Genetic characteristics influence the phenotype of marine macroalga Fucus vesiculosus (Phaeophyceae).

Intraspecific variation is an important component of heterogeneity in biological systems that can manifest at the genotypic and phenotypic level. This study investigates the influence of genetic characteristics on the phenotype of free-living Fucus vesiculosus using traditional morphological measures and microsatellite genotyping. Two sympatric morphotypes were observed to be significantly genetically and morphologically differentiated despite experiencing analogous local environmental conditions; indicating a genetic element to F. vesiculosus morphology. Additionally, the observed intraclonal variation established divergent morphology within some genets. This demonstrated that clonal lineages have the ability to alter morphological traits by either a plastic response or somatic mutations. We provide support for the potential occurrence of the Gigas effect (cellular/organ enlargement through genome duplication) in the Fucus genus, with polyploidization appearing to correlate with a general increase in the size of morphological features. Phenotypic traits, as designated by morphology within the study, of F. vesiculosus are partially controlled by the genetic characteristics of the thalli. This study suggests that largely asexually reproducing algal populations may have the potential to adapt to changing environmental conditions through genome changes or phenotypic plasticity.

Positive contribution of macrofaunal biodiversity to secondary production and seagrass carbon metabolism.

Coastal vegetated habitats such as seagrasses are known to play a critical role in carbon cycling and the potential to mitigate climate change, as blue carbon habitats have been repeatedly highlighted. However, little information is known about the role of associated macrofauna communities on the dynamics of critical processes of seagrass carbon metabolism (e.g., respiration, turnover, and production). We conducted a field study across a spatial gradient of seagrass meadows involving variable environmental conditions and macrobenthic diversity to investigate (1) the relationship between macrofauna biodiversity and secondary production (i.e., consumer incorporation of organic matter per time unit), and (2) the role of macrofauna communities in seagrass organic carbon metabolism (i.e., respiration and primary production). We show that, although several environmental factors influence secondary production, macrofauna biodiversity controls the range of local seagrass secondary production. We demonstrate that macrofauna respiration rates were responsible for almost 40% of the overall seafloor community respiration. Macrofauna represented on average >25% of the total benthic organic C stocks, high secondary production that is likely to become available to upper trophic levels of the coastal food web. Our findings support the role of macrofauna biodiversity in maintaining productive ecosystems, implying that biodiversity loss due to ongoing environmental change yields less productive seagrass ecosystems. Therefore, the assessment of carbon dynamics in coastal habitats should include associated macrofauna biodiversity elements if we aim to obtain robust estimates of global carbon budgets required to implement management actions for the sustainable functioning of the world's coasts.

Variable contributions of seafloor communities to ecosystem metabolism across a gradient of habitat-forming species.

The contributions of habitat-forming species to the biodiversity and ecosystem processes of marine and terrestrial ecosystems are widely recognized. Aquatic plants are considered foundation species in shallow ecosystems, as they maintain biodiversity and sustain many ecosystem functions such as primary production and respiration. Despite the increasing amount of biodiversity-ecosystem functioning experiments in seagrass habitats, the effects of benthic variability on ecosystem functioning are rarely investigated across spatially variable aquatic plant habitats. Here, we quantitatively link seasonal variability in seafloor metabolism (i.e. gross primary production and community respiration) with major benthic community components (i.e. microphytobenthos, aquatic plants and macrofauna) across a structural complexity gradient of habitat-forming species (in terms of shoot density and biomass), ranging from bare sand, to a sparse mixture of plants to a dense monospecific seagrass meadow. The increasing complexity gradient enhanced the magnitude of the relationships between benthic community and seafloor metabolism. The daily average seafloor metabolism per season at the bare site was similar to the sparse site, highlighting the role of microphytobenthos for seafloor metabolism in shallow unvegetated sediments. The contribution of the associated macrofauna to the seafloor respiration was similar to the aquatic plant community contribution. Infauna was the main macrofaunal component significantly explaining the seasonal variability of seafloor respiration. However, benthic community-metabolism relationships were stronger within the plant community than within the macrofauna community (i.e. steepest slopes and lowest p-values). Understanding these relationships are a priority since climate change and biodiversity loss are reducing habitat complexity around the world, jeopardizing valuable ecosystem functions and services.

Towards a sampling design for characterizing habitat-specific benthic biodiversity related to oxygen flux dynamics using Aquatic Eddy Covariance.

