risk associated with the presence of ticks in private yards, encompassing the occurrence of ticks, their pathogens, and human exposure. The project also seeks to identify the local and landscape features related to this risk. To reach these objectives, citizen scientists followed a standardized protocol to collect ticks in their yard and completed surveys about the attributes of their yard and their tick bites. In this paper, we aim to characterize the presence of ticks in private yards and determine whether habitat characteristics at various scales, ranging from the local scale to the landscape scale, were associated with the occurrence of ticks. This study mainly focuses on \u003cem\u003eIxodes ricinus\u003c/em\u003e, i.e. the most widely distributed tick species in Europe, the primary vector of Lyme borreliosis, and a vector of many other human and animal diseases (e.g. tick-borne encephalitis, Babesiosis, piroplasmosis, and anaplasmosis) [\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e1\u003c/span\u003e, \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e47\u003c/span\u003e].\u003c/p\u003e"},{"header":"Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy site\u003c/h2\u003e \u003cp\u003eThis study took place within a 35 km radius of Nancy (48\u0026deg;41\u0026prime; 31.3\u0026prime;\u0026prime; N, 6\u0026deg;11\u0026prime; 3.9\u0026prime;\u0026prime; E), a city located in northeastern France. Nancy (104 403 inhabitants) is part of a metropolitan community that covers 142 km\u003csup\u003e2\u003c/sup\u003e and has ~\u0026thinsp;255 000 inhabitants [\u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e48\u003c/span\u003e]. The climate is semi-continental with hot summers (mean daily maximum temperature during July is 25.8\u0026deg;C) and cold winters (mean daily temperature during January is 2.6\u0026deg;C) (1991\u0026thinsp;\u0026minus;\u0026thinsp;2020). The mean annual temperature is 11.0\u0026deg;C and mean annual precipitation is 746 mm (1991\u0026thinsp;\u0026minus;\u0026thinsp;2020) [\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e]. The landscape is composed of agricultural areas (35% of the territory), non-forest vegetated areas (e.g. grassland and herbaceous areas, permeable surfaces in urban areas, and meadows), and forest areas (both representing 29% of the territory). Artificial surfaces (i.e. impervious surfaces only), and water bodies and wetlands cover 4% and 2% of the territory, respectively. In the metropolitan community, artificial surfaces cover 27% of the territory (based on data provided by DataGrand Est [\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e]).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eTick sampling\u003c/h2\u003e \u003cp\u003eA massive local media coverage of the project has been organized each year to recruit participating citizens. Ticks were initially sampled in 213 private yards by the residents in 2020 (n\u0026thinsp;=\u0026thinsp;28), 2021 (n\u0026thinsp;=\u0026thinsp;73), and 2022 (n\u0026thinsp;=\u0026thinsp;112). However, due to missing yard features or key information to carry our analysis, we only kept data from 185 yards (n\u0026thinsp;=\u0026thinsp;26 in 2020, 59 in 2021, and 100 in 2022) (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). A yard was defined as a collective or an individual vegetated surface of at least 100 m\u003csup\u003e2\u003c/sup\u003e adjacent or away from the dwelling maintained by the residents. Each yard was sampled once from May to mid-July, i.e. the period of peak activity of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs in the forest in northeastern France [\u003cspan citationid=\"CR51\" class=\"CitationRef\"\u003e51\u003c/span\u003e, \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e52\u003c/span\u003e]. Citizen scientists conducted the sampling following a standardized protocol using a \u0026ldquo;tick collection kit\u0026rdquo;, containing all the material to sample, identify, handle, and store ticks (see supplementary material S1 for details). Participants dragged a 1 m\u003csup\u003e2\u003c/sup\u003e white cloth across the vegetation surface or over the leaf litter (e.g. [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e]). To maximize tick detection probability, participants had to avoid tick sampling when vegetation was wet (ticks attach less well to a wet drag cloth) and sampling between 11 a.m. and 4 p.m. (\u003cem\u003eI. ricinus\u003c/em\u003e activity decreases when temperature increases and/or relative humidity decreases [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e]). Participants had to sample 12 transects of 10 m long (~\u0026thinsp;10 steps), four along property edges, and eight in the core of the yard. After each transect, they inspected the drag cloth (and their clothing to prevent tick bites and missing ticks) and transferred any tick (or what they thought might be a tick) into a labeled ethanol tube. Date and time of tick sampling were recorded. An expert later counted and morphologically identified the ticks to the species level and developmental stage according to Estrada-Pe\u0026ntilde;a et al. [\u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e53\u003c/span\u003e] and P\u0026eacute;rez-Eid [\u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e54\u003c/span\u003e]. This study mainly focuses on \u003cem\u003eI. ricinus\u003c/em\u003e nymphs due to the higher number of individuals collected compared to other stages and species (see result section) and that humans are predominantly bitten by \u003cem\u003eI. ricinus\u003c/em\u003e nymphs [\u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e55\u003c/span\u003e, \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e56\u003c/span\u003e]. Given the low variability in the abundance of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs per yard (see result section), count data were converted into occurrence data.\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eTo enhance tick presence assessment and to account for the single sampling event per yard, participants noted if they had been bitten (or likely bitten) by ticks in their yard during the last three years.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eHabitat characteristics\u003c/h2\u003e \u003cp\u003eTo assess the influence of habitat characteristics on tick occurrence, we focused on three spatial scales: the transect scale, the yard scale, and the landscape scale.\u003c/p\u003e \u003cp\u003eTransect features\u003c/p\u003e \u003cp\u003eFor each transect, citizen scientists recorded the transect location (edge vs core of the yard), grass height (\u0026lt;\u0026thinsp;10 cm, \u0026gt; 10 cm, and mixed, such as litter or various grass heights), shading conditions (partly under trees/bushes vs no tree branches/bushes above the transect), and approximate length (m), since it was not always 10 m long.\u003c/p\u003e \u003cp\u003eYard features\u003c/p\u003e \u003cp\u003eWe computed 13 yard features potentially influencing habitat suitability for \u003cem\u003eI. ricinus\u003c/em\u003e or its hosts based mainly on the responses of participants to a questionnaire (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). These included the presence/absence of a wood/brush pile, a stone wall or a pile of stone, a vegetable garden, a compost, an unmanaged long grass area, a bird feeder, and nut-producing trees (i.e. oaks, beech, hazel, and/or walnut). Participants also reported the presence of six fruit-producing group species and we computed the \u0026ldquo;number of fruit-producing species groups\u0026rdquo; per yard. Groups were based on the type of vegetation (i.e. trees vs. brushes and shrubs), and the period of harvesting (Table \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e). Citizen scientists also indicated whether their yard was fully closed (preventing medium and large-sized mammals from entering the yard) or not. The mowing intensity was computed as the cumulative number of mowing events in spring, summer, and autumn based on participants\u0026rsquo; reported lawn mowing frequency per season. Using the location of the yard, satellite images from Google Earth (including CNES/Airbus, GeoContent, and Maxar technologies images), and cadastral information [\u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e57\u003c/span\u003e], we computed the \u0026ldquo;vegetated surface of the yard\u0026rdquo; as the property surface excluding buildings (e.g. house) and unvegetated areas (e.g. pool and paved alleys). To characterize host presence, participants reported deer sightings or signs of their presence (e.g. feces and footprints) near their property (i.e. inside or within 5 m from the edges of their property). Participants also reported if they had a dog and if cats were present in their yard. Only the variable \u0026ldquo;presence of a dog\u0026rdquo; was considered due to the high prevalence of cats (96% of yards in 2021 and 2022, n\u0026thinsp;=\u0026thinsp;159 yards) and missing information for 2020. The presence of free-ranging chicken was not considered due to the rarity of its occurrence (7% of the yards).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eExpected relationships between yard features computed and the occurrence of \u003cem\u003eI. ricinus\u003c/em\u003e in yards.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eYard features\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eExpected relationship with \u003cem\u003eI. ricinus\u003c/em\u003e occurrence in yards\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eExplanation\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003eHabitat suitability for tick hosts\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePresence of\u003c/p\u003e \u003cp\u003e\u0026bull; a wood/brush pile\u003c/p\u003e \u003cp\u003e\u0026bull; a stone wall or a pile of stone\u003c/p\u003e \u003cp\u003e\u0026bull; a vegetable garden\u003c/p\u003e \u003cp\u003e\u0026bull; a compost\u003c/p\u003e \u003cp\u003e\u0026bull; a bird feeder\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eThese attributes can provide a shelter (i.e. the presence of a wood/brush pile, a stone wall or pile of stone) or food for tick hosts (i.e. the presence of a vegetable garden, a compost, and a bird feeder) leading to an increase in host richness or abundance [\u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e89\u003c/span\u003e, \u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e90\u003c/span\u003e, \u003cspan citationid=\"CR99\" class=\"CitationRef\"\u003e99\u003c/span\u003e]. Previous studies have thus shown a positive relationship between the presence of these features and tick occurrence in yards or recreational sites [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e, \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e86\u003c/span\u003e, \u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e87\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eNo. of fruit-producing species groups\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eFruit or nut-producing trees can provide food for tick host species (mainly small mammals and birds) influencing their abundance (e.g. [\u003cspan additionalcitationids=\"CR101\" citationid=\"CR100\" class=\"CitationRef\"\u003e100\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR102\" class=\"CitationRef\"\u003e102\u003c/span\u003e]). Host species abundance may in turn influence tick abundance [\u003cspan citationid=\"CR100\" class=\"CitationRef\"\u003e100\u003c/span\u003e]. The abundance of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs in pasture has been shown to be positively related to the presence of fleshy fruit trees at pasture edges [\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePresence of nut-producing trees\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eVegetated surface of the yard (m\u003csup\u003e2\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eThe relative abundance of tick host species can increase with yard size [\u003cspan citationid=\"CR89\" class=\"CitationRef\"\u003e89\u003c/span\u003e] and tick abundance has been shown to increase with the vegetated surface of the yard [\u003cspan citationid=\"CR103\" class=\"CitationRef\"\u003e103\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eYard accessibility and presence of tick hosts\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eYard closure\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eYard fencing can reduce the diversity and abundance of host species by preventing medium and large-sized mammals from entering a yard [\u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e90\u003c/span\u003e, \u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e104\u003c/span\u003e, \u003cspan citationid=\"CR105\" class=\"CitationRef\"\u003e105\u003c/span\u003e]. Full fencing around the yards has been shown to decrease the odds of finding \u003cem\u003eI. scapularis\u003c/em\u003e [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e] (but see: [\u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e87\u003c/span\u003e]).