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Human activities degrade and fragment coastal marine habitats, reducing their structural complexity and making habitat edges a prevalent seascape feature. Though habitat edges frequently are implicated in reduced faunal survival and biodiversity, results of experiments on edge effects have been inconsistent, calling for a mechanistic approach to the study of edges that explicitly includes indirect and interactive effects of habitat alteration at multiple scales across biogeographic gradients. We used an experimental network spanning 17 eelgrass (Zostera marina) sites across the Atlantic and Pacific oceans and the Mediterranean Sea to determine (1) if eelgrass edges consistently increase faunal predation risk, (2) whether edge effects on predation risk are altered by habitat degradation (shoot thinning), and (3) whether variation in the strength of edge effects among sites can be explained by biogeographical variability in covarying eelgrass habitat features. Contrary to expectations, at most sites, predation risk for tethered crustaceans (crabs or shrimps) was lower along patch edges than in patch interiors, regardless of the extent of habitat degradation. However, the extent to which edges reduced predation risk, compared to the patch interior, was correlated with the extent to which edges supported higher eelgrass structural complexity and prey biomass compared to patch interiors. This suggests an indirect component to edge effects in which the impact of edge proximity on predation risk is mediated by the effect of edges on other key biotic factors. Our results suggest that studies on edge effects should consider structural characteristics of patch edges, which may vary geographically, and multiple ways that humans degrade habitats.

Spatially complex habitats provide refuge for prey and mediate many predator–prey interactions. Increasing anthropogenic pressures are eroding such habitats, reducing their complexity and potentially altering ecosystem stability on a global scale. Yet, we have only a rudimentary understanding of how structurally complex habitats create ecological refuges for most ecosystems. Better informed management decisions require an understanding of the mechanisms underpinning the provision of physical refuge and this may be linked to prey size, predator size and predator identity in priority habitats. We tested each of these factors empirically in a model biogenic reef system. Specifically, we tested whether mortality rates of blue mussels (Mytilus edulis) of different sizes differed among: (i) different forms of reef structural distribution (represented as ‘clumped’, ‘patchy’ and ‘sparse’); (ii) predator species identity (shore crab, Carcinus maenas and starfish, Asterias rubens); and (iii) predator size. The survival rate of small mussels was greatest in the clumped experimental habitat and larger predators generally consumed more prey regardless of the structural organisation of treatment. Small mussels were protected from larger A. rubens but not from larger C. maenas in the clumped habitats. The distribution pattern of structural objects, therefore, may be considered a useful proxy for reef complexity when assessing predator–prey interactions, and optimal organisations should be considered based on both prey and predator sizes. These findings are essential to understand ecological processes underpinning predation rates in structurally complex habitats and to inform future restoration and ecological engineering practices.

Similar environmental driving forces can produce similarity among geographically distant ecosystems. Coastal oceanic upwelling, for example, has been associated with elevated biomass and abundance patterns of certain functional groups, e.g., corticated macroalgae. In the upwelling system of Northern Chile, we examined measures of intertidal macrobenthic composition, structure and trophic ecology across eighteen shores varying in their proximity to two coastal upwelling centres, in a hierarchical sampling design (spatial scales of >1 and >10 km). The influence of coastal upwelling on intertidal communities was confirmed by the stable isotope values (δ13C and δ15N) of consumers, including a dominant suspension feeder, grazers, and their putative resources of POM, epilithic biofilm, and macroalgae. We highlight the utility of muscle δ15N from the suspension feeding mussel, Perumytilus purpuratus, as a proxy for upwelling, supported by satellite data and previous studies. Where possible, we used corrections for broader-scale trends, spatial autocorrelation, ontogenetic dietary shifts and spatial baseline isotopic variation prior to analysis. Our results showed macroalgal assemblage composition, and benthic consumer assemblage structure, varied significantly with the intertidal influence of coastal upwelling, especially contrasting bays and coastal headlands. Coastal topography also separated differences in consumer resource use. This suggested that coastal upwelling, itself driven by coastline topography, influences intertidal communities by advecting nearshore phytoplankton populations offshore and cooling coastal water temperatures. We recommend the isotopic values of benthic organisms, specifically long-lived suspension feeders, as in situ alternatives to offshore measurements of upwelling influence.

