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Anthropogenic activities can lead to changes in animal behavior. Predicting population consequences of these behavioral changes requires integrating short-term individual responses into models that forecast population dynamics across multiple generations. This is especially challenging for long-lived animals, because of the different time scales involved. Beaked whales are a group of deep-diving odontocete whales that respond behaviorally when exposed to military mid-frequency active sonar (MFAS), but the effect of these nonlethal responses on beaked whale populations is unknown. Population consequences of aggregate exposure to MFAS was assessed for two beaked whale populations that are regularly present on U.S. Navy training ranges where MFAS is frequently used. Our approach integrates a wide range of data sources, including telemetry data, information on spatial variation in habitat quality, passive acoustic data on the temporal pattern of sonar use and its relationship to beaked whale foraging activity, into an individual-based model with a dynamic bioenergetic module that governs individual life history. The predicted effect of disturbance from MFAS on population abundance ranged between population extinction to a slight increase in population abundance. These effects were driven by the interaction between the temporal pattern of MFAS use, baseline movement patterns, the spatial distribution of prey, the nature of beaked whale behavioral response to MFAS and the top-down impact of whale foraging on prey abundance. Based on these findings, we provide recommendations for monitoring of marine mammal populations and highlight key uncertainties to help guide future directions for assessing population impacts of nonlethal disturbance for these and other long-lived animals., Competing Interests: I have read the journal’s policy and the authors of this manuscript have the following competing interests: This research was funded by the Office of Naval Research (ONR), a division of the US Navy. None of the analysis nor reporting involved or was influenced by the sponsors of the research. Due to security reasons, the sonar data supporting this research are not accessible to the public., (Copyright: This is an open access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.)

To measure organismal coherence in a pelagic ecosystem, we used moored sensors to describe the vertical dynamics of each step in the food chain in shelf waters off the west shore of Oahu, Hawaii. Horizontally extensive, intense aggregations of phytoplankton, zooplankton, and micronekton exhibited strong diel patterns in abundance and vertical distribution, resulting in a highly variable potential for interaction amongst trophic levels. Only around dusk did zooplankton layers overlap with phytoplankton layers. Shortly after sunset, micronekton ascended from the deep, aggregating on the island's shelf. Short-lived departures in migration patterns were detected in depth, vertical distribution, density, and total abundance of micronekton when zooplankton layers were present with typical patterns resuming within one hour. Layers of zooplankton began to disappear within 20 minutes of the arrival of micronekton with no layers present after 50 minutes. The effects of zooplankton layers cascaded even further up the food chain, affecting many behaviors of dolphins observed at dusk including their depth, group size, and inter-individual spacing. As a result of these changes in behavior, during a 30-minute window just after dusk, the number of feeding events observed for each dolphin and consequently the feeding time for each individual more than doubled when zooplankton layers were present. Dusk is a critical period for interactions amongst species in this system from phytoplankton to top predators. Our observations that short time windows can drive the structure and function of a complex suite of organisms highlight the importance of explicitly adding a temporal dimension at a scale relevant to individual organisms to our descriptions of heterogeneity in ocean ecosystems.

Spatial coherence between predators and prey has rarely been observed in pelagic marine ecosystems. We used measures of the environment, prey abundance, prey quality, and prey distribution to explain the observed distributions of three co-occurring predator species breeding on islands in the southeastern Bering Sea: black-legged kittiwakes (Rissa tridactyla), thick-billed murres (Uria lomvia), and northern fur seals (Callorhinus ursinus). Predictions of statistical models were tested using movement patterns obtained from satellite-tracked individual animals. With the most commonly used measures to quantify prey distributions--areal biomass, density, and numerical abundance--we were unable to find a spatial relationship between predators and their prey. We instead found that habitat use by all three predators was predicted most strongly by prey patch characteristics such as depth and local density within spatial aggregations. Additional prey patch characteristics and physical habitat also contributed significantly to characterizing predator patterns. Our results indicate that the small-scale prey patch characteristics are critical to how predators perceive the quality of their food supply and the mechanisms they use to exploit it, regardless of time of day, sampling year, or source colony. The three focal predator species had different constraints and employed different foraging strategies--a shallow diver that makes trips of moderate distance (kittiwakes), a deep diver that makes trip of short distances (murres), and a deep diver that makes extensive trips (fur seals). However, all three were similarly linked by patchiness of prey rather than by the distribution of overall biomass. This supports the hypothesis that patchiness may be critical for understanding predator-prey relationships in pelagic marine systems more generally.