Marine and terrestrial ecosystems are connected via transfers of nutrients and organic matter in river discharges. In coastal seas, such freshwater outflows create prominent turbidity plumes. These plumes are areas of high biological activity in the pelagos, of which zooplankton is a key element. Conceptually, the increased biomass of zooplankton consumers in plumes can be supported by two alternative trophic pathways-consumption of fresh marine phytoplankton production stimulated by riverine nutrients, or direct trophic subsidies through the uptake of terrestrial and estuarine organic matter flushed to sea. The relative importance of these two pathways has not been established previously. Isotopic tracing (carbon and nitrogen) was used to measure the extent of incorporation of marine versus terrestrial matter into mesozooplankton consumers in the plumes off a small estuary in eastern Australia. Replicate zooplankton samples were taken during baseflow conditions with minimal freshwater influence to the sea, and during pulsed discharge events that generated turbidity plumes in coastal waters. Food sources utilized by zooplankton differed among locations and with the strength of freshwater flow. Terrestrial and estuarine carbon only made a sizeable contribution (47%) to the carbon demands of zooplankton in the lower estuary during pulsed freshwater flows. By contrast, in plumes that developed in nearshore marine waters, phytoplankton supplied up to 90% of the dietary carbon of zooplankton feeding in the plumes. Overall, it was ''fresh'' carbon, fixed by marine phytoplankton, the growth of which became stimulated by fluvial nutrient exports, that dominated energy flows in plume regions. The trophic role of terrestrial and estuarine organic exports was comparatively minor. The trophic dynamics of plankton in small coastal plumes is closely linked to variations in freshwater flow, but this coupling operates mainly through the enhancement of in-situ phytoplankton production rather than cross-boundary transfers of organic matter to marine food webs in the pelagos.

Freshwater discharge from rivers is a powerful forcing agent in coastal ecosystems. It not only generates strong ecological effects in estuaries, but also drives the dynamics of nearshore marine waters where prominent river plumes form biogeochemical hot spots in coastal seas worldwide. Large plumes from major rivers exert important controls on pelagic processes. The majority of estuaries are smaller, however, and the importance of the smaller plumes they generate is unknown. We measured the degree of coupling between freshwater flow and inshore zooplankton in such a plume from a subtropical estuary on the east coast of Australia. Flow regimes encompassed long periods of low freshwater input, punctuated by pulsed freshets that initiated the formation of buoyant, lower-salinity plumes in the nearshore marine zone. Plumes stimulated phytoplankton biomass in the receiving waters, and ultimately changes in zooplankton assemblages. Zooplankton responded strongly to river discharge: (1) in the absence of substantial freshwater flows and plumes, zooplankton was broadly similar in density and biomass across the estuarine-marine gradient; (2) freshets that generated significant plumes strongly modified hydrological conditions and lowered zooplankton in the estuarine and nearshore waters, and (3) after the initial freshet, zooplankton in the residual plume was at a higher density in nearshore than shelf waters. We demonstrate that coupling between riverine and coastal pelagic systems operates in small plumes, but that there is substantial temporal variance linked to fluctuations in freshwater delivery. IntroductionFreshwater inflow is one of the fundamental drivers of coastal ecosystems. The amount and temporal modes of freshwater discharge profoundly influence a wide range of biological processes at multiple levels of ecological organisation in estuaries, ranging from shifts * Corresponding author 642 T. A. SCHLACHER et al.

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