Understanding the transmission dynamics and persistence of avian influenza viruses (AIVs) in the wild is an important scientific and public health challenge because this system represents both a reservoir for recombination and a source of novel, potentially humanpathogenic strains. The current paradigm locates all important transmission events on the nearly direct fecal/oral bird-to-bird pathway. In this article, on the basis of overlooked evidence, we propose that an environmental virus reservoir gives rise to indirect transmission. This transmission mode could play an important epidemiological role. Using a stochastic model, we demonstrate how neglecting environmentally generated transmission chains could underestimate the explosiveness and duration of AIV epidemics. We show the important pathogen invasion implications of this phenomenon: the nonnegligible probability of outbreak even when direct transmission is absent, the long-term infectivity of locations of prior outbreaks, and the role of environmental heterogeneity in risk.stochastic model | mathematical model | demographic stochasticity | waterfowl | epidemic A vian influenza viruses (AIVs) are an important class of infectious agents, both as a model for the influenza viruses that infect millions of people each year and as a generator of the genetic variation that might give rise to a future pandemic strain (1, 2). In contrast to the dominant human strains (3-5), the dynamics, control, and management of transmission remain poorly understood even in historically prevalent low pathogenic avian influenza viruses (LPAIVs) (2, 6). Given the recent emergence of H5N1 highly pathogenic avian influenza virus (HPAIV) and its continued introduction into new territories with attendant impacts on domestic waterfowl, poultry, and human populations, a thorough understanding of influenza evolution and epidemiology takes on a new urgency (6, 7).For many infectious diseases, transmission theory assumes that the majority of infections is caused by direct interactions between infectious and susceptible individuals (8, 9). The presence of additional transmission modes, particularly environmental transmission, gives rise to mechanisms that alter the conditions for pathogen invasion and persistence (10). Based on a number of lines of reasoning, we believe environmental transmission of LPAIVs occurs in natural populations:1. Environmental persistence of LPAIVs. LPAIV persistence in incubations intended to mimic aquatic environments may last many months, depending on environmental conditions ( Fig. 1; (22) have argued that neither density-nor frequency-dependent direct transmission captured the observed pattern of infections during an outbreak of LPAIV in the Rhône delta, France. By contrast, they considered the predictions of a model including water-borne transmission and the data in strong agreement. For these reasons, environmental transmission could be important in AIV epidemiology (23). It is known that indirect transmission chains, which the standard susceptibles-infectiv...