Unpunctual in diversity: The effect of stand species richness on spring phenology of deciduous tree stands varies among species and years.

Climatic drivers alone do not adequately explain the regional variation in budburst timing in deciduous forests across Europe. Stand-level factors, such as tree species richness, might affect budburst timing by creating different microclimates under the same site macroclimate. We assessed different phases of the spring phenology (start, midpoint, end, and overall duration of the budburst period) of four important European tree species (Betula pendula, Fagus sylvatica, Quercus robur and Tilia cordata) in monocultures and four-species mixture stands of a common garden tree biodiversity experiment in Belgium (FORBIO) in 2021 and 2022. Microclimatic differences between the stands in terms of bud chilling, temperature forcing, and soil temperature were considerable, with four-species mixtures being generally colder than monocultures in spring, but not in winter. In the colder spring of 2021, at the stand level, the end of the budburst period was advanced, and its overall duration shortened, in the four-species mixtures. At species level, this response was significant for F. sylvatica. In the warmer spring of 2022, advances in spring phenology in four-species stands were observed again in F. sylvatica and, less markedly, in B. pendula but without a general response at the stand level. Q. robur showed specific patterns with delayed budburst start in 2021 in the four-species mixtures and very short budburst duration for all stands in 2022. Phenological differences between monocultures and four-species mixtures were linked to microclimatic differences in light availability rather than in temperature as even comparatively colder microclimates showed an advanced phenology. Compared to weather conditions, tree species richness had a lower impact on budburst timing, but this impact can be of importance for key species like F. sylvatica and colder springs. These results indicate that forest biodiversity can affect budburst phenology, with wider implications, especially for forest- and land surface models.

Body-size shifts in aquatic and terrestrial urban communities.

Body size is intrinsically linked to metabolic rate and life-history traits, and is a crucial determinant of food webs and community dynamics1,2. The increased temperatures associated with the urban-heat-island effect result in increased metabolic costs and are expected to drive shifts to smaller body sizes 3 . Urban environments are, however, also characterized by substantial habitat fragmentation 4 , which favours mobile species. Here, using a replicated, spatially nested sampling design across ten animal taxonomic groups, we show that urban communities generally consist of smaller species. In addition, although we show urban warming for three habitat types and associated reduced community-weighted mean body sizes for four taxa, three taxa display a shift to larger species along the urbanization gradients. Our results show that the general trend towards smaller-sized species is overruled by filtering for larger species when there is positive covariation between size and dispersal, a process that can mitigate the low connectivity of ecological resources in urban settings 5 . We thus demonstrate that the urban-heat-island effect and urban habitat fragmentation are associated with contrasting community-level shifts in body size that critically depend on the association between body size and dispersal. Because body size determines the structure and dynamics of ecological networks 1 , such shifts may affect urban ecosystem function.

Heterogeneous selection on exploration behavior within and among West European populations of a passerine bird.

Heterogeneous selection is often proposed as a key mechanism maintaining repeatable behavioral variation ("animal personality") in wild populations. Previous studies largely focused on temporal variation in selection within single populations. The relative importance of spatial versus temporal variation remains unexplored, despite these processes having distinct effects on local adaptation. Using data from >3,500 great tits (Parus major) and 35 nest box plots situated within five West-European populations monitored over 4 to 18 y, we show that selection on exploration behavior varies primarily spatially, across populations, and study plots within populations. Exploration was, simultaneously, selectively neutral in the average population and year. These findings imply that spatial variation in selection may represent a primary mechanism maintaining animal personalities, likely promoting the evolution of local adaptation, phenotype-dependent dispersal, and nonrandom settlement. Selection also varied within populations among years, which may counteract local adaptation. Our study underlines the importance of combining multiple spatiotemporal scales in the study of behavioral adaptation.

Connecting the data landscape of long-term ecological studies: The SPI-Birds data hub.