\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eSigns of deer\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDeer play a key role in the persistence of \u003cem\u003eI. ricinus\u003c/em\u003e populations as adult ticks mainly feed and copulate on deer [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]. Previous studies showed that the density of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs in forests was positively associated with deer presence [\u003cspan citationid=\"CR106\" class=\"CitationRef\"\u003e106\u003c/span\u003e]. In green spaces (mainly parks), \u003cem\u003eI. ricinus\u003c/em\u003e density increased with the connectivity to a known population of roe deer (\u003cem\u003eCapreolus capreolus\u003c/em\u003e) [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePresence of a dog\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e+/-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDogs can bring back ticks into the yard after a walk outside the yard (e.g. in park [\u003cspan citationid=\"CR107\" class=\"CitationRef\"\u003e107\u003c/span\u003e]) or serve as hosts for feeding ticks in yards. Alternatively, a free dog in a yard can prevent some hosts from entering the yard [\u003cspan citationid=\"CR104\" class=\"CitationRef\"\u003e104\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eHabitat suitability for ticks\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMowing intensity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eIncreasing mowing intensity and the presence of an unmanaged long grass area are expected to decrease and increase the availability of moist microhabitats for ticks, respectively. As low humidity and high temperatures usually have a detrimental effect on the survival of \u003cem\u003eIxodes\u003c/em\u003e ticks [\u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e16\u003c/span\u003e, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e, \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e19\u003c/span\u003e], these features are expected to affect tick occurrence or abundance. However, the influence of mowing intensity and the presence of an unmanaged long grass area on tick abundance or occurrence has been seldom tested appropriately. A study showed that a single mowing event did not affect the abundance of \u003cem\u003eI. scapularis\u003c/em\u003e and \u003cem\u003eDermacentor variabilis\u003c/em\u003e on recreational hiking trails [\u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e85\u003c/span\u003e], while another demonstrated that tick density was not associated with grass height in pasture edges [\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePresence of an unmanaged\u003c/p\u003e \u003cp\u003elong grass area\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eLandscape features\u003c/p\u003e \u003cp\u003eWe used land cover data obtained from high-resolution orthophotos [\u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e50\u003c/span\u003e] to classify land cover into five classes: 1) artificial surfaces (i.e. impervious surfaces only), 2) non-forest vegetated areas (e.g. grassland and herbaceous areas, permeable surfaces in developed areas, and abandoned agricultural areas), 3) forest areas (e.g. deciduous forest, clearcuts, and young plantations), 4) agricultural areas (e.g. annual and perennial crops), and 5) open water and wetlands. Subsequently, we derived eleven features describing the landscape composition and spatial configuration that can influence host availability and the availability of suitable habitats for ticks (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). The six spatial configuration variables were the density of forest patches, the shortest distance to a forest patch, the effective mesh size of vegetation areas (i.e. combining forest and non-forest vegetated areas), the effective mesh size of vegetation and agricultural areas combined, the total edge density, and the edge density of vegetation areas. The effective mesh size (MESH) is a fragmentation index and represents the average size (ha) of areas (i.e. vegetation areas or vegetation and agricultural areas combined) that an organism is connected to in a landscape starting from a randomly chosen point [\u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e58\u003c/span\u003e]. The more barriers in the landscape (e.g. artificial surfaces), the less vegetation areas organisms will have access to, and the lower the effective mesh size. Except for the shortest distance to a forest patch, we computed these variables within buffer sizes of 300, 400, and 500 m to maximize variability and avoid missing values. Indeed, when the landscape consists of only one vegetation patch, the MESH cannot be computed. To describe the landscape composition, we computed five variables, the percentage of artificial surfaces, non-forest vegetated areas, forest areas, and agricultural areas within the three buffer sizes, and the percentage of vegetation areas within a 50-m buffer, to characterize the immediate environment surrounding the yard (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eExpected relationships between landscape features and the occurrence of \u003cem\u003eI. ricinus\u003c/em\u003e in yards.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"4\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eLandscape features\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eExpected relationship with \u003cem\u003eI. ricinus\u003c/em\u003e occurrence in yards\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eExplanation\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"5\" rowspan=\"6\"\u003e \u003cp\u003eLandscape composition\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003ePercentage of\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026bull; artificial surfaces\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eDue to the lack of a litter layer and unsuitable microclimatic conditions, artificial surfaces are unsuitable habitats for ticks. Moreover, they can impede the movement of hosts, especially large-sized mammals which are key hosts of adults \u003cem\u003eI. ricinus\u003c/em\u003e [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e], and isolate the yard from putative sources of ticks and hosts such as forests. Consistently, densities of \u003cem\u003eI. ricinus\u003c/em\u003e (all stages) in green spaces in Belgium decreased with increasing urbanization (i.e. the proportion of built-up and paved areas [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e]).\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026bull; agricultural areas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e+/-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eOn one hand, crops are less suitable habitats for ticks (compared to forests) due to the lack of a litter layer and less suitable microclimatic conditions [\u003cspan citationid=\"CR108\" class=\"CitationRef\"\u003e108\u003c/span\u003e]. On the other hand, agricultural practices can alter host movements, and habitat use, with some mammals being more (or less) abundant in crops compared to mature forests [\u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e109\u003c/span\u003e, \u003cspan citationid=\"CR110\" class=\"CitationRef\"\u003e110\u003c/span\u003e]. Previous studies showed that the densities of \u003cem\u003eI. ricinus\u003c/em\u003e larvae and adults (but not nymphs) in green spaces in Belgium decreased with the percentage of agricultural areas [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026bull; Non-forest vegetated areas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003eNon-forest vegetated areas (e.g. parks and pasture) can be suitable habitats for ticks [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e, \u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e] and are used by some tick hosts [\u003cspan citationid=\"CR109\" class=\"CitationRef\"\u003e109\u003c/span\u003e, \u003cspan citationid=\"CR111\" class=\"CitationRef\"\u003e111\u003c/span\u003e]. In urban green spaces (e.g. parks, natural and amenity green spaces), the density of \u003cem\u003eI. ricinus\u003c/em\u003e nymph was positively associated with the proportion of vegetation areas in the landscape (i.e. urban forests, open green spaces, and green corridors such as wildflower verges and hedgerows [\u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e73\u003c/span\u003e]), while in forests it decreased with the percentage of pastures in the landscape [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026bull; Vegetation areas (forest and non-forest vegetated areas combined)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026bull; forest areas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\" morerows=\"1\" rowspan=\"2\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eGiven that the abundance of some host species increases with the percentage of forest areas in the landscape [\u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e38\u003c/span\u003e] and that \u003cem\u003eI. ricinus\u003c/em\u003e is most abundant in forests than in other habitat types [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e], many studies reported an increase in \u003cem\u003eI. ricinus\u003c/em\u003e abundance or occurrence in forests or green spaces with the proportion of forests or the number of forest patches [\u003cspan additionalcitationids=\"CR37 CR38 CR39\" citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]. The density and occurrence of nymphal \u003cem\u003eI. ricinus\u003c/em\u003e in green spaces were negatively related to the distance to woodland [\u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e73\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\" morerows=\"3\" rowspan=\"4\"\u003e \u003cp\u003eLandscape configuration\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eDensity of forest patch (number.100 ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMin. distance to the forest (m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEffective mesh size (MESH, ha)\u003c/p\u003e \u003cp\u003e\u0026bull; of vegetation areas\u003c/p\u003e \u003cp\u003e\u0026bull; of combined vegetation and agricultural areas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e+\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eFragmentation can alter host availability through changes in the host community composition, host movement, and habitat use. Highly fragmented habitats may hold populations of smaller and/or more mobile tick host species (e.g. rodents and birds), but not of larger animals (e.g. deer) [\u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e13\u003c/span\u003e]. Previous studies in both urban environments [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e] and rural environments [\u003cspan additionalcitationids=\"CR35\" citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e] have demonstrated that connectivity between suitable patches for ticks and their hosts, as well as connectivity to source populations of large propagation hosts (i.e. forest), are important for \u003cem\u003eI. ricinus\u003c/em\u003e persistence.\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEdge density (m. ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003cp\u003e\u0026bull; of landscape\u003c/p\u003e \u003cp\u003e\u0026bull; of vegetation areas\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e+/-\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003eOn one hand, some mammal species can be more abundant at habitat edges, i.e. zones of transition between adjacent ecological systems, than in the habitat interior (e.g. forest edges) [\u003cspan citationid=\"CR112\" class=\"CitationRef\"\u003e112\u003c/span\u003e, \u003cspan citationid=\"CR113\" class=\"CitationRef\"\u003e113\u003c/span\u003e]. On the other hand, increasing edge density implies increased habitat fragmentation, which could be detrimental to the movement of tick host species. The abundance of adult \u003cem\u003eI. ricinus\u003c/em\u003e has been shown to increase with the forest edge density, while that of nymph was not related to the edge density of forest [\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e36\u003c/span\u003e].\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eMeteorological data\u003c/h2\u003e \u003cp\u003eTo account for variation in meteorological conditions during sampling, we computed the average of daily saturation deficit per yard during the five days preceding tick sampling. Indeed, the saturation deficit affects the questing activity of \u003cem\u003eI. ricinus\u003c/em\u003e [\u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e21\u003c/span\u003e, \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e22\u003c/span\u003e, \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e59\u003c/span\u003e] and could thus affect the likelihood of finding ticks. For each yard and each of the five days preceding tick sampling, we computed the daily saturation deficit (mmHg) using the daily average of relative humidity (%), the daily average of temperature (\u0026deg;C), and the formula provided by Randolph and Storey [\u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e59\u003c/span\u003e]. We then averaged those five values. For each yard, we used hourly temperature and relative humidity data from the nearest weather station (mean distance between yards and the weather stations: 7770 m, range: 724\u0026thinsp;\u0026minus;\u0026thinsp;26835 m, n\u0026thinsp;=\u0026thinsp;185) [\u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e49\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eIn this paper, we investigated the relationships between the occurrence of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs and habitat characteristics at transect, yard, and landscape levels. Moreover, we examined how incorporating tick bites in the assessment of tick occurrence could influence the relationships between tick occurrence and environmental variables. We thus also explored the relationships between the occurrence of ticks, and the corrected occurrence of ticks (0 when no ticks were found, and the household reported no tick bites; otherwise, 1) and yard and landscape features. For all the following analyses, we used mixed-effect generalized linear regression models (GLMMs) with a binomial error distribution (logit link).