A challenge facing ecologists trying to predict responses to climate change is the few recent analogous conditions to use for comparison. For example, negative relationships between ectotherm body size and temperature are common both across natural thermal gradients and in small‐scale experiments. However, it is unknown if short‐term body size responses are representative of long‐term responses. Moreover, to understand population responses to warming, we must recognize that individual responses to temperature may vary over ontogeny. To enable predictions of how climate warming may affect natural populations, we therefore ask how body size and growth may shift in response to increased temperature over life history, and whether short‐ and long‐term growth responses differ. We addressed these questions using a unique setup with multidecadal artificial heating of an enclosed coastal bay in the Baltic Sea and an adjacent reference area (both with unexploited populations), using before‐after control‐impact paired time‐series analyses. We assembled individual growth trajectories of ~13,000 unique individuals of Eurasian perch and found that body growth increased substantially after warming, but the extent depended on body size: Only among small‐bodied perch did growth increase with temperature. Moreover, the strength of this response gradually increased over the 24 year warming period. Our study offers a unique example of how warming can affect fish populations over multiple generations, resulting in gradual changes in body growth, varying as organisms develop. Although increased juvenile growth rates are in line with predictions of the temperature–size rule, the fact that a larger body size at age was maintained over life history contrasts to that same rule. Because the artificially heated area is a contemporary system mimicking a warmer sea, our findings can aid predictions of fish responses to further warming, taking into account that growth responses may vary both over an individual's life history and over time.

Trophic cascades occur in many ecosystems, but the factors regulating them are still elusive. We suggest that an overlooked factor is that trophic interactions (TIs) are often scale-dependent and possibly interact across spatial scales. To explore the role of spatial scale for trophic cascades, and particularly the occurrence of cross-scale interactions (CSIs), we collected and analysed food-web data from 139 stations across 32 bays in the Baltic Sea. We found evidence of a four-level trophic cascade linking TIs across two spatial scales: at bay scale, piscivores (perch and pike) controlled mesopredators (three-spined stickleback), which in turn negatively affected epifaunal grazers. At station scale (within bays), grazers on average suppressed epiphytic algae, and indirectly benefitted habitat-forming vegetation. Moreover, the direction and strength of the grazer-algae relationship at station scale depended on the piscivore biomass at bay scale, indicating a cross-scale interaction effect, potentially caused by a shift in grazer assemblage composition. In summary, the trophic cascade from piscivores to algae appears to involve TIs that occur at, but also interact across, different spatial scales. Considering scale-dependence in general, and CSIs in particular, could therefore enhance our understanding of trophic cascades.

Citation: Jacobson, P., A. G ardmark, J. € Ostergren, M. Casini, and M. Huss. 2018. Size-dependent prey availability affects diet and performance of predatory fish at sea: a case study of Atlantic salmon. Ecosphere 9(1):e02081. 10.1002/ecs2.2081Abstract. Identifying factors determining the performance of individuals is an essential part of resolving what drives population dynamics. For species undergoing ontogenetic shifts in resource and habitat use, this entails assessing individual performance in all habitats used. Whereas survival and growth of anadromous Atlantic salmon, Salmo salar L., in its juvenile, river habitat are known to depend on size-dependent foraging and food availability, individual performance of salmon in the growth habitat out at sea is commonly explained only by abiotic factors. Still, individuals undergo this habitat shift to grow large, suggesting performance should be food-dependent also in the growth habitat. Because fish communities are highly size-structured, the link between predators and their prey may depend on their respective body sizes. Here, we study whether the performance of Baltic Sea salmon in its growth habitat is food-and sizedependent, by combining extensive diet and body size data of Baltic salmon with spatially resolved monitoring data on abundance and size distribution of their main prey, herring, Clupea harengus L., and sprat, Sprattus sprattus L. We found that both the species and size composition of prey in the diet varied with salmon body size. By accounting for this size-dependent predation and the spatially varying size distribution of prey species, we could explain the variation in salmon diet composition among salmon individuals in different Baltic Sea basins and of different length. The proportion of sprat in diet of salmon was better explained by size-specific prey availability (SSP) than total prey biomass, especially for small salmon. Further, salmon body condition increased with SSP, whereas total prey biomass could not explain variation in the condition of salmon. These findings demonstrate that food-and size-dependent processes indeed can influence the performance of anadromous fish also in large marine systems. Thus, we argue that consideration of these processes, stretching across habitats, is important for understanding performance and dynamics of predatory fish in open aquatic systems, as well as for successful management of species such as Atlantic salmon.

Population-specific assessment and management of anadromous fish at sea requires detailed information about the distribution at sea over ontogeny for each population.However, despite a long history of mixed-stock sea fisheries on Atlantic salmon, Salmo salar, migration studies showing that some salmon populations feed in different regions of the Baltic Sea and variation in dynamics occurs among populations feeding in the Baltic Sea, such information is often lacking. Also, current assessment of Baltic salmon assumes equal distribution at sea and therefore equal responses to changes in off-shore sea fisheries. Here, we test for differences in distribution at sea among and within ten Atlantic salmon Salmo salar populations originating from ten river-specific hatcheries along the Swedish Baltic Sea coast, using individual data from >125,000 tagged salmon, recaptured over five decades. We show strong population and size-specific differences in distribution at sea, varying between year classes and between individuals within year classes. This suggests that Atlantic salmon in the Baltic Sea experience great variation in environmental conditions and exploitation rates over ontogeny depending on origin and that current assessment assumptions about equal exploitation rates in the offshore fisheries and a shared environment at sea are not valid. Thus, our results provide additional arguments and necessary information for implementing population-specific management of salmon, also when targeting life stages at sea. K E Y W O R D Sanadromy, Atlantic salmon, Baltic Sea, body size, population-specific management, spatial distribution

Understanding the processes shaping the dynamics of anadromous fish populations is essential for their management and conservation. Yet, little is known about how variation in performance at sea affects their population dynamics. Here we show that variation in body growth at sea contributes to explaining variation in the reproductive potential for 2 Atlantic salmon Salmo salar populations, but to a varying extent. To this end, we assembled data collected during 50 yr for 2 Baltic salmon populations of hatchery origin, including annually released smolts, survival at sea estimates, size-specific growth at sea, annual length distributions of returning adult females and their reproductive potential. The regression models fitted to explain the reproductive potential of our 2 study populations improved when growth at sea was included as an explanatory variable, in addition to smolt year class abundance and estimates of their survival at sea. This link between body growth at sea and population-level reproductive potential suggests that growth at sea can be important to consider when resolving variation in recovery and dynamics among salmon populations sharing the same sea.