Floodplains are among the world’s economically-most-valuable, environmentally-most-threatened, and yet conceptually-least-understood ecosystems. Drawing on concepts from existing riverine and wetland models, and empirical data from floodplains of Atlantic Coast rivers in the Southeastern US (and elsewhere when possible), we introduce a conceptual model to explain a continuum of longitudinal variation in floodplain ecosystem functions with a particular focus on biotic change. Our hypothesis maintains that major controls on floodplain ecology are either external (ecotonal interactions with uplands or stream/river channels) or internal (wetland-specific functions), and the relative importance of these controls changes progressively from headwater to mid-river to lower-river floodplains. Inputs of water, sediments, nutrients, flora, and fauna from uplands-to-floodplains decrease, while the impacts of wetland biogeochemistry and obligate wetland plants and animals within-floodplains increase, along the length of a river floodplain. Inputs of water, sediment, nutrients, and fauna from river/stream channels to floodplains are greatest mid-river, and lower either up- or down-stream. While the floodplain continuum we develop is regional in scope, we review how aspects may apply more broadly. Management of coupled floodplain-river ecosystems would be improved by accounting for how factors controlling the floodplain ecosystem progressively change along longitudinal riverine gradients.