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International audience; Seagrass meadows and their associated biodiverse assemblages have declined globally due to environmental and anthropogenic stressors. Restoration of these critical habitats has the potential to reverse coastal biodiversity loss. Here, we tested the role of patch size (which can affect recruitment, food availability, and/or predation) in driving faunal colonization in an eelgrass (Zostera marina) restoration trial in Sweden. Eelgrass shoots were transplanted in plots with different configurations (continuous vs. checkerboard patterns with three patch sizes), and we followed invertebrate colonization (biodiversity and functional diversity) during the first two growing seasons. We found rapid faunal colonization following the transplantation of eelgrass shoots in all plots with invertebrate densities reaching 50-80% of the reference meadow after only one growing season (3 months). After two growing seasons (15 months), the faunal density, biodiversity, and functional diversity were similar to the reference meadow, despite eelgrass density and biomass still being lower than the reference meadow. Biodiversity, functional diversity, and community structure were similar among the different planted plots, that is, there was no indication that patch size influenced faunal colonization. We therefore consider that smaller patches embedded within larger restoration plots can be as effective for promoting biodiversity as continuous patches, with reduced costs and fewer shoots required. We also noted high natural variability between years both in the reference meadow and planted plots, showing the dynamic nature of seagrass ecosystems, and the importance of a well-planned monitoring scheme that considers the reference area and restored area within the same temporal scale.

Distribution of Earth's biomes is structured by the match between climate and plant traits, which in turn shape associated communities and ecosystem processes and services. However, that climate-trait match can be disrupted by historical events, with lasting ecosystem impacts. As Earth's environment changes faster than at any time in human history, critical questions are whether and how organismal traits and ecosystems can adjust to altered conditions. We quantified the relative importance of current environmental forcing versus evolutionary history in shaping the growth form (stature and biomass) and associated community of eelgrass (Zostera marina), a widespread foundation plant of marine ecosystems along Northern Hemisphere coastlines, which experienced major shifts in distribution and genetic composition during the Pleistocene. We found that eelgrass stature and biomass retain a legacy of the Pleistocene colonization of the Atlantic from the ancestral Pacific range and of more recent within-basin bottlenecks and genetic differentiation. This evolutionary legacy in turn influences the biomass of associated algae and invertebrates that fuel coastal food webs, with effects comparable to or stronger than effects of current environmental forcing. Such historical lags in phenotypic acclimatization may constrain ecosystem adjustments to rapid anthropogenic climate change, thus altering predictions about the future functioning of ecosystems., This work was supported by the US NSF (OCE-1031061, OCE-1336206, OCE0-1336741, OCE-1336905) and the Smithsonian Institution. F.T. was supported by José Castillejo Award CAS14/00177. A.H.E. was supported by the FCT (Foundation for Science and Technology) through Project UIDB/04326/2020 and Contract CEECINST/00114/2018. This is Contribution 106 from the Smithsonian’s MarineGEO and Tennenbaum Marine Observatories Network and Contribution 4105 of the Virginia Institute of Marine Science, College of William & Mary.

While considerable evidence exists of biogeographic patterns in the intensity of species interactions, the influence of these patterns on variation in community structure is less clear. Studying how the distributions of traits in communities vary along global gradients can inform how variation in interactions and other factors contribute to the process of community assembly. Using a model selection approach on measures of trait dispersion in crustaceans associated with eelgrass (Zostera marina) spanning 30° of latitude in two oceans, we found that dispersion strongly increased with increasing predation and decreasing latitude. Ocean and epiphyte load appeared as secondary predictors; Pacific communities were more overdispersed while Atlantic communities were more clustered, and increasing epiphytes were associated with increased clustering. By examining how species interactions and environmental filters influence community structure across biogeographic regions, we demonstrate how both latitudinal variation in species interactions and historical contingency shape these responses. Community trait distributions have implications for ecosystem stability and functioning, and integrating large-scale observations of environmental filters, species interactions and traits can help us predict how communities may respond to environmental change., This research was funded by National Science Foundation grants to J.E.D., J.J.S. and K.A.H. (NSF-OCE 1336206, OCE 1336905, and OCE 1336741). C.B. was funded by the Åbo Akademi University Foundation.

According to the EU Habitats directive, the Water Framework Directive, and the Marine Strategy Framework Directive, member states are required to map, monitor, and evaluate changes in quality and areal distribution of different marine habitats and biotopes to protect the marine environment more effectively. Submerged aquatic vegetation (SAV) is a key indicator of the ecological status of coastal ecosystems and is therefore widely used in reporting related to these directives. Environmental monitoring of the areal distribution of SAV is lacking in Sweden due to the challenges of large-scale monitoring using traditional small-scale methods. To address this gap, in 2020, we embarked on a project to combine Copernicus Sentinel-2 satellite imagery, novel machine learning (ML) techniques, and advanced data processing in a cloud-based web application that enables users to create up-to-date SAV classifications. At the same time, the approach was used to derive the first high-resolution SAV map for the entire coastline of Sweden, where an area of 1550 km2 was mapped as SAV. Quantitative evaluation of the accuracy of the classification using independent field data from three different regions along the Swedish coast demonstrated relative high accuracy within shallower areas, particularly where water transparency was high (average total accuracy per region 0.60-0.77). However, the classification missed large proportions of vegetation growing in deeper water (on average 31%-50%) and performed poorly in areas with fragmented or mixed vegetation and poor water quality, challenges that should be addressed in the development of the mapping methods towards integration into monitoring frameworks such as the EU directives. In this article, we present the results of the first satellite-derived SAV classification for the entire Swedish coast and show the implementation of a cloud-based SAV mapping application (prototype) developed within the frame of the project. Integr Environ Assess Manag 2021;00:1-12. © 2021 The Authors. Integrated Environmental Assessment and Management published by Wiley Periodicals LLC on behalf of Society of Environmental Toxicology & Chemistry (SETAC).

