To investigate the cumulative effects of colonial waterbird predation on fish mortality and to determine what proportion of all sources of fish mortality (1 − survival) was due to bird predation, we conducted a mark–recapture–recovery study with upper Columbia River steelhead Oncorhynchus mykiss that were PIT‐tagged and released (N = 78,409) at Rock Island Dam on the Columbia River, USA. We used a state–space Bayesian model that incorporated live detections and dead recoveries of tagged fish to jointly estimate predation and survival probabilities during smolt out‐migration to the Pacific Ocean over an 11‐year study period. Estimated cumulative (all colonies combined) avian predation probabilities ranged from 0.31 (95% credible interval [CRI] = 0.27–0.38) to 0.53 (95% CRI = 0.42–0.64) annually, indicating that avian predation was a substantial source of mortality. Of the predator species evaluated, predation by Caspian terns Hydroprogne caspia was often the highest, with predation probabilities ranging from 0.11 (95% CRI = 0.09–0.14) to 0.38 (95% CRI = 0.29–0.47). Probabilities of predation by double‐crested cormorants Phalacrocorax auritus and mixed colonies of California gulls Larus californicus and ring‐billed gulls L. delawarensis were generally lower than the probabilities for terns but were also substantial, with upwards of 0.04 (95% CRI = 0.03–0.07; cormorants) and 0.31 (95% CRI = 0.25–0.39; gulls) of steelhead consumed. Comparisons of total smolt mortality with mortality due to avian predation indicated that avian predation accounted for 42% (95% CRI = 30–56%) to 70% (95% CRI = 53–87%) of total mortality, suggesting that more steelhead were consumed by avian predators than died from all other mortality sources combined. Results indicate that avian predation, although not the original cause of steelhead declines in the basin, is now a factor limiting the survival of upper Columbia River steelhead. Using the analytical framework developed in this study, future studies can consider the cumulative impact of multiple mortality sources across large spatial and temporal scales to more fully understand the extent to which they limit fish survival.

We evaluated the impact of predation on juvenile steelhead Oncorhynchus mykiss and yearling and subyearling Chinook Salmon O. tshawytscha by piscivorous waterbirds from 11 different breeding colonies in the Columbia River basin during 2012 and 2014. Fish were tagged with both acoustic tags and PIT tags and were tracked via a network of hydrophone arrays to estimate total smolt mortality (1 – survival) at various spatial and temporal scales during out‐migration. Recoveries of PIT tags on bird colonies, coupled with the last known detections of live fish passing hydrophone arrays, were used to estimate the impact of avian predation relative to total smolt mortality. Results indicated that avian predation was a substantial source of steelhead mortality, with predation probability (proportion of available fish consumed by birds) ranging from 0.06 to 0.28 for fish traveling through the lower Snake River and the lower and middle Columbia River. Predation probability estimates ranged from 0.03 to 0.09 for available tagged yearling Chinook Salmon and from 0.01 to 0.05 for subyearlings. Smolt predation by gulls Larus spp. was concentrated near hydroelectric dams, while predation by Caspian terns Hydroprogne caspia was concentrated within reservoirs. No concentrated areas of predation were identified for double‐crested cormorants Phalacrocorax auritus or American white pelicans Pelecanus erythrorhynchos. Comparisons of total smolt mortality relative to mortality from colonial waterbirds indicated that avian predation was one of the greatest sources of mortality for steelhead and yearling Chinook Salmon during out‐migration. In contrast, avian predation on subyearling Chinook Salmon was generally low and constituted a minor component of total mortality. Our results demonstrate that acoustic and PIT tag technologies can be combined to quantify where and when smolt mortality occurs and the fraction of mortality that is due to colonial waterbird predation relative to non‐avian mortality sources. Received November 4, 2015; accepted February 1, 2016 Published online June 27, 2016

We evaluated predation on Lost River Suckers Deltistes luxatus and Shortnose Suckers Chasmistes brevirostris by American white pelicans Pelecanus erythrorhynchos and double‐crested cormorants Phalacrocorax auritus nesting at mixed‐species colonies in the Upper Klamath Basin of Oregon and California during 2009–2014. Predation was evaluated by recovering (detecting) PIT tags from tagged fish on bird colonies and calculating minimum predation rates, as the percentage of available suckers consumed, adjusted for PIT tag detection probabilities but not deposition probabilities (i.e., probability an egested tag was deposited on‐ or off‐colony). Results indicate that impacts of avian predation varied by sucker species, age‐class (adult, juvenile), bird colony location, and year, demonstrating dynamic predator–prey interactions. Tagged suckers ranging in size from 72 to 730 mm were susceptible to cormorant or pelican predation; all but the largest Lost River Suckers were susceptible to bird predation. Minimum predation rate estimates ranged annually from <0.1% to 4.6% of the available PIT‐tagged Lost River Suckers and from <0.1% to 4.2% of the available Shortnose Suckers, and predation rates were consistently higher on suckers in Clear Lake Reservoir, California, than on suckers in Upper Klamath Lake, Oregon. There was evidence that bird predation on juvenile suckers (species unknown) in Upper Klamath Lake was higher than on adult suckers in Upper Klamath Lake, where minimum predation rates ranged annually from 5.7% to 8.4% of available juveniles. Results suggest that avian predation is a factor limiting the recovery of populations of Lost River and Shortnose suckers, particularly juvenile suckers in Upper Klamath Lake and adult suckers in Clear Lake Reservoir. Additional research is needed to measure predator‐specific PIT tag deposition probabilities (which, based on other published studies, could increase predation rates presented herein by a factor of roughly 2.0) and to better understand biotic and abiotic factors that regulate sucker susceptibility to bird predation. Received January 19, 2016; accepted June 25, 2016 Published online October 7, 2016

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