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.Peer reviewe

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

Funding Information: We thank A. Toikkanen and N. Uotila for their valuable work in the laboratory, and I. McDonald for field assistance. We thank two anonymous reviewers for their very constructive comments that helped improve this paper. The study was funded by the Academy of Finland (Project ID 294853) and the University of Helsinki and Stockholm University strategic fund for collaborative research (The Baltic Bridge initiative). DL and ST were supported by a senior visiting research fellowship funded by the Walter and Andrée de Nottbeck Foundation. Publisher Copyright: © 2022 The Authors. Ecology published by Wiley Periodicals LLC on behalf of The Ecological Society of America.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.Peer reviewe

Seasonal ecosystem metabolism across shallow benthic habitats measured by aquatic eddy covariance

Abstract Shallow benthic habitats are hotspots for carbon cycling and energy flow, but metabolism (primary production and respiration) dynamics and habitat-specific differences remain poorly understood. We investigated daily, seasonal, and annual metabolism in six key benthic habitats in the Baltic Sea using ~?2900?h of in situ aquatic eddy covariance oxygen flux measurements. Rocky substrates had the highest metabolism rates. Habitat-specific annual primary production per m2 was in the order Fucus vesiculosus canopy?>?Mytilus trossulus reef?>?Zostera marina canopy?>?mixed macrophytes canopy?>?sands, whereas respiration was in the order M. trossulus?>?F. vesiculosus?>?Z. marina?>?mixed macrophytes?> sands?>?aphotic sediments. Winter metabolism contributed 22?31% of annual rates. Spatial upscaling revealed that benthic habitats drive >?90% of ecosystem metabolism in waters ≤5?m depth, highlighting their central role in carbon and nutrient cycling in shallow waters.Non peer reviewe

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

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

Abstract 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

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

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

Macrofauna communities across a seascape of seagrass meadows : environmental drivers, biodiversity patterns and conservation implications

Similar to other coastal biogenic habitats (e.g. tidal marshes, kelp forests, mangroves and coral reefs), a key function of seagrass meadows is the enhancement of biodiversity. Variability at multiple spatial scales is a driver of biodiversity, but our understanding of the response of macrofauna communities to variability of seagrass meadows is limited. We examined the macrofauna community structure (abundance and biomass) and diversity patterns (alpha- and beta-diversity) across a seascape gradient of eleven seagrass meadows differing in the number, composition and density of plant species. The variability of the macrobenthic communities was regulated by a combination of sedimentary (mainly for the infauna) and macrophyte (mainly for the epifauna) predictors. We demonstrate that the natural occurrence of drifting algae trapped in the aboveground complexity of the meadows benefits seagrass macrofauna. Seagrass-associated macrofauna showed a clear increase in abundance and alpha-diversity metrics with increasing habitat complexity attributes (i.e. shoot density, plant biomass and canopy height). Furthermore, partitioning of beta-diversity (i.e. the variation of species composition between sites) implied the replacement of some species by others between sites (i.e. spatial turnover) instead of a process of species loss (or gain) from site to site (i.e. nestedness). Therefore, the enhancement of macrofauna diversity across an increasing gradient of seagrass complexity, and the dominance of the turnover component suggest that devoting conservation efforts on many different types of meadows, including the less diverse, should be a priority for coastal habitat-management.Peer reviewe

Biplots showing the relationship between chlorophyll <i>a</i> vs. macrofauna abundance, and the main feeding guilds.

<p>The panels show the general relationship across the sampling area. Chlorophyll <i>a</i> (µg g⁻¹ sediment), total macrofauna abundance (number of individuals), main feeding guilds (abundance of deposit and suspension feeders; DF and SF, respectively). Plots are raw data. (Treatments: Organic Matter, Calcium carbonate, Mix, Control).</p