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            <funder>Most tick observatories were supported by the CC-EID and Climatick projects: "Adaptation of Agriculture and Forests to Climate Change" metaprogramme of the French National Research Institute for Agriculture, Food and Environment (INRAE). The observatory of Les Bordes was installed and equipped by the Centre Val-de-Loire region, the Loiret and the French National Forest Office. We also would like to thank the TEMPO French Network (https://tempo.pheno.fr/) for providing resources to support this study. The Les Bordes site belongs to the French national research infrastructure, ANAEE-F (http://www.anaee-france.fr/fr/), and is also included in the SOERE TEMPO (https://tempo.pheno.fr/).The Les Bordes site is also part of the framework of the ZAL (LTSER Zone Atelier Loire) and the GIS Coop network (https://www6.inra.fr/giscoop/), which is supported by the French Ministry for Agriculture and Food. This study was partially funded by EU grant 874850 MOOD and is catalogued as MOOD 040. The contents of this publication are the sole responsibility of the authors and do not necessarily reflect the views of the European Commission. Finally, PW is grateful for a grant from the Association Nationale de la Recherche et de la Technologie, bourse CIFRE (no 2019/0393).</funder>
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                <term xml:lang="en">Phenological patterns</term>
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              <p>Ixodes ricinus ticks (Acari: Ixodidae) are the most important vector for Lyme borreliosis in Europe. As environmental and climate changes might affect the distribution and activity of tick populations, this study aimed to determine the effects of environmental factors, i.e., meteorological, bioclimatic, and habitat characteristics on questing activity of I. ricinus nymphs across diverse climatic types in France over an 8-year period. Questing activity was observed using a repeated removal sampling design with a cloth-dragging technique in 11 sampling sites from 7 tick observatories from 2014 to 2021 at approximately 1-month intervals, involving 631 sampling campaigns. Three phenological patterns were observed, potentially following a climatic gradient. The mixed-effects negative binomial regression revealed that observed nymph counts were driven by different interval-average meteorological variables, including one-month average temperature, previous-season average temperature, and half-year average minimum relative humidity. The interaction effects indicated that the phenology in colder climates peaked differently from that of warmer climates. Also, the land cover characteristics that support the highest baseline abundance were moderate forest fragmentation with transition borders with agricultural areas. Finally, our model could potentially be used to predict seasonal human-tick exposure risks in France that could contribute to mitigating Lyme borreliosis risks.</p>
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