The Late Devonian Red Hill locality in north-central Pennsylvania contains an Archaeopteris-dominated plant fossil assemblage, a diverse fossil fauna, and an extensive sedimentary sequence ideal for investigating the landscapes and biotic associations of the earliest forest ecosystems. Sedimentological analysis of the main plant-fossil bearing layer at Red Hill indicates that it was a flood-plain pond. A seasonal wet-and-dry climate is indicated by well-developed paleovertisols. The presence of charcoal interspersed with plant fossils indicates that fires occurred in this landscape. Fires appear to have primarily affected the fern Rhacophyton. The specificity of the fires, the distribution profile of the plant remains deposited in the pond, and additional taphonomic evidence all support a model of niche partitioning of the Late Devonian landscape by plants at a high taxonomic level. At Red Hill, Archaeopteris was growing on the well-drained areas; Rhacophyton was growing in widespread monotypic stands; cormose lycopsids grew along the pond edge; and gymnosperms and Gillespiea were possibly opportunists following disturbances. Tetrapod fossils have been described from Red Hill—therefore, this paleoecological analysis is the first systematic interpretation of a specific site that reflects the type of wetland environment within which the earliest tetrapods evolved.