Estuaries are important nursery areas for many species and these habitats are often affected by anthropogenic activities. We investigate possible negative effects of pesticides and metals on penaeid shrimps in Maputo Bay, Mozambique. Shrimps and water samples were collected in three estuaries and one coastal area for biomarker and chemical analysis. Acetylcholinesterase (AChE) and glutathione-S transferase activities were analysed as biomarkers for pollutants. 37 different pesticides were analysed in water samples and shrimp muscle tissue was analysed for 10 metals. Risk assessment showed that the environmental thresholds were exceeded for several herbicides in three of four of the assessed nursery areas. Lower AChE activities were detected in shrimps captured close to an agriculture area and this location had the lowest shrimp densities. Metal analysis in shrimp showed low levels. Despite localized effects, results highlight the need to improve the regulation of pollutants in the Espirito Santo estuary in Maputo bay.

The eelgrass Zostera marina is an important foundation species of coastal areas in the Northern Hemisphere, but is continuing to decline, despite management actions. The development of new management tools is therefore urgent in order to prioritize limited resources for protecting meadows most vulnerable to local extinctions and identifying most valuable present and historic meadows to protect and restore, respectively. We assessed 377 eelgrass meadows along the complex coastlines of two fjord regions on the Swedish west coast-one is currently healthy and the other is substantially degraded. Shoot dispersal for all meadows was assessed with Lagrangian biophysical modeling (scale: 100-1,000 m) and used for barrier analysis and clustering; a subset (n = 22) was also assessed with population genetic methods (20 microsatellites) including diversity, structure, and network connectivity. Both approaches were in very good agreement, resulting in seven subpopulation groupings or management units (MUs). The MUs correspond to a spatial scale appropriate for coastal management of "waterbodies" used in the European Water Framework Directive. Adding demographic modeling based on the genetic and biophysical data as a third approach, we are able to assess past, present, and future metapopulation dynamics to identify especially vulnerable and valuable meadows. In a further application, we show how the biophysical approach, using eigenvalue perturbation theory (EPT) and distribution records from the 1980s, can be used to identify lost meadows where restoration would best benefit the present metapopulation. The combination of methods, presented here as a toolbox, allows the assessment of different temporal and spatial scales at the same time, as well as ranking of specific meadows according to key genetic, demographic and ecological metrics. It could be applied to any species or region, and we exemplify its versatility as a management guide for eelgrass along the Swedish west coast.

Maintaining and enabling evolutionary processes within meta‐populations are critical to resistance, resilience and adaptive potential. Knowledge about which populations act as sources or sinks, and the direction of gene flow, can help to focus conservation efforts more effectively and forecast how populations might respond to future anthropogenic and environmental pressures. As a foundation species and habitat provider, Zostera marina (eelgrass) is of critical importance to ecosystem functions including fisheries. Here, we estimate connectivity of Z. marina in the Skagerrak–Kattegat region of the North Sea based on genetic and biophysical modelling. Genetic diversity, population structure and migration were analysed at 23 locations using 20 microsatellite loci and a suite of analytical approaches. Oceanographic connectivity was analysed using Lagrangian dispersal simulations based on contemporary and historical distribution data dating back to the late 19th century. Population clusters, barriers and networks of connectivity were found to be very similar based on either genetic or oceanographic analyses. A single‐generation model of dispersal was not realistic, whereas multigeneration models that integrate stepping‐stone dispersal and extant and historic distribution data were able to capture and model genetic connectivity patterns well. Passive rafting of flowering shoots along oceanographic currents is the main driver of gene flow at this spatial–temporal scale, and extant genetic connectivity strongly reflects the “ghost of dispersal past“ sensu Benzie, 1999. The identification of distinct clusters, connectivity hotspots and areas where connectivity has become limited over the last century is critical information for spatial management, conservation and restoration of eelgrass.

Along the Swedish northwest coast, over 60% of the eelgrass meadows have been lost since the 1980s. Despite improved water quality, no recovery has occurred, and restoration is presently considered to mitigate historical losses. However, the factors preventing natural recovery of eelgrass are not known, and it is not clear if conditions would allow restoration. Here, we present the results from 5 years of field studies with the aim of identifying the key processes affecting eelgrass growth and survival at historical eelgrass areas. Continuous light measurements and comparison with historic eelgrass distribution indicate that maximum depth distribution has decreased locally with 1.5–2.3 m in areas that have lost large eelgrass beds in the last 10–30 years. Field studies suggest that wind-driven local resuspension of sediments that are no longer stabilized by eelgrass beds is the main cause behind the deteriorated light conditions. Field experiments show that a combination of low light condition and disturbance from drifting algal mats prevents eelgrass recovery in these areas, whereas the sulfide intrusion from the sediment and dislodgement of shoots by waves had little effect on growth and survival. These results suggest that local regime shifts acting on a scale of 40–200 ha have occurred after the loss of eelgrass beds, where increased sediment resuspension and proliferation of drifting algal mats act as feedback mechanisms that prevent both natural recovery and restoration of eelgrass. The feedbacks appear to be interacting and causing an accelerating loss of eelgrass that is presently spreading to neighboring areas.

Form–function relationships in plants underlie their ecosystem roles in supporting higher trophic levels through primary production, detrital pathways, and habitat provision. For widespread, phenotypically-variable plants, productivity may differ not only across abiotic conditions, but also from distinct morphological or demographic traits. A single foundation species, eelgrass Zostera marina, typically dominates north temperate seagrass meadows, which we studied across 14 sites spanning 32–61°N latitude and two ocean basins. Body size varied by nearly two orders of magnitude through this range, and was largest at mid-latitudes and in the Pacific Ocean. At the global scale, neither latitude, site-level environmental conditions, nor body size helped predict productivity (relative growth rate 1–2% day-1 at most sites), suggesting a remarkable capacity of Z. marina to achieve similar productivity in summer. Furthermore, among a suite of stressors applied within sites, only ambient leaf damage reduced productivity; grazer reduction and nutrient addition had no effect on eelgrass size or growth. Scale-dependence was evident in different allometric relationships within and across sites for productivity and for modules (leaf count) relative to size. Zostera marina provides a range of ecosystem functions related to both body size (habitat provision, water flow) and growth rates (food, carbon dynamics). Our observed decoupling of body size and maximum production suggests that geographic variation in these ecosystem functions may be independent, with a future need to resolve how local adaptation or plasticity of body size might actually enable more consistent peak productivity across disparate environmental conditions.

