Research showing that salmon carcasses support the productivity and biodiversity of 3 aquatic and riparian ecosystems has been conducted over a variety of spatial and temporal scales. 4In some studies, carcasses were manipulated in a single pulse or loading rate or manipulations 5 occurred during summer and early fall, rather than simulating the natural dynamic of an extended 6 spawning period, a gradient of loading rates, or testing carcass effects in late fall-early winter 7 when some salmon stocks in the US Pacific Northwest spawn. To address these discrepancies, period. We found little evidence that carcasses influenced primary producer biomass or fish 13 growth; however, nutrients and some primary consumer populations increased with loading rate. 14 These effects varied through time, however. We hypothesize that the variable effects of carcasses 15 were a result of ambient abiotic condition, such as light, temperature and disturbance that 16 constrained trophic response. There was some evidence to suggest peak responses for primary 17 producers and consumers occurred at a loading rate of ~1.0 -2.0 kg/m 2 , which was similar to 18 other experimental studies conducted during summer. 19 20
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