Patterns of spatial variation of nutrient content,epiphyte load and shoot size of Posidonia oceanica seagrass meadows (Mediterranean Sea)

Knowledge of patterns of spatial variability of vegetative development, epiphyte load and nutrient availability in seagrass meadows is essential for the adequate design of research and environmental monitoring programmes. Differences in shoot size, epiphyte load and nutrient content of leaves and epiphytes of the Mediterranean endemic seagrass Posidonia oceanica at spatial scales ranging from metres to hundreds of metres are evaluated using a hierarchical nested sampling design. The size and epiphyte load of P. oceanica shoots and the nitrogen and phosphorus content of leaves and epiphytes were different in most of the spatial scales considered. Sampling efforts concentrated at the metre scale incorporated most of the variability in size, epiphyte load and nutrient content of the leaves and epiphytes of P. oceanica shoots. Epiphyte load showed no correlation with nutrient content in the epiphytes or in the leaves. However, epiphyte load and shoot size were negatively correlated, which suggests that light penetration in the canopy may be a main determinant of epiphyte load.     

Harpacticoid copepod response to epiphyte load variations in Posidonia oceanica (L.) Delile meadows

We conducted a field experiment to assess the response of phytal harpacticoids to nutrient‐driven increases of epiphyte load in Posidonia oceanica meadows. First, we evaluated differences in species richness, diversity and assemblage structure of phytal harpacticoids in P. oceanica meadows with differing epiphyte loads. Secondly, we conducted a field experiment where epiphyte load was increased through an in situ addition of nutrients to the water column and evaluated the responses of the harpacticoid assemblages. We predicted that there would be changes in the harpacticoid assemblages as a result of nutrient‐driven increases of epiphyte load, and that these changes would be of a larger magnitude in meadows of low epiphyte load. Our results show that the harpacticoid fauna (>500 μm) present in P. oceanica meadows in the Bay of Palma comprised taxa which are considered phytal and other less abundant ones previously described as sediment dwellers or commensal on other invertebrate species. Nutrient addition had an overall significant effect on epiphyte biomass and on harpacticoid abundance, diversity and assemblage structure, possibly as a response to the increased resources and habitat complexity provided by epiphytes. The abundance of dominant species at each location was favoured by nutrient addition and in some cases correlated with epiphytic biomass, although never strongly. This may indicate that structural complexity or diversity of the epiphytic cover might be more important than the actual epiphytic biomass for the harpacticoid species investigated. More species‐specific studies are necessary to ascertain this and clarify the relationships between harpacticoids and epiphytes in seagrass meadows. To our knowledge, this is the first account of harpacticoid species associated with P. oceanica leaves and the epiphytic community they harbour in the Mediterranean Sea.     

Temporal variations in the spatial distribution of shoot density in a Posidonia oceanica meadow and patterns of genetic diversity

The meadows of the Mediterranean endemic seagrass Posidonia oceanica exhibit relatively high variations of structural and biometric features at various spatial scales. An investigation performed in 1992 in the meadow off Lacco Ameno (Island of Ischia, Gulf of Naples, Italy) detected peculiar spatial patterns of plant distribution, characterized by nestlike structures with radially increasing or decreasing shoot densities. Eight years later (2000), geo‐referenced collections at selected points were repeated to trace the temporal variations of shoot density and investigate the recurrence of the density cores previously detected. In addition, shoots for molecular analyses were collected to check the hypothesis that nestlike patterns exhibit highest levels of genetic variability, due to the confluence of several genetically distinct stolons. The 2000 survey confirmed the presence of the main density cores detected in 1992, although their spatial distribution was slightly shifted and a general decrease of spatial anisotropy was observed, probably due to an increased disturbance, mainly due to pleasure boat anchoring. Patterns of genetic diversity showed a more complex picture, well related to the shoot density spatial pattern, especially when compared with the previous 1992 survey. Patterns of genetic diversity confirmed our previous hypotheses on the genesis of shoot density cores, suggesting they are produced over long time, due to a slow stolonization process and a convergence of different genotypes. Regression of the meadow and decrease of density may lead, in short periods, to a homogenization of the density patterns, while genetic diversity cores represent a long‐term ‘memory’ of their previous distribution.     

