A Pleistocene legacy structures variation in modern seagrass ecosystems.

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.

The biogeography of community assembly: latitude and predation drive variation in community trait distribution in a guild of epifaunal crustaceans.

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.

Harnessing positive species interactions as a tool against climate-driven loss of coastal biodiversity.

Habitat-forming species sustain biodiversity and ecosystem functioning in harsh environments through the amelioration of physical stress. Nonetheless, their role in shaping patterns of species distribution under future climate scenarios is generally overlooked. Focusing on coastal systems, we assess how habitat-forming species can influence the ability of stress-sensitive species to exhibit plastic responses, adapt to novel environmental conditions, or track suitable climates. Here, we argue that habitat-former populations could be managed as a nature-based solution against climate-driven loss of biodiversity. Drawing from different ecological and biological disciplines, we identify a series of actions to sustain the resilience of marine habitat-forming species to climate change, as well as their effectiveness and reliability in rescuing stress-sensitive species from increasingly adverse environmental conditions.

Dispersal mitigates bacterial dominance over microalgal competitor in metacommunities.

Ecological theory suggests that a combination of local and regional factors regulate biodiversity and community functioning in metacommunities. The relative importance of different factors structuring communities likely changes over successional time, but to date this concept is scarcely documented. In addition, the few studies describing successional dynamics in metacommunity regulation have only focused on a single group of organisms. Here, we report results of an experimental study testing the effect size of initial local community composition and dispersal between local patches on community dynamics of benthic microalgae and their associated bacteria over community succession. Our results show that over time dispersal outweighed initial effects of community composition on microalgal evenness and biomass, microalgal ß-diversity, and the ratio of bacteria to microalgae. At the end of the experiment (ca. 20 microalgae generations), dispersal significantly decreased microalgal evenness and ß-diversity by promoting one regionally superior competitor. Dispersal also decreased the ratio of bacteria to microalgae, while it significantly increased microalgal biomass. These results suggest that the dispersal-mediated establishment of a dominant and superior microalgae species prevented bacteria from gaining competitive advantage over the autotrophs in these metacommunities, ultimately maintaining the provision of autotrophic biomass. Our study emphasizes the importance of time for dispersal to be a relevant community-structuring mechanism. Moreover, we highlight the need for considering multiple competitors in complex metacommunity systems to properly pinpoint the consequences of local change in dominance through dispersal for metacommunity function.

Latitude, temperature, and habitat complexity predict predation pressure in eelgrass beds across the Northern Hemisphere.

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.

Biodiversity mediates top-down control in eelgrass ecosystems: a global comparative-experimental approach.

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.

Predation and spatial connectivity interact to shape ecosystem resilience to an ongoing regime shift.

Ecosystem regime shifts can have severe ecological and economic consequences, making it a top priority to understand how to make systems more resilient. Theory predicts that spatial connectivity and the local environment interact to shape resilience, but empirical studies are scarce. Here, we use >7000 fish samplings from the Baltic Sea coast to test this prediction in an ongoing, spatially propagating shift in dominance from predatory fish to an opportunistic mesopredator, with cascading effects throughout the food web. After controlling for the influence of other drivers (including increasing mesopredator densities), we find that predatory fish habitat connectivity increases resilience to the shift, but only when densities of fish-eating top predators (seals, cormorants) are low. Resilience also increases with temperature, likely through boosted predatory fish growth and recruitment. These findings confirm theoretical predictions that spatial connectivity and the local environment can together shape resilience to regime shifts.

Cross-habitat interactions among bivalve species control community structure on intertidal flats.

Increasing evidence shows that spatial interactions between sedentary organisms can structure communities and promote landscape complexity in many ecosystems. Here we tested the hypothesis that reef-forming mussels (Mytilus edulis L.), a dominant intertidal ecosystem engineer in the Wadden Sea, promote abundances of the burrowing bivalve Cerastoderma edule L. (cockle) in neighboring habitats at relatively long distances coastward from mussel beds. Field surveys within and around three mussel beds showed a peak in cockle densities at 50-100 m toward the coast from the mussel bed, while cockle abundances elsewhere in the study area were very low. Field transplantation of cockles showed higher survival of young cockles (2-3 years old) and increased spat fall coastward of the mussel bed compared to within the bed and to areas without mussels, whereas growth decreased within and coastward of the mussel bed. Our measurements suggest that the observed spatial patterns in cockle numbers resulted from (1) inhibition effects by the mussels close to the beds due to preemptive algal depletion and deteriorated sediment conditions and (2) facilitation effects by the mussels farther away from the beds due to reduction of wave energy. Our results imply that these spatial, scale-dependent interactions between reef-forming ecosystem engineers and surrounding communities of sedentary benthic organisms can be an important determinant of the large-scale community structure in intertidal ecosystems. Understanding this interplay between neighboring communities of sedentary species is therefore essential for effective conservation and restoration of soft-bottom intertidal communities.

