Hidden cost of pH variability in seagrass beds on marine calcifiers under ocean acidification.

Coastal ecosystems experience large environmental variability leading to local adaptation. The key role of variability and adaptation in modulating the biological sensitivity to ocean acidification is increasingly acknowledged. Monitoring and understanding the ecological niche at the right spatio-temporal scale is key to understand the sensitivity of any organism and ecosystems. However, the role of the variability in relevant carbonate chemistry parameters as a driver is often overlooked. For example, the balance between photosynthesis and respiration over the day/night cycle is leading to high pH/pCO2 variability in seagrass beds. We hypothesized that (i) the calcifying larvae of the sea urchin Echinus esculentus exposed to seagrass-driven variability would have some physiological mechanisms to respond to such variability; and (ii) these mechanisms would reach their limit under ocean acidification. We compared the presence and absence of the seagrass Zostera marina in flow through mesocosms fed with seawater with 4 pHs. The carbonate chemistry was monitored and biological response of a sea urchin larvae was documented over 3 weeks. Growth and net calcification rates were measured twice a day to encompass diurnal variability. Our results show that larvae growth rate significantly decreased with decreasing average pHT in both absence and presence of seagrass. Moreover, sea urchin larvae showed a slower growth rate in presence of seagrass, only visible in the lowest pH conditions. In addition, larvae raised in presence of seagrass, maximized calcification during the day, and lower their calcification during the night. In contrast, no significant difference was observed between day and night for the net calcification rate in larvae raised in absence of seagrass. Our results demonstrate the limit of local adaptation to the present range of variability under ocean acidification conditions. It also demonstrates that photosynthetic ecosystems such as seagrass may not play a role of refuge against future ocean acidification.

The role epiphytes play in particle capture of seagrass canopies.

Seagrass epiphytic communities act as ecological indicators of the quality status of vegetated coastal environments. This study aims to determine the effect leaf epiphytes has on the sediment capture and distribution from outside sources. Thirteen laboratory experiments were conducted under a wave frequency of 0.5 Hz. Three epiphyte models were attached to a Zostera marina canopy of 100 plants/m2 density. The sediment deposited to the seabed, captured by the epiphytic leaf surface, and remaining in suspension within the canopy were quantified. This study demonstrated that the amount of epiphytes impacts on the sediment stocks. Zostera marina canopies with high epiphytic areas and long effective leaf heights may increase the sediment captured on the epiphyte surfaces. Also, reducing suspended sediment and increasing the deposition to the seabed, therefore enhancing the clarity of the water column. For largest epiphytic areas, a 34.5% increase of captured sediment mass is observed. The sediment trapped on the leaves can be 10 times greater for canopies with the highest epiphytic areas than those without epiphytes. Therefore, both the effective leaf length and the level of epiphytic colonization are found to determine the seagrass canopy ability at distributing sediment.

Novel approach to large-scale monitoring of submerged aquatic vegetation: A nationwide example from Sweden.

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 2022;18:909-920. © 2021 The Authors. Integrated Environmental Assessment and Management published by Wiley Periodicals LLC on behalf of Society of Environmental Toxicology & Chemistry (SETAC).

Microplastic retention by marine vegetated canopies: Simulations with seagrass meadows in a hydraulic flume.

Marine canopies formed by seagrass and other coastal vegetated ecosystems could act as sinks of microplastics for being efficient particle traps. Here we investigated for the first time the occurrence of microplastic retention by marine canopies in a hydraulic flume under unidirectional flow velocities from 2 to 30 cm s-1. We used as model canopy-forming species the seagrass Zostera marina with four canopy shoot density (0, 50, 100, 200 shoots m-2), and we used as microplastic particles industrial pristine pellets with specific densities from 0.90 to 1.34 g cm-3 (polypropylene PP; polystyrene PS; polyamide 6 PA; and polyethylene terephthalate PET). Overall, microplastics particles transported with the flow were retained in the seagrass canopies but not in bare sand. While seagrass canopies retained floating microplastics (PP) only at low velocities (<12 cm s-1) due to a barrier created by the canopy touching the water surface, the retention of sinking particles (PS, PA, PET) occurred across a wider range of flow velocities. Our simulations revealed that less dense sinking particles (PS) might escape from the canopy at high velocities, while denser sinking particles can be trapped in scouring areas created by erosive processes around the eelgrass shoots. Our results show that marine canopies might act as potential barriers or sinks for microplastics at certain bio-physical conditions, with the probability of retention generally increasing with the seagrass shoot density and polymer specific density and decreasing with the flow velocity. We conclude that seagrass meadows, and other aquatic canopy-forming ecosystems, should be prioritized habitats in assessment of microplastic exposure and impact on coastal areas since they may accumulate high concentration of microplastic particles that could affect associated fauna.

The influence of hydrodynamic exposure on carbon storage and nutrient retention in eelgrass (Zostera marina L.) meadows on the Swedish Skagerrak coast.

Cold-temperate seagrass (Zostera marina) meadows provide several important ecosystem services, including trapping and storage of sedimentary organic carbon and nutrients. However, seagrass meadows are rapidly decreasing worldwide and there is a pressing need for protective management of the meadows and the organic matter sinks they create. Their carbon and nutrient storage potential must be properly evaluated, both at present situation and under future climate change impacts. In this study, we assessed the effect of wave exposure on sedimentary carbon and nitrogen accumulation using existing data from 53 Z. marina meadows at the Swedish west coast. We found that meadows with higher hydrodynamic exposure had larger absolute organic carbon and nitrogen stocks (at 0-25 cm depth). This can be explained by a hydrodynamically induced sediment compaction in more exposed sites, resulting in increased sediment density and higher accumulation (per unit volume) of sedimentary organic carbon and nitrogen. With higher sediment density, the erosion threshold is assumed to increase, and as climate change-induced storms are predicted to be more common, we suggest that wave exposed meadows can be more resilient toward storms and might therefore be even more important as carbon- and nutrient sinks in the future.