The Aquatic Eddy Covariance (AEC) technique has emerged as an important method to quantify in situ seafloor metabolism over large areas of heterogeneous benthic communities, enabling cross-habitat comparisons of seafloor productivity. However, the lack of a corresponding sampling protocol to perform biodiversity comparisons across habitats is impeding a full assessment of marine ecosystem metabolism. Here, we study a range of coastal benthic habitats, from rocky-bed communities defined by either perennial macroalgae or blue mussel beds to soft-sediment communities comprised of either seagrass, patches of different macrophyte species or bare sand. We estimated that the maximum contribution to the AEC metabolic flux can be found for a seafloor area of approximately 80 m2 with a 5 meter upstream distance of the instrument across all the habitats. We conducted a sampling approach to characterize and quantify the dominant features of biodiversity (i.e., community biomass) within the main seafloor area of maximum metabolic contribution (i.e., gross primary production and community respiration) measured by the AEC. We documented a high biomass contribution of the macroalgal Fucus vesiculosus, the seagrass Zostera marina and the macroinvertebrate Mytilus edulis to the net ecosystem metabolism of the habitats. We also documented a significant role of the bare sediments for primary productivity compared to vegetated canopies of the soft sediments. The AEC also provided insight into dynamic short-term drivers of productivity such as PAR availability and water flow velocity for the productivity estimate. We regard this study as an important step forward, setting a framework for upcoming research focusing on linking biodiversity metrics and AEC flux measurements across habitats.

Warming intensify CO2 flux and nutrient release from algal wrack subsidies on sandy beaches.

Algal wrack subsidies underpin most of the food web structure of exposed sandy beaches and are responsible of important biogeochemical processes that link marine and terrestrial ecosystems. The response in decomposition of algal wrack deposits to global warming has not been studied in ocean-exposed sandy beaches to date. With this aim, passive open top chambers (OTCs) were used to increase soil temperature within the range predicted by the IPCC for western Europe (between 0.5 and 1.5°C), following the hypothesis that the biogeochemical processing of macroalgal wrack subsidies would accelerate in response to temperature increase. The effect of temperature manipulation on three target substrates: fresh and aged macroalgae, and bare sand, was tested. Results indicated that a small warming (<0.5°C) affected the wrack decomposition process through traceable increases in soil respiration through CO2 flux, inorganic nutrients within the interstitial environment (N and P), sediment organic contents measured through the amount of proteins and microbial pool through the total soil DNA. The different responses of soil variables in the studied substrates indicated that the decomposition stage of stranded macroalgae influences the biogeochemical processing of organic matter in sandy beaches. Thus, CO2 fluxes, releases of organic and inorganic nutrients and microbial activity intensify in aged wrack deposits. Our results predict that expected global warming will increase the release of inorganic nutrients to the coastal ocean by 30% for the N (21 Gg/year) and 5.9% for P (14 Gg/year); that increase for the flow of C to the atmosphere as CO2 was estimated in 8.2% (523 Gg/year). This study confirms the key role of sandy beaches in recycling ocean-derived organic matter, highlighting their sensitivity to a changing scenario of global warming that predicts significant increases in temperature over the next few decades.

The role of dispersal mode and habitat specialization for metacommunity structure of shallow beach invertebrates.

Metacommunity ecology recognizes the interplay between local and regional patterns in contributing to spatial variation in community structure. In aquatic systems, the relative importance of such patterns depends mainly on the potential connectivity of the specific system. Thus, connectivity is expected to increase in relation to the degree of water movement, and to depend on the specific traits of the study organism. We examined the role of environmental and spatial factors in structuring benthic communities from a highly connected shallow beach network using a metacommunity approach. Both factors contributed to a varying degree to the structure of the local communities suggesting that environmental filters and dispersal-related mechanisms played key roles in determining abundance patterns. We categorized benthic taxa according to their dispersal mode (passive vs. active) and habitat specialization (generalist vs. specialist) to understand the relative importance of environment and dispersal related processes for shallow beach metacommunities. Passive dispersers were predicted by a combination of environmental and spatial factors, whereas active dispersers were not spatially structured and responded only to local environmental factors. Generalists were predicted primarily by spatial factors, while specialists were only predicted by local environmental factors. The results suggest that the role of the spatial component in metacommunity organization is greater in open coastal waters, such as shallow beaches, compared to less-connected environmentally controlled aquatic systems. Our results also reveal a strong environmental role in structuring the benthic metacommunity of shallow beaches. Specifically, we highlight the sensitivity of shallow beach macrofauna to environmental factors related to eutrophication proxies.

Disentangling the effects of solar radiation, wrack macroalgae and beach macrofauna on associated bacterial assemblages.

