BACKGROUND: Anaplasma phagocytophilum is currently regarded as a single species. However, molecular studies indicate that it can be subdivided into ecotypes, each with distinct but overlapping transmission cycle. Here, we evaluate the interactions between and within clusters of haplotypes of the bacterium isolated from vertebrates and ticks, using phylogenetic and network-based methods. METHODS: The presence of A. phagocytophilum DNA was determined in ticks and vertebrate tissue samples. A fragment of the groEl gene was amplified and sequenced from qPCR-positive lysates. Additional groEl sequences from ticks and vertebrate reservoirs were obtained from GenBank and through literature searches, resulting in a dataset consisting of 1623 A. phagocytophilum field isolates. Phylogenetic analyses were used to infer clusters of haplotypes and to assess phylogenetic clustering of A. phagocytophilum in vertebrates or ticks. Network-based methods were used to resolve host-vector interactions and their relative importance in the segregating communities of haplotypes. RESULTS: Phylogenetic analyses resulted in 199 haplotypes within eight network-derived clusters, which were allocated to four ecotypes. The interactions of haplotypes between ticks, vertebrates and geographical origin, were visualized and quantified from networks. A high number of haplotypes were recorded in the tick Ixodes ricinus. Communities of A. phagocytophilum recorded from Korea, Japan, Far Eastern Russia, as well as those associated with rodents had no links with the larger set of isolates associated with I. ricinus, suggesting different evolutionary pressures. Rodents appeared to have a range of haplotypes associated with either Ixodes trianguliceps or Ixodes persulcatus and Ixodes pavlovskyi. Haplotypes found in rodents in Russia had low similarities with those recorded in rodents in other regions and shaped separate communities. CONCLUSIONS: The groEl gene fragment of A. phagocytophilum provides information about spatial segregation and associations of haplotypes to particular vector-host interactions. Further research is needed to understand the circulation of this bacterium in the gap between Europe and Asia before the overview of the speciation features of this bacterium is complete. Environmental traits may also play a role in the evolution of A. phagocytophilum in ecotypes through yet unknown relationships.

Biology Centre Institute of Parasitology Czech Academy of Sciences České Budějovice Czech Republic Department of Parasitology Faculty of Science Charles University Prague Czech Republic

Centre for Ecological and Evolutionary Synthesis Department of Biosciences University of Oslo P O Box 1066 Blindern 0316 Oslo Norway

Centre for Infectious Disease Control Bilthoven The Netherlands

Centre for Infectious Disease Control Bilthoven The Netherlands Laboratory of Entomology Wageningen University Wageningen The Netherlands

Department for Bacteriology and Parasitology Croatian Veterinary Institute Savska Cesta 143 Zagreb Croatia

Department of Parasitology and Zoology University of Veterinary Medicine Budapest Budapest Hungary Evolutionary Systems Research Group Centre for Ecological Research Hungarian Academy of Sciences Tihany Hungary

Department of Parasitology Faculty of Veterinary Medicine University of Zaragoza Zaragoza Spain Emerging Zoonoses Research Group Instituto Agroalimentario de Aragón Zaragoza Spain

Department of Pathology and Parasitology Faculty of Veterinary Medicine University of Veterinary and Pharmaceutical Sciences Brno Czech Republic Biology Centre Institute of Parasitology Czech Academy of Sciences České Budějovice Czech Republic

Department of Veterinary Sciences University of Turin Via L da Vinci 44 Grugliasco 10095 Turin Italy

Department of Wildlife Fish and Environmental Studies Swedish University of Agricultural Sciences Umeå Sweden

Diergeneeskundig Centrum Zuid Oost Drenthe Coevorden The Netherlands

Dutch Wildlife Health Centre Utrecht University Utrecht The Netherlands

Friedrich Loeffler Institut Federal Research Institute for Animal Health Greifswald Insel Riems Germany

Institute of Vertebrate Biology v v i Medical Zoology Laboratory Academy of Sciences of the Czech Republic Brno Czech Republic

Institute of Zoology Slovak Academy of Sciences Bratislava Slovakia

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