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003e\u003cem\u003eAnalysis of the occurrence of nymphal I. ricinus at the transect-level\u003c/em\u003e\u003c/h2\u003e \u003cp\u003eWe assessed the relationships between transect characteristics and the presence of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs at the transect-level. In this analysis, we only considered yards where nymphs were found, to only include suitable conditions for tick presence and avoid background noise due to unsuitable landscape characteristics. Fixed effects included transect location (edge \u003cem\u003evs\u003c/em\u003e core), grass height (\u0026lt;\u0026thinsp;10 cm, \u0026gt; 10 cm, and other), and shading (shaded \u003cem\u003evs\u003c/em\u003e not shaded). The length of the transect, log-transformed, was included as an offset variable to adjust models for differences in sampling effort among transects. The offset variable makes model adjustments with its regression coefficient being fixed at 1 [\u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e60\u003c/span\u003e]. To account for variations in sampling conditions between years and repeated measures within yards, we included the yard ID nested in the sampling year as a random effect. For this analysis, we only kept data from yards for which information was available for all transects (i.e. length, location, grass height, and shading), i.e. 434 transects (n\u0026thinsp;=\u0026thinsp;48 in 2020, 226 in 2021, and 160 in 2022) across 37 yards (4 in 2020, 19 in 2021, and 14 in 2022).\u003c/p\u003e \u003cp\u003e \u003cem\u003eAnalysis of the occurrence of nymphal I. ricinus and ticks, and the corrected occurrence of ticks at the yard level\u003c/em\u003e \u003c/p\u003e \u003cp\u003eTo assess the relationships between yard and landscape features, and the occurrence of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs, ticks, and the corrected occurrence of ticks, we used occurrence data at the yard level. The total length of transects sampled per yard (log-transformed) was included as an offset variable, to account for variations in sampling effort among yards, and the sampling year as a random effect. Fixed effects included 13 yard features (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e), 11 landscape features (Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e) (see below for the selected buffer size), and the 5-day average of daily saturation deficit to account for temporal variability of meteorological conditions. These analyses rely on data from 185 yards (n\u0026thinsp;=\u0026thinsp;26 in 2020, 59 in 2021, and 100 in 2022).\u003c/p\u003e \u003cp\u003eFor each of the three response variables, we built models that included different combinations of yard and landscape features. To avoid collinearity issues, we did not build candidate models that included variables with an absolute value of the Pearson correlation coefficient greater than 0.6. Model selection was based on the Akaike information criterion (AIC). We selected models within ΔAICc\u0026thinsp;\u0026lt;\u0026thinsp;2, as there is substantial evidence to support them as the best models to explain the observed patterns in the data [\u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e61\u003c/span\u003e]. To gain insight into the respective influence of yard and landscape features, the same model selection procedure was applied using combinations of either yard or landscape features. Model accuracy was evaluated using the AIC, AUC, sensitivity, and specificity. For each best model, the ROC curve was used to compute the AUC and determine the threshold that optimizes both sensitivity and specificity (Youden\u0026rsquo;s Index, [\u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e62\u003c/span\u003e]). A predicted occurrence greater than the cut-off threshold was assigned as tick (corrected or not) or nymphs were present. The sensitivity and specificity were computed using the optimal cut-off.\u003c/p\u003e \u003cp\u003eBefore exploring the relationships between environmental factors and the three response variables, we identified the optimal buffer size at which the relationships between landscape features and the occurrence of ticks were the strongest. Indeed, variables computed with different buffer sizes were highly correlated (Fig. \u003cspan refid=\"MOESM1\" class=\"InternalRef\"\u003eS1\u003c/span\u003e) which can cause multicollinearity. For each of the three buffer sizes, we built candidate models that included as fixed effects different combinations of the nine variables computed at various buffer sizes. We included the total length of transects sampled per yard (log-transformed) as an offset variable and the sampling year as a random effect. The best model for each buffer size was selected based on the AIC and the parsimony principle, as previously described. We then compared the AIC of the three best models and chose the buffer size at which the AIC was the lowest. A buffer size of 500 m was the best fit for tick occurrence (AIC\u0026thinsp;=\u0026thinsp;205.71 compared to AIC\u0026thinsp;=\u0026thinsp;206.71 and AIC\u0026thinsp;=\u0026thinsp;207.50 for buffers of 300 and 400 m, respectively). As a result, we kept landscape features computed within a 500 m buffer.\u003c/p\u003e \u003cp\u003eAll analyses were carried out in R v 4.3.0 [\u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e63\u003c/span\u003e] using the packages g\u003cem\u003elmmTMB\u003c/em\u003e [\u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e64\u003c/span\u003e] and \u003cem\u003eMuMIN\u003c/em\u003e [\u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e65\u003c/span\u003e] for GLMMs and model selection. Collinearity was assessed using the variance inflation factor (VIF) (package \u003cem\u003eperformance\u003c/em\u003e, [\u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e66\u003c/span\u003e]. The DHARMA package [\u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e67\u003c/span\u003e] was used to check for spatial autocorrelation (Moran\u0026rsquo;s I test) and for patterns in the residuals using simulation-based standardized residuals. The package \u003cem\u003ecutpointr\u003c/em\u003e [\u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e68\u003c/span\u003e] was used to find the optimal cutpoint and compute the accuracy metrics (i.e. AUC, sensitivity, and specificity) using these thresholds. The 95% CI of the accuracy metrics was computed using the package \u003cem\u003epROC\u003c/em\u003e [\u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e69\u003c/span\u003e]. Predicted odd ratios and their confidence intervals were obtained with the \u003cem\u003esjPlot\u003c/em\u003e package [\u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e70\u003c/span\u003e].\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003eDescription of sampled yards\u003c/h2\u003e \u003cp\u003eFollowing the classification of a territory according to its degree of urbanization [\u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e71\u003c/span\u003e, \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e72\u003c/span\u003e], 33% of sampled private yards with complete data (n\u0026thinsp;=\u0026thinsp;185) were in densely populated urban areas (cities), 29% in urban areas of intermediate density (towns and suburbs), and 38% in thinly populated rural areas. These yards were almost exclusively private individual yards adjacent to or surrounding to dwelling (96.2%). Others were individual yards away from the dwelling (2.2%), or collective yards (1.6%). Yards differed in their yard and landscape features (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). On average, yards had a vegetated surface of 953 m\u003csup\u003e2\u003c/sup\u003e (SD\u0026thinsp;=\u0026thinsp;1246) and were located 310 m from a forest patch (SD\u0026thinsp;=\u0026thinsp;318). The three most common yard features were the presence of a compost, the presence of a vegetable garden, and the presence of a bird feeder (83%,76%, and 71% of the yards, respectively). Additionally, over half of the yards had a wood/brush pile, a stone wall or pile of stone, or nut-producing trees while having a dog, a fully closed yard, and signs of deer were less common. On average participants mowed their lawn approximately twice a month in spring, summer, and autumn. Quickly, within a 500 m buffer surrounding the yard, the mean percentages of artificial surfaces and non-forest vegetated areas were 28% (SD\u0026thinsp;=\u0026thinsp;17) and 50% (SD\u0026thinsp;=\u0026thinsp;12), respectively, while the mean density of forest patches was 4.0 patches.100 ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e (SD\u0026thinsp;=\u0026thinsp;3.7).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eVariation of the yard and landscape features in private yards (n\u0026thinsp;=\u0026thinsp;185).\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"4\" nameend=\"c7\" namest=\"c4\"\u003e \u003cp\u003eFreq.\u0026nbsp;of occurrence of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs in yards when the feature is\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eContinuous variables\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eMean\u0026thinsp;\u0026plusmn;\u0026thinsp;SD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eRange\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003eBelow average\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003eAbove average\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMowing intensity\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e18.3\u0026thinsp;\u0026plusmn;\u0026thinsp;9.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2\u0026thinsp;\u0026minus;\u0026thinsp;36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e22%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e26%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eVegetated surface of the yard (m\u003csup\u003e2\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e953\u0026thinsp;\u0026plusmn;\u0026thinsp;1246\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e91\u0026thinsp;\u0026minus;\u0026thinsp;8846\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e19%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e35%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNo. of fruit-producing species groups\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e3.4\u0026thinsp;\u0026plusmn;\u0026thinsp;1.8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026thinsp;\u0026minus;\u0026thinsp;6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e13%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e34%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e5-day avg. daily saturation deficit (mmHg)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.7\u0026thinsp;\u0026plusmn;\u0026thinsp;2.0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1.3\u0026thinsp;\u0026minus;\u0026thinsp;11.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e26%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e22%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePercentage of artificial surfaces\u003csup\u003e500\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e28\u0026thinsp;\u0026plusmn;\u0026thinsp;17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2\u0026thinsp;\u0026minus;\u0026thinsp;70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e32%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e12%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePercentage of agricultural area\u003csup\u003e500\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9\u0026thinsp;\u0026plusmn;\u0026thinsp;13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026thinsp;\u0026minus;\u0026thinsp;51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e28%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e16%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePercentage of forest areas\u003csup\u003e500\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10\u0026thinsp;\u0026plusmn;\u0026thinsp;13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026thinsp;\u0026minus;\u0026thinsp;57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e14%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e43%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePercentage of non-forest vegetated areas\u003csup\u003e500\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e50\u0026thinsp;\u0026plusmn;\u0026thinsp;12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e21\u0026thinsp;\u0026minus;\u0026thinsp;80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e27%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e20%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePercentage of vegetation areas\u003csup\u003e50\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e59\u0026thinsp;\u0026plusmn;\u0026thinsp;16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e11\u0026thinsp;\u0026minus;\u0026thinsp;100\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e14%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e33%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEdge density of landscape\u003csup\u003e500\u003c/sup\u003e (m. ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e354\u0026thinsp;\u0026plusmn;\u0026thinsp;118\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e82\u0026thinsp;\u0026minus;\u0026thinsp;652\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e29%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e18%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEdge density of vegetation areas\u003csup\u003e500\u003c/sup\u003e (m.ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e346\u0026thinsp;\u0026plusmn;\u0026thinsp;131\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e65\u0026ndash;651\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e30%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e17%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDensity of forest patch\u003csup\u003e500\u003c/sup\u003e (number. 100 ha\u003csup\u003e\u0026minus;\u0026thinsp;1\u003c/sup\u003e)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e4.0\u0026thinsp;\u0026plusmn;\u0026thinsp;3.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.0\u0026ndash;18.3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e20%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e30%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMESH of vegetation areas\u003csup\u003e500\u003c/sup\u003e (ha)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e9.6\u0026thinsp;\u0026plusmn;\u0026thinsp;10.