Connections between hydrodynamics, benthic landscape and associated fauna in the Balearic Islands, western Mediterranean

Linking the abiotic and biotic traits of ecosystems is a critical step towards understanding their structure and functioning. Here we attempt to determine the connections between the hydrodynamics, benthic landscape and the associated fish communities on the coastal continental shelf off the Balearic Islands (western Mediterranean). Specifically we investigate the role the hydrodynamics play in shaping the benthic landscape, and whether the hydrodynamics affect the composition and structure of demersal fish communities. A realistic numerical model was used to establish the hydrodynamic characteristics of the area. The study area showed high hydrodynamic variability on a medium spatial scale (tens of km) in terms of mean water velocity (ū). Principal component analysis was used to determine the main gradients of macro-epibenthic variability. Redundancy analysis (RDA) was used to model the effect of the hydrodynamics on macro-epibenthic species. RDA was also used to model the effect of the hydrodynamics and macro-epibenthos on the abundance of the associated fish fauna, and on its biomass at a community level using biomass spectra classes. The results showed that the hydrodynamics had a significant influence on the distribution of both macro-epibenthic species and the associated fish species. The latter was also influenced by the macro-epibenthos. Fish size appeared to be a key attribute for the distribution of species across gradients of ū and macro-epibenthic change. Our findings can be applied in ecosystem-based fisheries management, as they show that it is necessary to take into account both the biotic and abiotic traits of the habitats when the habitat use and requirements of the associated species are defined.     

Early warning indicators for river nutrient and sediment loads in tropical seagrass beds: a benchmark from a near-pristine archipelago in Indonesia

In remote, tropical areas human influences increase, potentially threatening pristine seagrass systems. We aim (i) to provide a bench-mark for a near-pristine seagrass system in an archipelago in East Kalimantan, by quantifying a large spectrum of abiotic and biotic properties in seagrass meadows and (ii) to identify early warning indicators for river sediment and nutrient loading, by comparing the seagrass meadow properties over a gradient with varying river influence. Abiotic properties of water column, pore water and sediment were less suitable indicators for increased sediment and nutrient loading than seagrass properties. Seagrass meadows strongly responded to higher sediment and nutrient loads and proximity to the coast by decreasing seagrass cover, standing stock, number of seagrass species, changing species composition and shifts in tissue contents. Our study confirms that nutrient loads are more important than water nutrient concentrations. We identify seagrass system variables that are suitable indicators for sediment and nutrient loading, also in rapid survey scenarios with once-only measurements.     

On the potential of plant species invasion influencing bio‐geomorphologic landscape formation in salt marshes

Species invasions are known to change biotic and abiotic ecosystem characteristics such as community structure, cycling of materials and dynamics of rivers. However, their ability to alter interactions between biotic and abiotic ecosystem components, in particular bio‐geomorphic feedbacks and the resulting landscape configuration in tidal wetlands, such as tidal channels have not yet been demonstrated. We studied the impact of altered bio‐geomorphic feedbacks on geomorphologic features (i.e. tidal wetland channels), by comparing proxies for channel network geometry (unchanneled flow lengths, fractal dimension) over time between non‐invaded and invaded salt marsh habitats. The non‐invaded habitats (the south of eastern Chongming Island, Yangtze estuary, China) show little change in network geometry over time with a tendency for an increased drainage density. The invaded site (salt marshes in the north of eastern Chongming Island invaded by the exotic plant species Spartina alterniflora) showed a decreasing tendency in channel drainage density throughout and after the species invasion. This suggests that species invasions might not only affect biotic ecosystem characteristics, but also their ability to change bio‐geomorphic feedback loops, potentially leading to changes in existing geomorphologic features and therefore landscape configuration. Our results further suggest that the species invasion also altered sediment composition. Based on observations we propose a mechanism explaining the change in channel drainage density by an alteration in plant properties. The physical and physiological characteristics of the invading species Spartina alterniflora clearly differ from the native species Scirpus mariqueter, inducing different bio‐geomorphic feedback loops leading to the observed change in salt marsh channel configuration. Copyright © 2016 John Wiley & Sons, Ltd.     

