Freshening rather than warming drives trematode transmission from periwinkles to mussels

In the Western Baltic Sea, climate change is happening at much faster rate than in most other seas and organisms are additionally exposed to a steep and variable salinity gradient. Climate change has previously been shown to affect parasite transmission in other marine ecosystems, yet little is known about potential effects of warming and desalination on parasite–host interactions. In laboratory experiments, we determined the combined effects of projected seawater warming and freshening on the emergence, activity, survival, and infectivity of cercariae (free-swimming infectious stage) of the trematode Himasthla elongata (Mehlis 1831), shed from its first intermediate host, the periwinkle Littorina littorea (Linnaeus 1758), in the Baltic Sea. We also assessed the susceptibility of the second intermediate host, the mussel Mytilus edulis Linnaeus, 1758, to cercarial infections. Generally, salinity was the main driver, particularly of cercarial activity, infectivity, and mussel susceptibility to infection. At the lowest salinity (13), cercariae were 50% less active compared to the highest salinity (19). Infection success and host susceptibility followed a similar pattern, with 47% and 43% less metacercariae (encysted stage) present at salinity 13 than at salinity 19, respectively. In contrast, effects of simulated warming were found only for cercarial survival, with cercarial longevity being higher at 19 than at 23 °C. No significant interactions between temperature and salinity were found. In contrast to the literature, the results suggest that a climate change-driven freshening (partly also warming) may lead to a general decline of marine trematodes, with possible beneficial effects for the involved hosts

Editorial: Influence of environmental variability on climate change impacts in marine ecosystems

multiple drivers, environmental variability, Climate change, marine heatwaves, stressmemory, Ecological memory, Thermal performance curves, acclimatio

Cyclic thermal fluctuations can be burden or relief for an ectotherm depending on fluctuations’ average and amplitude

1. Predicting the implications of ongoing ocean climate warming demands a better understanding of how short-term thermal variability impacts marine ectotherms, particularly at beyond-optimal average conditions during summer heatwaves. 2. Using a globally important model species, the blue mussel Mytilus, in a 5-week-long experiment, we (a) assessed growth performance traits under 12 scenarios, consisting of four thermal averages (18.5, 21, 23.5 and 26℃) imposed as constant or daily fluctuating regimes with amplitudes of 2 or 4℃. Additionally, we conducted a short-term assay using different mussel individuals to (b) test for the species capacity for suppression and recovery of metabolic performance traits (feeding and aerobic respiration) when exposed to a 1-day thermal fluctuation regime (16.8–30.5℃). Using this high-resolution data, we (c) generated short-term thermal metabolic performance curves to predict and explain growth responses observed in the long-term experiment. 3. We found that daily high-amplitude thermal cycles (4℃) improved mussel growth when fluctuations were imposed around an extreme average temperature of 26℃, representing end-of-century heatwaves. In contrast, thermal cycles negatively affected mussel growth at a less extreme average temperature of 23.5℃, resembling current peak summer temperature scenarios. These results suggest that fluctuations ameliorate heat stress impacts only at critically high average temperatures. The short-term assay demonstrated that during the warming phase, animals stopped feeding between 24 and 30℃ while gradually suppressing respiration. In the subsequent cooling phase, feeding and respiration partially and fully recovered to pre-heating rates respectively. Furthermore, nonlinear averaging of short-term feeding responses (upscaling) well-predicted longer term growth responses to fluctuations. 4. Our findings suggest that fluctuations can be beneficial to or detrimental for the long-term performance of ectothermic animals, depending on the fluctuations' average and amplitude. Furthermore, the observed effects can be linked to fluctuation-mediated metabolic suppression and recovery. In a general framework, we propose various hypothetical scenarios of fluctuation impacts on ectotherm performance considering inter- or intra-species variability in heat sensitivity. Our research highlights the need for studying metabolic performance in relation to cyclic abiotic fluctuations to advance the understanding of climate change impacts on aquatic systems

