Ecological consequences of body size reduction under warming.

Body size reduction is a universal response to warming, but its ecological consequences across biological levels, from individuals to ecosystems, remain poorly understood. Most biological processes scale with body size, and warming-induced changes in body size can therefore have important ecological consequences. To understand these consequences, we propose a unifying, hierarchical framework for the ecological impacts of intraspecific body size reductions due to thermal plasticity that explicitly builds on three key pathways: morphological constraints, bioenergetic constraints and surface-to-volume ratio. Using this framework, we synthesize key consequences of warming-induced body size reductions at multiple levels of biological organization. We outline how this trait-based framework can improve our understanding, detection and generalization of the ecological impacts of warming.

Local thermal adaption mediates the sensitivity of Daphnia magna to nanoplastics under global warming scenarios.

The toxicity of nanoplastics at environmentally relevant concentrations has received widespread attention in the context of global warming. Despite numerous studies on the impact of mean temperature (MT), the effects of daily temperature fluctuations (DTFs) on the ecotoxicity of nanoplastics remains largely unexplored. Moreover, the role of evolutionary adaptation in assessing long-term ecological risks is unclear. Here, we investigated the effects of polystyrene nanoplastics (5 µg L-1) on Daphnia magna under varying MT (20 °C and 24 °C) and DTFs (0 °C, 5 °C, and 10 °C). Capitalizing on a space-for-time substitution approach, we further assessed how local thermal adaptation affect the sensitivity of Daphnia to nanoplastics under global warming. Our results indicated that nanoplastics exposure in general reduced heartbeat rate, thoracic limb activity and feeding rate, and increased CytP450, ETS activity and Hgb concentrations. Higher MT and DTFs enhanced these effects. Notably, clones originating from their respective sites performed better under their native temperature conditions, indicating local thermal adaptation. Warm-adapted low-latitude D. magna showed stronger nanoplastics-induced increases in CytP450, ETS activity and Hgb concentrations under local MT 24 °C, while cold-adapted high-latitude D. magna showed stronger nanoplastics-induced decreases in heartbeat rate, thoracic limb activity and feeding rate under high MT than under low MT.

Pollution offsets the rapid evolution of increased heat tolerance in a natural population.

Despite the increasing evidence for rapid thermal evolution in natural populations, evolutionary rescue under global warming may be constrained by the presence of other stressors. Highly relevant in our polluted planet, is the largely ignored evolutionary trade-off between heat tolerance and tolerance to pollutants. By using two subpopulations (separated 40 years in time) from a resurrected natural population of the water flea Daphnia magna that experienced a threefold increase in heat wave frequency during this period, we tested whether rapid evolution of heat tolerance resulted in reduced tolerance to the widespread metal zinc and whether this would affect heat tolerance upon exposure to the pollutant. Our results revealed rapid evolution of increased heat tolerance in the recent subpopulation. Notably, the sensitivity to the metal tended to be stronger (reduction in net energy budget) or was only present (reductions in heat tolerance and in sugar content) in the recent subpopulation. As a result, the rapidly evolved higher heat tolerance of the recent subpopulation was fully offset when exposed to zinc. Our results highlight that the many reports of evolutionary rescue to global change stressors may give a too optimistic view as our warming planet is polluted by metals and other pollutants.

Rapid evolution of consumptive and non-consumptive predator effects on prey population densities, bioenergetics and stoichiometry.

Predators can strongly influence prey populations not only through consumptive effects (CE) but also through non-consumptive effects (NCE) imposed by predation risk. Yet, the impact of NCE on bioenergetic and stoichiometric body contents of prey, traits that are shaping life histories, population and food web dynamics, is largely unknown. Moreover, the degree to which NCE can evolve and can drive evolution in prey populations is rarely studied. A 6-week outdoor mesocosm experiment with Caged-Fish (NCE) and Free-Ranging-Fish (CE and NCE) treatments was conducted to quantify and compare the effects of CE and NCE on population densities, bioenergetic and stoichiometric body contents of Daphnia magna, a keystone species in freshwater ecosystems. We tested for evolution of CE and NCE by using experimental populations consisting of D. magna clones from two periods of a resurrected natural pond population: a pre-fish period without fish and a high-fish period with high predation pressure. Both Caged-Fish and Free-Ranging-Fish treatments decreased the body size and population densities, especially in Daphnia from the high-fish period. Only the Free-Ranging-Fish treatment affected bioenergetic variables, while both the Caged-Fish and Free-Ranging-Fish treatments shaped body stoichiometry. The effects of CE and NCE were different between both periods indicating their rapid evolution in the natural resurrected population. Both the Caged-Fish and Free-Ranging-Fish treatments changed the clonal frequencies of the experimental Daphnia populations of the pre-fish as well as the high-fish period, indicating that not only CE but also NCE induced clonal sorting, hence rapid evolution during the mesocosm experiment in both periods. Our results demonstrate that CE as well as NCE have the potential to change not only the body size and population density but also the bioenergetic and stoichiometric characteristics of prey populations. Moreover, we show that these responses not only evolved in the studied resurrected population, but that CE and NCE also caused differential rapid evolution in a time frame of 6 weeks (ca. four to six generations). As NCE can evolve as well as can drive evolution, they may play an important role in shaping eco-evolutionary dynamics in predator-prey interactions.

Microgeographic differentiation in thermal and antipredator responses and their carry-over effects across life stages in a damselfly.

