Fish life histories in a warming climate: a mechanistic basis of change and a community context

Body size dependent interactions structure food webs, and these are changing with climate warming. We cannot yet predict how warming affects many aspects of life history evolution and species ecology, despite a longstanding interest in the structuring effects of temperature and body size in food webs. This is in part due to not recognizing the temperature dependence two aspects, namely that 1) withinspecies differences govern species interactions and 2) processes of adaptation depend on body size. In this thesis, I assess how body size dependent effects of temperature govern such interactions and processes using theoretical models of individual growth and reproduction. First, I study effects of warming on the energy allocation trade-off between somatic growth and energy reserves and find that warming favours allocation to reserves and reproduction through increasing importance of early life history processes. Specifically, failing to adapt to warming winters compromises viability of population through juvenile mortality. Second, I study how effects of warming on consumer-resource systems depend on energy allocation strategies. Here, energy allocation can modulate temperature dependent competition for food between stages, but competition mediated by diet is the main determinant of effects of warming. Last, I show how effects of warming affect the feedback mechanism of stage dependent competition and predation on interacting species and thus prevent adults from cultivating a low competition environment for their young. I conclude that linking underlying individual body size dependent physiological responses to warming to effects in population and communities provides novel mechanistic understanding of adaptation and food web processes. While these mechanistic predictions form a basis for, and require, empirical tests, I propose that diversity and function of aquatic food webs are at stake

Effects of Warming on Intraguild Predator Communities with Ontogenetic Diet Shifts

Species interactions mediate how warming affects community composition via individual growth and population size structure. While predictions on how warming affects composition of size- or stage-structured communities have so far focused on linear (food chain) communities, mixed competition-predation interactions, such as intraguild predation, are common. Intraguild predation often results from changes in diet over ontogeny ("ontogenetic diet shifts") and strongly affects community composition and dynamics. Here, we study how warming affects a community of intraguild predators with ontogenetic diet shifts, consumers, and shared prey by analyzing a stage-structured bioenergetics multispecies model with temperature- and body size-dependent individual-level rates. We find that warming can strengthen competition and decrease predation, leading to a loss of a cultivation mechanism (the feedback between predation on and competition with consumers exerted by predators) and ultimately predator collapse. Furthermore, we show that the effect of warming on community composition depends on the extent of the ontogenetic diet shift and that warming can cause a sequence of community reconfigurations in species with partial diet shifts. Our findings contrast previous predictions concerning individual growth of predators and the mechanisms behind predator loss in warmer environments and highlight how feedbacks between temperature and intraspecific size structure are important for understanding such effects on community composition

Optimal energy allocation trade-off driven by size-dependent physiological and demographic responses to warming

Body size-dependent physiological effects of temperature influence individual growth, reproduction, and survival, which govern animal population responses to global warming. Considerable knowledge has been established on how such effects can affect population growth and size structure, but less is known of their potential role in temperature-driven adaptation in life-history traits. In this study, we ask how warming affects the optimal allocation of energy between growth and reproduction and disentangle the underlying fitness trade-offs. To this end, we develop a novel dynamic energy budget integral projection model (DEB-IPM), linking individuals' size- and temperature-dependent consumption and maintenance via somatic growth, reproduction, and size-dependent energy allocation to emergent population responses. At the population level, we calculate the long-term population growth rate (fitness) and stable size structure emerging from demographic processes. Applying the model to an example of pike (Esox lucius), we find that optimal energy allocation to growth decreases with warming. Furthermore, we demonstrate how growth, fecundity, and survival contribute to this change in optimal allocation. Higher energy allocation to somatic growth at low temperatures increases fitness through survival of small individuals and through the reproduction of larger individuals. In contrast, at high temperatures, increased allocation to reproduction is favored because warming induces faster somatic growth of small individuals and increased fecundity but reduced growth and higher mortality of larger individuals. Reduced optimum allocation to growth leads to further reductions in body size and an increasingly truncated population size structure with warming. Our study demonstrates how, by incorporating general physiological mechanisms driving the temperature dependence of life-history traits, the DEB-IPM framework is useful for investigating the adaptation of size-structured organisms to warming

Compensatory Feeding in Eastern Baltic Cod (Gadus morhua) : Recent Shifts in Otolith Growth and Nitrogen Content Suggest Unprecedented Metabolic Changes

The productivity of the Eastern Baltic cod (EBC) has been severely reduced over the last 25 years, for reasons that remain unclear. The size distribution of EBC has become increasingly truncated, condition and health status have deteriorated, and sexual maturation has started to occur at increasingly smaller sizes. Despite an increasing trend in recruitment during this period, reduced growth or increased mortality rates after the recruitment phase have resulted in decreasing landing levels and low profitability in the cod fishery, whereas the scientific community has difficulties in disentangling the causes of the decline of EBC. We studied changes in metabolic status in EBC between the capture years of 1995 and 2015, by investigating two aspects of fish metabolism that can be extracted retrospectively from otolith (earstone) morphometry and nitrogen content. Changes in relative otolith size to fish size are related to the metabolic history of the individual fish, and the otolith nitrogen content reveals the level of protein synthesis and feeding rate. Because otoliths accrue continuously on their surface and are biological stable (inert), the chemical content of the otolith trajectory reflects the timeline of the fish. We measured the N/Ca ratio as a proxy for protein content in EBC otolith along distal radius traverses from the core to the edge of the otolith by using secondary ion mass spectrometry (SIMS). Here we show that the otoliths have become smaller at a given fish size, and the ratio of N/Ca has increased over the studied period. These proxies reveal significant metabolic changes during the same period as the condition, and stock productivity has declined. We discuss potential mechanisms behind the metabolic changes, including elevated temperature and compensatory feeding due to nutrient deficiencies. Such changes in food quality may, in turn, relate to still unrecognized but on-going ecosystem shifts, where climate change could be the ultimate driver.Correction in: Frontiers in Marine Sciende, vol. 10, article ID 1154309DOI: 10.3389/fmars.2023.1154309</p