Eelgrass

Herbivory is a fundamental process that controls primary producer abundance and regulates energy and nutrient flows to higher trophic levels. Despite the recent proliferation of small-scale studies on herbivore effects on aquatic plants, there remains limited understanding of the factors that control consumer regulation of vascular plants in aquatic ecosystems. Our current knowledge of the regulation of primary producers has hindered efforts to understand the structure and functioning of aquatic ecosystems, and to manage such ecosystems effectively. We conducted a global meta-analysis of the outcomes of plant–herbivore interactions using a data set comprised of 326 values from 163 studies, in order to test two mechanistic hypotheses: first, that greater negative changes in plant abundance would be associated with higher herbivore biomass densities; second, that the magnitude of changes in plant abundance would vary with herbivore taxonomic identity. We found evidence that plant abundance declined with increased herbivore density, with plants eliminated at high densities. Significant between-taxa differences in impact were detected, with insects associated with smaller reductions in plant abundance than all other taxa. Similarly, birds caused smaller reductions in plant abundance than echinoderms, fish, or molluscs. Furthermore, larger reductions in plant abundance were detected for fish relative to crustaceans. We found a positive relationship between herbivore species richness and change in plant abundance, with the strongest reductions in plant abundance reported for low herbivore species richness, suggesting that greater herbivore diversity may protect against large reductions in plant abundance. Finally, we found that herbivore–plant nativeness was a key factor affecting the magnitude of herbivore impacts on plant abundance across a wide range of species assemblages. Assemblages comprised of invasive herbivores and native plant assemblages were associated with greater reductions in plant abundance compared with invasive herbivores and invasive plants, native herbivores and invasive plants, native herbivores and mixed-nativeness plants, and native herbivores and native plants. By contrast, assemblages comprised of native herbivores and invasive plants were associated with lower reductions in plant abundance compared with both mixed-nativeness herbivores and native plants, and native herbivores and native plants. However, the effects of herbivore–plant nativeness on changes in plant abundance were reduced at high herbivore densities. Our mean reductions in aquatic plant abundance are greater than those reported in the literature for terrestrial plants, but lower than aquatic algae. Our findings highlight the need for a substantial shift in how biologists incorporate plant–herbivore interactions into theories of aquatic ecosystem structure and functioning. Currently, the failure to incorporate top-down effects continues to hinder our capacity to understand and manage the ecological dynamics of habitats that contain aquatic plants.

Aquatic plants fulfil a wide range of ecological roles, and make a substantial contribution to the structure, function and service provision of aquatic ecosystems. Given their well-documented importance in aquatic ecosystems, research into aquatic plants continues to blossom. The 14th International Symposium on Aquatic Plants, held in Edinburgh in September 2015, brought together 120 delegates from 28 countries and six continents. This special issue of Hydrobiologia includes a select number of papers on aspects of aquatic plants, covering a wide range of species, systems and issues. In this paper, we present an overview of current trends and future directions in aquatic plant research in the early twenty first century. Our understanding of aquatic plant biology, the range of scientific issues being addressed and the range of techniques available to researchers have all arguably never been greater; however, substantial challenges exist to the conservation and management of both aquatic plants and the ecosystems in which they are found. The range of countries and continents represented by conference delegates and authors of papers in the special issue illustrates the global relevance of aquatic plant research in the early twenty first century but also the many challenges that this burgeoning scientific discipline must address.

Seagrasses and bare sediment represent alternative stable states, with sediment resuspension being a key driver of system stability via the Seagrass-Sediment-Light (SSL) feedback. We explore the SSL feedback by quantifying the sediment stabilization by seagrass, and using these measurements to calculate under which conditions seagrass ends up in a turbid environment. We quantified in-situ sediment resuspension velocity thresholds (u cr) for Zostera marina growing in medium to fine sand, using a field flume inducing near-bed wave motion. u cr was determined for full length shoots, shoots clipped to 0.08 m, and removed shoots. We found that rhizomes did not influence u cr of the top sediment layer. Overall, u cr was linearly related to blade area, which became independent for sediment type when normalizing u cr for the resuspension threshold after shoot removal. Comparing measured u cr against natural wave conditions showed that the seagrass meadow at the study site is currently stable. Exploring the effects of changing hydrodynamic conditions revealed that effects of increasing storminess has limited influence on sediment resuspension and thus the SSL-feedback. Increasing mean wind velocity had a stronger influence on SSLfeedback dynamics by causing more frequent exceedance of u cr. The response of seagrasses to increasing wind pressure depends on bay topography. A fully exposed Z. marina meadow under low initial turbidity pressure trended toward bistability, as turbidity pressure increased mainly on bare sediments. The study site and a fully exposed Z. marina meadow under high initial turbidity pressure saw an increase in turbidity across all blade areas.