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.2\u0026ndash;60.6\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e15%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e44%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMESH of combined vegetation and agricultural areas\u003csup\u003e500\u003c/sup\u003e (ha)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e14.6\u0026thinsp;\u0026plusmn;\u0026thinsp;13.7\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.2\u0026thinsp;\u0026minus;\u0026thinsp;74.5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e16%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e35%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eMin. distance to the forest (m)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e310\u0026thinsp;\u0026plusmn;\u0026thinsp;318\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0\u0026thinsp;\u0026minus;\u0026thinsp;1856\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"3\" nameend=\"c6\" namest=\"c4\"\u003e \u003cp\u003e31%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e11%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDichotomous variables\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003ePercentage of yards with the feature (number)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"4\" nameend=\"c7\" namest=\"c4\"\u003e \u003cp\u003eFreq.\u0026nbsp;of occurrence of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs in yards\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003eWith the feature\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003eWithout the feature\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCompost\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e83% (153)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e27%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e9%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eVegetable garden\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e76% (140)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e25%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e20%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eBird feeder\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e71% (132)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e22%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e28%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eNut-producing trees\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e61% (112)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e31%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e12%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eStone wall or pile of stone\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e57% (105)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e22%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e26%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eWood/brush pile\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e55% (102)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e33%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e12%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eUnmanaged long grass area\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e41% (75)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e29%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e20%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eDog\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e22% (41)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e20%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e25%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSigns of deer\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e20% (37)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e57%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e16%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYard closure (fully closed)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c3\" namest=\"c2\"\u003e \u003cp\u003e18% (34)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c5\" namest=\"c4\"\u003e \u003cp\u003e18%\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c7\" namest=\"c6\"\u003e \u003cp\u003e25%\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"7\"\u003eThese variables were used to explore the relationships between environmental factors and the occurrence of nymphal \u003cem\u003eI. ricinus\u003c/em\u003e, ticks, and the corrected occurrence of ticks. Variables were computed within a 500-m (\u003csup\u003e500\u003c/sup\u003e) or a 50-m buffer (\u003csup\u003e50\u003c/sup\u003e) from the edges of the property. MESH: effective mesh size. Vegetation areas are composed of forest areas and non-forest vegetated areas.\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eTick collection and tick bites\u003c/h2\u003e \u003cp\u003eCitizen scientists sampled a total of 2030 transects in 185 yards. As indicated in the protocol, tick sampling took place at the appropriate time of day (i.e. before 11 a.m. or after 4 p.m.) 93% of the time. On average, citizen scientists sampled 11 transects per yard (SD\u0026thinsp;=\u0026thinsp;2.0, range: 4\u0026thinsp;\u0026minus;\u0026thinsp;12), covering an average area of 109 m\u003csup\u003e2\u003c/sup\u003e (SD\u0026thinsp;=\u0026thinsp;22.7, range: 36\u0026thinsp;\u0026minus;\u0026thinsp;200 m\u003csup\u003e2\u003c/sup\u003e). 84% of citizen scientists sampled at least 10 transects. Yards with fewer than ten transects had a mean vegetated surface of 339 m\u003csup\u003e2\u003c/sup\u003e (SD\u0026thinsp;=\u0026thinsp;295), compared to a mean of 1072 m\u003csup\u003e2\u003c/sup\u003e (SD\u0026thinsp;=\u0026thinsp;1323) for yards with at least ten transects. Citizen scientists, collected 501 ticks (112 larvae, 365 nymphs, and 24 adults) belonging to four species: \u003cem\u003eIxodes ricinus\u003c/em\u003e, \u003cem\u003eIxodes frontalis\u003c/em\u003e, \u003cem\u003eDermacentor marginatus\u003c/em\u003e, and \u003cem\u003eDermacentor reticulatus\u003c/em\u003e. \u003cem\u003eIxodes ricinus\u003c/em\u003e was by far the most collected species (96% of the ticks collected) and \u003cem\u003eI. ricinus\u003c/em\u003e nymphs (hereafter \u0026ldquo;\u003cem\u003enymphs\u003c/em\u003e\u0026rdquo;) represented 71% of the collected ticks (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). Ticks occurred in 8.6% of the transects and 32% of sampled yards (Table S2). At least one nymph was found in 24% of yards (Table S2), with an occurrence of 18% in yards in densely populated urban areas, 30% in urban areas of intermediate density, and 24% in thinly populated rural areas. Other species and life stages occurred in less than 6% of the yards, except for \u003cem\u003eI. frontalis\u003c/em\u003e larvae in 2020 (15% of the yards, 4 yards) and \u003cem\u003eI. ricinus\u003c/em\u003e male in 2021 (8% of yards, 5 yards) (Table S2). The average density of nymphs was 1.6 individuals per 100 m\u003csup\u003e2\u003c/sup\u003e (SD\u0026thinsp;=\u0026thinsp;6.3, range: 0\u0026thinsp;\u0026minus;\u0026thinsp;62.9). In yards with nymphs (n\u0026thinsp;=\u0026thinsp;44), the median density was 2.5 nymphs per 100 m\u003csup\u003e2\u003c/sup\u003e (mean: 6.7, SD\u0026thinsp;=\u0026thinsp;11.7, range: 0.8\u0026thinsp;\u0026minus;\u0026thinsp;62.9).\u003c/p\u003e \u003cp\u003eAmong households (n\u0026thinsp;=\u0026thinsp;185), 29% indicated that at least one family member had been (very likely) bitten by ticks in their yard in the last three years. In the last three years, 48% of family members reported being bitten by ticks and 18% reported being bitten by ticks in the yard (n\u0026thinsp;=\u0026thinsp;485, data only available for 2021 and 2022). When corrected by tick bites in the yard, ticks potentially occur in 45% of the yards (35% in 2020, 56% in 2021, and 41% in 2022).\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eFrequency of each tick species, stage, and sex (number collected) per sampling year.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"5\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSpecies\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eStage and sex\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003e2020\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2021\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003e2022\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIxodes ricinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003elarvae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e52% (96)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e2% (4)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5% (4)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIxodes ricinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003enymph\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e41% (75)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e91% (224)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e81% (59)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIxodes ricinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eadult female\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1% (2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1% (3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3% (2)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIxodes ricinus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eadult male\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e1% (2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e3% (7)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3% (2)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIxodes frontalis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003elarvae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4% (8)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0% (0)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0% (0)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eIxodes frontalis\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003enymph\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0% (0)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1% (3)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5% (4)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eDermacentor marginatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eadult female\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0% (0)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0% (0)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1% (1)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eDermacentor reticulatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eadult female\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0% (0)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1% (2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0% (0)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003e\u003cem\u003eDermacentor reticulatus\u003c/em\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eadult male\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0% (0)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e1% (2)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e1% (1)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e100% (183)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e100% (245)\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e100% (73)\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003eTick collection took place in 185 private yards between from May to mid-and July 2020 (n\u0026thinsp;=\u0026thinsp;26), 2021 (n\u0026thinsp;=\u0026thinsp;59), and 2022 (n\u0026thinsp;=\u0026thinsp;100) within a 35 km radius of Nancy, a city located in the northeastern of France.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003e\u003cem\u003eRelationships between transect features and the occurrence of nymphal I. ricinus at the transect-level\u003c/em\u003e\u003c/h2\u003e \u003cp\u003eIn yards with nymphal \u003cem\u003eI. ricinus\u003c/em\u003e and for which information was available for all transects (n\u0026thinsp;=\u0026thinsp;434 transects in 37 yards), nymphs occurred in 27% of the transects (SD\u0026thinsp;=\u0026thinsp;24). In these yards, nymphs occurred in 34% of shaded transects compared to 17% of unshaded transects (Table S3). The likelihood of finding at least one nymph increased by 2.75 (95% CI: 1.52\u0026ndash;4.97) when the transect was shaded by vegetation compared to not shaded transect. Grass height and the location of the transect (edge \u003cem\u003evs\u003c/em\u003e core) were not related to the likelihood of observing nymphs.