Environmental compensation should address negative impacts from human activities on nature, including loss of biodiversity and ecosystem services. However, successful compensation, achieving no net loss, requires broad quantitative information on different types of losses and gains. We find that the scope of compensatory schemes varies in what is considered compensable, which makes it challenging to apply a conceptual approach consistently across schemes with different needs. We propose a flexible yet structured framework for determining which values should be compensated and how. Our framework focuses specifically on habitat deterioration and is illustrated with a case study involving loss of eelgrass habitat. The framework helps identify compensation needs and selects among suitable compensation options, merging science-based information with normative issues and local concerns. By integrating the ecosystem services cascade model, it encompasses aspects from biodiversity structure to human wellbeing. The framework prefers in-kind compensation because this targets the structure level and thus meets compensation needs in all subsequent levels of the cascade model; further, it is more likely to capture non-instrumental values (i.e. in nature) and reduce exposure to uncertainty. We highlight the importance of spatial aspects of ecosystem functions, services and their subsequent impacts on wellbeing. Although our selection hierarchy assumes a “similar and nearby” principle for habitat restoration (preference for in-kind/on-site), this criterion is not universal. We underscore the hierarchy’s implicit normative assumptions and suggest that apparent disagreement about who should benefit may be traced to an unresolved conflict between egalitarianism and utilitarianism.

More than 50% of eelgrass habitats have disappeared from the Swedish NW coast in the last 30 years. Restoration is being proposed to assist recovery but little is known regarding methods suitable under Scandinavian conditions; e.g. short growing seasons and scouring by ice. In the present study we evaluated different restoration methods using shoots and seeds in a Swedish fjord and assessed if eelgrass could be successfully transplanted between sites with different depth and exposure. The study demonstrates that both shoot- and seed methods can be successfully used to restore eelgrass at this latitude. Survival and growth of unanchored single shoots, transplanted without sediment in shallow habitats (1.0-1.5 m) was very high (>500% increase in shoot density after 14 months). This restoration method showed 2-3.5 times higher growth rate and was 2-2.5 times faster compared with shoots anchored in the sediment and shoots transplanted in sediment cores, respectively, and is recommended for shallow habitats in Sweden. Growth within deeper habitats (3.0-4.5 m) was substantially tower (40% loss to 50% increase) due to light limitations and high winter mortality. Restoration using seeds distributed from mesh-bags showed very low seedling establishment rates (approximately 1%) making this method less cost-effective than transplanting single shoots in shallow habitats. However, growth of seedlings was high and this method is recommended for deep habitats with soft sediment where shoot transplantation is difficult. Despite dramatic differences in eelgrass morphology between habitats with different depth and exposure, all shoots within a planting site had the same morphology at the end of the study, independent of origin. A baseline genetic survey supported that the observed changes in morphology of transplants were due to a plastic response, suggesting that donor populations do not have to exactly match the morphology of the plants targeted for restoration. (C) 2016 Elsevier B.V. All rights reserved.

Along the northwest coast of Sweden, over 50% of Zostera marina L. (eelgrass) meadows have vanished since the 1980s. With the improvement of conditions, there is a growing interest to restore lost habitats, but methods are lacking for restoration of eelgrass beds at high latitudes where long winters create special challenges. We assessed if seed planting could be used for large-scale restoration, with the aim to identify the major causes of seed and seedling loss and to determine which planting method best increases restoration success. In the laboratory, we identified optimal conditions for long-term seed storage and demonstrated that eelgrass seeds can be successfully stored for 8 mo before being planted in the spring. However, field studies did not find an increased seedling establishment in seeds planted in the spring of 2013 compared to those planted in the fall of 2012. Field results suggest that the main causes for the seed loss were seed predation from crabs, seed transport by currents and bioturbation by lugworms, while the main processes affecting shoot development were light availability and physical disturbance. Covering the seeds with a layer of sand increased seedling establishment 2 to 6 times compared to uncovered seeds, probably through the reduction of seed predation and seed transport, and could potentially be developed as a method to reduce losses during restoration. In addition, positive feedback mechanisms (i.e. sediment resuspension and drifting macroalgae mats) may also prevent natural recovery and restoration success. However, high seed loss (on average 98.6%) and high shoot mortality pose a challenge that need to be addressed before restoration using seeds can be recommended for large-scale restoration.

Seagrass meadows are able to store significant amounts of organic carbon in their underlying sediment but global estimates are uncertain partly due to spatio-temporal heterogeneity between areas and species. In order to provide robust estimates, there is a need to better understand the fate of, and mechanisms behind, organic carbon storage. In this observational study, we analyse a suite of biotic and abiotic parameters in sediment cores from 47 different eelgrass (Zostera marina) beds spanning the distributional range of the Northern Hemisphere. Depth profiles revealed three patterns of vertical distribution where POC either increased, decreased or showed no distinct pattern with sediment depth. These categories exhibited distinct profiles of δ13C and C:N ratios, where high POC profiles had a proportionally larger storage of eelgrass-derived material whereas low POC profiles were dominated by phytoplanktonic and macroalgal material. However, high POC did not always translate into high carbon density. Though high POC did not always translate into high carbon density, this large-scale dataset evidences that the variability in organic matter source in response to natural and anthropogenic environmental changes affects the potential role of eelgrass beds as POC sinks, particularly where eelgrass decline is observed.

10 pages, 7 figures, 1 table, supplementary material https://doi.org/10.1016/j.pocean.2018.10.008, In Antarctica, summer is a time of extreme environmental shifts resulting in large coastal phytoplankton blooms fueling the food web. Despite the importance of the microbial loop in remineralizing biomass from primary production, studies of how microzooplankton communities respond to such blooms in the Southern Ocean are rather scarce. Microzooplankton (ciliate and dinoflagellate) communities were investigated combining microscopy and 18S rRNA sequencing analyses in the Amundsen Sea Polynya during an extensive summer bloom of Phaeocystis antarctica. The succession of microzooplankton was further assessed during a 15-day induced bloom microcosm experiment. Dinoflagellates accounted for up to 59 % of the microzooplankton biomass in situ with Gymnodinium spp., Protoperidium spp. and Gyrodinium spp. constituting 89 % of the dinoflagellate biomass. Strobilidium spp., Strombidium spp. and tintinids represented 90 % of the ciliate biomass. Gymnodinium, Gyrodinium and tintinnids are known grazers of Phaeocystis, suggesting that this prymnesiophyte selected for the key microzooplankton taxa. Availability of other potential prey, such as diatoms, heterotrophic nanoflagellates and bacteria, also correlated to changes in microzooplankton community structure. Overall, both heterotrophy and mixotrophy appeared to be key trophic strategies of the dominant microzooplankton observed, suggesting that they influence carbon flow in the microbial food web through top-down control on the phytoplankton community, This work was supported by the Swedish Research Council [grant 2008-6430] to S. Bertilsson and L. Riemann and [grant 824-2008-6429] to P.-O. Moksnes and J. Havenhand, and by the US National Science Foundation through the ASPIRE project [NSF OPP-0839069] to P. Yager

