Size-conditional smolting and the response of Carmel River steelhead to two decades of conservation efforts

Threshold effects are common in ecosystems and can generate counterintuitive outcomes in management interventions. A threshold effect proposed for steelhead trout (Oncorhynchus mykiss) is size-conditional smolting and marine survival. Steelhead are anadromous, maturing in the ocean but migrating to freshwater to spawn, where their offspring reside for one or more years before smolting-physiologically transforming to a saltwater form-and migrating to the ocean. In conditional smolting, juveniles transform only if growth exceeds a threshold body size prior to migration season, and subsequent marine survival correlates with size at ocean entry. Conditional smolting suggests that efforts to improve freshwater survival of juveniles may reduce smolt success if they increase competition and reduce growth. Using model-selection techniques, we asked if this effect explained declining numbers of adult Carmel River steelhead. This threatened population has been the focus of two decades of habitat restoration, as well as active translocation and captive-rearing of juveniles stranded in seasonally dewatered channels. In the top-ranked model selected by information-theoretic criteria, adult decline was linked to reduced juvenile growth rates in the lower river, consistent with the conditional smolting hypothesis. According to model inference, since 2005 most returning adult steelhead were captively-reared. However, a lower-ranked model without conditional smolting also had modest support, and suggested a negative effect of captive rearing. Translocations of juvenile fish to perennial reaches may have reduced the steelhead run slightly by raising competition, but this effect is confounded in the data with effects of river flow on growth. Efforts to recover Carmel River steelhead will probably be more successful if they focus on conditions promoting rapid growth in the river. Our analysis clearly favored a role for size-conditional smolting and marine survival in the decline of the population, but did not definitively rule out alternative explanations.