Influence of wave climate on architecture and landscape characteristics of Posidonia oceanica meadows

Seagrass meadow characteristics, including distribution, shape, size and within‐meadow architectural features, may be influenced by various physical factors, including hydrodynamic forces. However, such influences have hardly been assessed for meadows of the ecologically important and endemic Mediterranean seagrass Posidonia oceanica. The distribution of P. oceanica meadows at five sites in the Maltese Islands was mapped to a depth of c. 15 m using a combination of aerial photography and SCUBA diving surveys. Estimates of wind‐generated wave energy and energy attenuated by depth were computed using the hydrodynamic model WEMo (Wave Exposure Model). Metrics for P. oceanica landscape features were calculated using FRAGSTATS for replicate 2500 m² subsamples taken from the seagrass habitat maps in order to explore the influence of wave dynamics at the landscape scale. Data on within‐meadow architectural attributes were collected from five sites and analysed for relationships with wave energy. The results indicate that landscape and architectural features of P. oceanica meadows located within the 6–11 m depth range are significantly influenced by wave climate. Posidonia oceanica meadows tend to be patchier and have low overall cover, more complex patch shapes and reduced within‐patch architectural complexity along a wave exposure gradient from low to high energy. The findings from the present study provide new insight into the influence of hydrodynamic factors on the natural dynamism of P. oceanica meadow landscape and architecture, which has implications for the conservation and management of the habitat.     

Experimental assessment of Posidonia oceanica‐associated gastropods grazing on an early‐successional biofilm community: nutrient availability and species‐specific effects

Although grazing is considered an essential process controlling epiphyte biomass on seagrass leaves, there is still a lack of fundamental knowledge about the species‐specific consumption rates of the most common grazers in Mediterranean meadows. This study experimentally assessed the effect of Posidonia oceanica‐associated gastropod grazing on early successional biofilm and the species‐specific relationship between biofilm consumption rates and biofilm biomass. Two biofilms on artificial substrata, both developed in situ (in a P. oceanica meadow), one under ambient conditions and the other under nutrient‐enriched conditions, were offered in aquaria assays to nine species of grazers found in P. oceanica meadows. Biofilm consumption rates and their association with biofilm biomass were assessed. It was found that: (i) there was a positive association between biofilm consumption and biofilm biomass up to 20 mg Chl a·m^?2 for Bittium reticulatum, Gibbula ardens, Jujubinus exasperatus and Tricolia pullus; (ii) Alvania montagui, B. reticulatum and Jujubinus striatus showed the highest consumption rates and are thus expected to be amongst the leading consumers in early‐successional epiphytic communities; (iii) there was not an increase of consumption rate when a substratum colonized under nutrient‐enriched conditions was offered to any of the nine studied species. This study provides species‐specific consumption rates knowledge that is useful for the assessment of the strength of grazer–epiphyte interactions and trophic fluxes in P. oceanica meadows.     

Effects of fish farming on seagrass (Posidonia oceanica) in a Mediterranean bay: seagrass decline after organic loading cessation

The effects of fish farming on a seagrass (Posidonia oceanica) meadow at Fornells Bay, Minorca (Balearic Islands) were studied. Changes in plant and meadow features (e.g. shoot morphology, shoot density, biomass, rhizome growth, nutrient and soluble sugars concentrations…) in three stations along a transect from a disturbed (organic pollution due to fish cultures) to an undisturbed site were assessed. The fish culture had ceased in 1991; however, seagrass decline, already reported in a previous study for the period 1988–1990, was still taking place at the time of sampling (July 1994). Differences between stations were very clear; the station closest to the fish cages showed reduced shoot density, shoot size, underground biomass, sucrose concentration and photosynthetic capacities. In contrast, shoots from the most polluted station showed higher P-concentration in tissues and higher epiphyte biomass than the other two. Since water conditions had recovered completely by the time of the sampling mission, it is proposed that the persistence of the seagrass decline was due to the excess organic matter remaining in the sediment.     

Impact of the brine from a desalination plant on a shallow seagrass (Posidonia oceanica) meadow

Although seawater desalination has increased significantly over recent decades, little attention has been paid to the impact of the main by-product (hypersaline water: brine) on ecosystems. In the Mediterranean, potentially the most affected ecosystems are meadows of the endemic seagrass Posidonia oceanica. We studied the effect of brine on a shallow P. oceanica meadow exposed to reverse osmosis brine discharge for more than 6 years. P. oceanica proved to be very sensitive to both eutrophication and high salinities derived from the brine discharge. Affected plants showed high epiphyte load and nitrogen content in the leaves, high frequencies of necrosis marks, low total non-structural carbohydrates and low glutamine synthetase activity, compared to control plants. However, there was no indication of extensive decline of the affected meadow. This is probably due to its very shallow situation, which results in high incident radiation as well as fast dilution and dispersion of the brine plume.     