The colonization success of an exotic Poaceae is related to native macrophyte richness, wind disturbance and riparian vegetation

Biotic and abiotic factors are filters that prevent invasions in aquatic and terrestrial ecosystems. In this investigation we tested the hypothesis that the success of a non-native Poaceae (Urochloa subquadripara) is positively correlated with the richness of native macrophytes and negatively correlated with wind disturbance (fetch) and presence of riparian vegetation on coarse spatial scales. Our samplings were carried out in a tropical reservoir (Rosana Reservoir, Brazil). We first compared competing models using the Akaike criterion to find the main combinations of explanatory variables (native macrophyte richness, fetch, and presence of riparian vegetation) associated with the success of U. subquadripara. Then, we applied multiple regressions to assess the coefficient of determination of the best models selected according to the Akaike criterion. The probability of occurrence of U. subquadripara increased significantly with increases in the number of native macrophyte species, but decreased with fetch and the presence of riparian vegetation. Stand width and maximum depth of occurrence (indicators of the success of this Poaceae) were also positively related with native richness and negatively with fetch and riparian vegetation. Our results supported our expectation that wave disturbance is an important variable explaining U. subquadripara success. Because the less exposed sites are also more favorable for colonization by natives, positive relationships between the success of non-native species and native diversity emerge at the coarse scale. Taken together, our results support the theory of “biotic acceptance”; that is, favorable sites are more prone to colonization by both native and non-native species.     

Towards a predictive model to assess the natural position of the Posidonia oceanica seagrass meadows upper limit

The upper portion of the meadows of the protected Mediterranean seagrass Posidonia oceanica occurs in the region of the seafloor mostly affected by surf-related effects. Evaluation of its status is part of monitoring programs, but proper conclusions are difficult to draw due to the lack of definite reference conditions. Comparing the position of the meadow upper limit with the beach morphodynamics (i.e. the distinctive type of beach produced by topography and wave climate) provided evidence that the natural landwards extension of meadows can be predicted. An innovative model was therefore developed in order to locate the region of the seafloor where the meadow upper limit should lie in natural conditions (i.e. those governed only by hydrodynamics, in absence of significant anthropogenic impact). This predictive model was validated in additional sites, which showed perfect agreement between predictions and observations. This makes the model a valuable tool for coastal management.     

Importance of habitat heterogeneity and biotic processes in the spatial distribution of a riparian herb (Carex remota L.): a point process approach

This study attempts to understand the dependence on abiotic factors and on the biotic process of the population development. We used three spatial point process models (Poisson, Area-Interaction and shot-noise Cox processes) in both homogenous and inhomogeneous versions to model the distribution of three Carex remota cohorts in wet zones of a temperate forest in the north of Spain. The cohorts studied were adults and seedlings born in two consecutive years. With the use of these models we are able to simulate separately and jointly the effect on plant distribution of a homogeneous or heterogeneous habitat, and the absence or presence of some biotic processes, as seed dispersal and/or density-dependent interactions. The result of the bivariate function analysis does not reveal sufficient evidences, but suggests a weak positive relation between adults and seedlings that survived a dry period in the first summer. Models from the three cohorts show a decreasing degree of clustering from seedlings to adults. Besides, the results show that the importance of the main factors that explain the population structure changes along the development of Carex stages. Compared to seedlings, the adults pattern shows an increasing dependence on abiotic factors.     

Factors driving the seasonal distribution of planktonic and epiphytic ciliates in a eutrophicated Mediterranean Lagoon

We studied the distribution of planktonic and epiphytic ciliates coupled with environmental factors and microalgae abundance at five stations in Ghar El Melh Lagoon (Tunisia). Planktonic ciliates were monitored for a year and epiphytic ciliates were sampled during summer 2011 in concordance with the proliferation of the seagrass Ruppia cirrhosa. Ciliate assemblage was largely dominated by Spirotrichea followed respectively by Tintinnida of and Strombidiida. No significant difference was found in the distribution of ciliate species among the stations. Redundancy analysis indicates that abiotic factors (temperature and nutriments) have a significant effect on the dynamics of certain ciliates. For epiphytic ciliates, 4 species were identified: Tintinnopsis campanula, Aspidisca sp., Strombidium acutum and Amphorides amphora. Based on PERMANOVA analyses, ciliates exhibit significant correlations among months and stations. According to ACP, epiphyte distribution follows roughly those of R. cirrhosa and pH. Significant correlations were found between harmful dinoflagellates and both planktonic and epiphytic ciliates.     