\u003c/p\u003e \u003cp\u003e \u003cem\u003eRelationships between yard and landscape features, and the occurrence of I. ricinus nymphs, ticks, and the corrected occurrence of ticks\u003c/em\u003e \u003c/p\u003e \u003cp\u003eNine, twelve, and ten models explaining variations in the occurrence of nymphs, ticks, and corrected occurrence of ticks, respectively, were within ΔAIC\u0026thinsp;\u0026lt;\u0026thinsp;2 (Tables S4, S5, S6). Among these models, one model best explained the occurrence of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs (Table S4), while two models best explained the occurrence of ticks and the corrected occurrence of ticks (Tables S5, S6). These models always included three explanatory variables. Overall, the presence of a wood/brush pile and signs of deer in/near the yard were included in all best models (except for a model for signs of deer) and were positively associated with all three occurrence variables (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In yards with a wood/brush pile, the odds of finding ticks or nymphs was 2.37 (95% CI: 1.18\u0026ndash;4.75) to 3.57 (95% CI: 1.45\u0026ndash;8.77) greater compared to yards without this feature. Nymphs were observed in 33% of yards with a wood/brush pile compared to 12% of yards without it (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Similarly, the presence of deer signs in/near the yard increased the odds of finding ticks or nymphs by 2.67 (95% CI: 1.02\u0026ndash;6.97) to 3.33 (95% CI: 1.36\u0026ndash;8.14) compared to yards without signs of deer (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). Nymphs were observed in 16% of the yard without signs of deer, while in contrast, they were found in 57% of the yard with signs of deer nearby (Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003eIn addition to being positively associated with signs of deer and the presence of a wood/brush pile, the odds of finding a nymph also increased by 1.09 (95% CI: 1.04\u0026ndash;1.15) with a one-unit increase in MESH of vegetation areas (Table S4, Fig. S2). Nymphs were observed in 15% of the yard with a MESH of vegetation areas less than 9.6 ha (average) compared to 44% when the MESH of vegetation areas was higher. One of the best models explaining the occurrence of ticks was similar to the best model explaining the occurrence of nymphs as it also included signs of deer, the presence of a wood/brush pile, and the MESH of vegetation areas. The second best model explaining the occurrence of ticks included the percentage of forest areas instead of the MESH of vegetation areas (Table S5, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). This model, which included signs of deer, the presence of a wood/brush pile, and the percentage of forest areas, was identical to one of the best models explaining the corrected occurrence of ticks (Table S6, Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). In these models, the odds of finding ticks or ticks corrected by tick bites increased by 1.05 (95% CI: 1.02\u0026ndash;1.08) and 1.07 (95% CI: 1.04\u0026ndash;1.11) with a one-unit increase in the percentage of forest areas (see Fig. S3 for the occurrence of ticks), respectively. In particular, ticks were observed in 20% of yards with less than 10% of the landscape covered by forest areas (i.e. a below-average percentage), while in contrast, they were found in 54% of the yard with an above-average percentage of forest. Finally, the second best model explaining the corrected occurrence of ticks included the presence of a compost, in addition to the presence of a wood/brush pile, and the percentage of forest areas (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e, Table S6), but its association with the corrected occurrence of ticks was not significant (OR\u0026thinsp;=\u0026thinsp;2.53, 95% CI: 0.98\u0026ndash;6.55; Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eFor the three response variables, the AIC was the lowest for the models including yard and landscape features, intermediate for models with only landscape features, and the highest for models with only yard features. AUC values (0.72\u0026ndash;0.84) were similar irrespective of yard and/or landscape features inclusion. Overall, the sensitivity and specificity of the best models was around 0.67 (range: 0.49\u0026thinsp;\u0026minus;\u0026thinsp;0.89) and 0.78 (range: 0.60\u0026thinsp;\u0026minus;\u0026thinsp;0.88), respectively (Fig. S5). For the occurrence of nymphs, the model with yard and landscape features had a similar specificity and sensibility (~\u0026thinsp;0.8), while these values differed in models with only yard or landscape features. Differences in specificity and sensitivity were also observed for all models explaining the variation in the occurrence of ticks or the corrected occurrence of ticks with yard features (Fig. S5).\u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eIn Europe, there has been limited research on the factors associated with ticks presence or abundance in vegetated areas other than forests (e.g. green spaces including parks [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e, \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e73\u003c/span\u003e]), with even fewer studies focusing on private yards [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e]. Using citizen science, we assessed, for the first time in France, the occurrence of ticks in private yards within and around a metropolitan community, and investigated the yard and landscape features related to tick presence. Our study reveals that ticks are frequent in private yards, irrespective of whether they are located in densely populated urban areas, urban areas of intermediate density, or thinly populated rural areas, with ticks detected in 32% of the sampled yards and potentially present in 45% of yards when considering collected ticks and reported tick bites. We also demonstrated that their occurrence was linked to both yard and landscape features. Consistent with previous studies conducted in yards in Germany [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e] and in urban green spaces in Europe (reviewed in [\u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e74\u003c/span\u003e]), nymph density in yards was low (mean of 1.6 individuals per 100 m\u003csup\u003e2\u003c/sup\u003e). This suggests that tick presence is primarily due to introductions by hosts that acquired ticks elsewhere (sink populations), and could be sporadic. Nonetheless, 29% of households and 18% of family members reported tick bites in their yard over the last three years, aligning with previous studies highlighting the risk of tick bites in yards [\u003cspan additionalcitationids=\"CR42 CR43\" citationid=\"CR41\" class=\"CitationRef\"\u003e41\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e44\u003c/span\u003e]. Although the participating households were not randomly selected across the study area, which can lead to some biases that limit the representativeness of our study, there was no apparent massive participation of citizens who were previously bitten by ticks in our study. Taken together our findings emphasize the importance of informing the public about the risk of tick exposure and tick bites in yards.\u003c/p\u003e \u003cp\u003eTick distribution depends on habitat suitability for ticks and their hosts [\u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e14\u003c/span\u003e, \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e15\u003c/span\u003e]. At the transect level, nymphs were found across all types of transects, regardless of vegetation shading, location, or grass height, indicating their ability to occur in habitats typically deemed less suitable due to less favorable microclimatic conditions. Moreover, in yards with nymphs, the likelihood of finding a nymph was similar in long and short grass areas, as well as in the core or the edge of the yard, whereas it was nearly three times higher in transects beneath trees, hedges, and shrubs compared to transects in open areas without vegetation shading. Compared to open areas, the cover provided by trees and shrubs can buffer temperature extremes and maintain a higher humidity [\u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e23\u003c/span\u003e], thereby enhancing tick survival [\u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e17\u003c/span\u003e, \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e18\u003c/span\u003e]. Moreover, trees and shrubs may provide food resources, shelters, breeding, or resting sites for hosts. The abundance or cover of trees and shrubs is an important predictor of \u003cem\u003eI. ricinus\u003c/em\u003e nymphs density in various environments, including woodlands [\u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e37\u003c/span\u003e, \u003cspan additionalcitationids=\"CR76 CR77\" citationid=\"CR75\" class=\"CitationRef\"\u003e75\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e78\u003c/span\u003e], agricultural areas [\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e], and urban parks and green spaces [\u003cspan additionalcitationids=\"CR81\" citationid=\"CR80\" class=\"CitationRef\"\u003e80\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e82\u003c/span\u003e]. However, contrary to our results, Richter et al. [\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e40\u003c/span\u003e] found no relationship between \u003cem\u003eI. ricinus\u003c/em\u003e occurrence in transects and transect shading in yards, suggesting that the effect of vegetation shading might be variable in yards according to other features (type of vegetation, etc.). The lower shading effectiveness of herb layer vegetation compared to trees and shrubs [\u003cspan citationid=\"CR83\" class=\"CitationRef\"\u003e83\u003c/span\u003e] may explain the lack of a relationship between nymph occurrence and grass height (see also: [\u003cspan citationid=\"CR84\" class=\"CitationRef\"\u003e84\u003c/span\u003e]). However, the influence of the herb layer on ticks may be context-dependent (e.g. depending on yard size and availability of suitable habitats nearby), explaining why some studies found an association between tick density and the herb layer (e.g. [\u003cspan citationid=\"CR80\" class=\"CitationRef\"\u003e80\u003c/span\u003e] in urban parks), while others have not (e.g. [\u003cspan citationid=\"CR79\" class=\"CitationRef\"\u003e79\u003c/span\u003e] in pasture edges, [\u003cspan citationid=\"CR82\" class=\"CitationRef\"\u003e82\u003c/span\u003e, \u003cspan citationid=\"CR85\" class=\"CitationRef\"\u003e85\u003c/span\u003e] in sub-urban and rural parks).\u003c/p\u003e \u003cp\u003eAt the yard level, models for the three occurrence variables (i.e. occurrence of nymphs and ticks, and the corrected occurrence of ticks) were similar, given that most ticks were nymphal \u003cem\u003eI. ricinus\u003c/em\u003e. In all models, the likelihood of finding ticks or nymphs in a yard increased with the presence of a wood/brush pile and signs of deer in/near the yard (Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e). This aligns with previous studies in the US showing a higher occurrence or abundance of three tick species (including \u003cem\u003eI. scapularis)\u003c/em\u003e in residential properties with a log or brush pile [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e, \u003cspan citationid=\"CR86\" class=\"CitationRef\"\u003e86\u003c/span\u003e] (but see: [\u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e87\u003c/span\u003e]). Despite the expectation that wood/brush piles attract birds and small mammals, primary hosts for immature \u003cem\u003eI. ricinus\u003c/em\u003e ticks, studies investigating its influence on tick hosts in yards remain scarce and have shown weak or no relationship with mammal species abundance or richness [\u003cspan additionalcitationids=\"CR89\" citationid=\"CR88\" class=\"CitationRef\"\u003e88\u003c/span\u003e\u0026ndash;\u003cspan citationid=\"CR90\" class=\"CitationRef\"\u003e90\u003c/span\u003e]. Therefore, the causal relationships between the presence of a wood/brush pile and tick occurrence should be further investigated. Cervids are vital for the persistence of \u003cem\u003eI. ricinus\u003c/em\u003e populations, as adult ticks mainly feed and copulate on deer [\u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e8\u003c/span\u003e]. While cervids may contribute to tick presence in some yards, signs of deer more likely indicate yard proximity to forests. Indeed, the likelihood of observing signs of deer decreased with the minimal distance to the forest (OR\u0026thinsp;=\u0026thinsp;0.99, 95% CI: 0.99\u0026ndash;1.0; mean minimal distance to the forest: 121 m [SD\u0026thinsp;=\u0026thinsp;157 m] for yards with signs of deer vs 357 m [SD\u0026thinsp;=\u0026thinsp;331 m] for yards without signs of deer). Mowing intensity, the presence of an unmanaged long grass area, a bird feeder, a vegetable garden, and yard closure were yard features absent from the top models (within ΔAIC\u0026thinsp;\u0026lt;\u0026thinsp;2 from the lowest scoring model). The lack of relationship between the occurrence variables and mowing intensity and the presence of an unmanaged long grass area aligns with our finding at the transect level, where nymph occurrence was not related to grass height. Surprisingly, yard closure was also not related to the occurrence variables (but see: [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e] in the USA). This might be explained by the fact that we relied on citizen\u0026rsquo;s perception and did not accurately estimate fence permeability based on explicit criteria (e.g. fence type, height, and structural integrity). Alternatively, this finding might indicate that ticks are primarily introduced by hosts little or not affected by the most common types of fences in our yards (i.e. hedge and chain link), such as small mammals and birds.\u003c/p\u003e \u003cp\u003eAt the landscape scale, we found evidence that the odds of at least one of the three tick occurrence variables were positively linked to the percentage of forest areas (tick occurrence and corrected occurrence of ticks) and the MESH of vegetation areas (occurrence of nymphs and corrected occurrence of ticks). This finding aligns with previous literature indicating that, in Europe, forests are more suitable habitats for ticks compared to non-forest land-use types [\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e30\u003c/span\u003e]. As previously shown for tick persistence in meadows, forests can also be considered as a source of ticks for yards, while more artificialized or open landscape types (e.g. crops, artificial surfaces, or even yard lawns) act as sinks [\u003cspan citationid=\"CR91\" class=\"CitationRef\"\u003e91\u003c/span\u003e]. The positive association between the MESH of vegetation areas (i.e. a decrease in the fragmentation) and the occurrence variables is unsurprising considering the positive correlation between the MESH of vegetation areas and the percentage of forest areas (r\u0026thinsp;=\u0026thinsp;0.66, Fig. S4). The importance of landscape connectivity for tick populations has already been identified in rural areas in Spain [\u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e34\u003c/span\u003e, \u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e35\u003c/span\u003e], as well as in green spaces in urban areas in Belgium [\u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e32\u003c/span\u003e], and the US [\u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e31\u003c/span\u003e, \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e33\u003c/span\u003e]. Given favorable conditions for tick survival in yards, enhanced connectivity between suitable habitats for ticks, their hosts, and potential source host populations (e.g. forest) could promote tick presence in yards.