Wild pedigrees inform mutation rates and historic abundance in baleen whales.

Phylogeny-based estimates suggesting a low germline mutation rate (µ) in baleen whales have influenced research ranging from assessments of whaling impacts to evolutionary cancer biology. We estimated µ directly from pedigrees in four baleen whale species for both the mitochondrial control region and nuclear genome. The results suggest values higher than those obtained through phylogeny-based estimates and similar to pedigree-based values for primates and toothed whales. Applying our estimate of µ reduces previous genetic-based estimates of preexploitation whale abundance by 86% and suggests that µ cannot explain low cancer rates in gigantic mammals. Our study shows that it is feasible to estimate µ directly from pedigrees in natural populations, with wide-ranging implications for ecological and evolutionary research.

The seafloor from a trait perspective. A comprehensive life history dataset of soft sediment macrozoobenthos.

Biological trait analysis (BTA) is a valuable tool for evaluating changes in community diversity and its link to ecosystem processes as well as environmental and anthropogenic perturbations. Trait-based analytical techniques like BTA rely on standardised datasets of species traits. However, there are currently only a limited number of datasets available for marine macrobenthos that contain trait data across multiple taxonomic groups. Here, we present an open-access dataset of 16 traits for 235 macrozoobenthic species recorded throughout multiple sampling campaigns of the Dutch Wadden Sea; a dynamic soft bottom system where humans have long played a substantial role in shaping the coastal environment. The trait categories included in this dataset cover a variety of life history strategies that are tightly linked to ecosystem functioning and the resilience of communities to (anthropogenic) perturbations and can advance our understanding of environmental changes and human impacts on the functioning of soft bottom systems.

Stress ecology in fucus: abiotic, biotic and genetic interactions.

Stress regimes defined as the synchronous or sequential action of abiotic and biotic stresses determine the performance and distribution of species. The natural patterns of stress to which species are more or less well adapted have recently started to shift and alter under the influence of global change. This was the motivation to review our knowledge on the stress ecology of a benthic key player, the macroalgal genus Fucus. We first provide a comprehensive review of the genus as an ecological model including what is currently known about the major lineages of Fucus species with respect to hybridization, ecotypic differentiation and speciation; as well as life history, population structure and geographic distribution. We then review our current understanding of both extrinsic (abiotic/biotic) and intrinsic (genetic) stress(es) on Fucus species and how they interact with each other. It is concluded that (i) interactive stress effects appear to be equally distributed over additive, antagonistic and synergistic categories at the level of single experiments, but are predominantly additive when averaged over all studies in a meta-analysis of 41 experiments; (ii) juvenile and adult responses to stress frequently differ and (iii) several species or particular populations of Fucus may be relatively unaffected by climate change as a consequence of pre-adapted ecotypes that collectively express wide physiological tolerences. Future research on Fucus should (i) include additional species, (ii) include marginal populations as models for responses to environmental stress; (iii) assess a wider range of stress combinations, including their temporal fluctuations; (iv) better differentiate between stress sensitivity of juvenile versus adult stages; (v) include a functional genomic component in order to better integrate Fucus' ecological and evolutionary responses to stress regimes and (vi) utilize a multivariate modelling approach in order to develop and understand interaction networks.

Effects of altered offshore food webs on coastal ecosystems emphasize the need for cross-ecosystem management.

By mainly targeting larger predatory fish, commercial fisheries have indirectly promoted rapid increases in densities of their prey; smaller predatory fish like sprat, stickleback and gobies. This process, known as mesopredator release, has effectively transformed many marine offshore basins into mesopredator-dominated ecosystems. In this article, we discuss recent indications of trophic cascades on the Atlantic and Baltic coasts of Sweden, where increased abundances of mesopredatory fish are linked to increased nearshore production and biomass of ephemeral algae. Based on synthesis of monitoring data, we suggest that offshore exploitation of larger predatory fish has contributed to the increase in mesopredator fish also along the coasts, with indirect negative effects on important benthic habitats and coastal water quality. The results emphasize the need to rebuild offshore and coastal populations of larger predatory fish to levels where they regain their control over lower trophic levels and important links between offshore and coastal systems are restored.