Wrack detritus plays a significant role in shaping community dynamics and food-webs on sandy beaches. Macroalgae is the most abundant beach wrack, and it is broken down by the combination of environmental processes, macrofauna grazing, and microbial degradation before returning to the sea as nutrients. The role of solar radiation, algal species and beach macrofauna as ecological drivers for bacterial assemblages associated to wrack was investigated by experimental manipulation of Laminaria ochroleuca and Sargassum muticum. We examined the effects of changes in solar radiation on wrack-associated bacterial assemblages by using cut-off filters: PAR + UVA + UVB (280-700 nm; PAB), PAR + UVA (320-700 nm; PA), PAR (400-700 nm; P), and a control with no filter (C). Results showed that moderate changes in UVR are capable to promote substantial differences on bacterial assemblages so that wrack patches exposed to full sunlight treatments (C and PAB) showed more similar assemblages among them than compared to patches exposed to treatments that blocked part of the solar radiation (P and PA). Our findings also suggested that specific algal nutrient quality-related variables (i.e. nitrogen, C:N ratio and phlorotannins) are main determinants of bacterial dynamics on wrack deposits. We showed a positive relationship between beach macrofauna, especially the most abundant and active wrack-users, the amphipod Talitrus saltator and the coleopteran Phaleria cadaverina, and both bacterial abundance and richness. Moderate variations in natural solar radiation and shifts in the algal species entering beach ecosystems can modify the role of wrack in the energy-flow of nearshore environments with unknown ecological implications for coastal ecosystems.

Detecting Subtle Shifts in Ecosystem Functioning in a Dynamic Estuarine Environment.

Identifying the effects of stressors before they impact ecosystem functioning can be challenging in dynamic, heterogeneous 'real-world' ecosystems. In aquatic systems, for example, reductions in water clarity can limit the light available for photosynthesis, with knock-on consequences for secondary consumers, though in naturally turbid wave-swept estuaries, detecting the effects of elevated turbidity can be difficult. The objective of this study was to investigate the effects of shading on ecosystem functions mediated by sandflat primary producers (microphytobenthos) and deep-dwelling surface-feeding macrofauna (Macomona liliana; Bivalvia, Veneroida, Tellinidae). Shade cloths (which reduced incident light intensity by ~80%) were deployed on an exposed, intertidal sandflat to experimentally stress the microphytobenthic community associated with the sediment surface. After 13 weeks, sediment properties, macrofauna and fluxes of oxygen and inorganic nutrients across the sediment-water interface were measured. A multivariate metric of ecosystem function (MF) was generated by combining flux-based response variables, and distance-based linear models were used to determine shifts in the drivers of ecosystem function between non-shaded and shaded plots. No significant differences in MF or in the constituent ecosystem function variables were detected between the shaded and non-shaded plots. However, shading reduced the total explained variation in MF (from 64% in non-shaded plots to 15% in shaded plots) and affected the relative influence of M. liliana and other explanatory variables on MF. This suggests that although shade stress may shift the drivers of ecosystem functioning (consistent with earlier investigations of shading effects on sandflat interaction networks), ecosystem functions appear to have a degree of resilience to those changes.

Responses of dune plant communities to continental uplift from a major earthquake: sudden releases from coastal squeeze.

Vegetated dunes are recognized as important natural barriers that shelter inland ecosystems and coastlines suffering daily erosive impacts of the sea and extreme events, such as tsunamis. However, societal responses to erosion and shoreline retreat often result in man-made coastal defence structures that cover part of the intertidal and upper shore zones causing coastal squeeze and habitat loss, especially for upper shore biota, such as dune plants. Coseismic uplift of up to 2.0 m on the Peninsula de Arauco (South central Chile, ca. 37.5º S) caused by the 2010 Maule earthquake drastically modified the coastal landscape, including major increases in the width of uplifted beaches and the immediate conversion of mid to low sandy intertidal habitat to supralittoral sandy habitat above the reach of average tides and waves. To investigate the early stage responses in species richness, cover and across-shore distribution of the hitherto absent dune plants, we surveyed two formerly intertidal armoured sites and a nearby intertidal unarmoured site on a sandy beach located on the uplifted coast of Llico (Peninsula de Arauco) over two years. Almost 2 years after the 2010 earthquake, dune plants began to recruit, then rapidly grew and produced dune hummocks in the new upper beach habitats created by uplift at the three sites. Initial vegetation responses were very similar among sites. However, over the course of the study, the emerging vegetated dunes of the armoured sites suffered a slowdown in the development of the spatial distribution process, and remained impoverished in species richness and cover compared to the unarmoured site. Our results suggest that when released from the effects of coastal squeeze, vegetated dunes can recover without restoration actions. However, subsequent human activities and management of newly created beach and dune habitats can significantly alter the trajectory of vegetated dune development. Management that integrates the effects of natural and human induced disturbances, and promotes the development of dune vegetation as natural barriers can provide societal and conservation benefits in coastal ecosystems.