\u003c/p\u003e \u003cp\u003eCitizen science is a valuable approach for advancing ecological knowledge, yet concerns persist regarding data quality [\u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e46\u003c/span\u003e, \u003cspan citationid=\"CR92\" class=\"CitationRef\"\u003e92\u003c/span\u003e]. We implemented several recommended measures to ensure data quality and reliability [\u003cspan citationid=\"CR93\" class=\"CitationRef\"\u003e93\u003c/span\u003e], including a standardized protocol for tick collection, available in written and video format, as well as a support for tick recognition. Additionally, we asked participants to record easily identifiable yard characteristics. Although we did not quantify it, the participation of citizens in this study likely increased their tick risk awareness and tick-associated literacy, based on their feedbacks. This may in turn increase the acceptance of methods to prevent tick bites (but see: [\u003cspan citationid=\"CR94\" class=\"CitationRef\"\u003e94\u003c/span\u003e]).\u003c/p\u003e \u003cp\u003eOne of the objectives of our project was to determine whether certain features of the yard or landscape which can be modified to reduce the risk of tick occurrence. However, our results should be interpreted with caution regarding public health. For example, tick occurrence was related to the percentage of forest areas in the landscape and the connectivity of vegetated areas. This suggests that initiatives aimed at enhancing landscape connectivity, and/or creating vegetated areas near yards may increase tick exposure risk, but final decisions must be balanced against other benefits, such as an increase in biodiversity or human well-being [\u003cspan citationid=\"CR95\" class=\"CitationRef\"\u003e95\u003c/span\u003e]. Additionally, tick occurrence was linked to vegetation shading, the presence of a wood/brush pile, and signs of deer in/near the yard. However, we caution against providing clear recommendations to yard owners based solely on these results. Indeed, further research is needed to establish causal relationships between these features and tick occurrence, as well as to assess the potential impact of yard management measures on fauna and the associated financial implications for yard owners. Nonetheless, yard owners should be particularly vigilant when using shaded areas (e.g. while trimming hedges or lying down a tree), as nymph occurrence was three times higher than in unshaded areas. In our study the risk of encountering ticks did not appear to be associated with the presence of a stone wall, a vegetable garden, a compost, a bird feeder, a dog, and nut-producing trees, the vegetated surface of the yard, yard closure, mowing intensity, grass height, and the number of fruit-producing trees. Moreover, it was similar in the core and the edge of the yard. Health authorities (e.g. USA and Canada) have recommended various property management measures to control tick populations (e.g. [\u003cspan citationid=\"CR96\" class=\"CitationRef\"\u003e96\u003c/span\u003e]), not supported by the results of this study, but often cited in scientific or popular science publications (e.g. [\u003cspan citationid=\"CR97\" class=\"CitationRef\"\u003e97\u003c/span\u003e, \u003cspan citationid=\"CR98\" class=\"CitationRef\"\u003e98\u003c/span\u003e]). As previously stated [\u003cspan citationid=\"CR87\" class=\"CitationRef\"\u003e87\u003c/span\u003e], we recommend that public health officials provide clear information on the scientific support and uncertainties surrounding yard management measures, as most of them seem to lack scientific support. European health authorities should also be careful when using results from North American studies, as yard features (especially the size), tick species, and host ecology, might differ greatly from European yards. Further studies are necessary to identify features associated with tick occurrence in yards and to implement management strategies adapted to the private yards to mitigate the risk of tick exposure.\u003c/p\u003e"},{"header":"Conclusions","content":"\u003cp\u003eTicks are frequent in private yards across an urbanization gradient in our study area in northeastern France and their presence is shaped by both yard and landscape features. Despite official recommendations on some yard measures expected to control ticks abundance or occurrence, we found no evidence linking \u003cem\u003eI. ricinus\u003c/em\u003e occurrence in yards with the mowing intensity and other relevant yard features. However, we found that vegetation shading, the presence of a wood/brush pile, and signs of deer in/near the yard were related to the occurrence of ticks. At the landscape level, tick occurrence were positively related to the percentage of forest areas, and the connectivity of vegetated areas. We suggest further studies are needed to implement adapted management strategies against tick risk in yards. To better understand the threat posed by ticks in yards and enhance management measures effectiveness, future research should investigate the prevalence of pathogens in the ticks present in yards, as well as identify the hosts responsible for tick presence in yards.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eEthics approval and consent to participate\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eStudy participation was voluntary and all participants provided signed informed consent before entering the study. This study was declared to the\u0026nbsp;National Commission on Informatics and Liberty\u003cem\u003e\u0026nbsp;(Commission Nationale de l'Informatique et des Libertés\u003c/em\u003e [CNIL]), the independent French administrative regulatory body responsible for data protection, which ensures that all information is kept confidential and anonymous, in compliance with the Declaration of Helsinki.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and materials\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe data used in this paper includes personal residential information. Therefore, to protect study participant privacy the data are not publicly available. The anonymized data collected are, however, available from the corresponding author upon reasonable request.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no competing interests.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis study was funded by the French Agency for Food, Environmental and Occupational Health Safety (ANSES), grant no.: ANSES-21-EST-050, as well as the French National Research Agency (ANR) as part of the “Investissements d'Avenir” program (ANR-11-LABX-0002-01, Lab of Excellence ARBRE), the project “Des Hommes et Des Arbres” (DHDA) supported by the French government through its program “Plan d'Investissement d'Avenir - Territoire d'Innovation”, and the Metropole du Grand Nancy (grant no. 7434).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthors’ contributions\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eProject conceptualization (L.B, J.D, P.FK, F.B, C.G, I.C). Design of the work (L.B, J.D, I.C, S.C); data acquisition (Citizen scientists, A.M, L.B, J.D, C.C, and S.C); data modelling (A.M with valuable support from J.D and L.B); interpretation of results (A.M, L.B, J.D) and manuscript preparation (A.M, J.D, and L.B). All authors read, revised, and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAcknowledgements\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe would like to acknowledge the contribution of staff from the Centre Permanent d’Initiatives pour l’Environnement Nancy Champenoux (particularly Marcel Klein), the Nancy Laboratory for Rabies and Wildlife of The French Agency for Food, Environmental and Occupational Health and Safety (ANSES), and the Tous Chercheurs Laboratory (French National Research Institute for Agriculture, Food and Environment, INRAE), as well as the Jean-Marie Pelt botanical garden (Villers-lès-Nancy), for their invaluable assistance in distributing the tick collection kit. Thanks are extended to all colleagues at the ANSES Nancy Laboratory for Rabies and Wildlife, especially Olivier Musquar and Jean-Marc Boucher, as well as Sandrine Warion and Lucero Petrowski from INRAE Tous Chercheurs laboratory for their help in preparing the tick collection kit. Special thanks to Gerald Umhang (ANSES Nancy Laboratory for Rabies and Wildlife) for his assistance in tick sampling and valuable feedback on the sampling protocol. We are grateful to the departments of communication and Institutional relations of INRAE and ANSES for their support in recruiting citizen scientists through media campaigns. Last, but not least, we sincerely thank all of the citizen scientists whose participation made this project a true success.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eFuente JDL. Overview: Ticks as vectors of pathogens that cause disease in humans and animals. Front Biosci. 2008;13:6938\u0026ndash;46. \u003c/li\u003e\n\u003cli\u003eJongejan F, Uilenberg G. The global importance of ticks. Parasitology. 2004;129:S3\u0026ndash;14. \u003c/li\u003e\n\u003cli\u003eMac S, Da Silva SR, Sander B. The economic burden of Lyme disease and the cost-effectiveness of Lyme disease interventions: a scoping review. PLOS ONE. 2019;14:1\u0026ndash;17. \u003c/li\u003e\n\u003cli\u003eRosenberg R, Lindsey NP, Fischer M, Gregory CJ, Hinckley AF, Mead PS, et al. \u003cem\u003eVital signs\u003c/em\u003e : Trends in reported vectorborne disease cases \u0026mdash; United States and territories, 2004\u0026ndash;2016. MMWR Morb Mortal Wkly Rep. 2018;67:496\u0026ndash;501. \u003c/li\u003e\n\u003cli\u003eSteere AC, Strle F, Wormser GP, Hu LT, Branda JA, Hovius JWR, et al. Lyme borreliosis. Nat Rev Dis Primer. 2016;2:1\u0026ndash;18. \u003c/li\u003e\n\u003cli\u003eJohnson N. Ticks: biology, ecology, and diseases. London, United Kingdom: Elsevier, Academic Press, an Imprint of Elsevier; 2023. \u003c/li\u003e\n\u003cli\u003eCastro MB, Wright SA. Vertebrate hosts of \u003cem\u003eIxodes pacificus\u003c/em\u003e (Acari: Ixodidae) in California. J Vector Ecol. 2007;32:140\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eHofmeester TR, Coipan EC, Van Wieren SE, Prins HHT, Takken W, Sprong H. Few vertebrate species dominate the \u003cem\u003eBorrelia burgdorferi\u003c/em\u003e s.l. life cycle. Environ Res Lett. 2016;11:1\u0026ndash;16. \u003c/li\u003e\n\u003cli\u003eKeirans JE, Hutcheson HJ, Durden LA, Klompen JSH. \u003cem\u003eIxodes (Ixodes) scapularis (Acari: Ixodidae)\u003c/em\u003e: redescription of all active stages, distribution, hosts, geographical variation, and medical and veterinary importance. J Med Entomol. 1996;33:297\u0026ndash;318. \u003c/li\u003e\n\u003cli\u003eWang S-S, Liu J-Y, Wang B-Y, Wang W-J, Cui X-M, Jiang J-F, et al. Geographical distribution of \u003cem\u003eIxodes persulcatus\u003c/em\u003e and associated pathogens: analysis of integrated data from a China field survey and global published data. One Health. 2023;16:1\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eUspensky I. The Taiga Tick \u003cem\u003eIxodes Persulcatus\u003c/em\u003e(Acari: Ixodidae), the main vector of \u003cem\u003eBorrelia Burgdorferi Sensu Lato\u003c/em\u003e in Eurasia. In: Capinera JL, editor. Encycl Entomol. 2nd ed. 2016. \u003c/li\u003e\n\u003cli\u003eFischhoff IR, Keesing F, Ostfeld RS. Risk factors for bites and diseases associated with black-legged ticks: a meta-analysis. Am J Epidemiol. 2019;188:1742\u0026ndash;50. \u003c/li\u003e\n\u003cli\u003eDiuk-Wasser MA, VanAcker MC, Fernandez MP. Impact of land use changes and habitat fragmentation on the eco-epidemiology of tick-borne diseases. J Med Entomol. 2021;58:1546\u0026ndash;64. \u003c/li\u003e\n\u003cli\u003ePf\u0026auml;ffle M, Littwin N, Muders SV, Petney TN. The ecology of tick-borne diseases. Int J Parasitol. 2013;43:1059\u0026ndash;77. \u003c/li\u003e\n\u003cli\u003eEstrada-Pe\u0026ntilde;a A, De La Fuente J. The ecology of ticks and epidemiology of tick-borne viral diseases. Antiviral Res. 2014;108:104\u0026ndash;28. \u003c/li\u003e\n\u003cli\u003eEisen RJ, Eisen L, Ogden NH, Beard CB. Linkages of weather and climate with \u003cem\u003eIxodes scapularis\u003c/em\u003e and \u003cem\u003eIxodes pacificus\u003c/em\u003e (Acari: Ixodidae), enzootic transmission of \u003cem\u003eBorrelia burgdorferi\u003c/em\u003e, and Lyme disease in North America. J Med Entomol. 2016;53:250\u0026ndash;61. \u003c/li\u003e\n\u003cli\u003eGrigoryeva LA. Influence of air humidity on the survival rate, lifetime, and development of \u003cem\u003eIxodes ricinus\u003c/em\u003e(L., 1758) and \u003cem\u003eIxodes persulcatus\u003c/em\u003e Schulze, 1930 (Acari: Ixodidae). Syst Appl Acarol. 2022;27:2241\u0026ndash;8. \u003c/li\u003e\n\u003cli\u003eHerrmann C, Gern L. Survival of \u003cem\u003eIxodes ricinus\u003c/em\u003e (Acari: Ixodidae) under challenging conditions of temperature and humidity is influenced by \u003cem\u003eBorrelia burgdorferi\u003c/em\u003e sensu lato infection. J Med Entomol. 2010;47:1196\u0026ndash;204. \u003c/li\u003e\n\u003cli\u003eLeal B, Zamora E, Fuentes A, Thomas DB, Dearth RK. Questing by tick larvae (Acari: Ixodidae): a review of the influences that affect off-host survival. Reddy GVP, editor. Ann Entomol Soc Am. 2020;113:425\u0026ndash;38. \u003c/li\u003e\n\u003cli\u003eOgden NH, Lindsay LR, Beauchamp G, Charron D, Maarouf A, O\u0026rsquo;Callaghan CJ, et al. Investigation of relationships between temperature and developmental rates of tick \u003cem\u003eIxodes scapularis\u003c/em\u003e (Acari: Ixodidae) in the laboratory and field. J Med Entomol. 