The Virioneuston: A Review on Viral⁻Bacterial Associations at Air⁻Water Interfaces

Vast biofilm-like habitats at air⁻water interfaces of marine and freshwater ecosystems harbor surface-dwelling microorganisms, which are commonly referred to as neuston. Viruses in the microlayer, i.e., the virioneuston, remain the most enigmatic biological entities in boundary surface layers due to their potential ecological impact on the microbial loop and major air⁻water exchange processes. To provide a broad picture of the viral⁻bacterial dynamics in surface microlayers, this review compiles insights on the challenges that viruses likely encounter at air⁻water interfaces. By considering viral abundance and morphology in surface microlayers, as well as dispersal and infection mechanisms as inferred from the relevant literature, this work highlights why studying the virioneuston in addition to the bacterioneuston is a worthwhile task. In this regard, major knowledge gaps and possible future research directions are discussed.