Birds are hosts for several zoonotic pathogens. Because of their high mobility, especially of longdistance migrants, birds can disperse these pathogens, affecting their distribution and phylogeography. We focused on Borrelia burgdorferi sensu lato, which includes the causative agents of Lyme borreliosis, as an example for tick-borne pathogens, to address the role of birds as propagation hosts of zoonotic agents at a large geographical scale. We collected ticks from passerine birds in 11 European countries. B. burgdorferi s.l. prevalence in Ixodes spp. was 37% and increased with latitude. The fieldfare Turdus pilaris and the blackbird T. merula carried ticks with the highest Borrelia prevalence (92 and 58%, respectively), whereas robin Erithacus rubecula ticks were the least infected (3.8%). Borrelia garinii was the most prevalent genospecies (61%), followed by B. valaisiana (24%), B. afzelii (9%), B. turdi (5%) and B. lusitaniae (0.5%). A novel Borrelia genospecies "Candidatus Borrelia aligera" was also detected. Multilocus sequence typing (MLST) analysis of B. garinii isolates together with the global collection of B. garinii genotypes obtained from the Borrelia MLST public database revealed that: (a) there was little overlap among genotypes from different continents, (b) there was no geographical structuring within Europe, and (c) there was no evident association pattern detectable among B. garinii genotypes from ticks feeding on birds, questing ticks or human isolates. These findings strengthen the hypothesis that the population structure and evolutionary biology of tick-borne pathogens are shaped by their host associations and the movement patterns of these hosts.

Behavioural Ecology Department of Biology Ludwig Maximilians University of Munich Planegg Martinsried Germany

Behavioural Ecology Group Department of Systematic Zoology and Ecology Eötvös Loránd University Budapest Hungary

Center for Vector and Infectious Diseases Research National Institute of Health Dr Ricardo Jorge Lisbon Portugal

CIBIO InBIO Research Center in Biodiversity and Genetic Resources University of Porto Porto Portugal

CNRS Department of Biometry and Evolutionary Biology University Lyon 1 University of Lyon Villeurbanne France

Czech Union for Nature Conservation Břeclav Czech Republic

Departamento de Ecología Evolutiva Museo Nacional de Ciencias Naturales Madrid Spain

Department of Anatomy Cell and Developmental Biology Eötvös Loránd University Budapest Hungary

Department of Animal Ecology Netherlands Institute of Ecology Wageningen The Netherlands

Department of Biodiversity Ecology and Evolution Universidad Complutense de Madrid Madrid Spain

Department of Biology and Wildlife Diseases Faculty of Veterinary Hygiene and Ecology University of Veterinary and Pharmaceutical Sciences Brno Brno Czech Republic

Department of Biology Faculty of Medicine Masaryk University Brno Czech Republic

Department of Biology Molecular Ecology and Evolution Lab University of Lund Lund Sweden

Department of Biology University of Turku Turku Finland

Department of Ecology and Evolution Animal Ecology Evolutionary Biology Centre Uppsala University Uppsala Sweden

Department of Ecology and Evolutionary Biology Princeton University Princeton NJ USA

Department of Ecology and Genetics University of Oulu Oulu Finland

Department of Life Sciences CFE Centre for Functional Ecology Science for People and the Planet University of Coimbra Coimbra Portugal

Department of Parasitology and Zoology University of Veterinary Medicine Budapest Hungary

Department of Zoology Palacky University Olomouc Czech Republic

Department of Zoology University of Tartu Tartu Estonia

Division of Evolutionary Biology Faculty of Biology LMU Munich Planegg Martinsried Germany

Evolutionary Physiology Laboratory Max Planck Institute for Ornithology Seewiesen Germany

Finnish Museum of Natural History University of Helsinki Helsinki Finland

Forest Research Institute Hellenic Agricultural Organization DEMETER Thesaloniki Greece

German National Reference Centre for Borrelia Oberschleissheim Germany

Hellenic Bird Ringing Center Athens Greece

Hungarian Biodiversity Research Society Budapest Hungary

Instituto Cavanilles de Biodiversidad y Biología Evolutiva Universidad de Valencia Valencia Spain

Interuniversity Institute for Biostatistics and Statistical Bioinformatics Hasselt University Diepenbeek Belgium

Laboratory of Parasitology and Parasitic Diseases Faculty of Health Sciences School of Veterinary Medicine Aristotle University of Thessaloniki Thessaloniki Greece

MARE Marine and Environmental Sciences Centre University of Coimbra Coimbra Portugal

Museum and Institute of Zoology Polish Academy of Sciences Warszawa Poland

Museum of the Moravian Wallachia Region Vsetín Czech Republic

National Institute of Public Health and Environment Laboratory for Zoonoses and Environmental Microbiology Bilthoven The Netherlands

Ócsa Bird Ringing Station Ócsa Hungary

School of Biological Sciences University of Aberdeen Aberdeen United Kingdom

Slovenian Museum of Natural History Ljubljana Slovenia

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