Latitudinal gradients in species interactions are widely cited as potential causes or consequences of global patterns of biodiversity. However, mechanistic studies documenting changes in interactions across broad geographic ranges are limited. We surveyed predation intensity on common prey (live amphipods and gastropods) in communities of eelgrass (Zostera marina) at 48 sites across its Northern Hemisphere range, encompassing over 37 degrees of latitude and four continental coastlines. Predation on amphipods declined with latitude on all coasts but declined more strongly along western ocean margins where temperature gradients are steeper. Whereas insitu water temperature at the time of the experiments was uncorrelated with predation, mean annual temperature strongly positively predicted predation, suggesting a more complex mechanism than simply increased metabolic activity at the time of predation. This large-scale biogeographic pattern was modified by local habitat characteristics; predation declined with higher shoot density both among and within sites. Predation rates on gastropods, by contrast, were uniformly low and varied little among sites. The high replication and geographic extent of our study not only provides additional evidence to support biogeographic variation in predation intensity, but also insight into the mechanisms that relate temperature and biogeographic gradients in species interactions. NSF BIO-OCE [1336206, 1336741, 1336905] info:eu-repo/semantics/publishedVersion

Using a combination of empirical and model studies we tested whether European shore crab larvae (Carcinus maenas) from environments with different tidal regimes in the North Sea area have different swimming behaviors, and whether this affects connectivity and settlement success of larvae. Laboratory studies demonstrated the presence of an inherited tidal migration rhythm in newly hatched crab larvae from the mesotidal Danish Wadden Sea, and field studies showed that postlarvae swam in surface water almost exclusively during flood tides, suggesting that larvae use selective tidal stream transport to control the dispersal process. In contrast, shore crab larvae from microtidal Skagerrak displayed a nocturnal vertical migration behavior that appeared to switch to a diurnal behavior at the end of the postlarval phase, indicating an adaptation to avoid visual predators and to use wind-driven transport to reach shallow settlement areas. A biophysical model showed that tidal-migrating larvae in the Wadden Sea had two times higher settlement success than larvae with a diel behavior. However, no differences in settlement success were found between the two larval behaviors in microtidal Skagerrak, where lower fitness is suggested for tidal-migrating larvae due to higher predation mortality from visual predators. We suggest that the differences in inherited larval behavior in larvae from meso- and microtidal regions reflect local adaptations maintained through natural selection of successful recruits. Consistent with recent population genetic studies, modeled connectivity of shore crabs indicated an oceanographic dispersal barrier to gene flow in Eastern Wadden Sea that may facilitate such adaptations.

Seagrass meadows are able to store significant amounts of organic carbon in their underlying sediment, but global estimates are uncertain partly owing to spatio-temporal heterogeneity between and within areas and species. In order to provide robust estimates, there is a need to better understand the fate of, and mechanisms behind, organic carbon storage. In this observational study, we analyse a suite of biotic and abiotic parameters in sediment cores from 47 different eelgrass ( Zostera marina ) beds spanning the distributional range of the Northern Hemisphere. Depth profiles of particulate organic carbon (POC) revealed three patterns of vertical distribution where POC either increased, decreased or showed no pattern with sediment depth. These categories exhibited distinct profiles of δ 13 C and C:N ratios, where high POC profiles had a proportionally larger storage of eelgrass-derived material whereas low POC profiles were dominated by phytoplanktonic and macroalgal material. However, high POC did not always translate into high carbon density. Nevertheless, this large-scale dataset provides evidence that the variability in organic matter source in response to natural and anthropogenic environmental changes affects the potential role of eelgrass beds as POC sinks, particularly where eelgrass decline is observed.

Blooming filamentous algae recurrently overgrow macroalgae and seagrass in many coastal ecosystems, leading to potential losses in the functions and services these habitats provide. The seasonal reoccurrence of algal blooms is partly caused by excessive input of nutrients into coastal ecosystems. At the same time, overfishing of top predators has led to increased densities of smaller predators that reduce herbivore abundance. Managers of coastal ecosystems are in urgent need of information on how overfishing may act synergistically with eutrophication to promote algal blooms. In the present study, we model the interaction between the opportunistic filamentous algae Ulva spp. and the effective mesograzer (small invertebrate herbivore) Gammarus locusta L. under different nutrient regimes and predation rates by intermediate-size fish (4 to 12 cm). The aim is to as- sess how productivity, herbivory and predation interact to regulate seasonal algal growth under a range of scenarios, including temporal matching or mismatching between trophic groups. The model is parameterised with nutrient and demographic data from a seagrass ecosystem on the west coast of Sweden. Model simulations show that both nutrient level and herbivore abundance regulate seasonal algal biomass in the seagrass meadow. Their relative importance is dynamic and sensitive to the arrival times of herbivores in relation to the start of algal development in spring. Thus, herbivore presence and timing are crucial factors for the control of algal biomass, particularly during times when resource levels are ambient to slightly elevated. Simulations also show that it is mainly the predation rate and timing of predators, rather than food limitation, that limit herbivore abundance. As predation rates by invertebrates and smaller fish can be linked to the presence of larger and commercially targeted fish, overfishing may have similar effects to eutrophication in the seagrass ecosystems.