Human influence on seagrass habitat fragmentation in NW Mediterranean Sea

Habitat fragmentation in meadows of Posidonia oceanica, the most important and abundant seagrass in the Mediterranean Sea, was investigated at a region-wide spatial scale using a synthetic ecological index, the Patchiness Index (PI). We tested the hypothesis that human impacts are the major factor responsible for habitat fragmentation in P. oceanica meadows contrasting fragmentation of meadows located in “anthropized” areas with that of meadows located in areas with low anthropization and considered as virtually “natural”. We also related fragmentation of meadow with the morphodynamic state of the submerged beach (i.e. distinctive types of beach produced by the topography, the wave climate and the sediment composition) in order to investigate the influence of one natural component on the seagrass meadow seascape. Results demonstrated that fragmentation in the P. oceanica meadows is strongly influenced by the human component, being lower in natural meadows than in anthropized ones, and that it is little influenced by the morphodynamic state of the coast. The use of landscape approaches to discriminate natural disturbance from human impacts that affect seagrass meadows is thus recommended for the proper management of coastal zones.     

Architectural characteristics of two bed types of the seagrass Posidonia oceanica over different spatial scales

Seagrass beds occur in various morphological forms, ranging from small patches to continuous meadows. The endemic Mediterranean seagrass Posidonia oceanica forms dense and extensive stands that occur in several different morphotypes, including reticulate (seagrass interspersed with a different habitat type, such as bare sand) and continuous beds. This study, undertaken in the Maltese Islands, examined whether reticulate and continuous P. oceanica beds, located adjacent to each other and at similar depths, had different within-bed architectural characteristics. Five commonly used architectural measures (shoot density, number of leaves per shoot, mean leaf length, mean leaf width and shoot biomass) were measured from P. oceanica shoots collected from the two bed types at three different spatial scales: (1) tens of metres (‘small’ scale); (2) hundreds of metres (‘medium’ scale); and (3) kilometres (‘large’ scale). Results of 2-factor ANOVA (factor 1=bed type; factor 2=sampling locality) carried out at the three spatial scales indicated significant differences between the two bed types in shoot density (P<0.01) and leaf length (P<0.05) at the small scale, and in leaf number (P<0.05) at the large scale. Significant interactions were also apparent for shoot density (at the large scale) and for shoot biomass (at the medium scale). However, the results obtained did not indicate consistent architectural differences between the two P. oceanica bed types over the spatial scales considered. Spatial variations in within-bed architectural characteristics observed were therefore thought to be attributable mainly to the influence of local environmental factors. The findings are discussed with reference to the conservation and management of P. oceanica habitat.     

Population dynamics and production of the seagrass Zostera noltii in colonizing versus established meadows

The dynamics of the seagrass Zostera noltii in established and colonizing meadows were assessed in Ria Formosa lagoon, Southern Portugal. Shoot weight, above:belowground biomass ratio, flowering shoot density, meadow production, and biomass–density relationships were investigated. Results indicate that the species population dynamics differ clearly in different development stages of the meadows. The overall mean of flowering shoot density was five times higher in the colonizing (83 flowering shoots m⁻²) than in the established meadow (16 flowering shoots m⁻²), revealing a greater contribution of sexual reproduction during the species colonization process. The temporal variation of the biomass–density relationship in the colonizing meadow showed a cyclic seasonal trajectory, a wider range of data, and a simultaneous peak of biomass and density, suggesting no space limitations constraining the internal packing of shoots during the growing season. In the established meadow, density peaked before biomass in agreement to the dominant role of the clonal architecture of seagrasses in the configuration of closed meadows, suggesting the occurrence of self-thinning and/or regulation of ramet formation. Slope of the biomass–density relationships was similar in the established and colonizing meadows, generally suggesting similar nutritional conditions, regardless of their muddy and sandy sediments. Plants of the colonizing meadow invest mainly on the belowground fraction (above:belowground biomass ratio <1), as meadow expansion is mainly controlled by the elongation of horizontal rhizomes. The annual total production (1163 g C m⁻² year⁻¹) and the biomass turnover (34.8 year⁻¹) were also higher in this meadow, corroborating the high investment of the species during the meadow expansion. The faster biomass turnover of the colonizing meadow implies a more limited capacity to accumulate biomass, indicating a greater exportation of organic carbon and nutrients to the coastal area. The different biomass turnover rates suggest different trophic and structural roles of Z. noltii communities in established and colonizing meadows.     