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 dislodge-ment 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.

Changes in climate, food abundance and disturbance from humans threaten the ability of species to successfully use stopover sites and migrate between non-breeding and breeding areas. To devise successful conservation strategies for migratory species we need to be able to predict how such changes will affect both individuals and populations. Such predictions should ideally be process-based, focusing on the mechanisms through which changes alter individual physiological state and behavior. In this study we use a process-based model to evaluate how Black Brant (Branta bernicla nigricans) foraging on common eelgrass (Zostera marina) at a stopover site (Humboldt Bay, USA), may be affected by changes in sea level, food abundance and disturbance. The model is individual-based, with empirically based parameters, and incorporates the immigration of birds into the site, tidal changes in availability of eelgrass, seasonal and depth-related changes in eelgrass biomass, foraging behavior and energetics of the birds, and their mass-dependent decisions to emigrate. The model is validated by comparing predictions to observations across a range of system properties including the time birds spent foraging, probability of birds emigrating, mean stopover duration, peak bird numbers, rates of mass gain and distribution of birds within the site: all 11 predictions were within 35% of the observed value, and 8 within 20%. The model predicted that the eelgrass within the site could potentially support up to five times as many birds as currently use the site. Future predictions indicated that the rate of mass gain and mean stopover duration were relatively insensitive to sea level rise over the next 100 years, primarily because eelgrass habitat could redistribute shoreward into intertidal mudflats within the site to compensate for higher sea levels. In contrast, the rate of mass gain and mean stopover duration were sensitive to changes in total eelgrass biomass and the percentage of time for which birds were disturbed. We discuss the consequences of these predictions for Black Brant conservation. A wide range of migratory species responses are expected in response to environmental change. Process-based models are potential tools to predict such responses and understand the mechanisms which underpin them.

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 (N 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 lower (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.

Eelgrass (Zostera marina) flowering development and seed production were assessed along a depth gradient at three sites during 2012 and 2013 to 1) describe the flowering seasonality in the Swedish west coast, and 2) evaluate methods using seeds for large-scale restoration, including harvesting, storage and separation of viable seeds using a vertical flume. Eelgrass flowering shoots were found in the field from June to October, reaching the highest densities in July and August (average 3.8 ± 0.5 shoots m −2). Flowering shoot densities decreased with depth, whereas shoot length, number of spathes, seeds/shoot and seed size increased with depth, resulting in the highest seed production at intermediate depths (2 m) in most bays and years. Because of low densities of flowering shoots, seed production in Sweden (on average 39–126 seeds m −2 , Jul–Sep) was an order of magnitude lower than in other studied areas. Results showed that seed production differed 2.5–3.4 times between meadows and years, mainly driven by variation in seed production per shoot. This variation was only partly explained by temperature over the growing season, suggesting that other factors such as light and the amount of filamentous algal mats might also be important in flower development. Results suggest that flowering shoots should be harvested when > 50% of the spathes have developing seeds, and that shoots should not be stored longer than 40 days in tanks to obtain an optimal release of viable seeds. A new mechanized method using a vertical flume to separate large amounts of viable seeds from the harvest is also presented.

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 ...

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 À1. We used as model canopy-forming species the seagrass Zostera marina with four canopy shoot density (0, 50, 100, 200 shoots m À2), and we used as microplastic particles industrial pristine pellets with specific densities from 0.90 to 1.34 g cm À3 (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 À1) 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.

Colonization of eelgrass (Zostera marina L.) by tunicates can lead to reduced plant growth and survival. Several of the tunicate species that are found on eelgrass in the northwest Atlantic are highly aggressive colonizers, and range expansions are predicted in association with climate-change induced increases in seawater temperature. In 2017, we surveyed tunicates within eelgrass meadows at 33 sites from New Jersey to Newfoundland. Eight tunicate species were identified colonizing eelgrass, of which four were non-native and one was cryptogenic. The most common species (Botrylloides violaceus and Botryllus schlosseri) occurred from New York to Atlantic Canada. Tunicate faunas attached to eelgrass were less diverse north of Cape Cod, Massachusetts. Artificial substrates in the vicinity of the eelgrass meadows generally supported more tunicate species than did the eelgrass, but fewer species co-occurred in northern sites than southern sites. The latitudinal gradient in tunicate diversity corresponded to gradients of summertime sea surface temperature and traditional biogeographical zones in the northwest Atlantic, where Cape Cod represents a transition between cold-water and warm-water invertebrate faunas. Tunicate density in the eelgrass meadows was low, ranging generally from 1-25% cover of eelgrass shoots, suggesting that space availability does not currently limit tunicate colonization of eelgrass. This survey, along with our 2013 survey, provide a baseline for identifying future changes in tunicate distribution and abundance in northwest Atlantic eelgrass meadows.