This study aims to improve estimates of dispersal by including information on larval traits, and in particular to explore how larval depth distribution affects connectivity and MPA (marine protected area) functionality in the Baltic Sea. A field survey showed that both invertebrates and fish differed in their larval depth distribution, ranging from surface waters to >100 m. A biophysical model of larval dispersal in the Baltic Sea showed that decreased depth distribution increased average dispersal distance 2.5-fold, decreased coastal retention and local recruitment, and substantially increased connectivity. Together with pelagic larval duration (PLD), depth distribution explained 80% of total variation in dispersal distance, whereas spawning season, and geographic and annual variations in circulation had only marginal effects. Median dispersal distances varied between 8 and 46 km, with 10% of simulated trajectories dispersing 30 to 160 km depending on drift depth and PLD. In the Baltic Sea, the majority of shallow Natura 2000 MPAs are 10% within MPAs of this size. Connectivity between MPAs was expected to be low for most larval trait combinations. Our simulations and the empirical data suggest that the MPA size within the Natura 2000 system is considerably below what is required for local recruitment of most sessile invertebrates and sedentary fish. Future designs of MPA networks would benefit from spatially explicit biophysical models that consider dispersal and connectivity for complex circulation patterns and informed larval traits.

The interaction of eutrophication and predation in structuring seagrass Zostera marina L. ecosystems was assessed in a field experiment in three regions along an estuarine salinity gradient, from southern Finland to the Skagerrak area of the Swedish west coast. All regions are considered to be affected by eutrophication and overfishing but differ in the abundance of intermediate predators (e.g., small fish, shrimp, and crabs), mesograzers, and the biomass of epiphytic algae. Using transplanted Zostera (eelgrass), nutrient levels and intermediate predator abundance were manipulated in a full-factorial cage experiment. On the Swedish west coast, where ambient densities of mesograzers are very low, epiphytic algae responded strongly to nutrient enrichment, resulting in significantly reduced growth of eelgrass. At the Baltic sites however, where ambient densities of mesograzers are high, no significant growth of epiphytic algae was detected, and only grazer biomass responded to nutrient enrichment. Predation from small fish, shrimp, and crabs decreased the biomass of mesograzers by . 98% on the Swedish west coast, but natural predators had no significant effect on mesograzers biomass at the Baltic sites. Predation and nutrient enrichment interacted to affect the growth of eelgrass by controlling the biomass of mesograzers and nuisance algae. The differing effect of nutrient enrichment and grazing in the three regions may therefore be a result of the prevailing low and high predation pressure on mesograzers in Zostera. This absence or presence of predation may derive from interregional changes in trophic interactions, possibly caused by a combination of eutrophication and overfishing.

Algal grazers are thought to play a critical role in preventing overgrowth by ephemeral algae in seagrass beds. We studied the interaction between the omnivorous grass shrimp Palaemon elegans, the algal grazer Gammarus locusta and the filamentous green alga Ulva sp. (syn. Entero- morpha sp.) to assess whether the shrimp has a positive or negative effect on the growth of ephemeral algae in eelgrass beds on the Swedish west coast. Laboratory experiments demonstrated that P. ele- gans were effective predators on gammarids 5 mm TL, but negatively affected when G. locusta was smaller. In a field cage experiment, enclosed P. elegans decreased the abundance of gammarids by on average 68%, causing a trophic cascade that increased the biomass of filamentous Ulva sp. 7-fold. These results suggest that P. elegans on the Swedish northwest coast feed mainly on amphipods, including the algal grazer G. locusta, resulting in a positive net trophic effect on the growth of filamentous Ulva sp. This study indicates that P. ele- gans may be an important link between overexploitation of cod in northwest Sweden and the recent increase of algal mats and loss of eelgrass in the area.

We compared eelgrass Zostera marina communities in 3 regions in Sweden believed to be affected by eutrophication and overfishing, to determine whether bottom-up or top-down processes control the biomass of epiphytic macroalgae and grazers. Nitrogen and carbon isotope signatures were analyzed to explore the food webs and to identify the grazing species feeding on filamentous algae and/or eelgrass. Mixing model (IsoSource version 1.3.1) analysis of the isotope signatures indicated that the amphipods Gammarus locusta and Microdeutopus gryllotalpa fed primarily on filamentous algae and that only 2 small gastropod species consumed eelgrass. Moreover, the grass shrimp Palaemon elegans and F adspersus were ca. 1 trophic level above amphipods and algae, but according to the mixing model played different trophic roles in the different areas. The highest biomass of filamentous algae was found in the west coast beds housing grazers with the lowest biomass and mean size (predominantly G. locusta and M. gryllotalpa, 0.5 to 3 mm). In contrast, the Baltic Sea beds had low algal biomass, but the grazers (mostly G. locusta and Idotea baltica) had higher biomass and were significantly larger (mean size ca. 10 mm). An overall negative correlation was found between grazer biomass and biomass of filamentous algae. The significantly smaller grazers and absence of isopod grazers on the west coast may be due to substantial consumption by small predatory fish. This supports the suggestions that, in Swedish eelgrass beds, grazers are top-down controlled, and overexploitation. of large predators and eutrophication play an important role in the recent increases in algal biomass.

We assessed the relative importance of bottom-up and top-down processes in structuring an eelgrass community in Sweden, a system impacted both by eutrophication and overfishing. Using artificial seagrass as substrate, we manipulated nutrient levels and predator abundance in a full-factorial cage-experiment. The results revealed a seagrass community dominated by strong top-down processes controlling the aggregate biomass of mesograzers and macroalgae. In the absence of predators the large amphipod Gammarus locusta became very abundant resulting in a leaf community with low biomass of algae and smaller mobile fauna. One enclosed gobid fish predator reduced the abundance of adult G. locusta by > 90%, causing a three to six times increase in the biomass of algae, smaller mesograzers and meiofauna. Numerous small predators in uncaged habitats reduced the biomass of G. locusta and other mesograzers by > 95% in comparison to the fish treatment, further increasing the biomass of epiphytic algae and meiofauna. Although water column nutrient enrichment caused a temporal bloom of the filamentous macroalgae Ulva spp., no significant nutrient-effects were found on the algal community at the end of the experiment. The only lasting nutrient-effect was a significant increase in the biomass of G. locusta, but only in the absence of ambient predators. These results demonstrate that mesograzers can respond to enhanced food supply, increase their biomass and control the algal growth when predation rates are low. However, in the assessed system, high predation rates appear to make mesograzers functionally extinct, causing a community-wide trophic cascade that promotes the growth of ephemeral algae. This top-down effect could penetrate down, despite a complex food-web because the interaction strength in the community was strongly skewed towards two functionally dominant algal and grazer species that were vulnerable to consumption. These results indicate that overexploitation of gadoid fish may be linked to increased macroalgal blooms and loss of eelgrass in the area through a trophic cascade affecting the abundance of mesograzers.