Crustacean decapod assemblages associated with fragmented Posidonia oceanica meadows in the Alboran Sea (Western Mediterranean Sea): composition,temporal dynamics and influence of meadow structure

The decapod assemblage associated with a Posidonia oceanica meadow located near its western limit of biogeographic distribution was studied over an annual cycle. Fauna samples were taken seasonally over a year (five replicates per season) in two sites located 7 km apart, using a non‐destructive sampling method (airlift sampler) for the seagrass. The dominant species of the assemblage, Pisidia longimana, Pilumnus hirtellus and Athanas nitescens, were associated with the protective rhizome stratum, which is mainly used as a nursery. The correlations between decapod assemblage structure and some phenological parameters of the seagrass shoots and wave height were negative or null, which reflects that species associated with the rhizome had a higher importance than those associated with the leaf stratum. The abundance and composition of the decapod assemblage as well as the ecological indexes displayed a seasonality trend with maximum values in summer‐autumn and minimum in winter‐spring, which were related to the seawater temperature and the recruitment periods of the dominant species. The spatial differences found in the structure and dynamics of the assemblages may be due to variations in the recruitment of the dominant species, probably as a result of the influence of local factors (e.g. temperature, currents) and the high dispersal ability of decapods, together with the patchy configuration and the surrounding habitats. The studied meadows are fragmented and are integrated within a mosaic of habitats (Cymodocea nodosa patches, algal meadows, rocky and sandy bottoms), which promotes the movement of individuals and species among them, maintaining a high species richness and evenness.     

The influence of coastal dynamics on the upper limit of the Posidonia oceanica meadow

The recognized ecological importance of Posidonia oceanica, the most important seagrass of the Mediterranean Sea, makes it crucial to assess the state of health of its meadows, discriminating natural from anthropogenic impacts. In this paper, the hydrodynamic conditions at the upper limit of P. oceanica meadows along the Ligurian coast (NW Mediterranean Sea) were investigated. A relationship between the distance of the upper limit of the meadow from the shoreline and the morphodynamic domain of the beach (i.e. distinctive types of beach produced by the topography, wave climate and sediment composition) was found. A zonation of the state of the shallow portions of the meadows down the submerged beach profile was identified. Zone a, from the shoreline to the breaking limit, is naturally critical for the development of the meadow. Zone b, from the breaking limit to the closure depth, is subjected to natural and human impacts. Zone c, below the closure depth, is little influenced by coastal dynamics. This study quantifies for the first time how much the status of the shallow portions of P. oceanica meadows is dependent on coastal dynamics, which is important for their proper management.     

Ontogenic variation and effect of collection procedure on leaf biomechanical properties of Mediterranean seagrass Posidonia oceanica (L.) Delile

Leaf mechanical traits are important to understand how aquatic plants fracture and deform when subjected to abiotic (currents or waves) or biotic (herbivory attack) mechanical forces. The likely occurrence of variation during leaf ontogeny in these traits may thus have implications for hydrodynamic performance and vulnerability to herbivory damage, and may be associated with changes in morphologic and chemical traits. Seagrasses, marine flowering plants, consist of shoot bundles holding several leaves with different developmental stages, in which outer older leaves protect inner younger leaves. In this study we examined the long‐lived seagrass Posidonia oceanica to determine ontogenic variation in mechanical traits across leaf position within a shoot, representing different developmental stages. Moreover, we investigated whether or not the collection procedure (classical uprooted shoot versus non‐destructive shoot method: cutting the shoot without a portion of rhizome) and time span after collection influence mechanical measurements. Neither collection procedure nor time elapsed within 48 h of collection affected measurements of leaf biomechanical traits when seagrass shoots were kept moist in dark cool conditions. Ontogenic variation in mechanical traits in P. oceanica leaves over intermediate and adult developmental stages was observed: leaves weakened and lost stiffness with aging, while mid‐aged leaves (the longest and thickest ones) were able to withstand higher breaking forces. In addition, younger leaves had higher nitrogen content and lower fiber content than older leaves. The observed patterns may explain fine‐scale within‐shoot ecological processes of leaves at different developmental stages, such as leaf shedding and herbivory consumption in P. oceanica.     

