Caused by Yersinia pestis, plague ravaged the world through three known pandemics: the First or the Justinianic (6th-8th century); the Second (beginning with the Black Death during c.1338-1353 and lasting until the 19th century); and the Third (which became global in 1894). It is debatable whether Y. pestis persisted in European wildlife reservoirs or was repeatedly introduced from outside Europe (as covered by European Union and the British Isles). Here, we analyze environmental data (soil characteristics and climate) from active Chinese plague reservoirs to assess whether such environmental conditions in Europe had ever supported "natural plague reservoirs". We have used new statistical methods which are validated through predicting the presence of modern plague reservoirs in the western United States. We find no support for persistent natural plague reservoirs in either historical or modern Europe. Two factors make Europe unfavorable for long-term plague reservoirs: 1) Soil texture and biochemistry and 2) low rodent diversity. By comparing rodent communities in Europe with those in China and the United States, we conclude that a lack of suitable host species might be the main reason for the absence of plague reservoirs in Europe today. These findings support the hypothesis that long-term plague reservoirs did not exist in Europe and therefore question the importance of wildlife rodent species as the primary plague hosts in Europe.

Department of Biosciences Centre for Ecological and Evolutionary Synthesis University of Oslo Oslo 0316 Norway

Department of Earth System Science Ministry of Education Key Laboratory for Earth System Modeling Tsinghua University Beijing 100084 China

Department of Environmental and Prevention Sciences University of Ferrara Ferrara 44121 Italy

Department of Geography Faculty of Science Masaryk University Brno 611 37 Czech Republic

Department of Geography University of Cambridge Cambridge CB2 3EN UK

Department of History University of Glasgow Glasgow G12 8QQ UK

Department of Industrial Engineering Center for Statistical Science Tsinghua University Beijing 100084 China

Department of Soil and Water Sciences China Agricultural University Beijing 100193 China

Division of History and Politics University of Stirling Stirling FK9 4LJ UK

Forensic Soil Science Group James Hutton Institute Aberdeen AB15 8QH UK

Global Change Research Institute Czech Academy of Sciences Brno 603 00 Czech Republic

Institute of Healthy China Tsinghua University Beijing 100084 China

London School of Economics London WC2A 2AE UK

School of Biosciences University of Nottingham Loughborough Leics LE12 5RD UK

School of Geography University of Nottingham University Park Nottingham NG7 2RD UK

State Key Laboratory of Pathogen and Biosecurity Beijing Institute of Microbiology and Epidemiology Beijing 100071 China

Swiss Federal Research Institute Birmensdorf 8903 Switzerland

The 1st Institute of Endemic Disease Control and Prevention of Jilin Province Baicheng 137000 Jilin Province China

Univ Lille CNRS Inserm CHU Lille Institut Pasteur Lille U1019 UMR 9017 CIIL Center for Infection and Immunity of Lille Lille F 59000 France

Vanke School of Public Health Tsinghua University Beijing 100084 China

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Reply to Alfani: Reconstructing past plague ecology to understand human history