In the 1980s, Danish coastal waters suffered from eutrophication and several nutrient management plans have been implemented during the years to improve ecological status. This study aims at giving a holistic ecosystem perspective on 25 years of mitigation measures. We report trends of nutrient inputs and the responses to these in various chemical and biological components. Nutrient inputs from land were reduced by ~50 % for nitrogen (N) and 56 % for phosphorus (P) since 1990. These reductions resulted in significant and parallel declines in nutrient concentrations, and initiated a shift in the dominance of primary producers towards reduced phytoplankton biomass (chlorophyll a concentration) and increased cover of macroalgae in deeper waters. In the last 5 years, eelgrass meadows have also expanded towards deeper waters, in response to improving water clarity. An expected improvement of bottom water oxygen conditions has not been observed, presumably because more frequent stratification and higher water temperatures have counteracted the expected positive effects of reduced nutrient inputs. The biomass of the benthic macrofauna decreased as expected, but it was composed of a drastic decline of filter feeders paralleled by a more moderate increase of deposit feeders. This shift was most likely induced by increasing stratification. The reduced benthic filtration along with the limited eelgrass cover probably kept relatively more particles in suspension, which can explain why improvements in the Secchi depths were modest. Overall, several ecosystem components demonstrated clear signs of improvement, suggesting that at least partial recovery is attainable. On this basis, we propose a conceptual scheme for recovery of shallow coastal ecosystems following marked reductions in nutrient inputs.

Carbon-to-chlorophyll a ratios (C:Chl a; weight : weight) were analyzed for 7578 coastal seawater samples collected from Danish waters from 1990 to 2014. The aim was to identify the seasonal and spatial dynamics relative to nutrient richness and to study the effect of reduced nitrogen loadings over time. C:Chl a values were lowest during winter, about 15 across all stations. During the spring, C:Chl a increased to summer values between 20 and 96, depending on the annual mean of total nitrogen concentration. An inter-annual sinusoidal model with monthly time steps described the seasonal C:Chl a pattern well. The amplitudes of the model varied inversely with the annual mean of total nitrogen. Data also showed that a reduction in nitrogen loadings to the area by ∼ 40% over the past 24 yr, resulted in a statistically significant increase in mean annual C:Chl a values of 0.8 ± 0.2 yr⁻¹. The patterns derived from this large data set can be used to predict C:Chl a values for temperate coastal phytoplankton. Use of the empirical relationships derived from the data set improves predictions of C:Chl a values and thereby, e.g., carbon based food-web calculations and carbon-based ecosystem models, which often are validated using chlorophyll measurements.

Chain formation is common among phytoplankton organisms but the underlying reasons and consequences are poorly understood. Here we show that chain formation is strongly impaired by waterborne cues from copepod grazers in the dinoflagellate Alexandrium tamarense . Chains of Alexandrium cells exposed to copepod cues responded by splitting into single cells or shorter chains. Motion analysis revealed significantly lower swimming velocities for single cells compared with chains, with two- to fivefold higher simulated predator encounter rates for two- and four-cell chains, respectively. In addition, the few remaining two-cell chains in grazed treatments were swimming at approximately half the speed of two-cell chains in treatments without grazers, which reduced encounter rates with grazers to values similar to that of single cells. Chain length plasticity and swimming behavior constitute unique mechanisms to reduce encounters with grazers. We argue that dinoflagellates can regulate the balance between motility and predator avoidance by adjusting chain length. The high predator encounter rate for motile chains may have contributed to the low prevalence of chain formation in motile phytoplankton compared with in nonmotile phytoplankton where chain formation is more common.

We describe the kinematics of escape jumps in three species of 0.3–3.0 mm-sized planktonic copepods. We find similar kinematics between species with periodically alternating power strokes and passive coasting and a resulting highly fluctuating escape velocity. By direct numerical simulations, we estimate the force and power output needed to accelerate and overcome drag. Both are very high compared with those of other organisms, as are the escape velocities in comparison to startle velocities of other aquatic animals. Thus, the maximum weight-specific force, which for muscle motors of other animals has been found to be near constant at 57 N (kg muscle) ⁻¹ , is more than an order of magnitude higher for the escaping copepods. We argue that this is feasible because most copepods have different systems for steady propulsion (feeding appendages) and intensive escapes (swimming legs), with the muscular arrangement of the latter probably adapted for high force production during short-lasting bursts. The resulting escape velocities scale with body length to power 0.65, different from the size-scaling of both similar sized and larger animals moving at constant velocity, but similar to that found for startle velocities in other aquatic organisms. The relative duration of the pauses between power strokes was observed to increase with organism size. We demonstrate that this is an inherent property of swimming by alternating power strokes and pauses. We finally show that the Strouhal number is in the range of peak propulsion efficiency, again suggesting that copepods are optimally designed for rapid escape jumps.

Invasive alien species are a major worldwide driver of biodiversity change. The current study lists verified records of non-indigenous species (NIS) in European marine waters until 2020, with the purpose of establishing a baseline, assessing trends, and discussing appropriate threshold values for good environmental status (GES) according to the relevant European legislation. All NIS records were verified by national experts and trends are presented in six-year assessment periods from 1970 to 2020 according to the European Union Marine Strategy Framework Directive. Altogether, 874 NIS have been introduced to European marine waters until 2020 with the Mediterranean Sea and North-East Atlantic Ocean hosting most of the introductions. Overall, the number of new introductions has steadily increased since 2000. The annual rate of new introductions reached 21 new NIS in European seas within the last six-year assessment period (2012–2017). This increase is likely due to increased human activities and research efforts that have intensified during the early 21st century within European Seas. As Europe seas are not environmentally, nor geographically homogenous, the setting of threshold values for assessing GES requires regional expertise. Further, once management measures are operational, pathway-specific threshold values would enable assessing the effectiveness of such measures.