The cestode Echinococcus multilocularis is the causative agent of alveolar echinococcosis, a fatal zoonotic parasitic disease of the northern hemisphere. Red foxes are the main reservoir hosts and, likely, the main drivers of the geographic spread of the disease in Europe. Knowledge of genetic relationships among E. multilocularis isolates at a European scale is key to understanding the dispersal characteristics of E. multilocularis. Hence, the present study aimed to describe the genetic diversity of E. multilocularis isolates obtained from different host species in 19 European countries. Based on the analysis of complete nucleotide sequences of the cob, atp6, nad2, nad1 and cox1 mitochondrial genes (4,968 bp), 43 haplotypes were inferred. Four haplotypes represented 62.56 % of the examined isolates (142/227), and one of these four haplotypes was found in each country investigated, except Svalbard, Norway. While the haplotypes from Svalbard were markedly different from all the others, mainland Europe appeared to be dominated by two main clusters, represented by most western, central and eastern European countries, and the Baltic countries and northeastern Poland, respectively. Moreover, one Asian-like haplotype was identified in Latvia and northeastern Poland. To better elucidate the presence of Asian genetic variants of E. multilocularis in Europe, and to obtain a more comprehensive Europe-wide coverage, further studies, including samples from endemic regions not investigated in the present study, especially some eastern European countries, are needed. Further, the present work proposes historical causes that may have contributed to shaping the current genetic variability of E. multilocularis in Europe.

Anses Nancy Laboratory for Rabies and Wildlife National Reference Laboratory Echinococcus spp 54220 Malzéville France

Centro Specialistico Fauna Selvatica SCT2 Belluno Istituto Zooprofilattico Sperimentale delle Venezie Via Cappellari 44 A 32100 Belluno Italy

Croatian Veterinary Institute Laboratory for Parasitology 10000 Zagreb Croatia

Danish Veterinary and Food Administration Ministry of Food Agriculture and Fisheries of Denmark Stationsparken 31 33 2600 Glostrup Denmark

Department of Genetic Research Institute for Biological Research Siniša Stanković National Institute of Republic of Serbia University of Belgrade Bulevar Despota Stefana 142 11000 Belgrade Serbia

Department of Veterinary Pathobiology Veterinary Academy Lithuanian University of Health Sciences Tilžės str 18 47181 Kaunas Lithuania

Department of Zoology Institute of Ecology and Earth Sciences University of Tartu J Liivi 2 50409 Tartu Estonia

European Union Reference Laboratory for Parasites Department of Infectious Diseases Istituto Superiore Di Sanità Viale Regina Elena 299 00161 Rome Italy

European Union Reference Laboratory for Parasites Department of Infectious Diseases Istituto Superiore Di Sanità Viale Regina Elena 299 00161 Rome Italy; WHO Collaborating Centre for the Epidemiology Detection and Control of Cystic and Alveolar Echinococcosis Department of Infectious Diseases Istituto Superiore Di Sanità Viale Regina Elena 299 00161 Rome Italy

Friedrich Loeffler Institut Federal Research Institute for Animal Health Institute of Epidemiology Südufer 10 17493 Greifswald‑Insel Riems Germany

Infectious Disease Preparedness Statens Serum Institut Artillerivej 5 DK 2300 Copenhagen S Denmark

Institute for Fish and Wildlife Health Department of Infectious Diseases and Pathobiology Vetsuisse Faculty University of Bern Längassstrasse 122 3012 Bern Switzerland

Institute for Veterinary Disease Control Innsbruck Austrian Agency for Health and Food Safety Technikerstraße 70 6020 Innsbruck Austria

Institute of Food Safety Animal Health and Environment BIOR Lejupes Street 3 Riga LV 1076 Latvia

Institute of Food Safety Animal Health and Environment BIOR Lejupes Street 3 Riga LV 1076 Latvia; Faculty of Biology University of Lavia Jelgavas Street 1 Riga LV 1004 Latvia

Institute of Microbiology and Parasitology Veterinary Faculty University of Ljubljana Gerbičeva 60 1000 Ljubljana Slovenia

Institute of Parasitology Slovak Academy of Sciences Košice Hlinkova 3 040 01 Košice Slovakia

Institute of Parasitology Vetsuisse Faculty University of Bern Länggassstrasse 122 CH 3012 Bern Switzerland

Institute of Parasitology Vetsuisse Faculty University of Bern Länggassstrasse 122 CH 3012 Bern Switzerland; Institute of Infectious Diseases Faculty of Medicine University of Bern CH 3012 Bern Switzerland

Institute of Parasitology Vetsuisse Faculty University of Bern Länggassstrasse 122 CH 3012 Bern Switzerland; Multidisciplinary Center for Infectious Diseases University of Bern Länggass Strasse 122 3012 Bern Switzerland

Institute of Tropical Medicine Department of Biomedical Sciences 155 Nationalestraat B 2000 Antwerp Belgium

Latvian State Forest Research Institute Silava Wildlife Management Research Group Salaspils Rigas Street 111 LV 2169 Salaspils Latvia

Medical Parasitology Institute of Specific Prophylaxis and Tropical Medicine Center of Pathophysiology Infectiology and Immunology Medical University Vienna Kinderspitalgasse 15 1090 Vienna Austria

National Centre for Laboratory Research and Risk Assessment Fr R Kreutzwaldi 30 Tartu Estonia

National Reference Laboratory of Medical Parasitology National Public Health Center Albert Flórián út 2 6 Budapest Hungary

National Veterinary Research Institute Department of Parasitology and Invasive Diseases Partyzantow Avenue 57 24 100 Pulawy Poland

Norwegian Polar Institute Fram Centre NO 9296 Tromsø Norway

Norwegian Veterinary Institute Holtvegen 66 9016 Tromsø Norway

Parasitology Unit Institute of Biology University of Hohenheim 70593 Stuttgart Germany

State Veterinary Institute Olomouc Jakoubka ze Stribra 1 779 00 Olomouc Czech Republic

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Helminth diversity of nutria (Myocastor coypus) across the Morava basin in the Czech Republic