Warming and temperature variability determine the performance of two invertebrate predators

in a warming ocean, temperature variability imposes intensified peak stress, but offers periods of stress release. While field observations on organismic responses to heatwaves are emerging, experimental evidence is rare and almost lacking for shorter-scale environmental variability. For two major invertebrate predators, we simulated sinusoidal temperature variability (±3 °C) around todays’ warm summer temperatures and around a future warming scenario (+4 °C) over two months, based on high-resolution 15-year temperature data that allowed implementation of realistic seasonal temperature shifts peaking midpoint. Warming decreased sea stars’ (Asterias rubens) energy uptake (Mytilus edulis consumption) and overall growth. Variability around the warming scenario imposed additional stress onto Asterias leading to an earlier collapse in feeding under sinusoidal fluctuations. High-peak temperatures prevented feeding, which was not compensated during phases of stress release (low-temperature peaks). In contrast, increased temperatures increased feeding on Mytilus but not growth rates of the recent invader Hemigrapsus takanoi, irrespective of the scale at which temperature variability was imposed. This study highlights species-specific impacts of warming and identifies temperature variability at the scale of days to weeks/months as important driver of thermal responses. When species’ thermal limits are exceeded, temperature variability represents an additional source of stress as seen from future warming scenarios

Population structure of the recent invader Hemigrapsus takanoi and prey size selection on Baltic Sea mussels

The shore crab Hemigrapsus takanoi Asakura and Watanabe, 2005, native to the Northwest Pacific, was recorded in European waters about 25 years ago and it was first found in the Baltic Sea in 2014. Information on population structure of invaders and their new niche is needed in order to understand their biological impact. Over one year, we assessed temporal changes in relative abundance, size-class and sex ratio, as well as breeding season of H. takanoi in the Kiel Fjord (Western Baltic Sea). In addition, prey size preference and consumption rates on mussels (Mytilus edulis Linnaeus, 1758) were experimentally assessed in spring, summer and autumn. A total of 596 individuals were collected with highest and lowest abundances in June and February, respectively. Females were dominant over males (sex ratio 1.4:1), but males grew to larger sizes. H. takanoi reproduced between June and August with ovigerous females representing 30% of the entire female abundance registered over the entire year. Males were able to open larger mussels (due to larger claws) and consumed twice as many mussels when compared to females of similar size. Consumption rates for males were 6 and 2 times higher in summer (seawater temperature of 19 °C) compared to spring (8 °C) and autumn (13 °C), respectively. Females consumed 3 times more mussels in autumn than in spring. H. takanoi is an active predator, capable of reproduction in stressful brackish water conditions. Due to large abundances and high feeding pressure, this recently introduced species could play a key role in structuring post-settlement population dynamics of the dominant habitat builder M. edulis

Response of foundation macrophytes to near‐natural simulated marine heatwaves

Marine heatwaves have been observed worldwide and are expected to increase in both frequency and intensity due to climate change. Such events may cause ecosystem reconfigurations arising from species range contraction or redistribution, with ecological, economic and social implications. Macrophytes such as the brown seaweed Fucus vesiculosus and the seagrass Zostera marina are foundation species in many coastal ecosystems of the temperate northern hemisphere. Hence, their response to extreme events can potentially determine the fate of associated ecosystems. Macrophyte functioning is intimately linked to the maintenance of photosynthesis, growth and reproduction, and resistance against pathogens, epibionts and grazers. We investigated morphological, physiological, pathological and chemical defence responses of western Baltic Sea F. vesiculosus and Z. marina populations to simulated near‐natural marine heatwaves. Along with (a) the control, which constituted no heatwave but natural stochastic temperature variability (0HW), two treatments were applied: (b) two late‐spring heatwaves (June, July) followed by a summer heatwave (August; 3HW) and (c) a summer heatwave only (1HW). The 3HW treatment was applied to test whether preconditioning events can modulate the potential sensitivity to the summer heatwave. Despite the variety of responses measured in both species, only Z. marina growth was impaired by the accumulative heat stress imposed by the 3HW treatment. Photosynthetic rate, however, remained high after the last heatwave indicating potential for recovery. Only epibacterial abundance was significantly affected in F. vesiculosus. Hence both macrophytes, and in particular F. vesiculosus, seem to be fairly tolerant to short‐term marine heatwaves at least at the intensities applied in this experiment (up to 5°C above mean temperature over a period of 9 days). This may partly be due to the fact that F. vesiculosus grows in a highly variable environment, and may have a high phenotypic plasticity

Environmental variability in aquatic ecosystems: avenues for future multifactorial experiments

The relevance of considering environmental variability for understanding and predicting biological responses to environmental changes has resulted in a recent surge in variability-focused ecological research. However, integration of findings that emerge across studies and identification of remaining knowledge gaps in aquatic ecosystems remain critical. Here, we address these aspects by: (1) summarizing relevant terms of variability research including the components (characteristics) of variability and key interactions when considering multiple environmental factors; (2) identifying conceptual frameworks for understanding the consequences of environmental variability in single and multi-factorial scenarios; (3) highlighting challenges for bridging theoretical and experimental studies involving transitioning from simple to more complex scenarios; (4) proposing improved approaches to overcome current mismatches between theoretical predictions and experimental observations; and (5) providing a guide for designing integrated experiments across multiple scales, degrees of control, and complexity in light of their specific strengths and limitations