Epiphyte loads on seagrasses and microphytobenthos abundance are not reliable indicators of nutrient availability in oligotrophic coastal ecosystems

Despite marked gradients in nutrient availability that control the abundance and species composition of seagrasses in south Florida, and the importance of nutrient availability in controlling abundance and composition of epiphytes on seagrasses in other locations, we did not find that epiphyte load on the dominant seagrass, Thalassia testudinum, or that the relative contribution of algal epiphytes to the epiphyte community, was positively correlated with nutrient availability in the water column or the sediment in oligotrophic seagrass beds. Further, the abundance of microphytobenthos, as indicated by Chlorophyll-a concentration in the sediments, was not directly correlated with concentrations of nutrients in the sediments. Our results suggest that epiphyte and microphytobenthos abundance are not unambiguous indicators of nutrient availability in relatively pristine seagrass environments, and therefore would make poor candidates for indicators of the status and trends of seagrass ecosystems in relatively low-nutrient environments like the Florida Keys.     

Comparison of methods for integrating biological and physical data for marine habitat mapping and classification

An important first step in marine spatial planning and ecosystem-based management efforts is the creation of benthic habitat maps that allow scientists and managers to understand the distribution of living and non-living resources on the seafloor. However, the location of boundaries between and composition of habitats is highly dependent on the approach taken to integrate abiotic and biotic information. The purpose of this study was to test “top-down” and “bottom-up” approaches for integrating physical and biological data derived from commonly used sub-tidal benthic mapping tools to create a habitat map compatible with the US Coastal and Marine Ecological Classification Standard (CMECS). We found that a top-down framework, where we tested for differences in macrofauna assemblages among side scan sonar facies, defined two broad-scale and general habitats. Using the bottom-up approach, where patterns in abiotic and biotic variables were examined with multivariate statistics (BEST, LINKTREE, ANOSIM, SIMPER), we generated seven biotopes based on the macrofauna abundance, percent sand, water depth, and backscatter standard deviation that corresponded well to, but provided more fine-scale detail than the top-down habitats. We were able to use the statistical relationship between abiotic variables and macrofauna assemblages in the LINKTREE to predict the spatial distribution of assemblages over ∼50% of the study area. We created a local catalogue of biotopes specific to our study area that contributes to the CMECS library. In addition, we were able to fully map CMECS Geoform, Surface Geology, and Biotic Cover Components. This mapping effort represented real progress toward reconciling the “data density mismatch” between physical and biological mapping methods, and it provided further evidence that using a bottom-up methodology preserves species–environment relationships.     

Seasonal variations of biomarker responses in the marine blue mussel (Mytilus spp.)

Biomarkers, which can detect changes at the biochemical level, have been used for many years as early warning tools in environmental monitoring. In order to distinguish between natural variability and the potential effects of anthropogenic pollution, it is essential to identify background levels and establish the potential influence of abiotic (season, temperature and salinity) and biotic (gametogenesis) factors. In this study, we examined various biomarkers of stress (glutathione S-transferase (GST)), reproduction (vitellin-like proteins) and damage (lipid peroxidation (LPO) and DNA damage (DNA)) in marine mussels (Mytilus spp.) from a known pristine hybrid zone. Levels of all biomarker expression varied between seasons and appeared to be linked to the reproductive cycle. Oxidative stress in winter, with low GST expression and a higher expression of LPO and DNA damage displayed could be explained by low food availability. These data provide vital baseline information for future ecotoxicological and environmental monitoring studies.     