Valuing nature’s benefits in monetary terms is necessary for policy-makers facing trade-offs in how to spend limited financial resources on environmental protection. We provide information to assess trade-offs associated with the management of seagrass beds, which provide a number of ecosystem services, but are presently impacted by many stressors. We develop an interdisciplinary framework for valuing multiple ecosystem services and apply it to the case of eelgrass (Zostera marina), a dominant seagrass species in the northern hemisphere. We identify and quantify links between three eelgrass functions (habitat for fish, carbon and nitrogen uptake) and economic goods in Sweden, quantify these using ecological endpoints, estimate the marginal average value of the impact of losing one hectare of eelgrass along the Swedish northwest coast on welfare in monetary terms, and aggregate these values while considering double-counting. Over a 20 to 50 year period we find that compared to unvegetated habitats, a hectare of eelgrass, including the organic material accumulated in the sediment, produces an additional 626 kg cod fishes and 7,535 wrasse individuals and sequesters 98.6 ton carbon and 466 kg nitrogen. We value the flow of future benefits associated with commercial fishing, avoided climate change damages, and reduced eutrophication at 170,000 SEK in 2014 (20,700 US$) or 11,000 SEK (1,300 US$) annualized at 4%. Fish production, which is the most commonly valued ecosystem service in the seagrass literature, only represented 25% of the total value whereas a conservative estimate of nitrogen regulation constituted 46%, suggesting that most seagrass beds are undervalued. Comparing these values with historic losses of eelgrass we show that the Swedish northwest coast has suffered a substantial reduction in fish production and mineral regulation. Future work should improve the understanding of the geographic scale of eelgrass functions, how local variables affect the value of these functions, and how to defensibly aggregate a multitude of economic values.

Nutrient pollution and reduced grazing each can stimulate algal blooms as shown by numerous experiments. But because experiments rarely incorporate natural variation in environmental factors and biodiversity, conditions determining the relative strength of bottom-up and top-down forcing remain unresolved. We factorially added nutrients and reduced grazing at 15 sites across the range of the marine foundation species eelgrass (Zostera marina) to quantify how top-down and bottom-up control interact with natural gradients in biodiversity and environmental forcing. Experiments confirmed modest top-down control of algae, whereas fertilisation had no general effect. Unexpectedly, grazer and algal biomass were better predicted by cross-site variation in grazer and eelgrass diversity than by global environmental gradients. Moreover, these large-scale patterns corresponded strikingly with prior small-scale experiments. Our results link global and local evidence that biodiversity and top-down control strongly influence functioning of threatened seagrass ecosystems, and suggest that biodiversity is comparably important to global change stressors. National Science Foundation [OCE-1031061] info:eu-repo/semantics/publishedVersion

We assessed the relative role of 3 potentially important processes affecting the distribu- tion of young juvenile blue crabs Callinectes sapidus Rathbun: (1) habitat selection at settlement, (2) selection of habitats by dispersing juveniles, and (3) habitat-specific predation rates, using cage experiments. The results suggest that active habitat selection by postlarvae and young juvenile crabs determines the habitat-specific distribution of juvenile blue crabs. Densities of blue crab settlers (megalopae and first instar crabs) in caged habitat patches were high and similar in artificial sea- grass, live shoal grass Halodule wrightii, and live oyster habitats (on average 69 to 82 settlers m -2 ), but significantly lower in mud (26 settlers m -2 ), indicating active selection for any structurally com- plex habitat at settlement. Second and third instar juvenile blue crabs (J2 and J3 crabs) also colonized the structurally complex habitats in higher numbers (34 to 115 crabs m -2 ), compared to mud (14 crabs m -2 ), demonstrating that young juvenile blue crabs are highly mobile and redistribute soon after metamorphosis. Densities of J2 and J3 blue crabs in uncaged treatments were significantly lower in all habitats than in caged treatments (on average 44% loss), suggesting high predation mortality. However, the loss of settlers and juvenile crabs was similar in all habitats, and had no significant effect on the juvenile crab distribution. Densities of potential predators were on average 5 times higher in the structurally complex habitats than in mud. An aggregation of predators in the refuge habitats, coupled with a refuge at low prey densities in unstructured habitats, appears to decrease the proximate effect of predation on the distribution of juvenile crabs. Concurrent high settlement of a small xanthid crab (cf. Eurypanopeus sp.) occurred almost exclusively in the oyster habitat, sug- gesting that species with a limited ability to redistribute after metamorphosis may display a more specific selection of the adult habitat at settlement.

Juvenile cannibalism may play an important role in population regulation of marine organisms with juvenile stages that aggregate in nursery habitats. I conducted a series of field and laboratory experiments to assess whether three potential density-depen- dent mechanisms could regulate local recruitment of juvenile shore crabs Carcinus maenas in Swedish nursery areas: (1) intracohort cannibalism, (2) juvenile cannibals' functional response to smaller conspecific prey densities, and (3) juvenile cannibals' numerical effect on prey survival. Juvenile cannibals in mesocosms displayed a type III (sigmoid) density-dependent func- tional response, resulting in significantly higher proportional mortality of smaller conspe- cifics at high prey densities. Consistent with the laboratory result, cage-enclosed juvenile cannibals in a field experiment eliminated conspecific settlement peaks in mussel habitats and decoupled the relationship between larval and settlement abundance within three days. Density-dependent interference between cannibals in mesocosms decreased per capita pre- dation rates, but conspecific prey mortality still increased with cannibal densities when the abundance of cannibals passed a critical threshold, suggesting that the recruitment success of consecutive cohorts may be negatively correlated. This was supported in a field exper- iment where the abundance of juvenile cannibals explained 71% of the variation in settle- ment loss from uncaged mussel habitats. Density-dependent cannibalism also occurred within juvenile cohorts even though food was provided in excess, but the resulting mortality was too low to eliminate initial density patterns over a 30-day period. The results demonstrate that cannibalism between juvenile shore crabs can cause high and strongly density-dependent mortality at natural densities, and that both a functional response and a numerical effect of larger cannibals can regulate the recruitment of new cohorts within days of settlement. A coupling of cannibal and settlement densities on a regional scale is expected to decrease recruitment variability within and between local populations. Moreover, the interaction between the type III functional response, mutual interference, and density-dependent dispersal of cannibals is predicted to reduce canni- balistic rates at low prey and high cannibal densities, respectively, and stabilize local population dynamics in relation to the abundance of juvenile habitats, consistent with field observations of shore crab populations in the study area.