The integration and synthesis of the data in different areas of science is drastically slowed and hindered by a lack of standards and networking programmes. Long-term studies of individually marked animals are not an exception. These studies are especially important as instrumental for understanding evolutionary and ecological processes in the wild. Furthermore, their number and global distribution provides a unique opportunity to assess the generality of patterns and to address broad-scale global issues (e.g. climate change). To solve data integration issues and enable a new scale of ecological and evolutionary research based on long-term studies of birds, we have created the SPI-Birds Network and Database (www.spibirds.org)-a large-scale initiative that connects data from, and researchers working on, studies of wild populations of individually recognizable (usually ringed) birds. Within year and a half since the establishment, SPI-Birds has recruited over 120 members, and currently hosts data on almost 1.5 million individual birds collected in 80 populations over 2,000 cumulative years, and counting. SPI-Birds acts as a data hub and a catalogue of studied populations. It prevents data loss, secures easy data finding, use and integration and thus facilitates collaboration and synthesis. We provide community-derived data and meta-data standards and improve data integrity guided by the principles of Findable, Accessible, Interoperable and Reusable (FAIR), and aligned with the existing metadata languages (e.g. ecological meta-data language). The encouraging community involvement stems from SPI-Bird's decentralized approach: research groups retain full control over data use and their way of data management, while SPI-Birds creates tailored pipelines to convert each unique data format into a standard format. We outline the lessons learned, so that other communities (e.g. those working on other taxa) can adapt our successful model. Creating community-specific hubs (such as ours, COMADRE for animal demography, etc.) will aid much-needed large-scale ecological data integration.

The effects of personality on survival and trappability in a wild mouse during a population cycle.

The pace-of-life syndrome (POLS) theory provides an evolutionary explanation for the existence of consistent among-individual variation in behaviour, or animal personality. Herein, individuals with a fast lifestyle are considered to be bolder and should take more risks resulting in a lower life expectancy compared to shyer individuals with a slower lifestyle. However, this assumption depends on the levels of intra-specific competition that the individuals experience which has rarely been tested in species that experience large changes in competition on a very short time scale. We used the multimammate mice (Mastomys natalensis) as a model system to study the POLS assumption by investigating the effects of two personality traits (exploration and stress-sensitivity) on survival, maturation (a proxy for reproductive investment) and recapture probability during one population cycle (Nindividuals = 201). Such a cycle consists of two phases in which the levels of intra-specific competition vary drastically. We found that only one personality trait, namely stress-sensitivity, had a negative effect on both survival and recapture probability but none of them affected maturation. This suggests that less stress-sensitive individuals take more risks in the wild and have a higher survival probability compared to high stress-sensitive individuals. However, the effect of personality on survival was only present during the population decrease phase, when the levels of intra-specific competition are high due to a scarcity of food. This suggests that seasonal changes in competition might be important in the evolution and maintenance of animal personalities in species whose population dynamics have a clear seasonal component.

Habitat fragmentation shapes natal dispersal and sociality in an Afrotropical cooperative breeder.

It remains poorly understood how effects of anthropogenic activity, such as large-scale habitat fragmentation, impact sociality in animals. In cooperatively breeding species, groups are mostly formed through delayed offspring dispersal, and habitat fragmentation can affect this process in two opposite directions. Increased habitat isolation may increase dispersal costs, promoting delayed dispersal. Alternatively, reduced patch size and quality may decrease benefits of philopatry, promoting dispersal. Here, we test both predictions in a cooperatively breeding bird (placid greenbul, Phyllastrephus placidus) from an Afrotropical cloud forest archipelago. Males born in fragmented forest dispersed about 1 year earlier than those born in continuous forest. Contrary to females, males also started to reproduce earlier and mostly settled within their natal patch. Females only rarely delayed their dispersal for more than 1 year, both in fragmented and continuous forests. Our results suggest that early male dispersal and reproduction is jointly driven by a decrease in the value of the natal territory and an increase in local breeding opportunities in fragmented forest. While plasticity in dispersal strategies of cooperative breeders in response to anthropogenic change is believed to optimize reproduction-survival trade-offs, to what extent it shapes the ability of species to respond to rapid environmental change remains to be studied.

