The macroecology of the COVID-19 pandemic in the Anthropocene.

Severe acute respiratory syndrome coronavirus 2, the virus that causes coronavirus disease 2019 (COVID-19), has expanded rapidly throughout the world. Thus, it is important to understand how global factors linked with the functioning of the Anthropocene are responsible for the COVID-19 outbreak. We tested hypotheses that the number of COVID-19 cases, number of deaths and growth rate of recorded infections: (1) are positively associated with population density as well as (2) proportion of the human population living in urban areas as a proxies of interpersonal contact rate, (3) age of the population in a given country as an indication of that population's susceptibility to COVID-19; (4) net migration rate and (5) number of tourists as proxies of infection pressure, and negatively associated with (5) gross domestic product which is a proxy of health care quality. Data at the country level were compiled from publicly available databases and analysed with gradient boosting regression trees after controlling for confounding factors (e.g. geographic location). We found a positive association between the number of COVID-19 cases in a given country and gross domestic product, number of tourists, and geographic longitude. The number of deaths was positively associated with gross domestic product, number of tourists in a country, and geographic longitude. The effects of gross domestic product and number of tourists were non-linear, with clear thresholds above which the number of COVID-19 cases and deaths increased rapidly. The growth rate of COVID-19 cases was positively linked to the number of tourists and gross domestic product. The growth rate of COVID-19 cases was negatively associated with the mean age of the population and geographic longitude. Growth was slower in less urbanised countries. This study demonstrates that the characteristics of the human population and high mobility, but not population density, may help explain the global spread of the virus. In addition, geography, possibly via climate, may play a role in the pandemic. The unexpected positive and strong association between gross domestic product and number of cases, deaths, and growth rate suggests that COVID-19 may be a new civilisation disease affecting rich economies.