The calendar of epidemics: Seasonal cycles of infectious diseases

Seasonal cyclicity is a ubiquitous feature of acute infectious diseases [1] and may be a ubiquitous feature of human infectious diseases in general, as illustrated in Tables 1–4. Each acute infectious disease has its own seasonal window of occurrence, which, importantly, may vary among geographic locations and differ from other diseases within the same location. Here we explore the concept of an epidemic calendar, which is the idea that seasonality is a unifying feature of epidemic-prone diseases and, in the absence of control measures, the local calendar can be marked by epidemics (Fig 1). A well-known example of a calendar marked by epidemics is that of the Northern Hemisphere, where influenza outbreaks occur each winter [2, 3] (hence the colloquial reference to winter as "the flu season"). In contrast, chickenpox outbreaks peak each spring [4, 5], and polio transmission historically occurred each summer [6]. Seasonal variation in infectious disease transmission plays an important role in determining when epidemics happen; however, it is not the sole determinant. For instance, some infectious diseases with known seasonal transmission, such as pertussis and measles, can display multi-annual outbreaks, meaning their epidemics occur in multi-year intervals, such as every two or four years, rather than annually. This is because the timing of these epidemics is determined by a combination of (i) seasonal transmission and (ii) different processes shaping the number of susceptible individuals in the population, a sufficient number of which is a prerequisite for an outbreak. Within the fields of infectious disease ecology and epidemic modeling, seasonal variation in transmission is known as seasonal forcing [78]. Over the past century, attention has been paid to detailing the cyclicity and mechanisms of seasonal forcing for a few diseases of public interest, such as measles, influenza, and cholera (e.g., see contemporary work by [3, 79, 80]). Despite these notable examples, disease seasonality has yet to be systemically and/or rigorously characterized for the majority of infections. Here, I aim to motivate future studies of disease seasonality by drawing attention to the importance of seasonality in public health, medicine, and biology. I will explore documented seasonal cycles in human infections, including notifiable and neglected tropical diseases. I also aim to present a holistic view of hypothesized drivers of seasonality for each disease, with the caveat that, for the majority of infections, the current state of the science is insufficient to draw conclusions about seasonal timing, seasonal magnitude, and geographic variation in incidence. Although published data regarding disease seasonality are limited for individual diseases, collectively the body of work on disease seasonality is vast and reveals that infections—which may differ enormously in their pathology and/or ecology—coalesce via underlying seasonal drivers