Diet contributes to urban-induced alterations in gut microbiota: experimental evidence from a wild passerine.

Urban sprawl increasingly affects the ecology of natural populations, including host-microbiota interactions, with observed differences in the gut microbiota between urban and rural hosts. While different mechanisms could explain this pattern, dietary uptake constitutes a likely candidate. To assess the contribution of diet in explaining urban-rural variation in gut microbiota, we performed an aviary experiment in which urban and rural house sparrows were fed with mimics of urban or rural diets. Before the experiment, rural sparrows hosted more diverse gut communities, with a higher relative abundance of Enterococcaceae and Staphylococcaceae and lower abundance of genes involved in xenobiotic degradation and lipid metabolism than their urban counterparts. The experimental diets significantly altered gut microbiota α- and ß-diversity and taxonomic composition, with the strongest shifts occurring in individuals exposed to contrasting diets. Overall, diet-induced shifts resembled initial differences between free-ranging urban and rural hosts. Furthermore, rural diet had a positive impact on urban host body mass but only in hosts with the highest initial gut diversity. Overall, our results indicate that diet constitutes an important factor contributing to differences in gut microbiota along the urbanization gradient and provide new insights on possible fitness consequences of a reduced gut diversity in urban settings.

Urbanization drives cross-taxon declines in abundance and diversity at multiple spatial scales.

The increasing urbanization process is hypothesized to drastically alter (semi-)natural environments with a concomitant major decline in species abundance and diversity. Yet, studies on this effect of urbanization, and the spatial scale at which it acts, are at present inconclusive due to the large heterogeneity in taxonomic groups and spatial scales at which this relationship has been investigated among studies. Comprehensive studies analysing this relationship across multiple animal groups and at multiple spatial scales are rare, hampering the assessment of how biodiversity generally responds to urbanization. We studied aquatic (cladocerans), limno-terrestrial (bdelloid rotifers) and terrestrial (butterflies, ground beetles, ground- and web spiders, macro-moths, orthopterans and snails) invertebrate groups using a hierarchical spatial design, wherein three local-scale (200 m × 200 m) urbanization levels were repeatedly sampled across three landscape-scale (3 km × 3 km) urbanization levels. We tested for local and landscape urbanization effects on abundance and species richness of each group, whereby total richness was partitioned into the average richness of local communities and the richness due to variation among local communities. Abundances of the terrestrial active dispersers declined in response to local urbanization, with reductions up to 85% for butterflies, while passive dispersers did not show any clear trend. Species richness also declined with increasing levels of urbanization, but responses were highly heterogeneous among the different groups with respect to the richness component and the spatial scale at which urbanization impacts richness. Depending on the group, species richness declined due to biotic homogenization and/or local species loss. This resulted in an overall decrease in total richness across groups in urban areas. These results provide strong support to the general negative impact of urbanization on abundance and species richness within habitat patches and highlight the importance of considering multiple spatial scales and taxa to assess the impacts of urbanization on biodiversity.

Getting under the birds' skin: tissue tropism of Borrelia burgdorferi s.l. in naturally and experimentally infected avian hosts.

