Ecology and pathology of amphibian ranaviruses.

Mass mortality of amphibians has occurred globally since at least the early 1990s from viral pathogens that are members of the genus Ranavirus, family Iridoviridae. The pathogen infects multiple amphibian hosts, larval and adult cohorts, and may persist in herpetofaunal and osteichthyan reservoirs. Environmental persistence of ranavirus virions outside a host may be several weeks or longer in aquatic systems. Transmission occurs by indirect and direct routes, and includes exposure to contaminated water or soil, casual or direct contact with infected individuals, and ingestion of infected tissue during predation, cannibalism, or necrophagy. Some gross lesions include swelling of the limbs or body, erythema, swollen friable livers, and hemorrhage. Susceptible amphibians usually die from chronic cell death in multiple organs, which can occur within a few days following infection or may take several weeks. Amphibian species differ in their susceptibility to ranaviruses, which may be related to their co-evolutionary history with the pathogen. The occurrence of recent widespread amphibian population die-offs from ranaviruses may be an interaction of suppressed and naïve host immunity, anthropogenic stressors, and novel strain introduction. This review summarizes the ecological research on amphibian ranaviruses, discusses possible drivers of emergence and conservation strategies, and presents ideas for future research directions. We also discuss common pathological signs of ranaviral disease, methods for diagnostic evaluation, and ranavirus surveillance methods. In as much as ranaviral disease is listed as a notifiable disease by the World Organization for Animal Health and is a threat to amphibian survival, we recommend that biosecurity precautions are implemented by nations to reduce the likelihood of transporting ranavirus virions among populations. Biosecurity precautions include disinfecting footwear and equipment that comes in contact with surface water inhabited by amphibians and testing commercially shipped amphibians for the pathogen. We also encourage natural resource organizations to establish routine surveillance programs for ranaviruses in wild amphibian populations.