Harvesting forage fish can prevent fishing-induced population collapses of large piscivorous fish.

Fisheries have reduced the abundances of large piscivores-such as gadids (cod, pollock, etc.) and tunas-in ecosystems around the world. Fisheries also target smaller species-such as herring, capelin, and sprat-that are important parts of the piscivores' diets. It has been suggested that harvesting of these so-called forage fish will harm piscivores. Multispecies models used for fisheries assessments typically ignore important facets of fish community dynamics, such as individual-level bioenergetics and/or size structure. We test the effects of fishing for both forage fish and piscivores using a dynamic, multitrophic, size-structured, bioenergetics model of the Baltic Sea. In addition, we analyze historical patterns in piscivore-biomass declines and fishing mortalities of piscivores and forage fish using global fish-stock assessment data. Our community-dynamics model shows that piscivores benefit from harvesting of their forage fish when piscivore fishing mortality is high. With substantial harvesting of forage fish, the piscivores can withstand higher fishing mortality. On the other hand, when piscivore fishing mortality is low, piscivore biomass decreases with more fishing of the forage fish. In accordance with these predictions, our statistical analysis of global fisheries data shows a positive interaction between the fishing mortalities of forage-fish stocks and piscivore stocks on the strength of piscivore-biomass declines. While overfishing of forage fish must be prevented, our study shows that reducing fishing pressures on forage fish may have unwanted negative side effects on piscivores. In some cases, decreasing forage-fish exploitation could cause declines, or even collapses, of piscivore stocks.

Population-level effects of acoustic disturbance in Atlantic cod: a size-structured analysis based on energy budgets.

Anthropogenic underwater noise may negatively affect marine animals. Yet, while fishes are highly sensitive to sounds, effects of acoustic disturbances on fishes have not been extensively studied at the population level. In this study, we use a size-structured model based on energy budgets to analyse potential population-level effects of anthropogenic noise on Atlantic cod (Gadus morhua). Using the model framework, we assess the impact of four possible effect pathways of disturbance on the cod population growth rate. Through increased stress, changes in foraging and movement behaviour, and effects on the auditory system, anthropogenic noise can lead to (i) increased energy expenditure, (ii) reduced food intake, (iii) increased mortality, and (iv) reduced reproductive output. Our results show that population growth rates are particularly sensitive to changes in energy expenditure and food intake because they indirectly affect the age of maturation, survival and fecundity. Sub-lethal effects of sound exposure may thus affect populations of cod and fishes with similar life histories more than lethal effects of sound exposure. Moreover, anthropogenic noise may negatively affect populations when causing persistent increases of energy expenditure or decreases of food intake. Effects of specific acoustic pollutants on energy acquisition and expenditure should therefore be further investigated.

Predator Persistence through Variability of Resource Productivity in Tritrophic Systems.

The trophic structure of species communities depends on the energy transfer between trophic levels. Primary productivity varies strongly through time, challenging the persistence of species at higher trophic levels. Yet resource variability has mostly been studied in systems with only one or two trophic levels. We test the effect of variability in resource productivity in a tritrophic model system including a resource, a size-structured consumer, and a size-specific predator. The model complies with fundamental principles of mass conservation and the body-size dependence of individual-level energetics and predator-prey interactions. Surprisingly, we find that resource variability may promote predator persistence. The positive effect of variability on the predator arises through periods with starvation mortality of juvenile prey, which reduces the intraspecific competition in the prey population. With increasing variability in productivity and starvation mortality in the juvenile prey, the prey availability increases in the size range preferred by the predator. The positive effect of prey mortality on the trophic transfer efficiency depends on the biologically realistic consideration of body size-dependent and food-dependent functions for growth and reproduction in our model. Our findings show that variability may promote the trophic transfer efficiency, indicating that environmental variability may sustain species at higher trophic levels in natural ecosystems.

Approximation of a physiologically structured population model with seasonal reproduction by a stage-structured biomass model.

Seasonal reproduction causes, due to the periodic inflow of young small individuals in the population, seasonal fluctuations in population size distributions. Seasonal reproduction furthermore implies that the energetic body condition of reproducing individuals varies over time. Through these mechanisms, seasonal reproduction likely affects population and community dynamics. While seasonal reproduction is often incorporated in population models using discrete time equations, these are not suitable for size-structured populations in which individuals grow continuously between reproductive events. Size-structured population models that consider seasonal reproduction, an explicit growing season and individual-level energetic processes exist in the form of physiologically structured population models. However, modeling large species ensembles with these models is virtually impossible. In this study, we therefore develop a simpler model framework by approximating a cohort-based size-structured population model with seasonal reproduction to a stage-structured biomass model of four ODEs. The model translates individual-level assumptions about food ingestion, bioenergetics, growth, investment in reproduction, storage of reproductive energy, and seasonal reproduction in stage-based processes at the population level. Numerical analysis of the two models shows similar values for the average biomass of juveniles, adults, and resource unless large-amplitude cycles with a single cohort dominating the population occur. The model framework can be extended by adding species or multiple juvenile and/or adult stages. This opens up possibilities to investigate population dynamics of interacting species while incorporating ontogenetic development and complex life histories in combination with seasonal reproduction.