Unifying research on the fragmentation of terrestrial and aquatic habitats: patches, connectivity and the matrix in riverscapes

1. While there is an increasing emphasis in terrestrial ecology on determining the influence of the area that surrounds habitat patches (the landscape 'matrix') relative to the characteristics of the patches themselves, research on these aspects in running waters is still rather underrepresented. 2. Here we outline conceptual foundations of matrix ecology for stream and river ecosystems ('riverscapes'). We discuss how a hierarchical, patch-based perspective is necessary for the delineation of habitat patches and the surrounding matrix, through which we may identify two classes of habitat edges in riverscapes (i.e. edges between the terrestrial-aquatic interface and edges within streams). 3. Under this conceptual framework, we discuss the role of the matrix in influencing between-patch movement, and resource quality and quantity within and among habitat patches in riverscapes. We also review types of empirical and modelling approaches which may advance our understanding of fragmentation effects in these systems. 4. We identify five key challenges for understanding fragmentation and matrix effects more completely: (i) defining populations and their status (i.e. quantifying the demographic contribution of habitat patches to metapopulation dynamics), (ii) scaling from metapopulations to metacommunities (particularly searching for generalities in species responses to landscape heterogeneity), (iii) scaling from metacommunities to metaecosystems (i.e. exploring the interactive role of the terrestrial-aquatic and within-stream matrix effects on the flow of material and energy at the network scale), (iv) understanding temporal dynamics in matrix permeability and (v) revealing the utility of different patch and matrix representations for modelling connectivity relationships. 5. Fragmentation of habitats is a critical issue in the conservation and management of stream networks across spatial scales. Although the effects of individual barriers (e.g. dams) are well documented, we argue that a more comprehensive patch-matrix landscape model will improve our understanding of fragmentation effects and improve management in riverscapes.