Severe shifts of Zostera marina epifauna: Comparative study between 1997 and 2018 on the Swedish Skagerrak coast.

The interaction between bottom-up and top-down processes in coastal ecosystems has been scarcely studied so far. Temporal changes in trophic interactions of Zostera marina along the Swedish west coast are relatively well studied, with the exception of epifaunal communities. Epifauna was used as a model study to explore resource (bottom-up) or predator (top-down) regulated in a vegetated ecosystem. We conducted a 21-year comparative study (1997 and 2018) using epifauna of 19 Zostera marina meadows along the Swedish Skagerrak coast. Large changes were observed in the composition of small (0.2-1 mm) and large (>1 mm) epifauna. In the small-sized epifauna, the nematode Southernia zosterae and harpacticoids showed an increase of 90% and a decrease of 50% of their abundances, respectively. In the large-sized epifauna, the polychaete Platynereis dumerilii and chironomid larvae were absent in 1997 but thrived in 2018 (>2000 ind. m-2). Mesoherbivores (Idoteids and gammarids) were locally very abundant in 1997 but disappeared in 2018. An 83% decline of mytilids settling in Zostera marina leaves was observed. Our results showed that epifauna is predominantly top-down regulated. An integrative framework of the study area is outlined to shed light on the causes and consequences of the environmental shifts reported in Zostera meadows from the northern Skagerrak area throughout the last three decades.

Mimicry of emergent traits amplifies coastal restoration success.

Restoration is becoming a vital tool to counteract coastal ecosystem degradation. Modifying transplant designs of habitat-forming organisms from dispersed to clumped can amplify coastal restoration yields as it generates self-facilitation from emergent traits, i.e. traits not expressed by individuals or small clones, but that emerge in clumped individuals or large clones. Here, we advance restoration science by mimicking key emergent traits that locally suppress physical stress using biodegradable establishment structures. Experiments across (sub)tropical and temperate seagrass and salt marsh systems demonstrate greatly enhanced yields when individuals are transplanted within structures mimicking emergent traits that suppress waves or sediment mobility. Specifically, belowground mimics of dense root mats most facilitate seagrasses via sediment stabilization, while mimics of aboveground plant structures most facilitate marsh grasses by reducing stem movement. Mimicking key emergent traits may allow upscaling of restoration in many ecosystems that depend on self-facilitation for persistence, by constraining biological material requirements and implementation costs.

The influence of hydrodynamics and ecosystem engineers on eelgrass seed trapping.

Propagule dispersal is an integral part of the life cycle of seagrasses; important for colonising unvegetated areas and increasing their spatial distribution. However, to understand recruitment success, seed dispersal and survival in habitats of different complexity remains to be quantified. We tested the single and synergistic effects of three commonly distributed ecosystem engineers-eelgrass (Zostera marina), oysters (Magellana gigas) and blue mussels (Mytilus edulis)-on trapping of Z. marina seeds in a hydraulic flume under currents. Our results suggest that seed retention increases with habitat complexity and further reveal insights into the underlying mechanisms. In eelgrass canopy, trapping occurred mostly through direct blocking of a seed's pathway, while trapping in bivalve patches was mainly related to altered hydrodynamics in the lee side, i.e. behind each specimen. With increasing flow velocity (24-30 cm s-1 in eelgrass canopy, 18-30 cm s-1 in bivalve patches), modifications of the sediment surface through increased turbulence and erosive processes became more important and resulted in high seed trapping rates. Furthermore, we show that while monospecific patches of seagrass and bivalves had different trapping optima depending on flow velocities, intermixing resulted in consistently high trapping rates throughout the investigated hydrodynamic gradient. Our results highlight the importance of positive interactions among ecosystem engineers for seed retention and patch emergence in eelgrass.

Seed Predation by the Shore Crab Carcinus maenas: A Positive Feedback Preventing Eelgrass Recovery?

There is an increasing interest to restore the ecosystem services that eelgrass provides, after their continuous worldwide decline. Most attempts to restore eelgrass using seeds are challenged by very high seed losses and the reasons for these losses are not all clear. We assess the impact of predation on seed loss and eelgrass establishment, and explore methods to decrease seed loss during restoration in the Swedish northwest coast. In a laboratory study we identified three previously undescribed seed predators, the shore crab Carcinus maenas, the hermit crab Pagurus bernhardus and the sea urchin Strongylocentrotus droebachiensis, of which shore crabs consumed 2-7 times more seeds than the other two species. The importance of shore crabs as seed predators was supported in field cage experiments where one enclosed crab caused 73% loss of seeds over a 1-week period on average (~ 21 seeds per day). Seedling establishment was significantly higher (14%) in cages that excluded predators over an 8-month period than in uncaged plots and cages that allowed predators but prevented seed-transport (0.5%), suggesting that seed predation constitutes a major source of seed loss in the study area. Burying the seeds 2 cm below the sediment surface prevented seed predation in the laboratory and decreased predation in the field, constituting a way to decrease seed loss during restoration. Shore crabs may act as a key feedback mechanism that prevent the return of eelgrass both by direct consumption of eelgrass seeds and as a predator of algal mesograzers, allowing algal mats to overgrow eelgrass beds. This shore crab feedback mechanism could become self-generating by promoting the growth of its own nursery habitat (algal mats) and by decreasing the nursery habitat (seagrass meadow) of its dominant predator (cod). This double feedback-loop is supported by a strong increase of shore crab abundance in the last decades and may partly explain the regime shift in vegetation observed along the Swedish west coast.