Migratory Fishes as Material and Process Subsidies in Riverine Ecosystems
Alexander S. Flecker, Peter B. McIntyre, Jonathan W. Moore, Jill T. Anderson, Brad W. Taylor, and Robert O. Hall, Jr.
Abstract.—Migratory fishes are common in freshwaters throughout the world and can fundamentally alter recipient ecosystems. We describe different types of fish migrations and consider their importance from the perspective of ecosystem subsidies—that is, landscape-scale flows of energy, materials, and organisms that are important in driving local food web and ecosystem dynamics. We distinguish between two general categories of subsidies, which we term here material subsidies and process subsidies. Material subsidies are the transfer of energy, nutrients, and other resources resulting in direct changes in resource pools within ecosystems. We posit that material subsidies occur under only a subset of life history strategies and ecological settings, and the potential for migratory fish to represent major material subsidies is greatest when (1) the biomass of migrants is high relative to recipient ecosystem size, (2) the availability of nutrients and energy is low in the recipient ecosystem (i.e., oligotrophic), and (3) there are effective mechanisms for both liberating nutrients and energy from migratory fishes and retaining those materials within the food web of the recipient ecosystem. Thus, anadromous semelparous Pacific salmon Oncorhynchus spp. with en masse programmed senescence in oligotrophic Pacific Northwest streams can be large material subsidies. In contrast, process subsidies arise from feeding or other activities of migratory species that directly affect process rates within recipient ecosystems. For example, the physical and chemical effects of grazing and sediment-feeding fishes such as prochilodontids, as well as seed dispersal by large-bodied frugivorous characins, represent potentially key process subsidies by migratory fishes in some of the great rivers of South America. We speculate that process subsidies are more widespread than material subsidies from migratory stream fishes because they are independent of the type of migration patterns, life history, and distance traveled. Nevertheless, the magnitude of process subsidies is likely to be greatest under a specific subset of ecological conditions, which can differ from those where material subsidies might be most important. In addition to migrant biomass, the potential for migratory fish to represent strong process subsidies is regulated by migrant interaction strength and the degree to which a migratory species is functionally unique in a particular ecological setting. Unlike material subsidies, which require high migrant biomass as conveyor belts of materials, migratory fishes can be crucial process subsidies, even when migrant biomass is low, if they are functionally unique and strong interactors. We provide specific examples of these different types of subsidies and outline key directions of research for furthering our understanding of the functional significance of migratory stream fishes. Our aim is to highlight the diversity of subsidies provided by migratory fishes in order to foster a more comprehensive perspective on fishes as essential components of riverine ecosystems.