Wild birds are frequently exposed to the zoonotic tick-borne bacteria Borrelia burgdorferi sensu lato (s.l.), and some bird species act as reservoirs for some Borrelia genospecies. Studying the tropism of Borrelia in the host, how it is sequestered in different organs, and whether it is maintained in circulation and/or in the host's skin is important to understand pathogenicity, infectivity to vector ticks and reservoir competency.We evaluated tissue dissemination of Borrelia in blackbirds (Turdus merula) and great tits (Parus major), naturally and experimentally infected with Borrelia genospecies from enzootic foci. We collected both minimally invasive biological samples (feathers, skin biopsies and blood) and skin, joint, brain and visceral tissues from necropsied birds. Infectiousness of the host was evaluated through xenodiagnoses and infection rates in fed and moulted ticks. Skin biopsies were the most reliable method for assessing avian hosts' Borrelia infectiousness, which was supported by the agreement of infection status results obtained from the analysis of chin and lore skin samples from necropsied birds and of their xenodiagnostic ticks, including a significant correlation between the estimated concentration of Borrelia genome copies in the skin and the Borrelia infection rate in the xenodiagnostic ticks. This confirms a dermatropism of Borrelia garinii, B. valaisiana and B. turdi in its avian hosts. However, time elapsed from exposure to Borrelia and interaction between host species and Borrelia genospecies may affect the reliability of skin biopsies. The blood was not useful to assess infectiousness of birds, even during the period of expected maximum spirochetaemia. From the tissues sampled (foot joint, liver, spleen, heart, kidney, gut and brain), Borrelia was detected only in the gut, which could be related with infection mode, genospecies competition, genospecies-specific seasonality and/or excretion processes.

Inter-individual variability in spring phenology of temperate deciduous trees depends on species, tree size and previous year autumn phenology.

We explored the inter-individual variability in bud-burst and its potential drivers, in homogeneous mature stands of temperate deciduous trees. Phenological observations of leaves and wood formation were performed weekly from summer 2017 to summer 2018 for pedunculate oak, European beech and silver birch in Belgium. The variability of bud-burst was correlated to previous' year autumn phenology (i.e. the onset of leaf senescence and the cessation of wood formation) and tree size but with important differences among species. In fact, variability of bud-burst was primarily related to onset of leaf senescence, cessation of wood formation and tree height for oak, beech and birch, respectively. The inter-individual variability of onset of leaf senescence was not related to the tree characteristics considered and was much larger than the inter-individual variability in bud-burst. Multi-species multivariate models could explain up to 66% of the bud-burst variability. These findings represent an important advance in our fundamental understanding and modelling of phenology and tree functioning of deciduous tree species.

Accounting for interspecific competition and age structure in demographic analyses of density dependence improves predictions of fluctuations in population size.

Understanding species coexistence has long been a major goal of ecology. Coexistence theory for two competing species posits that intraspecific density dependence should be stronger than interspecific density dependence. Great tits and blue tits are two bird species that compete for food resources and nesting cavities. On the basis of long-term monitoring of these two competing species at sites across Europe, combining observational and manipulative approaches, we show that the strength of density regulation is similar for both species, and that individuals have contrasting abilities to compete depending on their age. For great tits, density regulation is driven mainly by intraspecific competition. In contrast, for blue tits, interspecific competition contributes as much as intraspecific competition, consistent with asymmetric competition between the two species. In addition, including age-specific effects of intra- and interspecific competition in density-dependence models improves predictions of fluctuations in population size by up to three times.

Lyme neuroborreliosis and bird populations in northern Europe.

Many vector-borne diseases are transmitted through complex pathogen-vector-host networks, which makes it challenging to identify the role of specific host groups in disease emergence. Lyme borreliosis in humans is now the most common vector-borne zoonosis in the Northern Hemisphere. The disease is caused by multiple genospecies of Borrelia burgdorferi sensu lato bacteria transmitted by ixodid (hard) ticks, and the major host groups transmit Borrelia genospecies with different pathogenicity, causing variable clinical symptoms in humans. The health impact of a given host group is a function of the number of ticks it infects as well as the pathogenicity of the genospecies it carries. Borrelia afzelii, with mainly small mammals as reservoirs, is the most common pathogen causing Lyme borreliosis, and it is often responsible for the largest proportion of infected host-seeking tick nymphs in Europe. The bird-borne Borrelia garinii, though less prevalent in nymphal ticks, is more likely to cause Lyme neuroborreliosis, but whether B. garinii causes disseminated disease more frequently has not been documented. Based on extensive data of annual disease incidence across Norway from 1995 to 2017, we show here that 69% of disseminated Lyme borreliosis cases were neuroborreliosis, which is three times higher than predicted from the infection prevalence of B. garinii in host-seeking ticks (21%). The population estimate of migratory birds, mainly of thrushes, explained part of the annual variation in cases of neuroborreliosis, with a one-year time lag. We highlight the important role of the genospecies' pathogenicity and the host associations for understanding the epidemiology of disseminated Lyme borreliosis.

