Environmental Fluctuation And Shifting Predation Pressure Contribute To Substantial Variation In Early Marine Survival Of Steelhead

Steelhead Oncorhynchus mykiss smolts must migrate through distinct freshwater, estuarine, and coastal habitats, each with unique biological and physical characteristics, on their route from natal streams to the ocean to complete their life cycle. Fewer than 15% of steelhead smolts survived their migration from the Nisqually River through Puget Sound to the Pacific Ocean from 2006 to 2009 and in 2014. Rapid smolt migration coupled with evidence of smolt mortality at harbor seal Phoca vitulina haulouts during 2014 indicated that predation by pinnipeds was an important source of mortality. When Puget Sound temperatures increased with the Northeast Pacific marine heat wave from late 2014 to 2016, steelhead smolt survival probabilities increased to 38% in 2016 and 2017, then decreased again as Puget Sound water temperatures cooled. Many ecological changes accompanied the increased marine temperatures, including the increased abundance of northern anchovy Engraulis mordax in Puget Sound. The years of lowest smolt mortality coincided with abundant larval and post-larval anchovy during the previous year; and we provide data from telemetered harbor seals and steelhead indicating that the resulting high abundance of age-1+ anchovy provided an alternative prey source for predators of migrating steelhead smolts. Identification of ecological mechanisms that drive patterns in the survival of Endangered Species Act-threatened steelhead provides critical understanding of the systems within which management strategies must operate.