Global warming and invasive species, separately or combined, can impose a large impact on the condition of native species. However, we know relatively little about how these two factors, individually and in combination, shape phenotypes in ectotherms across life stages and how this can differ between populations. We investigated the non-consumptive predator effects (NCEs) imposed by native (perch) and invasive (signal crayfish) predators experienced only during the egg stage or during both the egg and larval stages in combination with warming on adult life history traits of the damselfly Ischnura elegans. To explore microgeographic differentiation, we compared two nearby populations differing in thermal conditions and predator history. In the absence of predator cues, warming positively affected damselfly survival, possibly because the warmer temperature was closer to the optimal temperature. In the presence of predator cues, warming decreased survival, indicating a synergistic effect of these two variables on survival. In one population, predator cues from perch led to increased survival, especially under the current temperature, likely because of predator stress acclimation phenomena. While warming decreased, predator cues increased larval development time with a proportionally stronger effect of signal crayfish cues experienced during the egg stage, indicating a negative carry-over effect from egg to larva. Warming and predator cues increased mass at emergence, with the predator effect driven mainly by exposure to signal crayfish cues during the egg stage, indicating a positive carry-over effect from egg to adult. Notably, warming and predator effects were not consistent across the two studied populations, suggesting a phenotypic signal of adaptation at a microgeographic scale to thermal conditions and predator history. We also observed pronounced shifts during ontogeny from synergistic (egg and early larval stage) toward additive (late larval stage up to emergence) effects between warming and predator stress. The results point out that population- and life-stage-specific responses in life-history traits to NCEs are needed to predict fitness consequences of exposure to native and invasive predators and warming in prey at a microgeographic scale.

When and how can we predict adaptive responses to climate change?

Predicting if, when, and how populations can adapt to climate change constitutes one of the greatest challenges in science today. Here, we build from contributions to the special issue on evolutionary adaptation to climate change, a survey of its authors, and recent literature to explore the limits and opportunities for predicting adaptive responses to climate change. We outline what might be predictable now, in the future, and perhaps never even with our best efforts. More accurate predictions are expected for traits characterized by a well-understood mapping between genotypes and phenotypes and traits experiencing strong, direct selection due to climate change. A meta-analysis revealed an overall moderate trait heritability and evolvability in studies performed under future climate conditions but indicated no significant change between current and future climate conditions, suggesting neither more nor less genetic variation for adapting to future climates. Predicting population persistence and evolutionary rescue remains uncertain, especially for the many species without sufficient ecological data. Still, when polled, authors contributing to this special issue were relatively optimistic about our ability to predict future evolutionary responses to climate change. Predictions will improve as we expand efforts to understand diverse organisms, their ecology, and their adaptive potential. Advancements in functional genomic resources, especially their extension to non-model species and the union of evolutionary experiments and "omics," should also enhance predictions. Although predicting evolutionary responses to climate change remains challenging, even small advances will reduce the substantial uncertainties surrounding future evolutionary responses to climate change.

Plasticity and associated epigenetic mechanisms play a role in thermal evolution during range expansion.

Due to global change, many species are shifting their distribution and are thereby confronted with novel thermal conditions at the moving range edges. Especially during the initial phases of exposure to a new environment, it has been hypothesized that plasticity and associated epigenetic mechanisms enable species to cope with environmental change. We tested this idea by capitalizing on the well-documented southward range expansion of the damselfly Ischnura elegans from France into Spain where the species invaded warmer regions in the 1950s in eastern Spain (old edge region) and in the 2010s in central Spain (new edge region). Using a common garden experiment at rearing temperatures matching the ancestral and invaded thermal regimes, we tested for evolutionary changes in (thermal plasticity in) larval life history and heat tolerance in these expansion zones. Through the use of de- and hypermethylating agents, we tested whether epigenetic mechanisms play a role in enabling heat tolerance during expansion. We used the phenotype of the native sister species in Spain, I. graellsii, as proxy for the locally adapted phenotype. New edge populations converged toward the phenotype of the native species through plastic thermal responses in life history and heat tolerance while old edge populations (partly) constitutively evolved a faster life history and higher heat tolerance than the core populations, thereby matching the native species. Only the heat tolerance of new edge populations increased significantly when exposed to the hypermethylating agent. This suggests that the DNA methylation machinery is more amenable to perturbation at the new edge and shows it is able to play a role in achieving a higher heat tolerance. Our results show that both (evolved) plasticity as well as associated epigenetic mechanisms are initially important when facing new thermal regimes but that their importance diminishes with time.

Microplastics increases the heat tolerance of Daphnia magna under global warming via hormetic effects.

The ecological risk assessment of microplastics under global warming receives increasing attention. Yet, such studies mostly focused on increased mean temperatures (MT), ignoring another key component of global warming, namely daily temperature fluctuations (DTF). Moreover, we know next to nothing about the combined effects of multigenerational exposure to microplastics and warming. In this study, Daphnia magna was exposed to an environmentally relevant concentration of polystyrene microplastics (5 µg L-1) under six thermal conditions (MT: 20 â„ƒ, 24 â„ƒ; DTF: 0 â„ƒ, 5 â„ƒ, 10 â„ƒ) over two generations to investigate the interactive effects of microplastics and global warming. Results showed that microplastics had no effects on Daphnia at standard thermal conditions (constant 20 °C). Yet, microplastics increased the fecundity, heat tolerance, amount of energy storage, net energy budget and cytochrome P450 activity, and decreased the energy consumption when tested under an increased MT or DTF, indicating a hormesis effect induced by microplastics under warming. The unexpected increase in heat tolerance upon exposure to microplastics could be partly explained by the reduced energy consumption and/or increased energy availability. Overall, the present study highlighted the importance of including DTF and multigenerational exposure to improve the ecological risk assessment of microplastics under global warming.