Diversifying forest communities may change Lyme disease risk: extra dimension to the dilution effect in Europe.

Lyme disease is caused by bacteria of the Borrelia burgdorferi genospecies complex and transmitted by Ixodid ticks. In North America only one pathogenic genospecies occurs, in Europe there are several. According to the dilution effect hypothesis (DEH), formulated in North America, nymphal infection prevalence (NIP) decreases with increasing host diversity since host species differ in transmission potential. We analysed Borrelia infection in nymphs from 94 forest stands in Belgium, which are part of a diversification gradient with a supposedly related increasing host diversity: from pine stands without to oak stands with a shrub layer. We expected changing tree species and forest structure to increase host diversity and decrease NIP. In contrast with the DEH, NIP did not differ between different forest types. Genospecies diversity however, and presumably also host diversity, was higher in oak than in pine stands. Infected nymphs tended to harbour Borrelia afzelii infection more often in pine stands while Borrelia garinii and Borrelia burgdorferi ss. infection appeared to be more prevalent in oak stands. This has important health consequences, since the latter two cause more severe disease manifestations. We show that the DEH must be nuanced for Europe and should consider the response of multiple pathogenic genospecies.

Testing for effects of climate change on competitive relationships and coexistence between two bird species.

Climate change is expected to have profound ecological effects, yet shifts in competitive abilities among species are rarely studied in this context. Blue tits (Cyanistes caeruleus) and great tits (Parus major) compete for food and roosting sites, yet coexist across much of their range. Climate change might thus change the competitive relationships and coexistence between these two species. Analysing four of the highest-quality, long-term datasets available on these species across Europe, we extend the textbook example of coexistence between competing species to include the dynamic effects of long-term climate variation. Using threshold time-series statistical modelling, we demonstrate that long-term climate variation affects species demography through different influences on density-dependent and density-independent processes. The competitive interaction between blue tits and great tits has shifted in one of the studied sites, creating conditions that alter the relative equilibrium densities between the two species, potentially disrupting long-term coexistence. Our analyses show that long-term climate change can, but does not always, generate local differences in the equilibrium conditions of spatially structured species assemblages. We demonstrate how long-term data can be used to better understand whether (and how), for instance, climate change might change the relationships between coexisting species. However, the studied populations are rather robust against competitive exclusion.

Ancestral origins and invasion pathways in a globally invasive bird correlate with climate and influences from bird trade.

Invasive species present a major threat to global biodiversity. Understanding genetic patterns and evolutionary processes that reinforce successful establishment is paramount for elucidating mechanisms underlying biological invasions. Among birds, the ring-necked parakeet (Psittacula krameri) is one of the most successful invasive species, established in over 35 countries. However, little is known about the evolutionary genetic origins of this species and what population genetic signatures tell us about patterns of invasion. We reveal the ancestral origins of populations across the invasive range and explore the potential influence of climate and propagule pressure from the pet trade on observed genetic patterns. Ring-necked parakeet samples representing the ancestral native range (n = 96) were collected from museum specimens, and modern samples from the invasive range (n = 855) were gathered from across Europe, Mauritius and Seychelles, and sequenced for two mitochondrial DNA markers comprising 868 bp of cytochrome b and control region, and genotyped at 10 microsatellite loci. Invasive populations comprise birds that originate predominantly from Pakistan and northern areas of India. Haplotypes associated with more northerly distribution limits in the ancestral native range were more prevalent in invasive populations in Europe, and the predominance of Asian haplotypes in Europe is consistent with the higher number of Asian birds transported by the pet trade outside the native range. Successful establishment of invasive species is likely to be underpinned by a combination of environmental and anthropogenic influences.