Differential kinetics and temperature dependence of abiotic and biotic processes controlling the environmental fate of TNT in simulated marine systems

This work seeks to understand how the balance of abiotic and biotic kinetic processes in sediments control the residual concentration of TNT in marine systems after release from ocean-dumped source. Kinetics of TNT disappearance were followed using marine sediments at different temperatures and under both biotic and presumably abiotic conditions (through sodium azide addition). Sediments exhibiting the highest rate of TNT disappearance under biotic conditions also exhibited the highest sorption affinity for TNT under abiotic conditions. Significant temperature dependence in the abiotic processes was observed in the diffusion coefficient of TNT and not sediment sorption affinity. At higher temperature, kinetics of biotic processes outpaced abiotic processes, but at low temperature, kinetics of abiotic processes were much more significant. We concluded that the differential influence of temperature on the kinetics of abiotic and biotic processes could provide distinguishing predictions for the potential residual concentration of TNT contamination in marine-sediment systems.     

Prediction of the consumption rates for juvenile Atlantic salmon (Salmo salar L.) on the scale of habitat patches: Development of empirical models based on stomach contents

Rivers and streams are unstable environments in which estimation of energetic costs and benefits of habitat utilization are the daunting exercise. Empirical models of food consumption may be used to estimate energetic benefits based on abiotic and biotic conditions in patches of habitat. We performed thirty daily surveys of fish stomach contents to estimate the consumption rates for juvenile Atlantic salmon (Salmo salar) in a river. The data were used to assess whether variations of daily consumption rates existed within the river, and to develop empirical models that could predict fish consumption rates using abiotic and biotic conditions as independent variables. Daily consumption rates based on stomach content surveys in the field (range: 0.15–1.49 g dry/(100 g wet day)) varied significantly depending on habitat patch (500–1000 m²), summer period, and sampling year. Variables such as water temperature, numerical density of salmon, water depth and moon phase explained 83–93% of the variations in daily food consumption rates. Daily consumption rates tended to increase with water temperature and depth, and were also higher near a full moon. However, they tended to decrease with the numerical density of salmon. Our work suggests that empirical models based on independent variables that are relatively simple to estimate in the field may be developed to predict fish consumption rates in different habitat patches in a river.     

Modelling the effects of Zostera noltei meadows on sediment dynamics: application to the Arcachon lagoon

A three-dimensional model has been modified to describe the complex interactions between hydrodynamics, sediment dynamics and biological parameters in the presence of Zostera noltei. The model treats seagrass leafs as flexible blades that bend under hydrodynamic forcing and alter the local momentum and turbulence fluxes and, therefore, the benthic shear conditions; these changes cause related changes to the mass balance at the boundary of the bed, in turn affecting the suspended matter in the column and ultimately primary productivity and the growth of the dwarf-grass. Modelling parameters related to the impact of Z. noltei to the local flow and to erosion and deposition rates were calibrated using flume experimental measurements; results from the calibration of the model are presented and discussed. The coupled model is applied in the Arcachon Bay, an area with high environmental significance and large abundance of dwarf-grass meadows. In the present paper, results from preliminary applications of the model are presented and discussed; the effectiveness of the coupled model is assessed comparing modelling results with available field measurements of suspended sediment concentrations and seagrass growth parameters. The model generally reproduces sediment dynamics and dwarf-grass seasonal growth in the domain efficiently. Investigations regarding the effects of the vegetation to the near-bed hydrodynamics and to the sediment suspension in the domain show that dwarf-grass meadows play an important part to velocity attenuation and to sediment stabilisation, with flow and suspended sediment concentrations damping, compared to an unvegetated state, to reach 35–50 and 65 %, respectively, at peak seagrass growth.     

Net ecosystem CO<Subscript>2</Subscript> exchange and controlling factors in a steppe—<Emphasis Type="Italic">Kobresia</Emphasis> meadow on the Tibetan Plateau