Effects of organic matter input from a fish farming facility on a Posidonia oceanica meadow

In the Mediterranean, the development of aquaculture along the coasts appears as a source of disturbance to the littoral ecosystems, and in particular to Posidonia oceanica seagrass meadows. Although the impact of fish farms in Northern Europe has been studied over the last few years, the data are more scarce in the Mediterranean. Thus, a number of physico-chemical and biological parameters have been examined here in order to evaluate the impact of a fish farm in a littoral bay of Corsica. The following values that were recorded in the vicinity of the fish farm are much higher than those at the reference station: organic content of the sediment (24–21 versus 2%), nitrogen concentrations (ammonium: 19.5–8.4 versus 1.8 μM) and phosphorous levels in the pore water (orthophosphates: 5.2–1.3 versus 1.7 μM). The seagrass meadow vitality also seems to be affected in the vicinity of cages, with densities that drop from 466 (reference station) to 108 shoots m⁻² (20 m from cages). Total primary production also varies from 1070.6 to 87.9 g m⁻² year⁻¹. The main impact factors seem to be the input of organic matter originating from the cages and the high epiphyte biomass caused by the nutrient enrichment. The high level of organic matter and the presence of mud seem to alter the physico-chemical characteristics of the bottom sediment; moreover, the plant/epiphyte competition seems to lead to a leaf fragility and, more importantly, to a decrease in available light.     

Impact of fish farming facilities on Posidonia oceanica meadows: a review

The impact of fish farming facilities on Posidonia oceanica meadows was assessed from studies of intensive facilities carried out over the last few years. The disturbances caused by these fish farms were measured by means of both abiotic (light, sediment, interstitial water) and biotic variables (meadow density, leaf biometry, lepidochronology, primary production, epiphytes, reserve carbohydrates in the rhizomes), in function of increasing distance from cages and/or inside a geographically close reference site. The results showed significant degradation of these seagrass meadows in all the sectors investigated. When fish farming cages were placed above a P. oceanica bed, the meadow was severely degraded or disappeared and the sediment showed a strong increase in organic matter that could lead to anoxia phenomena. The irreversible impact of fish farming projects on P. oceanica meadows requires the application of the precautionary principle. Several recommendations (site selection, preliminary studies and monitoring over time) are suggested in order to enable piscicultural activities to be incorporated in a global process of Integrated Coastal Zone Management.     

Comparison of the fish assemblages associated with Posidonia oceanica after the partial loss and consequent fragmentation of the meadow

An extensive Posidonia oceanica meadow was partially destroyed by excavation, resulting in areas of seagrass habitat of equal complexity (shoot density) but different heterogeneity (degree of fragmentation). The fish assemblages associated with differently fragmented beds were compared from a landscape perspective. Differences in the fish assemblages were detected, with several species showing different patterns: (1) species that increased their abundance along with the degree of bed fragmentation, (2) species that were more abundant in fragmented beds, but did not show differences between more or less fragmented beds, and (3) species that were mostly abundant in large seagrass patches or in the continuous meadow. The fish assemblages were also affected by depth, but further research is needed to determine properly this effect. Some ecological mechanisms are suggested to operate in the interaction between P. oceanica and the held fish assemblage.     

Comparison between epiphyte assemblages of leaves and rhizomes of the seagrass Posidonia oceanica subjected to different levels of anthropogenic eutrophication

This paper aimed to compare epiphyte assemblages of leaves and rhizomes of Posidonia oceanica exposed to different levels of concentration of nutrients. The same design including a potentially impacted meadow and two reference meadows was used in each of two locations, characterized by the presence of a city or of suspended cages of a fish farm, respectively. This allowed to test for the consistency of responses of epiphytic assemblages to different sources of eutrophication. In both studies, results documented differences in patterns of composition and abundance of epiphytic assemblages on leaves between disturbed and reference meadows, while assemblages on rhizomes did not appear sensitive to this kind of disturbance. Moreover, in potentially impacted meadows, both assemblages showed different patterns of spatial variability compared to reference assemblages. Species composition and abundance of epiphyte assemblages seemed suitable for detecting moderate nutrient increases, even if adequate sampling designs are needed to separate patterns related to the large natural spatial variability of these systems from those related to changes in environmental conditions.     