Exploration of the relationship between species diversity and ecological stability has occupied a prominent place in ecological research for decades. Yet, a key component of this puzzle—the contributions of individual species to the overall stability of ecosystems—remains largely unknown. Here, we show that individual species simultaneously stabilize and destabilize ecosystems along different dimensions of stability, and also that their contributions to functional (biomass) and compositional stability are largely independent. By simulating experimentally the extinction of three consumer species (the limpet Patella , the periwinkle Littorina and the topshell Gibbula ) from a coastal rocky shore, we found that the capacity to predict the combined contribution of species to stability from the sum of their individual contributions varied among stability dimensions. This implies that the nature of the diversity–stability relationship depends upon the dimension of stability under consideration, and may be additive, synergistic or antagonistic. We conclude that, although the profoundly multifaceted and context-dependent consequences of species loss pose a significant challenge, the predictability of cumulative species contributions to some dimensions of stability provide a way forward for ecologists trying to conserve ecosystems and manage their stability under global change. By simulating experimentally the extinction of three key grazer species from an intertidal community, the authors show that the contribution of individual species to different dimensions of ecological stability is highly context dependent, and may simultaneously be positive or negative.

Coastal habitats dominated by marine macroalgae typically exhibit high rates of primary productivity and play a key role in local and regional carbon cycles and stores. In temperate regions, large brown algae (i.e. kelps and fucoids) contribute significantly to macroalgal primary production, most of which is exported from source habitats as detritus. The ultimate fate of this detritus and the processes controlling detrital pathways into food webs and carbon cycles remain poorly understood. Based on field surveys, we quantified the biomass of kelp-derived detritus (wrack) at sandy and pebble-dominated shores in Ireland and conducted a manipulative field experiment to test for inter-specific differences in detritus degradation rates and the effect of macroinvertebrate detritivores. Overall, accumulated wrack biomass was similar on all shores but varied temporally depending on habitat type. Degradation rates and the nutritional (C:N) and chemical (polyphenol concentrations) properties differed among kelp species. Interestingly, exclusion of macroinvertebrate detritivores did not affect kelp degradation rates, C:N ratios or polyphenol content. Our findings show that rates of macroalgal breakdown differ among kelp species and that, in contrast to other aquatic systems, macroinvertebrates appear to play a very limited role in the breakdown of these marine detrital subsidies, suggesting a key role for meiofauna and microbes in detritus processing. Increasing recognition for the role of detritus in coastal food webs and carbon cycles warrants a better understanding of the mechanisms underpinning degradation rates.

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° 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 in situ 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.

Studies of trait-mediated indirect interactions (TMIIs) typically focus on effects higher predators have on per capita consumption by intermediate consumers of a third, basal prey resource. TMIIs are usually evidenced by changes in feeding rates of intermediate consumers and/or differences in densities of this third species. However, understanding and predicting effects of TMIIs on population stability of such basal species requires examination of the type and magnitude of the functional responses exhibited towards them. Here, in a marine intertidal system consisting of a higher-order fish predator, the shanny Lipophrys pholis, an intermediate predator, the amphipod Echinogammarus marinus, and a basal prey resource, the isopod Jaera nordmanni, we detected TMIIs, demonstrating the importance of habitat complexity in such interactions, by deriving functional responses and exploring consequences for prey population stability. Echinogammarus marinus reacted to fish predator diet cues by reducing activity, a typical anti-predator response, but did not alter habitat use. Basal prey, Jaera nordmanni, did not respond to fish diet cues with respect to activity, distribution or aggregation behaviour. Echinogammarus marinus exhibited type II functional responses towards J. nordmanni in simple habitat, but type III functional responses in complex habitat. However, while predator cue decreased the magnitude of the type II functional response in simple habitat, it increased the magnitude of the type III functional response in complex habitat. These findings indicate that, in simple habitats, TMIIs may drive down consumption rates within type II responses, however, this interaction may remain de-stabilising for prey populations. Conversely, in complex habitats, TMIIs may strengthen regulatory influences of intermediate consumers on prey populations, whilst potentially maintaining prey population stability. We thus highlight that TMIIs can have unexpected and complex ramifications throughout communities, but can be unravelled by considering effects on intermediate predator functional response types and magnitudes.

