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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How predators respond to changes in prey abundance (i.e., functional responses) is foundational to consumer-resource interactions, predator-prey dynamics, and the stability of predator-prey systems. Predation by piscivorous waterbirds on out-migrating juvenile steelhead trout (Oncorhynchus mykiss) is considered a factor affecting the recovery of multiple Endangered Species Act-listed steelhead populations in the Columbia River basin. Waterbird functional responses, however, may vary by predator species and location, with important implications to predator management strategies. We used a 13-year dataset on waterbird abundance across seven breeding colonies (three Caspian tern [Hydroprogne caspia], two double-crested cormorant [Nannopterum auritum], and two California and ring-billed gull [Larus californicus and Larus delawarensis] colonies) and steelhead tag-recovery data (>645,000 tagged and >32,000 recovered steelhead) to quantify weekly predation probabilities and functional responses across waterbird species, colonies, and years. Weekly predation probabilities were highly variable, ranging from 0.01 to 0.30 at tern colonies, 0.01 to 0.20 at cormorant colonies, and 0.03 to 0.13 at gull colonies.Per capita predation probabilities were an order of magnitude higher at inland tern and cormorant colonies relative to estuary colonies of the same species.Terns displayed Type II functional responses across colonies and years, where predation probabilities peaked at low steelhead abundances and declined as steelhead abundance increased (i.e., predator swamping). Cormorants nesting at the large estuary colony (several thousand birds) displayed a Type III functional response, but cormorants nesting at the smaller inland colony (several hundred birds) displayed a Type II response. Consumption probabilities of steelhead by gulls remained consistent across a large range of steelhead availability, suggesting a Type I or a Type III functional response, but a lack of colony abundance data prevented quantifying functional responses. The level of tern predation combined with Type II functional responses indicate possible

We developed a state-space mark–recapture–recovery model that incorporates multiple recovery types and state uncertainty to estimate survival of an anadromous fish species. We apply the model to a dataset of outmigrating juvenile steelhead trout (Oncorhynchus mykiss (Walbaum, 1792)) tagged with passive integrated transponders, recaptured during outmigration, and recovered on bird colonies in the Columbia River basin (2008–2014). Recoveries on bird colonies are often ignored in survival studies because the river reach of mortality is often unknown, which we model as a form of state uncertainty. Median outmigration survival from release to the lower river (river kilometre 729 to 75) ranged from 0.27 to 0.35, depending on year. Recovery probabilities were frequently ≥0.20 in the first river reach following tagging, indicating that one out of five fish that died in that reach was recovered on a bird colony. Integrating dead recovery data provided increased parameter precision, estimation of where birds consumed fish, and survival estimates across larger spatial scales. More generally, these modeling approaches provide a flexible framework to integrate multiple sources of tag recovery data into mark–recapture studies.

The degree to which predation is an additive vs. compensatory source of mortality is fundamental to understanding the effects of predation on prey populations and evaluating the efficacy of predator management actions. In the Columbia River basin, USA, predation by Caspian Terns (Hydroprogne caspia) on U.S. Endangered Species Act (ESA)listed juvenile salmonids (smolts; Oncorhynchus spp.) has led to predator management actions to reduce predation; however, the assumption that reduced predation translates into greater salmonid survival, either within the life stage where predation occurs or across their lifetime, has remained untested. To address this critical uncertainty, we analyzed a long-term (2008-2018) mark-recapture-recovery data set of ESA-listed steelhead trout (O. mykiss) that were tagged (n = 78,409) and subsequently exposed to predation during smolt out-migration through multiple river reaches (spatial scales), jointly estimating weekly probabilities of steelhead survival, mortality due to bird predation, and mortality due to other causes. This concurrent estimation across time-stratified cohorts allowed for the direct measurement of the strength, magnitude, and direction of relationships between survival and Caspian Tern predation. Estimates of Tern predation on steelhead were substantial in most years, with cumulative annual estimates ranging from 0.075 (95% creditable interval = 0.058-0.099) to 0.375 (0.290-0.461). Increases in Tern predation probabilities were associated with statistically significant decreases in steelhead survival probabilities in all evaluated years and salmonid life stages (smolt out-migration and smolt-to-adult returns). Results provide novel evidence that predation by Caspian Terns may have been a super-additive source of mortality during the smolt life stage and a partially additive source of mortality to the adult life stage. Annual estimates of the difference between observed survival and baseline survival (i.e., in the absence of Tern predation) ranged from 0.052 (0.017-0.103) to 0.314 (0.172-0.459) during the steelhead smolt life stage and from 0.011 (0.001-0.029) to 0.049 (0.025-0.078) to the adult life stage. The estimated levels of compensation have important implications for predator management actions aimed at increasing the survival of endangered salmonids, and the modeling approach developed herein provides a framework to directly quantify the impacts of source-specific mortality factors on prey populations.