Predator biomass and vegetation influence the coastal distribution of threespine stickleback morphotypes.

Intraspecific niche differentiation can contribute to population persistence in changing environments. Following declines in large predatory fish, eutrophication, and climate change, there has been a major increase in the abundance of threespine stickleback (Gasterosteus aculeatus) in the Baltic Sea. Two morphotype groups with different levels of body armor-completely plated and incompletely plated-are common in coastal Baltic Sea habitats. The morphotypes are similar in shape, size, and other morphological characteristics and live as one apparently intermixed population. Variation in resource use between the groups could indicate a degree of niche segregation that could aid population persistence in the face of further environmental change. To assess whether morphotypes exhibit niche segregation associated with resource and/or habitat exploitation and predator avoidance, we conducted a field survey of stickleback morphotypes, and biotic and abiotic ecosystem structure, in two habitat types within shallow coastal bays in the Baltic Sea: deeper central waters and shallow near-shore waters. In the deeper waters, the proportion of completely plated stickleback was greater in habitats with greater biomass of two piscivorous fish: perch (Perca fluviatilis) and pike (Esox lucius). In the shallow waters, the proportion of completely plated stickleback was greater in habitats with greater coverage of habitat-forming vegetation. Our results suggest niche segregation between morphotypes, which may contribute to the continued success of stickleback in coastal Baltic Sea habitats.

A spatial regime shift from predator to prey dominance in a large coastal ecosystem.

Regime shifts in ecosystem structure and processes are typically studied from a temporal perspective. Yet, theory predicts that in large ecosystems with environmental gradients, shifts should start locally and gradually spread through space. Here we empirically document a spatially propagating shift in the trophic structure of a large aquatic ecosystem, from dominance of large predatory fish (perch, pike) to the small prey fish, the three-spined stickleback. Fish surveys in 486 shallow bays along the 1200 km western Baltic Sea coast during 1979-2017 show that the shift started in wave-exposed archipelago areas near the open sea, but gradually spread towards the wave-sheltered mainland coast. Ecosystem surveys in 32 bays in 2014 show that stickleback predation on juvenile predators (predator-prey reversal) generates a feedback mechanism that appears to reinforce the shift. In summary, managers must account for spatial heterogeneity and dispersal to better predict, detect and confront regime shifts within large ecosystems.

Declines in predatory fish promote bloom-forming macroalgae.

In the Baltic Sea, increased dominance of ephemeral and bloom-forming algae is presently attributed to increased nutrient loads. Simultaneously, coastal predatory fish are in strong decline. Using field data from nine areas covering a 700-km coastline, we examined whether formation of macroalgal blooms could be linked to the composition of the fish community. We then tested whether predator or nutrient availability could explain the field patterns in two small-scale field experiments, by comparing joint effects on algal net production from nutrient enrichment with agricultural fertilizer and exclusion of larger predatory fish with cages. We also manipulated the presence of invertebrate grazers. The abundance of piscivorous fish had a strong negative correlation with the large-scale distribution of bloom-forming macroalgae. Areas with depleted top-predator communities displayed massive increases in their prey, small-bodied fish, and high covers of ephemeral algae. Combining the results from the two experiments showed that excluding larger piscivorous fish: (1) increased the abundance of small-bodied predatory fish; (2) changed the size distribution of the dominating grazers, decreasing the smaller gastropod scrapers; and (3) increased the net production of ephemeral macroalgae. Effects of removing top predators and nutrient enrichment were similar and additive, together increasing the abundance of ephemeral algae many times. Predator effects depended on invertebrate grazers; in the absence of invertebrates there were no significant effects of predator exclusion on algal production. Our results provide strong support for regional declines of larger predatory fish in the Baltic Sea promoting algal production by decreasing invertebrate grazer control. This highlights the importance of trophic interactions for ecosystem responses to eutrophication. The view emerges that to achieve management goals for water quality we need to consider the interplay between top-down and bottom-up processes in future ecosystem management of marine resources.

The Macroalgal Holobiont in a Changing Sea.