2004;41:622\u0026ndash;33. \u003c/li\u003e\n\u003cli\u003ePerret JL, Rais O, Gern L. Influence of climate on the proportion of \u003cem\u003eIxodes ricinus\u003c/em\u003e nymphs and adults questing in a tick population. J Med Entomol. 2004;41:361\u0026ndash;5. \u003c/li\u003e\n\u003cli\u003ePerret J-L, Guerin PM, Diehl PA, Vlimant M, Gern L. Darkness induces mobility, and saturation deficit limits questing duration, in the tick \u003cem\u003eIxodes ricinus\u003c/em\u003e. J Exp Biol. 2003;206:1809\u0026ndash;15. \u003c/li\u003e\n\u003cli\u003eDe Frenne P, Lenoir J, Luoto M, Scheffers BR, Zellweger F, Aalto J, et al. Forest microclimates and climate change: importance, drivers and future research agenda. Glob Change Biol. 2021;27:2279\u0026ndash;97. \u003c/li\u003e\n\u003cli\u003eMilling CR, Rachlow JL, Olsoy PJ, Chappell MA, Johnson TR, Forbey JS, et al. Habitat structure modifies microclimate: an approach for mapping fine‐scale thermal refuge. Mahon A, editor. Methods Ecol Evol. 2018;9:1648\u0026ndash;57. \u003c/li\u003e\n\u003cli\u003eMathisson DC, Kross SM, Palmer MI, Diuk-Wasser MA. Effect of vegetation on the abundance of tick vectors in the northeastern United States: a review of the literature. Ginsberg H, editor. J Med Entomol. 2021;58:2030\u0026ndash;7. \u003c/li\u003e\n\u003cli\u003eMedlock JM, Shuttleworth H, Copley V, Hansford KM, Leach S. Woodland biodiversity management as a tool for reducing human exposure to \u003cem\u003eIxodes ricinus\u003c/em\u003e ticks: a preliminary study in an English woodland. J Vector Ecol. 2012;37:307\u0026ndash;15. \u003c/li\u003e\n\u003cli\u003eSchulze TL, Jordan RA. Influence of meso- and microscale habitat structure on focal distribution of sympatric \u003cem\u003eIxodes scapularis\u003c/em\u003e and \u003cem\u003eAmblyomma americanum\u003c/em\u003e (Acari: Ixodidae). J Med Entomol. 2005;42:285\u0026ndash;94. \u003c/li\u003e\n\u003cli\u003eEcke F, L\u0026ouml;fgren O, S\u0026ouml;rlin D. Population dynamics of small mammals in relation to forest age and structural habitat factors in northern Sweden. J Appl Ecol. 2002;39:781\u0026ndash;92. \u003c/li\u003e\n\u003cli\u003eFerger SW, Schleuning M, Hemp A, Howell KM, B\u0026ouml;hning‐Gaese K. Food resources and vegetation structure mediate climatic effects on species richness of birds. Glob Ecol Biogeogr. 2014;23:541\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eBourdin A, Dokhelar T, Bord S, Van Halder I, Stemmelen A, Scherer-Lorenzen M, et al. Forests harbor more ticks than other habitats: a meta-analysis. For Ecol Manag. 2023;541:1\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eGregory N, Fernandez MP, Diuk-Wasser M. Risk of tick-borne pathogen spillover into urban yards in New York City. Parasit Vectors. 2022;15:1\u0026ndash;14. \u003c/li\u003e\n\u003cli\u003eHeylen D, Lasters R, Adriaensen F, Fonville M, Sprong H, Matthysen E. Ticks and tick-borne diseases in the city: role of landscape connectivity and green space characteristics in a metropolitan area. Sci Total Environ. 2019;670:941\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eVanAcker MC, Little EAH, Molaei G, Bajwa WI, Diuk-Wasser MA. Enhancement of risk for Lyme disease by landscape connectivity, New York, New York, USA. Emerg Infect Dis. 2019;25:1136\u0026ndash;43. \u003c/li\u003e\n\u003cli\u003eEstrada-Pe\u0026ntilde;a A. The relationships between habitat topology, critical scales of connectivity and tick abundance \u003cem\u003eIxodes ricinus\u003c/em\u003e in a heterogeneous landscape in northern Spain. Ecography. 2003;26:661\u0026ndash;71. \u003c/li\u003e\n\u003cli\u003eEstrada-Pe\u0026ntilde;a A. Effects of habitat suitability and landscape patterns on tick (Acarina) metapopulation processes. Landsc Ecol. 2005;20:529\u0026ndash;41. \u003c/li\u003e\n\u003cli\u003eLi S, Heyman P, Cochez C, Simons L, Vanwambeke SO. A multi-level analysis of the relationship between environmental factors and questing \u003cem\u003eIxodes ricinus\u003c/em\u003e dynamics in Belgium. Parasit Vectors. 2012;5:1\u0026ndash;11. \u003c/li\u003e\n\u003cli\u003eEhrmann S, Liira J, G\u0026auml;rtner S, Hansen K, Brunet J, Cousins SAO, et al. Environmental drivers of \u003cem\u003eIxodes ricinus\u003c/em\u003e abundance in forest fragments of rural European landscapes. BMC Ecol. 2017;17:1\u0026ndash;14. \u003c/li\u003e\n\u003cli\u003ePerez G, Bastian S, Agoulon A, Bouju A, Durand A, Faille F, et al. Effect of landscape features on the relationship between \u003cem\u003eIxodes ricinus\u003c/em\u003e ticks and their small mammal hosts. Parasit Vectors. 2016;9:1\u0026ndash;18. \u003c/li\u003e\n\u003cli\u003eTalbot B, Slatculescu A, Thickstun CR, Koffi JK, Leighton PA, McKay R, et al. Landscape determinants of density of blacklegged ticks, vectors of Lyme disease, at the northern edge of their distribution in Canada. Sci Rep. 2019;9:1\u0026ndash;12. \u003c/li\u003e\n\u003cli\u003eRichter D, Schneider A-K, Schibalski A, Dahlkamp A, Schr\u0026ouml;der B. Features in and around residential gardens affecting the presence and abundance of questing \u003cem\u003eIxodes ricinus\u003c/em\u003e ticks. Infect Ecol Epidemiol. 2023;13:1\u0026ndash;12. \u003c/li\u003e\n\u003cli\u003eHassler L, Durand J, Frey-Klett P. \u003cem\u003eCiTIQUE-Tracker\u003c/em\u003e [Internet]. INRAE, SK8; 2024 [cited 2024 May 17]. Available from: https://ci-tique-tracker.sk8.inrae.fr/\u003c/li\u003e\n\u003cli\u003eMulder S, Van Vliet AJH, Bron WA, Gassner F, Takken W. High risk of tick bites in Dutch gardens. Vector-Borne Zoonotic Dis. 2013;13:865\u0026ndash;71. \u003c/li\u003e\n\u003cli\u003eRaguet S, Couturier E. \u0026Eacute;tude ALSA(CE)TIQUE 2014-2015. Principaux r\u0026eacute;sultats descriptifs. Saint-Maurice: Sant\u0026eacute; Publique France; 2016 p. 1\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eSciensano. TIQUESNET 2020, surveillance des morsures de tiques en Belgique. \u0026Eacute;pid\u0026eacute;miologie des maladies infectieuses. Bruxelles; 2021 p. 1\u0026ndash;12. \u003c/li\u003e\n\u003cli\u003eSeifert VA, Wilson S, Toivonen S, Clarke B, Prunuske A. Community partnership designed to promote Lyme disease prevention and engagement in citizen science. J Microbiol Biol Educ. 2016;17:63\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eEisen L, Eisen RJ. Benefits and drawbacks of citizen science to complement traditional data gathering approaches for medically important hard ticks (Acari: Ixodidae) in the United States. J Med Entomol. 2021;58:1\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eHeyman P, Cochez C, Hofhuis A, Van Der Giessen J, Sprong H, Porter SR, et al. A clear and present danger: tick-borne diseases in Europe. Expert Rev Anti Infect Ther. 2010;8:33\u0026ndash;50. \u003c/li\u003e\n\u003cli\u003eINSEE. Growth and structure of the population in 2020 inter-communality metropolis of M\u0026eacute;tropole du Grand Nancy (245400676) [Internet]. 2023 Jun. Available from: https://www.insee.fr/fr/statistiques/1405599?geo=EPCI-245400676\u003c/li\u003e\n\u003cli\u003eM\u0026eacute;t\u0026eacute;o France. Fiche climatologique; indicatif : 54526001 [Internet]. 2023 [cited 2024 May 6]. Available from: https://donneespubliques.meteofrance.fr/FichesClim/FICHECLIM_54526001.pdf\u003c/li\u003e\n\u003cli\u003eDataGrand Est. Occupation du sol \u0026agrave; grande \u0026eacute;chelle en r\u0026eacute;gion Grand Est [Internet]. 2022 [cited 2024 May 6]. Available from: https://www.datagrandest.fr/portail/fr/projets/occupation-du-sol\u003c/li\u003e\n\u003cli\u003eWongnak P, Bord S, Jacquot M, Agoulon A, Beugnet F, Bournez L, et al. Meteorological and climatic variables predict the phenology of \u003cem\u003eIxodes ricinus\u003c/em\u003e nymph activity in France, accounting for habitat heterogeneity. Sci Rep. 2022;12:1\u0026ndash;16. \u003c/li\u003e\n\u003cli\u003ePerez G, Bournez L, Boulanger N, Fite J, Livoreil B, McCoy KD, et al. The distribution, phenology, host range and pathogen prevalence of \u003cem\u003eIxodes ricinus\u003c/em\u003e in France: a systematic map and narrative review. Peer Community J. 2023;3:1\u0026ndash;40. \u003c/li\u003e\n\u003cli\u003eEstrada-Pe\u0026ntilde;a A, Mihalca AD, Petney T, Domşa C, Gragera Artal J, editors. Ticks of Europe and North Africa: a guide to species identification. Cham: Springer Switzerland; 2017. \u003c/li\u003e\n\u003cli\u003eP\u0026eacute;rez-Eid C. Les tiques : identification, biologie, importance medicale et veterinaire. Lavoisier, Editions m\u0026eacute;dicales internationales; 2007. \u003c/li\u003e\n\u003cli\u003eDurand J, Bournez L, Marchand J, Schmid C, Carravieri I, Palin B, et al. Are orienteers protected enough against tick bites? Estimating human exposure to tick bites through a participative science survey during an orienteering competition. Int J Environ Res Public Health. 2021;18:1\u0026ndash;19. \u003c/li\u003e\n\u003cli\u003eWilhelmsson P, Lindblom P, Fryland L, Nyman D, Jaenson TG, Forsberg P, et al. \u003cem\u003eIxodes ricinus\u003c/em\u003e ticks removed from humans in Northern Europe: seasonal pattern of infestation, attachment sites and duration of feeding. Parasit Vectors. 2013;6:1\u0026ndash;11. \u003c/li\u003e\n\u003cli\u003eETALAB. Cadastre data of France [Internet]. 2022 [cited 2023 Aug 2]. Available from: https://cadastre.data.gouv.fr/data/etalab-cadastre/2020-10-01/shp/departements/\u003c/li\u003e\n\u003cli\u003eJaeger JAG. Landscape division, splitting index, and effective mesh size: new measures of landscape fragmentation. Landsc Ecol. 2000;15:115\u0026ndash;30. \u003c/li\u003e\n\u003cli\u003eRandolph SE, Storey K. Impact of microclimate on immature tick-rodent host interactions (Acari: Ixodidae): implications for parasite transmission. J Med Entomol. 1999;36:741\u0026ndash;8. \u003c/li\u003e\n\u003cli\u003eHilbe JM. Negative binomial regression. 2nd ed. Cambridge, UK ; New York: Cambridge University Press; 2011. \u003c/li\u003e\n\u003cli\u003eBurnham KP, Anderson DR, editors. Model selection and multimodel inference. New York, NY: Springer New York; 2004. \u003c/li\u003e\n\u003cli\u003eYouden WJ. Index for rating diagnostic tests. Cancer. 1950;3:32\u0026ndash;5. \u003c/li\u003e\n\u003cli\u003eR Core Team. R: a language and environment for statistical computing [Internet]. Vienna, Austria: R Foundation for Statistical Computing; 2023. Available from: https://www.R-project.org/\u003c/li\u003e\n\u003cli\u003eBrooks ME, Kristensen K, van Benthem KJ, Magnusson A, Berg CW, Nielsen A, et al. glmmTMB balances speed and flexibility among packages for zero-inflated generalized linear mixed modeling. 2017; Available from: http://hdl.handle.net/20.500.11850/242692\u003c/li\u003e\n\u003cli\u003eBartoń K. MuMIn: Multi-Model Inference. 2023; Available from: https://CRAN.R-project.org/package=MuMIn\u003c/li\u003e\n\u003cli\u003eL\u0026uuml;decke D, Ben-Shachar MS, Patil I, Waggoner P, Makowski D. An R package for assessment, comparison and testing of statistical models. J Open Source Softw. 2021;6:1\u0026ndash;8. \u003c/li\u003e\n\u003cli\u003eHartig F. DHARMa: Residual Diagnostics for Hierarchical (Multi-Level / Mixed) Regression Models. 2022; Available from: https://CRAN.R-project.org/package=DHARMa\u003c/li\u003e\n\u003cli\u003eThiele C, Hirschfeld G. cutpointr: improved estimation and validation of optimal cutpoints in \u003cem\u003eR\u003c/em\u003e. J Stat Softw. 2021;98:1\u0026ndash;27. \u003c/li\u003e\n\u003cli\u003eRobin X, Turck N, Hainard A, Tiberti N, Lisacek F, Sanchez J-C, et al. pROC: an open-source package for R and S+ to analyze and compare ROC curves. BMC Bioinformatics. 2011;12:1\u0026ndash;8. \u003c/li\u003e\n\u003cli\u003eL\u0026uuml;decke D. sjPlot: data visualization for statistics in social science. 2023; Available from: https://CRAN.R-project.org/package=sjPlot\u003c/li\u003e\n\u003cli\u003eDijkstra L, Poelman H. A harmonised definition of cities and rural areas: the new degree of urbanisation. 2021;European Commission Working Paper WP 01/2014. \u003c/li\u003e\n\u003cli\u003eINSEE. La grille communale de densit\u0026eacute; \u0026agrave; 7 niveaux [Internet]. 2023 [cited 2024 May 6]. Available from: https://www.insee.fr/fr/information/6439600\u003c/li\u003e\n\u003cli\u003eHansford KM, Gillingham EL, Vaux AGC, Cull B, McGinley L, Catton M, et al. Impact of green space connectivity on urban tick presence, density and \u003cem\u003eBorrelia\u003c/em\u003e infected ticks in different habitats and seasons in three cities in southern England. Ticks Tick-Borne Dis. 2023;14:1\u0026ndash;11. \u003c/li\u003e\n\u003cli\u003eHansford KM, Wheeler BW, Tschirren B, Medlock JM. Questing \u003cem\u003eIxodes ricinus\u003c/em\u003e ticks and \u003cem\u003eBorrelia\u003c/em\u003e spp. in urban green space across Europe: A review. Zoonoses Public Health. 2022;69:153\u0026ndash;66. \u003c/li\u003e\n\u003cli\u003eBourdin A, Bord S, Durand J, Galon C, Moutailler S, Scherer-Lorenzen M, et al. Forest diversity reduces the prevalence of pathogens transmitted by the tick \u003cem\u003eIxodes ricinus\u003c/em\u003e. Front Ecol Evol. 2022;10:1\u0026ndash;13. \u003c/li\u003e\n\u003cli\u003eKiewra D, Stefańska-Krzaczek E, Szymanowski M, Szczepańska A. Local-scale spatio-temporal distribution of questing \u003cem\u003eIxodes ricinus\u003c/em\u003e L. (Acari: Ixodidae)-A case study from a riparian urban forest in Wrocław, SW Poland. Ticks Tick-Borne Dis. 2017;8:362\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eTack W, Madder M, Baeten L, Vanhellemont M, Gruwez R, Verheyen K. Local habitat and landscape affect \u003cem\u003eIxodes ricinus\u003c/em\u003e tick abundances in forests on poor, sandy soils. For Ecol Manag. 2012;265:30\u0026ndash;6. \u003c/li\u003e\n\u003cli\u003eTack W, Madder M, Baeten L, Verheyen K. The abundance of \u003cem\u003eIxodes ricinus\u003c/em\u003e ticks depends on tree species composition and shrub cover. Parasitology. 