Songbirds as general transmitters but selective amplifiers of Borrelia burgdorferi sensu lato genotypes in Ixodes rinicus ticks.

We investigated to what extent a European songbird (Parus major) selectively transmits and amplifies Borrelia burgdorferi s.l. bacteria. Borrelia-naïve birds were recurrently exposed to Ixodes ricinus nymphs carrying a community of more than 34 5S-23S genotypes belonging to five genospecies (Borrelia garinii, Borrelia valaisiana, Borrelia afzelii, B. burgdorferi s.s. and Borrelia spielmanii). Fed ticks were screened for Borrelia after moulting. We found evidence for co-feeding transmission of avian and possibly also mammalian genotypes. Throughout the course of infestations, the infection rate of B. garinii and B. valaisiana increased, indicating successful amplification and transmission, while the infection rate for B. afzelii, B. burgdorferi s.s and B. spielmanii tended to decrease. Within the B. garinii and B. valaisiana genotype communities, certain genotypes were transmitted more than others. Moreover, birds were able to host mixed infections of B. garinii and B. valaisiana, as well as mixed infections of genotypes of the same genospecies. We experimentally show that resident songbirds transmit a broad range of Borrelia genotypes, but selectively amplify certain genotypes, and that one bird can transmit simultaneously several genotypes. Our results highlight the need to explicitly consider the association between genotypes and hosts, which may offer opportunities to point out which hosts are most responsible for the Borrelia presence in questing ticks.

Are the specialized bird ticks, Ixodes arboricola and I. frontalis, competent vectors for Borrelia burgdorferi sensu lato?

Our study tested whether two European bird-specialized ticks, Ixodes arboricola and I. frontalis, can act as vectors in the transmission cycles of Borrelia burgdorferi s.l. The ticks have contrasting ecologies but share songbird hosts (such as the great tit, Parus major) with the generalist I. ricinus which may therefore act as a bridging vector. In the first phase of the experiment, we obtained Borrelia-infected ornithophilic nymphs by exposing larvae to great tits that had previously been exposed to I. ricinus nymphs carrying a community of genospecies (Borrelia garinii, valaisiana, afzelii, burgdorferi s.s., spielmanii). Skin samples showed that birds selectively amplified B. garinii and B. valaisiana. The spirochetes were transmitted to the ornithophilic ticks and survived moulting, leading to infection rates of 16% and 27% in nymphs of I. arboricola and I. frontalis respectively. In the second phase, pathogen-free great tits were exposed to the Borrelia-infected ornithophilic nymphs. None of these ticks were able to infect the birds, as indicated by the tissue samples. Analysis of xenodiagnostic I. ricinus larvae found no evidence for co-feeding or systemic transmission of B. burgdorferi s.l. These outcomes do not support the occurrence of enzootic cycles of Borrelia burgdorferi s.l. involving songbirds and their specialized ornithophilic ticks.

Seasonal- and sex-specific correlations between dispersal and exploratory behaviour in the great tit.

While the importance of personality in explaining individual dispersal strategies is increasingly recognized, limited information is still available on how patterns of personality-dependent dispersal may develop, hampering our understanding of the ecological significance of behavioural dispersal syndromes. Here, we examine the relative importance of personality at different stages of dispersal in the great tit, by analysing the sex-specific relationship between exploratory behaviour (EB; quantified in a novel environment) and dispersal distances in different seasons over the course of the first year of life (summer, autumn, winter, and until the first breeding attempt). In females, we found that EB was an important predictor of dispersal distances in summer and autumn, but only a weak to moderate correlation remained for females captured in winter or for natal dispersal distances based on first breeding records. We obtained a contrasting pattern at the population level, whereby male (but not female) immigrants captured in summer and autumn had higher EB scores than locally born birds, while this was not the case in birds captured in winter and those recruited as breeders into the population. In addition to providing further evidence for the existence of a behaviour dispersal syndrome in birds, our results show that correlations between EB and dispersal appear strongest at the early stages of the dispersal process, rather than being developed gradually. These findings show the importance of analysing the effect of phenotypic attributes on dispersal across different stages of the dispersal phenomenon and in each sex separately.

