Frozen out: unanswered questions about winter biology

Winter conditions impose dramatic constraints on temperate, boreal, and polar ecosystems, and shape the abiotic and biotic interactions underpinning these systems. At high latitudes, winter can last longer than the growing season and may have a disproportionately large impact on organisms and ecosystems. Even so, our understanding of the ecological implications of winter is often lacking. Indeed, even what exactly defines winter is currently unclear, and boundaries that delineate this season are blurred across marine, freshwater, and terrestrial realms and fields of biology. Here, we discuss the complexity of defining winter, and highlight the importance of maintaining the capacity to test hypotheses across seasons, realms, and domains of life. We then outline questions drawn from diverse fields of research that address current gaps in our understanding of winter ecology and how winter influences multiple levels of biological organization, from individuals to ecosystems. Finally, we highlight the potential consequences of changes to both the length and severity of winter due to climate change, and discuss the role winter may play in mediating ecosystem function in the future.