荣成马山里海域海草床分布现状及其生态特征

海草床是滨海三大典型生态系统之一,具有极其重要的环境改善、资源养护和减灾防灾等生态功能,亦是全球重要的碳库。2016年8月,通过对荣成马山里海域的现场调查,发现了面积为58.26 hm2的海草床,其海草的种类为红纤维虾形草（Phyllospadix iwatensis）和丛生鳗草（Zostera caespitosa）。红纤维虾形草分布面积为54.50 hm2,占该海域海草床总面积的93.5%,分为北部和南部2个带状区域,平均茎枝密度为（368.0±18.2）shoots/m2,平均生物量（干重,下同）为（297.0±41.5）g/m2。丛生鳗草分布面积为3.76 hm2,占海草床总面积的6.5%,呈现斑块状分布,与红纤维虾形草交错而生,平均茎枝密度为（691.2±17.1）shoots/m2,平均生物量为（534.0±70.7）g/m2。马山里海域海草床主要分布在平均水深为（2.8±0.3） m的以岩礁为主的底质上。海草的生长状况存在显著的空间差异,与水温呈现显著的正相关,与水深和陆源污染物存在显著的负相关。结合历史资料,发现该海域海草床退化现象较严重,其主要威胁因素是人为干扰,主要包括渔业生产、养殖活动和陆源污染输入。建议合理规划周边海域的养殖规模和密度、加强陆源污染物管控和开展海草床生态修复工程,以期为温带海草床的有效保护和科学管理提供参考。     

Epiphyte assemblages of the Mediterranean seagrass Posidonia oceanica: an overview

This paper reviews the main studies dealing with epiphytes of both leaves and rhizomes of Posidonia oceanica, the main seagrass found in the Mediterranean Sea. A total of 660 epiphyte species has been recorded, including 430 on leaves and 437 on rhizomes. Of these, 205 are Rhodophyta, 59 Ochrophyta, 43 Chlorophyta, 78 Porifera, 82 Cnidaria, 19 Annelida, 155 Bryozoa and 19 Tunicata. The epiphyte assemblages demonstrate a gradient of dissimilarity from west to east, with the eastern area being the most unlike the others. The differences can be attributed mostly to rare species that differ amongst the Mediterranean regions. Many of the dominant taxa have been found in all meadows studied. Data concerning species composition, abundance, and patterns of spatial and temporal variability are summarized, and biotic and abiotic factors controlling the structure of assemblages are discussed. Responses of epiphyte assemblages to anthropogenic stressors, such as nutrient enrichment and biological invasions, are discussed in relation to the use of epiphytes as ecological indicators.     

The relationship between seagrass (Posidonia oceanica) decline and sulfide porewater concentration in carbonate sediments

In this study we test the hypothesized negative relationship between seagrass status and porewater hydrogen sulfide (H₂S) levels, through a comparative analysis within a range of seven Posidonia oceanica meadows growing over carbonate sediments in the NW Mediterranean Sea around Mallorca Island. The studied meadows range from meadows growing on sediments with very low sulfide porewater concentrations (4.6 μM) to those growing over higher sulfide conditions (33.5 μM). Organic matter content, sulfate reduction rates and sulfide porewater concentrations in the sediments were determined concurrently with the assessment of demographic plant dynamics (specific mortality and net population growth rates). Sulfide porewater concentration increased with increasing organic matter content in the sediment, while net population growth decreased significantly with low increases of sulfide concentrations. Our results confirm the previously suspected vulnerability of seagrass meadows growing on carbonate sediments to increased sulfide levels. An excess of 10 μmols H₂S L⁻¹ porewater is identified to already conduce P. oceanica meadows to decline, which this study identifies, particularly, as strongly sensitive to sulfides. The results reported here suggest that even moderate increases in organic carbon inputs may lead to enhancement of dissolved sulfides and may be an important factor for seagrass status in these iron-depleted carbonate sediments from the Mediterranean Sea.