Xenodiagnosis in the wild: A methodology to investigate infectiousness for tick-borne bacteria in a songbird reservoir.

A crucial factor to predict the persistence and spread of infections in natural systems is the capacity of reservoir hosts to maintain the infection and transmit it to others. This is known to greatly vary within and between species and through time, although the latter part of the variation is often less well understood in the wild. Borrelia garinii is one of the causal agents of Lyme disease in humans and is transmitted among avian hosts by the hard tick Ixodes ricinus. Great tits are known to be a reservoir in Europe for B. garinii. For tick-borne pathogens like B. garinii, infectiousness or host-to-vector transmission can be measured using xenodiagnosis where pathogen-free vectors are fed on a host, and the blood-fed vectors are subsequently tested for the pathogen. Here we describe and evaluate a methodology to quantify infectiousness for tick-borne pathogens in individual wild great tits (Parus major), involving captures and recaptures of targeted individuals. The methodology can potentially be applied to other species where recapturing is sufficiently guaranteed. We successfully recaptured most of the infested great tits two to three days after initial infestation (i.e. just before ticks have fully fed) with sufficient numbers of I. ricinus larval ticks, which were subsequently screened for B. garinii using a newly developed B. garinii-specific real-time PCR assay. Higher larval tick numbers were recovered from birds during the breeding seasons than during the winter months. Our novel B. garinii-qPCR performed well, and greatly reduced the amount of Sanger sequencing needed. Preliminary results suggest both seasonal and individual variation in infectiousness; heterogeneity that needs to be unravelled to further understand the contribution of resident birds to the epidemiology of B. garinii.

Recreational hazard: Vegetation and host habitat use correlate with changes in tick-borne disease hazard at infrastructure within forest stands.

Studies on density and pathogen prevalence of Ixodes ricinus indicate that vegetation and local host community drive much of their variation between green spaces. Contrarily, micro-geographic variation is understudied, although its understanding could reduce disease risk. We studied the density of infectious nymphal Ixodes sp. ("DIN", proxy for disease hazard), density of questing nymphs ("DON") and nymphal infection prevalence ("NIP") near recreational forest infrastructure. Drag sampling within forest stands and at adjacent benches and trails was combined with vegetation surveys, camera trapping hosts and pathogen screening of ticks. We analysed Borrelia burgdorferi s.l. and its genospecies, with complementary analyses on Rickettsia sp., Anaplasma phagocytophilum, Neoehrlichia mikurensis and Borrelia miyamotoi. DIN was highest in forest interior and at trails enclosed by forest. Lower disease hazard was observed at benches and trails at forest edges. This infrastructure effect can be attributed to variation in vegetation characteristics and the habitat use of tick hosts, specifically roe deer, rodents and songbirds. DON is the main driver of DIN at micro-geographic scale and negatively affected by infrastructure and forest edges. A positive association with vegetation cover in understorey and canopy was observed, as were positive trends for local rodent and songbird abundance. NIP of different pathogens was affected by different drivers. Lower B. burgdorferi s.l. prevalence in the interior of forest stands, driven by its most prevalent genospecies B. afzelii, points towards higher density of uninfected hosts there. B. afzelii was positively associated with understorey containing tall species and with high canopy cover, whereas local bird community composition predicts B. garinii prevalence. A positive effect of songbird abundance and a negative effect of pigeons were observed. Our findings support amplification and inhibition mechanisms within forest stands and highlight that the effect of established drivers of DIN may differ based on the considered spatial scale.