Meta-ecosystems and biological energy transport from ocean to coast: the ecological importance of herring migration.

Ecosystems are not closed, but receive resource subsidies from other ecosystems. Energy, material and organisms are moved between systems by physical vectors, but migrating animals also transport resources between systems. We report on large scale energy transport from ocean to coast by a migrating fish population, the Norwegian spring-spawning (NSS) herring Clupea harengus. We observe a rapid body mass increase during parts of the annual, oceanic feeding migration and we use a bioenergetics model to quantify energy consumption. The model predicts strong seasonal variation in food consumption with a marked peak in late May to July. The copepod Calanus finmarchicus is the most important prey and 23 x 10(6) tones (wet weight) of C. finmarchicus is consumed annually. The annual consumption-biomass ratio is 5.2. During the feeding migration 17% of consumed energy is converted to body mass. The biomass transported to the coast and left as reproductive output is estimated from gonad weight and is about 1.3 x 10(6) tones for the current population. This transport is to our knowledge the world's largest flux of energy caused by a single population. We demonstrate marked temporal variation in transport during the last century and discuss the effects of NSS herring in the ocean, as a major consumer, and at the coast, where eggs and larvae are important for coastal predators. In particular, we suggest that the rapid decline of lobster Homarus gammarus landings in Western Norway during the 1960s was related to the collapse of NSS herring. We also discuss spatial variation in energy transport caused by changed migration patterns. Both climate and fisheries probably triggered historical changes in the migration patterns of NSS herring. New migration routes emerge at the level of individuals, which in turn determines where resources are gathered and delivered, and therefore, how meta-ecosystems function.