2012;139:1273\u0026ndash;81. \u003c/li\u003e\n\u003cli\u003eBoyard C, Barnouin J, Bord S, Gasqui P, Vourc\u0026rsquo;h G. Reproducibility of local environmental factors for the abundance of questing \u003cem\u003eIxodes ricinus\u003c/em\u003e nymphs on pastures. Ticks Tick-Borne Dis. 2011;2:104\u0026ndash;10. \u003c/li\u003e\n\u003cli\u003eHansford KM, McGinley L, Wilkinson S, Gillingham EL, Cull B, Gandy S, et al. \u003cem\u003eIxodes ricinus\u003c/em\u003e and \u003cem\u003eBorrelia burgdorferi\u003c/em\u003e sensu lato in the Royal Parks of London, UK. Exp Appl Acarol. 2021;84:593\u0026ndash;606. \u003c/li\u003e\n\u003cli\u003eMathews-Martin L, Nam\u0026egrave;che M, Vourc\u0026rsquo;h G, Gasser S, Lebert I, Poux V, et al. Questing tick abundance in urban and peri-urban parks in the French city of Lyon. Parasit Vectors. 2020;13:1\u0026ndash;9. \u003c/li\u003e\n\u003cli\u003eVan Gestel M, Verheyen K, Matthysen E, Heylen D. Danger on the track? Tick densities near recreation infrastructures in forests. Urban For Urban Green. 2021;59:1\u0026ndash;7. \u003c/li\u003e\n\u003cli\u003eSodoudi S, Zhang H, Chi X, M\u0026uuml;ller F, Li H. The influence of spatial configuration of green areas on microclimate and thermal comfort. Urban For Urban Green. 2018;34:85\u0026ndash;96. \u003c/li\u003e\n\u003cli\u003eVan Gestel M, Matthysen E, Heylen D, Verheyen K. Survival in the understorey: testing direct and indirect effects of microclimatological changes on \u003cem\u003eIxodes ricinus\u003c/em\u003e. Ticks Tick-Borne Dis. 2022;13:1\u0026ndash;7. \u003c/li\u003e\n\u003cli\u003eLee X, Maxson G-A, Paskewitz S. Single mowing event does not reduce abundance of \u003cem\u003eIxodes scapularis\u003c/em\u003e (Acari: Ixodidae) and \u003cem\u003eDermacentor variabilis\u003c/em\u003e (Acari: Ixodidae) on recreational hiking trails. J Med Entomol. 2023;60:228\u0026ndash;34. \u003c/li\u003e\n\u003cli\u003eHahn MB, Bjork JKH, Neitzel DF, Dorr FM, Whitemarsh T, Boegler KA, et al. Evaluating acarological risk for exposure to \u003cem\u003eIxodes scapularis\u003c/em\u003e and \u003cem\u003eIxodes scapularis\u003c/em\u003e-borne pathogens in recreational and residential settings in Washington County, Minnesota. Ticks Tick-Borne Dis. 2018;9:340\u0026ndash;8. \u003c/li\u003e\n\u003cli\u003eFischhoff IR, Keesing F, Pendleton J, DePietro D, Teator M, Duerr STK, et al. Assessing effectiveness of recommended residential yard management measures against ticks. J Med Entomol. 2019;56:1420\u0026ndash;7. \u003c/li\u003e\n\u003cli\u003eGrade AM, Warren PS, Lerman SB. Managing yards for mammals: mammal species richness peaks in the suburbs. Landsc Urban Plan. 2022;220:1\u0026ndash;11. \u003c/li\u003e\n\u003cli\u003eHansen CP, Parsons AW, Kays R, Millspaugh JJ. Does Use of backyard resources explain the abundance of urban wildlife? Front Ecol Evol. 2020;8:1\u0026ndash;12. \u003c/li\u003e\n\u003cli\u003eJohansson EP, DeGregorio BA. The effects of landscape and yard features on mammal diversity in residential yards within Northwest Arkansas, USA. Urban Ecosyst. 2024;27:275\u0026ndash;87. \u003c/li\u003e\n\u003cli\u003eHoch T, Monnet Y, Agoulon A. Influence of host migration between woodland and pasture on the population dynamics of the tick \u003cem\u003eIxodes ricinus\u003c/em\u003e: a modelling approach. Ecol Model. 2010;221:1798\u0026ndash;806. \u003c/li\u003e\n\u003cli\u003eFraisl D, Hager G, Bedessem B, Gold M, Hsing P-Y, Danielsen F, et al. Citizen science in environmental and ecological sciences. Nat Rev Methods Primer. 2022;2:1\u0026ndash;20. \u003c/li\u003e\n\u003cli\u003eKosmala M, Wiggins A, Swanson A, Simmons B. Assessing data quality in citizen science. Front Ecol Environ. 2016;14:551\u0026ndash;60. \u003c/li\u003e\n\u003cli\u003ePotes L, Bouchard C, Rocheleau J-P, Richard L, Leighton P, Pelletier J, et al. Evaluation of a community-based \u003cem\u003eOne Health\u003c/em\u003e intervention to reduce the risk of Lyme disease in a high-incidence municipality. CABI One Health. 2023;2:1\u0026ndash;12. \u003c/li\u003e\n\u003cli\u003eEcosystems and human well-being: wetlands and water synthesis: a report of the Millennium Ecosystem Assessment. Washington, DC: World Resources Institute; 2005. \u003c/li\u003e\n\u003cli\u003eStafford KC. Tick management handbook; an integrated guide for homeowners, pest control operators, and public health officials for the prevention of tick-associated disease. New Haven, CT: The Connecticut Agricultural Experiment Station; 2007. \u003c/li\u003e\n\u003cli\u003eČern\u0026yacute; J, Lynn G, Hrnkov\u0026aacute; J, Golovchenko M, Rudenko N, Grubhoffer L. Management options for \u003cem\u003eIxodes ricinus\u003c/em\u003e-associated pathogens: a review of prevention strategies. Int J Environ Res Public Health. 2020;17:1\u0026ndash;18. \u003c/li\u003e\n\u003cli\u003eHansen J. How to kill and prevent ticks in your lawn [Internet]. [cited 2024 May 6]. Available from: https://www.gardentech.com/blog/pest-id-and-prevention/protecting-your-lawn-and-garden-against-ticks\u003c/li\u003e\n\u003cli\u003eReed JH, Bonter DN. Supplementing non-target taxa: bird feeding alters the local distribution of mammals. Ecol Appl. 2018;28:761\u0026ndash;70. \u003c/li\u003e\n\u003cli\u003eClotfelter ED, Pedersen AB, Cranford JA, Ram N, Snajdr EA, Nolan V, et al. Acorn mast drives long-term dynamics of rodent and songbird populations. Oecologia. 2007;154:493\u0026ndash;503. \u003c/li\u003e\n\u003cli\u003eOstfeld RS, Levi T, Keesing F, Oggenfuss K, Canham CD. Tick‐borne disease risk in a forest food web. Ecology. 2018;99:1562\u0026ndash;73. \u003c/li\u003e\n\u003cli\u003eSchnurr JL, Ostfeld RS, Canham CD. Direct and indirect effects of masting on rodent populations and tree seed survival. Oikos. 2002;96:402\u0026ndash;10. \u003c/li\u003e\n\u003cli\u003eMaupin GO, Fish D, Zultowsky J, Campos EG, Piesman J. Landscape ecology of Lyme disease in a residential area of Westchester County, New York. Am J Epidemiol. 1991;133:1105\u0026ndash;13. \u003c/li\u003e\n\u003cli\u003eKays R, Parsons AW. Mammals in and around suburban yards, and the attraction of chicken coops. Urban Ecosyst. 2014;17:691\u0026ndash;705. \u003c/li\u003e\n\u003cli\u003eVercauteren KC, Vandeelen TR, Lavelle MJ, Hall WH. Assessment of abilities of white‐tailed deer to jump fences. J Wildl Manag. 2010;74:1378\u0026ndash;81. \u003c/li\u003e\n\u003cli\u003eHofmeester TR, Sprong H, Jansen PA, Prins HHT, van Wieren SE. Deer presence rather than abundance determines the population density of the sheep tick, \u003cem\u003eIxodes ricinus\u003c/em\u003e, in Dutch forests. Parasit Vectors. 2017;10:1\u0026ndash;8. \u003c/li\u003e\n\u003cli\u003eJennett AL, Smith FD, Wall R. Tick infestation risk for dogs in a peri-urban park. Parasit Vectors. 2013;6:1\u0026ndash;10. \u003c/li\u003e\n\u003cli\u003eMedlock JM, Vaux AGC, Hansford KM, Pietzsch ME, Gillingham EL. Ticks in the ecotone: the impact of agri‐environment field margins on the presence and intensity of \u003cem\u003eIxodes ricinus\u003c/em\u003e ticks (Acari: Ixodidae) in farmland in southern England. Med Vet Entomol. 2020;34:175\u0026ndash;83. \u003c/li\u003e\n\u003cli\u003eGehring TM, Swihart RK. Body size, niche breadth, and ecologically scaled responses to habitat fragmentation: mammalian predators in an agricultural landscape. Biol Conserv. 2003;109:283\u0026ndash;95. \u003c/li\u003e\n\u003cli\u003ePanzacchi M, Linnell JDC, Melis C, Odden M, Odden J, Gorini L, et al. Effect of land-use on small mammal abundance and diversity in a forest\u0026ndash;farmland mosaic landscape in south-eastern Norway. For Ecol Manag. 2010;259:1536\u0026ndash;45. \u003c/li\u003e\n\u003cli\u003eBaker PJ, Harris S. Urban mammals: what does the future hold? An analysis of the factors affecting patterns of use of residential gardens in Great Britain. Mammal Rev. 2007;37:297\u0026ndash;315. \u003c/li\u003e\n\u003cli\u003eAnderson CS, Cady AB, Meikle DB. Effects of vegetation structure and edge habitat on the density and distribution of white-footed mice (\u003cem\u003ePeromyscus leucopus\u003c/em\u003e) in small and large forest patches. Can J Zool. 2003;81:897\u0026ndash;904. \u003c/li\u003e\n\u003cli\u003eRies L, Fletcher RJ, Battin J, Sisk TD. Ecological responses to habitat edges: mechanisms, models, and variability explained. Annu Rev Ecol Evol Syst. 2004;35:491\u0026ndash;522. \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":"international-journal-of-health-geographics","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ijhg","sideBox":"Learn more about [International Journal of Health Geographics](http://ij-healthgeographics.biomedcentral.com/)","snPcode":"12942","submissionUrl":"https://submission.nature.com/new-submission/12942/3","title":"International Journal of Health Geographics","twitterHandle":"@IJHGeo","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Ixodes ricinus, tick, occurrence, yard, landscape, citizen science, tick-borne disease prevention","lastPublishedDoi":"10.21203/rs.3.rs-4453852/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-4453852/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003e\u003cstrong\u003eBackground: \u003c/strong\u003eLyme borreliosis is the most frequent zoonotic disease in the northern hemisphere and is transmitted by ticks of the genus \u003cem\u003eIxodes\u003c/em\u003e. Although many people are bitten by ticks in private yards, our understanding of the factors associated with their presence in these areas remains limited. To address this gap, we used a citizen science approach to identify the local and landscape features associated with tick presence in yards.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMethods: \u003c/strong\u003eThis study was conducted near Nancy, a city in northeastern France, from 2020 to 2022. Citizen scientists collected ticks in their yard on a single event (n = 185) and measured 13 yard features. Additionally, we computed 11 features related to the landscape composition and spatial configuration surrounding these yards. Using mixed-effect generalized linear models, we determined the yard and landscape features associated with the presence of ticks and nymphal \u003cem\u003eIxodes ricinus \u003c/em\u003e(hereafter nymphs), the life stage, and species that mostly bite humans.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eResults:\u003c/strong\u003e Despite a low density, ticks were found in 32% of the yards, including yards in urbanized areas. At the transect level, the likelihood of finding a nymph was nearly three times higher in transects shaded by vegetation compared to those in open areas, with no relationship between nymph occurrence and transect location or grass height. At the yard level, the occurrence of ticks and nymphs was related to both yard and landscape characteristics. Nymph and tick occurrence were more than twice as high in yards with signs of deer and a wood/brush pile compared to those without these characteristics, and increased with the connectivity of vegetation areas and the percentage of forest areas in the landscape.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConclusions:\u003c/strong\u003e Our study reveals that private yards across an urbanization gradient are locations of tick exposure with tick presence linked to both yard and landscape factors. These findings emphasize the importance of public awareness regarding tick exposure in yards and provide crucial insights for future public health prevention campaigns.\u003c/p\u003e","manuscriptTitle":"Understanding Ixodes ricinus occurrence in private yards: influence of yard and landscape features","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2024-06-03 11:06:01","doi":"10.21203/rs.3.rs-4453852/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2024-07-05T21:05:54+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-06-26T09:39:30+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2024-06-04T11:30:37+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"7439464960514451169875394324074767626","date":"2024-05-29T07:30:37+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"131537913021862932139496575215298767998","date":"2024-05-28T16:05:56+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2024-05-28T14:33:24+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2024-05-22T01:59:34+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2024-05-22T01:59:34+00:00","index":"","fulltext":""},{"type":"submitted","content":"International Journal of Health Geographics","date":"2024-05-21T09:47:42+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"international-journal-of-health-geographics","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"ijhg","sideBox":"Learn more about [International Journal of Health Geographics](http://ij-healthgeographics.biomedcentral.com/)","snPcode":"12942","submissionUrl":"https://submission.nature.com/new-submission/12942/3","title":"International Journal of Health Geographics","twitterHandle":"@IJHGeo","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"BMC/SO AJ","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"80cc916d-71ba-4c6f-aad1-3d9d93d40fca","owner":[],"postedDate":"June 3rd, 2024","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2024-10-14T16:00:58+00:00","versionOfRecord":{"articleIdentity":"rs-4453852","link":"https://doi.org/10.1186/s12942-024-00380-9","journal":{"identity":"international-journal-of-health-geographics","isVorOnly":false,"title":"International Journal of Health Geographics"},"publishedOn":"2024-10-10 15:57:23","publishedOnDateReadable":"October 10th, 2024"},"versionCreatedAt":"2024-06-03 11:06:01","video":"","vorDoi":"10.1186/s12942-024-00380-9","vorDoiUrl":"https://doi.org/10.1186/s12942-024-00380-9","workflowStages":[]},"version":"v1","identity":"rs-4453852","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-4453852","identity":"rs-4453852","version":["v1"]},"buildId":"qtupq5eGEP_6zYnWcrvyt","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}