Response of foundation macrophytes to near‐natural simulated marine heatwaves

Marine heatwaves have been observed worldwide and are expected to increase in both frequency and intensity due to climate change. Such events may cause ecosystem reconfigurations arising from species range contraction or redistribution, with ecological, economic and social implications. Macrophytes such as the brown seaweed Fucus vesiculosus and the seagrass Zostera marina are foundation species in many coastal ecosystems of the temperate northern hemisphere. Hence, their response to extreme events can potentially determine the fate of associated ecosystems. Macrophyte functioning is intimately linked to the maintenance of photosynthesis, growth and reproduction, and resistance against pathogens, epibionts and grazers. We investigated morphological, physiological, pathological and chemical defence responses of western Baltic Sea F. vesiculosus and Z. marina populations to simulated near‐natural marine heatwaves. Along with (a) the control, which constituted no heatwave but natural stochastic temperature variability (0HW), two treatments were applied: (b) two late‐spring heatwaves (June, July) followed by a summer heatwave (August; 3HW) and (c) a summer heatwave only (1HW). The 3HW treatment was applied to test whether preconditioning events can modulate the potential sensitivity to the summer heatwave. Despite the variety of responses measured in both species, only Z. marina growth was impaired by the accumulative heat stress imposed by the 3HW treatment. Photosynthetic rate, however, remained high after the last heatwave indicating potential for recovery. Only epibacterial abundance was significantly affected in F. vesiculosus. Hence both macrophytes, and in particular F. vesiculosus, seem to be fairly tolerant to short‐term marine heatwaves at least at the intensities applied in this experiment (up to 5°C above mean temperature over a period of 9 days). This may partly be due to the fact that F. vesiculosus grows in a highly variable environment, and may have a high phenotypic plasticit

Extreme variations of pCO2 and pH in a macrophyte meadow of the Baltic Sea in summer: evidence of the effect of photosynthesis and local upwelling

The impact of ocean acidification on benthic habitats is a major preoccupation of the scientific community. However, the natural variability of pCO2 and pH in those habitats remains understudied, especially in temperate areas. In this study we investigated temporal variations of the carbonate system in nearshore macrophyte meadows of the western Baltic Sea. These are key benthic ecosystems, providing spawning and nursery areas as well as food to numerous commercially important species. In situ pCO2, pH (total scale), salinity and PAR irradiance were measured with a continuous recording sensor package dropped in a shallow macrophyte meadow (Eckernförde bay, western Baltic Sea) during three different weeks in July (pCO2 and PAR only), August and September 2011.The mean (± SD) pCO2 in July was 383±117 µatm. The mean (± SD) pCO2 and pHtot in August were 239±20 µatm and 8.22±0.1, respectively. The mean (± SD) pCO2 and pHtot in September were 1082±711 µatm and 7.83±0.40, respectively. Daily variations of pCO2 due to photosynthesis and respiration (difference between daily maximum and minimum) were of the same order of magnitude: 281±88 µatm, 219±89 μatm and 1488±574 µatm in July, August and September respectively. The observed variations of pCO2 were explained through a statistical model considering wind direction and speed together with PAR irradiance. At a time scale of days to weeks, local upwelling of elevated pCO2 water masses with offshore winds drives the variation. Within days, primary production is responsible. The results demonstrate the high variability of the carbonate system in nearshore macrophyte meadows depending on meteorology and biological activities. We highlight the need to incorporate these variations in future pCO2 scenarios and experimental designs for nearshore habitats

Environmental variability in aquatic ecosystems: Avenues for future multifactorial experiments

The relevance of considering environmental variability for understanding and predicting biological responses to environmental changes has resulted in a recent surge in variability-focused ecological research. However, integration of findings that emerge across studies and identification of remaining knowledge gaps in aquatic ecosystems remain critical. Here, we address these aspects by: (1) summarizing relevant terms of variability research including the components (characteristics) of variability and key interactions when considering multiple environmental factors; (2) identifying conceptual frameworks for understanding the consequences of environmental variability in single and multifactorial scenarios; (3) highlighting challenges for bridging theoretical and experimental studies involving transitioning from simple to more complex scenarios; (4) proposing improved approaches to overcome current mismatches between theoretical predictions and experimental observations; and (5) providing a guide for designing integrated experiments across multiple scales, degrees of control, and complexity in light of their specific strengths and limitations