Experimenting with ecosystem interaction networks in search of threshold potentials in real-world marine ecosystems.

Thresholds profoundly affect our understanding and management of ecosystem dynamics, but we have yet to develop practical techniques to assess the risk that thresholds will be crossed. Combining ecological knowledge of critical system interdependencies with a large-scale experiment, we tested for breaks in the ecosystem interaction network to identify threshold potential in real-world ecosystem dynamics. Our experiment with the bivalves Macomona liliana and Austrovenus stutchburyi on marine sandflats in New Zealand demonstrated that reductions in incident sunlight changed the interaction network between sediment biogeochemical fluxes, productivity, and macrofauna. By demonstrating loss of positive feedbacks and changes in the architecture of the network, we provide mechanistic evidence that stressors lead to break points in dynamics, which theory predicts predispose a system to a critical transition.

Sensitivity of heterogeneous marine benthic habitats to subtle stressors.

It is important to understand the consequences of low level disturbances on the functioning of ecological communities because of the pervasiveness and frequency of this type of environmental change. In this study we investigated the response of a heterogeneous, subtidal, soft-sediment habitat to small experimental additions of organic matter and calcium carbonate to examine the sensitivity of benthic ecosystem functioning to changes in sediment characteristics that relate to the environmental threats of coastal eutrophication and ocean acidification. Our results documented significant changes between key biogeochemical and sedimentary variables such as gross primary production, ammonium uptake and dissolved reactive phosphorus flux following treatment additions. Moreover, the application of treatments affected relationships between macrofauna communities, sediment characteristics (e.g., chlorophyll a content) and biogeochemical processes (oxygen and nutrient fluxes). In this experiment organic matter and calcium carbonate showed persistent opposing effects on sedimentary processes, and we demonstrated that highly heterogeneous sediment habitats can be surprisingly sensitive to subtle perturbations. Our results have important biological implications in a world with relentless anthropogenic inputs of atmospheric CO2 and nutrients in coastal waters.

Biogenic habitat transitions influence facilitation in a marine soft-sediment ecosystem.

Habitats are often defined by the presence of key species and biogenic features. However, the ecological consequences of interactions among distinct habitat-forming species in transition zones where their habitats overlap remain poorly understood. We investigated transition zone interactions by conducting experiments at three locations in Mahurangi Harbour, New Zealand, where the abundance of two habitat-forming marine species naturally varied. The two key species differed in form and function: One was a sessile suspension-feeding bivalve that protruded from the sediment (Atrina zelandica; Pinnidae); the other was a mobile infaunal urchin that bioturbated sediment (Echinocardium cordatum; Spatangoida). The experimental treatments established at each site reflected the natural densities of the species across sites (Atrina only, Echinocardium only, Atrina and Echinocardium together, and plots with neither species present). We identified the individual and combined effects of the two key species on sediment characteristics and co-occurring macrofauna. After five months, we documented significant treatment effects, including the highest abundance of co-occurring macrofauna in the Atrina-only treatments. However, the facilitation of macrofauna by Atrina (relative to removal treatments) was entirely negated in the presence of Echinocardium at densities >10 individuals/m2. The transitional areas in Mahurangi Harbour composed of co-occurring Atrina and Echinocardium are currently widespread and are probably more common now than monospecific patches of either individual species, due to the thinning of dense Atrina patches into sparser mixed zones during the last 10-15 years. Thus, although some ecologists avoid ecotones and habitat edges when designing experiments, suspecting that it will skew the extrapolation of results, this study increased our understanding of benthic community dynamics across larger proportions of the seascape and provided insights into temporal changes in community structure associated with patch dynamics. Particularly in situations where non-abrupt habitat transitions are commonplace, documentation of community dynamics in individual biogenic habitats and in mixed transition zones is required in order to scale-up and generalize results.

Detecting shifts in ecosystem functioning: the decoupling of fundamental relationships with increased pollutant stress on sandflats.