Knowledge of seasonal variation of net ecosystem CO₂ exchange (NEE) and its biotic and abiotic controllers will further our understanding of carbon cycling process, mechanism and large-scale modelling. Eddy covariance technique was used to measure NEE, biotic and abiotic factors for nearly 3 years in the hinterland alpine steppe—Korbresia meadow grassland on the Tibetan Plateau, the present highest fluxnet station in the world. The main objectives are to investigate dynamics of NEE and its components and to determine the major controlling factors. Maximum carbon assimilation took place in August and maximum carbon loss occurred in November. In June, rainfall amount due to monsoon climate played a great role in grass greening and consequently influenced interannual variation of ecosystem carbon gain. From July through September, monthly NEE presented net carbon assimilation. In other months, ecosystem exhibited carbon loss. In growing season, daytime NEE was mainly controlled by photosynthetically active radiation (PAR). In addition, leaf area index (LAI) interacted with PAR and together modulated NEE rates. Ecosystem respiration was controlled mainly by soil temperature and simultaneously by soil moisture. Q ₁₀ was negatively correlated with soil temperature but positively correlated with soil moisture. Large daily range of air temperature is not necessary to enhance carbon gain. Standard respiration rate at referenced 10°C (R ₁₀) was positively correlated with soil moisture, soil temperature, LAI and aboveground biomass. Rainfall patterns in growing season markedly influenced soil moisture and therefore soil moisture controlled seasonal change of ecosystem respiration. Pulse rainfall in the beginning and at the end of growing season induced great ecosystem respiration and consequently a great amount of carbon was lost. Short growing season and relative low temperature restrained alpine grass vegetation development. The results suggested that LAI be usually in a low level and carbon uptake be relatively low. Rainfall patterns in the growing season and pulse rainfall in the beginning and at end of growing season control ecosystem respiration and consequently influence carbon balance of ecosystem.     

Guidelines for seagrass restoration: Importance of habitat selection and donor population, spreading of risks, and ecosystem engineering effects

Large-scale losses of seagrass beds have been reported for decades and lead to numerous restoration programs. From worldwide scientific literature and 20 years of seagrass restoration research in the Wadden Sea, we review and evaluate the traditional guidelines and propose new guidelines for seagrass restoration.Habitat and donor selection are crucial: large differences in survival were found among habitats and among donor populations. The need to preferably transplant in historically confirmed seagrass habitats, and to collect donor material from comparable habitats, were underlined by our results. The importance of sufficient genetic variation of donor material and prevention of genetic isolation by distance was reviewed. The spreading of risks among transplantation sites, which differed in habitat characteristics (or among replicate sites), was positively evaluated. The importance of ecosystem engineering was shown in two ways: seagrass self-facilitation and facilitation by shellfish reefs. Seagrass self-facilitative properties may require a large transplantation scale or additional measures.     

Spatial patterns of sub-tidal seagrasses and their tissue nutrients in the Torres Strait,northern Australia: Implications for management

The distribution and nutritional profiles of sub-tidal seagrasses from the Torres Strait were surveyed and mapped across an area of 31,000 km². Benthic sediment composition, water depth, seagrass species type and nutrients were sampled at 168 points selected in a stratified representative pattern. Eleven species of seagrass were present at 56 (33.3%) of the sample points. Halophila spinulosa, Halophila ovalis, Cymodocea serrulata and Syringodium isoetifolium were the most common species and these were nutrient profiled. Sub-tidal seagrass distribution (and associated seagrass nutrient concentrations) was generally confined to northern-central and south-western regions of the survey area (<longitude 142.60), where mean water depth was relatively shallow (approximately 13 m below MSL) and where sediments were comprised primarily muddy sand to gravelly sand. Seagrass nitrogen and starch content, the most important nutrients for marine herbivores, were significantly correlated with species and with the plant component (above or below ground). For all seagrass species, the above-ground component (shoots and leaves) possessed greater nitrogen concentrations than the below-ground component (roots and rhizomes), which possessed greater starch concentrations. S. isoetifolium had the highest total nitrogen concentrations (1.40±0.05% DW). However, it also had higher fibre concentrations (38.2±0.68% DW) relative to the other four species. H. ovalis possessed the highest starch concentrations (2.76±0.12% DW) and highest digestibility (83.24±0.66% DW) as well as the lowest fibre (27.2±0.66% DW). The high relative abundance (found at 55% of the sites that had seagrass) and nutrient quality characteristics of H. ovalis make it an important source of energy to marine herbivores that forage sub-tidally in the Torres Strait. There were two regions in Torres Strait (north-central and south-western) where sub-tidal seagrass meadows were prevalent and of relatively higher nutritional value. This spatial and nutritional information can be used by local agencies to manage and to protect the ecological, economic and cultural values of the sub-tidal seagrass ecosystems and associated fisheries of the Torres Strait.     