To understand the consequences of biodiversity loss, it is necessary to test how biodiversity-ecosystem functioning relationships may vary with predicted environmental change. In particular, our understanding will be advanced by studies addressing the interactive effects of multiple stressors on the role of biodiversity across trophic levels. Predicted increases in wave disturbance and ocean warming, together with climate-driven range shifts of key consumer species, are likely to have profound impacts on the dynamics of coastal marine communities. We tested whether wave action and temperature modified the effects of gastropod grazer diversity ( Patella vulgata , Littorina littorea , and Gibbula umbilicalis ) on algal assemblages in experimental rock pools. The presence or absence of L. littorea appeared to drive changes in microalgal and macroalgal biomass and macroalgal assemblage structure. Macroalgal biomass also decreased with increasing grazer species richness, but only when wave action was enhanced. Further, independently of grazer diversity, wave action and temperature had interactive effects on macroalgal assemblage structure. Warming also led to a reversal of grazer-macroalgal interaction strengths from negative to positive, but only when there was no wave action. Our results show that hydrodynamic disturbance can exacerbate the effects of changing consumer diversity, and may also disrupt the influence of other environmental stressors on key consumer-resource interactions. These findings suggest that the combined effects of anticipated abiotic and biotic change on the functioning of coastal marine ecosystems, although difficult to predict, may be substantial.

The mean and variance of ecological variables are dependent on sampling attributes such as the coverage of environmental heterogeneity (sampling extent) and spatial scale. Trophic niche width is often approximated by bulk tissue stable isotopes of C and N, that is, the population isotopic niche. However, recent studies suggest that environmental heterogeneity (experienced by individuals) may be more important in defining the isotopic niche width than trophic variability. We hypothesised that isotopic niche width will increase monotonically with spatial scale, largely produced by environmental variation, for example, nutrient source. To refine this hypothesis, by describing the shapes of isotope scaling curves, we explored a previously published dataset describing three Chilean intertidal species representing different feeding guilds (grazing snails, suspension‐feeding mussel). We tested these hypotheses on a new, larger dataset describing three functionally analogous intertidal species from Northern Ireland. We generated isotopic variance–area curves from a spatially explicit bootstrap and investigated the scale dependency of environment–isotope relationships, including wave exposure and sub‐habitat heterogeneity. Spatial scale explained 50% of the variance in population isotopic niche widths (bivariate C‐N ellipse area) by simple, nonlinear relationships. Finer scales (<1–10 km lag) accounted for most variance. Scale dependence was strong for δ15N variance, of which >40% was explained by modelling linear coefficients. A δ15N baseline gradient, or isoscape, dominated δ15N variance scaling patterns, from sheltered, terrestrially influenced embayments to exposed, pelagic‐dominated coastline. Consumer δ13C variance had a weaker scale dependence, plateauing at mesoscales (>20 km lag). We show that isotopic niche width is strongly dependent on sampling spatial extent, which controls the environmental heterogeneity experienced by individual consumers. Environmental heterogeneity must be accounted for before isotopic niche width can be considered to accurately represent trophic niche width. Studies conducted at different spatial scales are likely to identify different environment–isotope relationships. We recommend that spatial scale should be incorporated into sampling designs explicitly, easiest by maintaining a consistent lag distance or area within which populations are sampled. Identified isoscapes can be de‐trended, where necessary. A plain language summary is available for this article. Plain Language Summary