When studying the effects of climate change on eukaryotic organisms we often oversee a major ecological process: the interaction with microbes. Eukaryotic hosts and microbes form functional units, termed holobionts, where microbes play crucial roles in host functioning. Environmental stress may disturb these complex mutualistic relations. Macroalgae form the foundation of coastal ecosystems worldwide and provide important ecosystem services - services they could likely not provide without their microbial associates. Still, today we do not know how environmental stress will affect the macroalgal holobiont in an increasingly changing ocean. In this review, we provide a conceptual framework that contributes to understanding the different levels at which the holobiont and environment interact, and we suggest a manipulative experimental approach as a guideline for future research.

Recreational boating degrades vegetation important for fish recruitment.

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.

Light induced intraspecific variability in response to thermal stress in the hard coral Stylophora pistillata.

Recent research suggests that prior exposure of several months to elevated irradiance induces enhanced thermal tolerance in scleractinian corals. While this tolerance has been reported at the species level, individual coral colonies may react differently due to individual variability in thermal tolerance. As thermal anomalies are predicted to become common in the upcoming future, intraspecific variation may be key to the survival of coral populations. In order to study light-history based thermal stress responses on individual colonies, we developed a preliminary microcosm experiment where three randomly chosen, aquacultured colonies of the model coral Stylophora pistillata were exposed to two irradiance treatments (200 and 400 µmol photons m-2 s-1) for 31 days, followed by artificially induced heat stress (∼33.4 °C). We found different responses to occur at both the intraspecific and the intracolonial levels, as indicated by either equal, less severe, delayed, and/or even non-necrotic responses of corals previously exposed to the irradiance of 400 compared to 200 µmol photons m-2 s-1. In addition, all individual colonies revealed light-enhanced calcification. Finally, elevated irradiance resulted in a lower chlorophyll a concentration in one colony compared to the control treatment, and the same colony displayed more rapid bleaching compared to the other ones. Taken together, this study highlights the potential importance of intra-individual variability in physiological responses of scleractinian corals and provides recommendations for improving methodological designs for future studies.

Biotic habitat complexity controls species diversity and nutrient effects on net biomass production.

Canopy-forming plants and algae commonly contribute to spatial variation in habitat complexity for associated organisms and thereby create a biotic patchiness of communities. In this study, we tested for interaction effects between biotic habitat complexity and resource availability on net biomass production and species diversity of understory macroalgae by factorial field manipulations of light, nutrients, and algal canopy cover in a subtidal rocky-shore community. Presence of algal canopy cover and/or artificial shadings limited net biomass production and facilitated species diversity. Artificial shadings reduced light to levels similar to those under canopy cover, and net biomass production was significantly and positively correlated to light availability. Considering the comparable and dependent experimental effects from shadings and canopy cover, the results strongly suggest that canopy cover controlled net biomass production and species diversity by limiting light and thereby limiting resource availability for community production. Canopy cover also controlled experimental nutrient effects by preventing a significant increase in net biomass production from nutrient enrichment recorded in ambient light (no shading). Changes in species diversity were mediated by changes in species dominance patterns and species evenness, where canopy cover and shadings facilitated slow-growing crust-forming species and suppressed spatial dominance by Fucus vesiculosus, which was the main contributor to net production of algal biomass. The demonstrated impacts of biotic habitat complexity on biomass production and local diversity contribute significantly to understanding the importance of functionally important species and biodiversity for ecosystem processes. In particular, this study demonstrates how loss of a dominant species and decreased habitat complexity change the response of the remaining assembly to resource loading. This is of potential significance for marine conservation since resource loading often promotes low habitat complexity and canopy species are among the first groups lost in degraded aquatic systems.

Boating and navigation activities influence the recruitment of fish in a Baltic Sea archipelago area.

We studied the effects of boating and navigation activities on the recruitment of coastal fish in the Stockholm archipelago in the NW Baltic proper. The impacts were quantified by sampling metamorphosed young-of-the-year (Y-O-Y) fish in inlets adjacent to i) routes for medium-sized passenger ferries; ii) berths (small marinas) with small boats; and iii) references. Species with high preference for vegetation were negatively influenced by boating and navigation activities and species with low preference positively influenced. Pike (Esox lucius) Y-O-Y were significantly more abundant in reference areas, while bleak (Alburnus alburnus) were more abundant in dredged marinas. No statistically significant patterns were identified for perch (Perca fluviatilis) although there was a trend of low abundance along ferry routes. Many species of nearshore fishes are dependent on submerged vegetation as spawning and larval substrate, structural refuge and feeding habitat. Our results suggest that the negative effects from boating and navigation activities on the coverage and height of vegetation, especially on species of Chara and Potamogeton spp., may contribute to changes in the Y-O-Y fish community.