A Review of Coronaviruses in Wild Birds and Opportunities for Future Research on Migratory Waterfowl

Coronaviruses (CoVs) were first described in poultry in the early 1930s and formally recognized as pathogens of both animal and human populations in the late 1960s. They are now considered among the most abundant viral families in the world. Though their distribution and diversity remain understudied in wild animals, representatives from 13 orders of wild birds worldwide have tested positive for CoVs of the gamma and delta genera over the last 25 years. Many of these wild bird species are in the orders Charadriiformes (shorebirds and their relatives) and Anseriformes (waterfowl including ducks, geese, and swans). Waterfowl are particularly concerning as potential reservoirs for CoVs because they are globally distributed; often congregate in large, mixed-species flocks; and may exist in close proximity to humans and domesticated animals. This review describes the history and current knowledge of CoVs in birds, provides an updated list of global detections of CoVs in 124 species of wild birds as reported in the peer-reviewed literature since 2000, and highlights topics for future research that would help elucidate the role of waterfowl in CoV transmission. Our review reiterates the need for continuous surveillance to detect and monitor CoVs across all bird species and for standardization in data reporting and analysis of both negative and positive results. Such information is critical to understand the potential role of free-ranging birds in the maintenance, evolution, and transmission of the virus. Further, we believe that research on the potential impacts of coronavirus infections and coinfections on avian demographics, especially reproduction in waterfowl, is warranted given known consequences in domestic poultry.