In this study, we investigated the influence of low level contamination by copper, lead and zinc on the functioning of estuarine sandflat ecosystems by comparing the strength and variability of relationships between benthic macrofauna and fluxes (oxygen and nutrients) at three clean and three mildly contaminated sites. Specifically, as indicators of ecosystem functioning, we examined relationships between bivalve biomass, total benthic respiration and ammonium release, and ammonium uptake and benthic primary production. Furthermore, a small amount of organic matter was added to experimental plots at all sites (35 g/0.2 m²) to evaluate stress-on-stress responses relative to controls. Relationships were strongest at the clean sites (steepest slopes, highest r² values, lowest p-values) and weakest at the mildly contaminated sites and in organically enriched plots. Our results suggest that changes in ecosystem functioning may be occurring at mild (

Exploring macroinvertebrate species distributions at regional and local scales across a sandy beach geographic continuum.

Exposed sandy beaches are highly dynamic ecosystems where macroinvertebrate species cope with extremely variable environmental conditions. The majority of the beach ecology studies present exposed beaches as physically dominated ecosystems where abiotic factors largely determine the structure and distribution of macrobenthic communities. However, beach species patterns at different scales can be modified by the interaction between different environmental variables, including biotic interactions. In this study, we examined the role of different environmental variables for describing the regional and local scale distributions of common macrobenthic species across 39 beaches along the North coast of Spain. The analyses were carried out using boosted regression trees, a relatively new technique from the field of machine learning. Our study showed that the macroinvertebrate community on exposed beaches is not structured by a single physical factor, but instead by a complex set of drivers including the biotic compound. Thus, at a regional scale the macrobenthic community, in terms of number of species and abundance, was mainly explained by surrogates of food availability, such as chlorophyll a. The results also revealed that the local scale is a feasible way to construct general predictive species-environmental models, since relationships derived from different beaches showed similar responses for most of the species. However, additional information on aspects of beach species distribution can be obtained with large scale models. This study showed that species-environmental models should be validated against changes in spatial extent, and also illustrates the utility of BRTs as a powerful analysis tool for ecology data insight.

Disturbance of sandflats by thin terrigenous sediment deposits: consequences for primary production and nutrient cycling.

The contributions of biodiversity to ecosystem functioning are increasingly recognized by ecologists, with biodiversity loss considered a significant threat to the maintenance of life-supporting ecosystem goods and services. Although ecologists have increased the amount of realism in biodiversity-ecosystem functioning (BEF) experiments, effects on functioning are rarely investigated in the field in conjunction with disturbances that affect biodiversity. Here, effects on functioning were investigated in situ following experimental disturbance of a realistic type and magnitude. Experimental deposits of terrigenous sediment (5 mm thickness) were applied at three intertidal sites in Whangapoua Harbour (Coromandel Peninsula, New Zealand), where soil erosion associated with logging is a serious problem. Experimental disturbance by terrigenous sediment impacted macrofaunal abundance and community composition (suspension and deposit feeders), and there were coincident shifts in ecosystem functioning at all three sites. Relative to controls, sediment-treated plots had higher rates of ammonium efflux (despite 50% fewer macrofaunal individuals) and lower rates of gross primary production and photosynthetic efficiency (despite similar sediment chlorophyll a concentrations). Judging from nutrient ratios in sediment pore water, microphyte primary producers living on the sediment surface in control plots were likely nitrogen limited (the normal situation for marine waters), whereas microphytes in sediment-treated plots were likely phosphate limited. Gross primary production and photosynthetic efficiency were significantly and positively correlated with ammonium uptake in control plots (r2 = 0.463 and 0.856, respectively) but not in treated plots (P > 0.05, r2 < 0.3). We suggest that the higher content of charged particles (clay) in the experimental deposits bound up phosphate, limiting its bio-availability, and shifted functional relationships between sandflat macrofauna and sediment primary productivity.

Limited impact of Sargassum muticum on native algal assemblages from rocky intertidal shores.

Invasive species represent a serious threat to natural ecosystems through a range of negative effects on native species in the region invaded. The invasive species Sargassum muticum has invaded several temperate regions worldwide including the Galician rocky shoreline (northwestern Spain) in Western Europe. The main aim of this study was to assess if colonization by S. muticum has any effect on native algal assemblages by experimental removal of S. muticum. We predicted that in those plots where S. muticum plants were removed, the structure of native algal assemblages would differ from that in plots where S. muticum plants were untouched. In addition, we predicted that the effect of Sargassum removal would be more important than other causes of variability at the small scale investigated. Results indicated limited impact of S. muticum on native assemblages. The impact was only evident on the total number of native taxa and two understory morpho-functional groups, filamentous and foliose algae, rather than on the entire macroalgal assemblages.