Trophic cascades occur in many ecosystems, but the factors regulating them are still elusive. We suggest that an overlooked factor is that trophic interactions (TIs) are often scale-dependent and possibly interact across spatial scales. To explore the role of spatial scale for trophic cascades, and particularly the occurrence of cross-scale interactions (CSIs), we collected and analysed food-web data from 139 stations across 32 bays in the Baltic Sea. We found evidence of a four-level trophic cascade linking TIs across two spatial scales: at bay scale, piscivores (perch and pike) controlled mesopredators (three-spined stickleback), which in turn negatively affected epifaunal grazers. At station scale (within bays), grazers on average suppressed epiphytic algae, and indirectly benefitted habitat-forming vegetation. Moreover, the direction and strength of the grazer-algae relationship at station scale depended on the piscivore biomass at bay scale, indicating a cross-scale interaction effect, potentially caused by a shift in grazer assemblage composition. In summary, the trophic cascade from piscivores to algae appears to involve TIs that occur at, but also interact across, different spatial scales. Considering scale-dependence in general, and CSIs in particular, could therefore enhance our understanding of trophic cascades.

Recreational boating increases globally and associated moorings are often placed in vegetated habitats important for fish recruitment. Meanwhile, assessments of the effects of boating on vegetation, and potential effects on associated fish assemblages are rare. Here, we analysed (i) the effect of small-boat marinas on vegetation structure, and (ii) juvenile fish abundance in relation to vegetation cover in shallow wave-sheltered coastal inlets. We found marinas to have lower vegetation cover and height, and a different species composition, compared to control inlets. This effect became stronger with increasing berth density. Moreover, there was a clear positive relationship between vegetation cover and fish abundance. We conclude that recreational boating and related moorings are associated with reduced cover of aquatic vegetation constituting important habitats for juvenile fish. We therefore recommend that coastal constructions and associated boating should be allocated to more disturbance tolerant environments (e.g. naturally wave-exposed shores), thereby minimizing negative environmental impacts.

BackgroundOrganism biomass is one of the most important variables in ecological studies, making biomass estimations one of the most common laboratory tasks. Biomass of small macroinvertebrates is usually estimated as dry mass or ash-free dry mass (hereafter ‘DM’ vs. ‘AFDM’) per sample; a laborious and time consuming process, that often can be speeded up using easily measured and reliable proxy variables like body size or wet (fresh) mass. Another common way of estimating AFDM (one of the most accurate but also time-consuming estimates of biologically active tissue mass) is the use of AFDM/DM ratios as conversion factors. So far, however, these ratios typically ignore the possibility that the relative mass of biologically active vs. non-active support tissue (e.g., protective exoskeleton or shell)—and therefore, also AFDM/DM ratios—may change with body size, as previously shown for taxa like spiders, vertebrates and trees.MethodsWe collected aquatic, epibenthic macroinvertebrates (>1 mm) in 32 shallow bays along a 360 km stretch of the Swedish coast along the Baltic Sea; one of the largest brackish water bodies on Earth. We then estimated statistical relationships between the body size (length or height in mm), body dry mass and ash-free dry mass for 14 of the most common taxa; five gastropods, three bivalves, three crustaceans and three insect larvae. Finally, we statistically estimated the potential influence of body size on the AFDM/DM ratio per taxon.ResultsFor most taxa, non-linear regression models describing the power relationship between body size and (i) DM and (ii) AFDM fit the data well (as indicated by low SE and high R2). Moreover, for more than half of the taxa studied (including the vast majority of the shelled molluscs), body size had a negative influence on organism AFDM/DM ratios.DiscussionThe good fit of the modelled power relationships suggests that the constants reported here can be used to quickly estimate organism dry- and ash-free dry mass based on body size, thereby freeing up considerable work resources. However, the considerable differences in constants between taxa emphasize the need for taxon-specific relationships, and the potential dangers associated with ignoring body size. The negative influence of body size on the AFDM/DM ratio found in a majority of the molluscs could be caused by increasingly thicker shells with organism age, and/or spawning-induced loss of biologically active tissue in adults. Consequently, future studies utilizing AFDM/DM (and presumably also AFDM/wet mass) ratios should carefully assess the potential influence of body size to ensure more reliable estimates of organism body mass.

Field surveys often show that high water turbidity limits cover of aquatic vegetation, while many small-scale experiments show that vegetation can reduce turbidity by decreasing water flow, stabilizing sediments, and competing with phytoplankton for nutrients. Here we bridged these two views by exploring the direction and strength of causal relationships between aquatic vegetation and turbidity across seasons (spring and late summer) and spatial scales (local and regional), using causal modeling based on data from a field survey along the central Swedish Baltic Sea coast. The two best-fitting regional-scale models both suggested that in spring, high cover of vegetation reduces water turbidity. In summer, the relationships differed between the two models; in the first model high vegetation cover reduced turbidity; while in the second model reduction of summer turbidity by high vegetation cover in spring had a positive effect on summer vegetation which suggests a positive feedback of vegetation on itself. Nitrogen load had a positive effect on turbidity in both seasons, which was comparable in strength to the effect of vegetation on turbidity. To assess whether the effect of vegetation was primarily caused by sediment stabilization or a reduction of phytoplankton, we also tested models where turbidity was replaced by phytoplankton fluorescence or sediment-driven turbidity. The best-fitting regional-scale models suggested that high sediment-driven turbidity in spring reduces vegetation cover in summer, which in turn has a negative effect on sediment-driven turbidity in summer, indicating a potential positive feedback of sediment-driven turbidity on itself. Using data at the local scale, few relationships were significant, likely due to the influence of unmeasured variables and/or spatial heterogeneity. In summary, causal modeling based on data from a large-scale field survey suggested that aquatic vegetation can reduce turbidity at regional scales, and that high vegetation cover vs. high sediment-driven turbidity may represent two self-enhancing, alternative states of shallow bay ecosystems.

Ecosystem multifunctionality is an increasingly popular concept used to approximate multifaceted ecosystem functioning, which in turn may help advance ecosystem-based management. However, while experimental studies have shown a positive effect of diversity on multifunctionality, observational studies from natural systems—particularly aquatic—are scarce. Here, we tested the relative importance of species richness and cover of rooted aquatic vegetation, as well as cover of the loose-lying form of the macroalgae bladderwrack (Fucus vesiculosus), for ecosystem multifunctionality in shallow bays along the western Baltic Sea coast. We estimated multifunctionality based on four indicators of functions that support ecosystem services: recruitment of large predatory fish, grazer biomass, inverted ‘nuisance’ algal biomass, and water clarity. Piecewise path analysis showed that multifunctionality was driven by high cover of rooted aquatic vegetation and bladderwrack, particularly when the two co-occurred. This synergistic effect was nearly three times as strong as a negative effect of land-derived nitrogen loading. Species richness of aquatic vegetation indirectly benefitted multifunctionality by increasing vegetation cover. Meanwhile, high bladderwrack cover tended to decrease vegetation species richness, indicating that bladderwrack has both positive and negative effects on multifunctionality. We conclude that managing for dense and diverse vegetation assemblages may mitigate effects of anthropogenic pressures (for example, eutrophication) and support healthy coastal ecosystems that provide a range of benefits. To balance the exploitation of coastal ecosystems and maintain their multiple processes and services, management therefore needs to go beyond estimation of vegetation cover and consider the diversity and functional types of aquatic vegetation.