In marine organisms with dispersing larval stages, current ecological theory suggests that recruitment of benthic juveniles may be limited by juvenile habitats and regulated by early post- settlement processes. However, few studies have attempted to identify the density-dependent mech- anisms responsible for these demographic bottlenecks. I conducted a series of experiments to assess if interference competition for space, within nursery habitats, could result in density-dependent juve- nile growth and dispersal from refuge habitats, and regulate local populations of juvenile shore crabs Carcinus maenas in shallow nursery areas in Sweden. In laboratory mesocosms, increased natural densities of similarly sized crabs resulted in decreased growth and increased per capita emigration from mussel habitats, even though food was provided in excess, suggesting that competition for space and mutual interference mediated the density-dependent growth and dispersal. This was supported in a field experiment where a density-dependent relation was found between the abundance of juve- nile crabs in nursery areas and their migration rates to experimental mussel patches. The results sug- gest that interference competition for space and density-dependent emigration from refuge habitats, coupled with habitat-specific mortality, is an important regulating mechanism for juvenile shore crab populations, in particular for older juvenile cohorts that have escaped inter-cohort cannibalism through a size refuge. Density-dependent effects on growth were found only at unusually high nat- ural densities and may, therefore, be less important as a regulating mechanism. However, negative effects on feeding rates and growth at high conspecific densities may represent an important selec- tive force in juvenile migration behavior. Density-dependent dispersal was found at juvenile densi- ties that are regularly found in nursery areas, suggesting that nursery habitats become saturated with juveniles during the recruitment season and represent a limiting resource for local populations, con- sistent with field observations of shore crab populations in the study area.

In contrast to general settlement models of invertebrate larvae, recent Swedish studies of settlement of the shore crab Carcinus maenas indicate that many postlarvae close to metamorphosis emigrate from their preferred settlement habitats. In the present study, we assessed physical and biological factors affecting shore crab settlement and emigration from nursery habitats, and explored possible ultimate factors in this behavior. Postlarval abundance in the field was not affected by the tidal phase, but varied significantly according to the light conditions. Settlement densities on artificial substrates were significantly higher during the day compared to at night. In contrast, densities of planktonic postlarvae were highest at night. This result suggests that the postlarvae cling to shallow benthic habitats during the day and swim in the plankton at night. Mesocosm experiments demonstrated that most postlarvae remained in the benthos during the day, even in suboptimal habitats, but that a large proportion (on average 58%) actively emigrated at dusk, irrespective of settlement conditions. Emigration rates were significantly higher and metamorphosis rates lower in open sand compared to mussel habitats, whereas food levels and settlement densities had only small effects on the settlement process. Emigrated postlarvae were at a significantly earlier developmental stage and metamorphosed on average 1 d later than the megalopae that remained in the benthic habitat. Post-larvae >2 d from metamorphosis emigrated even from optimal settlement conditions. These results suggest that settlement in shore crabs is not an irreversible process once a late-stage postlarva has found a settlement habitat, but involves 2 separate behavioral components: (1) the selection of a benthic habitat during the day, which is mainly affected by the availability of refuges from predation; and (2) the choice to stay or emigrate the following night, where emigration is induced by a decrease in light and determined mainly by the development stage of the postlarvae. We propose a conceptual model of settlement in the shore crab where the dynamic behavior of the postlarvae is an adaptation to diel differences in predation risk between the settlement habitat and the plankton.

Cold-water conditions have excluded durophagous (skeleton-breaking) predators from the Antarctic seafloor for millions of years. Rapidly warming seas off the western Antarctic Peninsula could now facilitate their return to the continental shelf, with profound consequences for the endemic fauna. Among the likely first arrivals are king crabs (Lithodidae), which were discovered recently on the adjacent continental slope. During the austral summer of 2010 ‒ 2011, we used underwater imagery to survey a slope-dwelling population of the lithodid Paralomis birsteini off Marguerite Bay, western Antarctic Peninsula for environmental or trophic impediments to shoreward expansion. The population density averaged ∼ 4.5 individuals × 1,000 m(-2) within a depth range of 1,100 ‒ 1,500 m (overall observed depth range 841-2,266 m). Images of juveniles, discarded molts, and precopulatory behavior, as well as gravid females in a trapping study, suggested a reproductively viable population on the slope. At the time of the survey, there was no thermal barrier to prevent the lithodids from expanding upward and emerging on the outer shelf (400- to 550-m depth); however, near-surface temperatures remained too cold for them to survive in inner-shelf and coastal environments (

Tide-related vertical migration behaviours which assist in horizontal transport of marine larvae have been documented in many species from meso-tidal systems; however, little is known about larval behaviour in areas where the tides do not produce predictable changes in water cur- rents. We assessed the vertical migration behaviour in larvae of the shore crab Carcinus maenas (L.) from the Swedish west coast, to explore possible transport mechanisms in a micro-tidal system. Repli- cated plankton samples were collected at a fixed station from 2 depths during flood and ebb tides, and analysed for effects of tides and light on temporal and vertical patterns in abundance. In a com- plementary experiment, first-stage zoeae were hatched in the laboratory from field-collected females and studied in constant darkness for endogenous circa-tidal or circa-diel rhythms. The abundance of different larval stages decreased from the first to the fourth zoeal stage by a factor 10, whereas mega- lopal densities were similar to those of the first zoea. This pattern in abundance indicates an off- shore/onshore transportation of larvae during ontogeny, suggesting a behavioural component in larval dispersal. However, contrary to what has been demonstrated in larvae from meso-tidal areas, none of the larval stages displayed a vertical or temporal pattern in abundance related to the tide, although a clear semi-diurnal tidal change of approximately 0.3 m was registered throughout the sampling periods. Instead, the highest densities of all larval stages were consistently sampled during low light conditions, indicating a diel pattern in swimming behaviour. This behaviour appears to be controlled by exogenous cues, since the laboratory experiment with first zoeae in constant darkness did not indicate any consistent tidal or diel rhythm in vertical migration. The lack of an endogenous circa-tidal vertical migration rhythm suggests that Swedish shore crabs are reproductively isolated from the British Isles populations, where this behaviour has been demonstrated to be inherited. The possibility that C. maenas larvae in micro-tidal systems may use predictable sea and land breezes as a mechanism for cross-shelf transport is discussed.

