![]() ![]() During the late medieval period in Europe, long-distance maritime trade routes were developed as a result of the commercial revival of Italy 26. The study timespan was determined by the availability of data before the Industrial Revolution in Europe (see SI Text for details). 25, this study examined the extent to which major trade routes shaped the dispersal of plague in Europe between AD13. Using the geo-referenced Old World trade route database prepared by Ciolek 24 and the historical human plague outbreak distribution in Europe by Büntgen et al. In an earlier study, the authors also hypothesized the possible role of trade routes in plague distribution through examining the linkage between navigable rivers and plague outbreaks 23. The idea that infectious diseases are spread by transportation routes has been supported by proof from various studies on infectious diseases 19– 22. Ironically, remarkably little research has been done to elucidate how plague spread through metastatic spatial domains and the mechanism behind its distribution. Given the nature of plague and its notorious history, the international community should be more prepared for the re-emergence of plague. ![]() Flashing back in history, plague caused three great pandemics, in which 200 million people perished 18. Considering the widespread wildlife reservoirs of plague foci, together with the quick spread, rapid clinical course, inherent communicability, and high mortality rate of plague, the risk of plague outbreak should never be underestimated, although the number of human plague cases is relatively low compared to other infectious diseases at present 17. Some researchers suggested that plague may reign over our planet again when global climate change makes some places on earth become wetter and hotter 14– 16. Over the past few decades, thousands of cases of human plague have occurred around the world, particularly in Africa 13. Furthermore, laboratory results illustrated the experimental possibility of oral route transmission of plague 8 and epidemiological records suggested plague infection through consumption of contaminated meat 9, 10.ĭespite improvements in sanitation and medical advancements in the course of human history, plague remains a major threat to human beings 11, 12. Yet, recent evidence revealed that not only rodent fleas ( Xenopsylla cheopis as a classic example) are to blame for plague transmission, human fleas ( Pulex irritans) and cat fleas ( Ctenocephalides felis) are also likely to play a role in disseminating Yersinia pestis 6, 7. Traditional thought suggested that the clustering of Yersinia pestis rarely happened on human fleas 5. If an infected flea attempts to feed on a human, it will transmit Yersinia pestis to the human and lead to human plague in the form of bubonic plague or pulmonary plague. When the infected flea jumps onto another mammal, preferably rodents, it will transmit the bacteria to the new host by regurgitating the clotted blood from the blockage of the alimentary canal 4. The bacterium will quickly multiply and cluster, leading to the blockage of the alimentary canal in the fleas’ guts 3. Normally, the bloodsucking fleas acquire Yersinia pestis from an infected rodent. ![]() Plague is initiated by the flea-borne bacterium Yersinia pestis, which circulates mainly on rodents and other mammal hosts through the rodent’s associated fleas 1, 2. Based on our findings, we propose the mechanism of plague transmission in historical Europe, which is imperative in demonstrating how pandemics were spread in recent human history. A case study in Germany indicates that plague penetrated further into Europe through the local trade route network. Major trade routes decided the major plague outbreak hotspots, while navigable rivers determined the geographic pattern of sporadic plague cases. Furthermore, the negative correlation between plague outbreaks and their distance from major trade ports indicates the absence of a permanent plague focus in the inland areas of Europe. Ordinary Least Square (OLS) estimation results show that major trade routes played a dominant role in spreading plague in pre-industrial Europe. Here, we resolve the hypothetical role of trade routes through statistical analysis on the geo-referenced major trade routes in the early modern period and the 6,656 geo-referenced plague outbreak records in AD1347–1760. Numerous historical works have mentioned that trade routes were to blame for the spread of plague in European history, yet this relationship has never been tested by quantitative evidence.
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