Seagrass tolerance to herbivory under increased ocean temperatures

Climate change is acknowledged as a major threat to marine ecosystems, but the effect of temperature on species interactions remains poorly understood. We quantified the effects of long-term warming on plant-herbivore interactions of a dominant seagrass, Zostera muelleri. Growth, herbivory and tolerance to damage were compared between a meadow warmed by the thermal plume from a power station for 30 years (2–3 °C above background temperatures) and three control locations. Leaf growth rates and tissue loss were spatially variable but unrelated to temperature regimes. Natural herbivory was generally low. Simulated herbivory experiments showed that the tolerance of Z. muelleri to defoliation did not differ between warm and unimpacted meadows, with damaged and undamaged plants maintaining similar growth rates irrespective of temperature. These results suggest that the ability of temperate Z. muelleri to tolerate herbivory is not strongly influenced by warming, and this species may be relatively resilient to future environmental change.     

Site-specific success of three transplanting methods and the effect of planting time on the establishment of Zostera marina transplants

Eelgrass, Zostera marina, is the most abundant seagrass species on the coast of Korea, but most large eelgrass meadows have disappeared since the 1970s due to human-induced disturbance. To restore disturbed seagrass habitats in die-off areas, seagrass transplanting has been attempted using various methods. Here, we assessed the feasibility and efficiency of seagrass transplanting methods depending on the sediment type at the planting site. Additionally, the effect of planting time on the establishment of transplant was examined to determine the optimal season for planting. We conducted an eelgrass transplanting experiment from fall 2003 to summer 2004 at three sites with different sediment types using three planting techniques. The staple method resulted in the highest transplant survival rate (77.1-93.8%) at all three sites, but was labor intensive. Transplanting Eelgrass Remotely with Frame Systems (TERFS) method also resulted in relatively a high survival rate (58.7-69.0%) at all sites. The shell method is a newer eelgrass transplanting method in which oyster shells are used as an anchoring device, and does not require SCUBA diving for subtidal transplanting. The shell method resulted in high survival rates in muddy (81.3%) and silty sediments (76.5%), but remarkably low survival rate in sandy sediments (5.0%). The TERFS, and shell methods reduced underwater labor; thus, these methods is suitable for large-scale seagrass restoration. Eelgrass transplants planted in summer had exhibited significant mortality due to high summer water temperatures. Although transplants planted in fall to spring had relatively high survival rates, transplanting and collection of vegetative shoots are difficult in winter and spring. Therefore, fall was suggested as the most effective transplanting season off the coast of Korea.     

Signal crayfish burrowing,bank retreat and sediment supply to rivers: A biophysical sediment budget

Burrowing into riverbanks by animals transfers sediment directly into river channels and has been hypothesised to accelerate bank erosion and promote mass failure. A field monitoring study on two UK rivers invaded by signal crayfish (Pacifastacus leniusculus) assessed the impact of burrowing on bank erosion processes. Erosion pins were installed in 17 riverbanks across a gradient of crayfish burrow densities and monitored for 22 months. Bank retreat increased significantly with crayfish burrow density. At the bank scale (<6 m river length), high crayfish burrow densities were associated with accelerated bank retreat of up to 253% and more than a doubling of the area of bank collapse compared with banks without burrows. Direct sediment supply by burrowing activity contributed 0.2% and 0.6% of total sediment at the reach (1.1 km) and local bank (<6 m) scales. However, accelerated bank retreat caused by burrows contributed 12.2% and 29.8% of the total sediment supply at the reach and bank scales. Together, burrowing and the associated acceleration of retreat and collapse supplied an additional 25.4 t km⁻¹ a⁻¹ of floodplain sediments at one site, demonstrating the substantial impact that signal crayfish can have on fine sediment supply. For the first time, an empirical relation linking animal burrow characteristics to riverbank retreat is presented. The study adds to a small number of sediment budget studies that compare sediment fluxes driven by biotic and abiotic energy but is unique in isolating and measuring the substantial interactive effect of the acceleration of abiotic bank erosion facilitated by biotic activity. Biotic energy expended through burrowing represents an energy surcharge to the river system that can augment sediment erosion by geophysical mechanisms.