In marine benthic organisms with a pelagic larval phase, assessment of recruitment reg- ulation necessitates accurate estimates of larval supply and initial settlement densities. We assessed 2 commonly used methods for estimating decapod larval supply, plankton net tows and artificial settle- ment substrates (ASS), together with a new technique using passive-migration traps. The aim was to evaluate how these methods estimate larval supply and settlement of 3 common decapod species (the shore crab Carcinus maenas L., the brown shrimp Crangon crangon L. and the grass shrimps Palae- mon spp.) in a microtidal nursery area on the Swedish west coast, and to assess how these estimates relate to the juvenile recruitment of these species. Discrete plankton net tows outside the shallow nursery area collected a higher proportion of larvae at an early development stage, compared to the other methods, and produced only a snapshot of the variable water-column abundance of shore crab and grass shrimp larvae that correlated poorly with numbers collected with ASS at the same location. Artificial settlement substrates appeared to produce good, integrated relative estimates of shore crab larval supply and settlement. The number of larvae collected from ASS correlated significantly with larval abundance estimated by passive-migration traps in shallow nursery areas, and reflected changes in shore crab settlement densities in caged mussel habitats. We did not detect any effect of predation on ASS placed in nursery areas. However, a large proportion (an estimated 83%) of shore crab postlarvae appeared to emigrate from ASS immersed for periods longer than 12 h, possibly during the night. All grass shrimp larvae appeared to emigrate from the collectors within 24 h, suggesting that ASS immersed longer than 12 h do not produce useful integrated estimates of larval supply for this species. These results demonstrate the importance of assessing postlarval emigration patterns from ASS to optimize immersion and collection time, and to avoid confounded estimates of larval abun- dance and settlement. A new method using replicated passive migration traps that fished continuously in opposite directions (on-shore and off-shore) gave promising integrated estimates of net fluxes (total number immigrating minus number emigrating per unit time) of both pelagic postlarvae and early benthic juvenile stages of crabs and shrimp. The majority of the brown shrimp recruits were young juveniles, demonstrating the importance of incorporating juvenile migration in recruitment studies of motile benthic species. High numbers of shore crab and grass shrimp larvae were found to emigrate from the bay, indicating that many decapod postlarvae found in nursery areas may be transitional. The emigrating larvae were on average in an earlier development stage than those migrating to the bay. These short-term experiments demonstrate the importance of assessing larval development stage and migratory behavior when estimating larval supply and settlement for recruitment studies.

Distribution and biomass of green algal mats were studied in marine shallow (0–1 m) soft-bottom areas on the Swedish west coast from 1994 to 1996, by combining aerial photography surveys with ground truth sampling. Filamentous green algae, dominated by species of the genera Cladophora and Enteromorpha, were generally present throughout the study area during July and August, and largely absent in late April and early May. These algae occurred at 60 to 90% of the locations investigated during the summer, and were estimated to cover between 30 and 50% of the total area of shallow soft bottoms of the Swedish Skagerrak archipelago. The distributional patterns were similar during the three years of the investigation and appeared unrelated to annual local nutrient inputs from point sources and river discharge. We postulate that the apparent lack of such a relationship is due to an altered state of nutrient dynamics throughout the archipelago. Mechanisms are likely to involve long-term, diffuse elevations in nutrient levels in coastal waters of the Skagerrak and the Kattegat over several decades leading to current eutrophic conditions, exceeding nutrient requirements for abundant filamentous algal growth. Patterns of algal abundance in our study were largely related to physical factors such as exposure to wind, waves and water exchange under conditions where nutrient loads among embayments seemed to be unlimited. Further, our results show that sediments covered by algal mats had higher carbon and nitrogen contents than unvegetated sediments. We hypothesise that sustained high nutrient loads, manifested in extensive biomass of filamentous algae during summer months, are re-mineralised via decay and sedimentation in the benthic realm. Hence, accumulated carbon and nutrients in the sediment could, in turn, constitute the basic pool for future algal mat production overlying soft bottoms in areas where tidal exchange is limited.

Settlement and early juvenile stages are considered a bottleneck in the Life history of many epibenthic organisms because of high predation mortality. Nursery habitats may play an important role in mitigating settlement and post-settlement mortality by providing refuge from predation. We examined these relationships in postlarvae and early juvenile stages of the shore crab Carcinus maenas L. in laboratory and field tethering experiments. We studied habitat and size related habitat mortality using postlarvae and young juvenile crabs as prey, and various predators, including juvenile conspecifics, in several habitats common in shallow (0 to 1 m) soft bottom nursery areas on the Swedish west coast. Settling mortality was high in open sand (80 to 90%), whereas a significant habitat refuge was obtained in mussel beds, eelgrass and filamentous green algae, the latter yielding the lowest mortality (13 to 14%). Small differences in structural complexity of ephemeral macroalgae dramatically affected predation mortality of first instar crabs, with a significant refuge obtained only in algae of medium complexity. Predation rate on tethered crabs in the field was high (52 to 67%) only on the smallest crabs (

In search of an explanation for some of the greenest waters ever seen in coastal Antarctica and their possible link to some of the fastest melting glaciers and declining summer sea ice, the Amundsen Sea Polynya International Research Expedition (ASPIRE) challenged the capabilities of USAP and the RVIB Nathaniel B. Palmer (NBP) during Austral summer 2010-2011. We were well rewarded by both an extraordinary research platform and a truly remarkable oceanic setting (Figure 1). Here we provide further insights to the key questions that motivated our sampling approach during ASPIRE, and present some preliminary findings, while highlighting the value of the NBP for accomplishing complex, multi-faceted oceanographic research in such a challenging environment.