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.     

Temporal and bathymetric variation of epiphyte cover and leaf biomass in a southern Posidonia oceanica (L.) meadow: the case of Mahdia coast,Tunisia

The aim of this research was to study spatial and temporal variation in epiphyte cover and leaf biomass of Posidonia oceanica in Eastern Tunisia. Sampling was conducted at four stations on the Mahdia coast during October and December 2010, and April and August 2011, which correspond respectively to autumn, winter, spring and summer in this area. Posidonia oceanica shoots were collected at two depths (5 and 10 m). Cover of macroinvertebrates and macroalgae was estimated on adult leaves. The results showed that leaf and epiphyte biomasses vary significantly according to sampling date, with the highest values recorded in August. We found a high diversity of epiphytic assemblages on the leaves of P. oceanica with clear qualitative and quantitative dominance of Rhodophyceae compared to other groups, followed by Phaeophyceae. Most epiphyte species on the leaves of P. oceanica in Eastern Tunisia are the same as those in other parts of the Mediterranean Sea. No bathymetrical variation in the epiphytic community was found in our study area, which can be explained by the high levels of water clarity in Mahdia.     

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.     

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.     

Composition of epiphytic leaf community of Posidonia oceanica as a tool for environmental biomonitoring

The demand for sensitive biological tools to assess the environmental quality of coastal waters at broad spatial scales is increasing. Many of the tools used are based on the taxonomic composition of biotic assemblages. They usually require a valuable taxonomic expertise while are unique reflecting the overall ecosystem integrity. Here, we evaluate the potential indicator value of several features of the epiphytic community (overall assemblage composition, species richness, and proportion of the main taxonomic groups) growing on the seagrass Posidonia oceanica leaves. We do so by empirically examining their changes along a disturbance gradient where multiple human activities have interactive and cumulative impacts, sampling at different spatial scales and at two different depths (5 and 15 m). Our results show that the specific composition of the epiphytic assemblages (i.e. species composition) closely reflects, in the deep meadows, the combined effects of different anthropogenic stressors along the gradient, showing an integrative and non-specific response. Similarly, an increase in the proportion of hydrozoans, and a decrease in the proportion of rhodophytes and chlorophytes are observed in deep meadows along the gradient. In shallow meadows, grazing and biotic features of the seagrass seem the main forcing factors determining species composition, and therefore masking the response of epiphytes to the deterioration gradient. After address the effect of natural sources of variability (water depth, within- and between-meadow heterogeneity), changes in epiphyte assemblages and in the proportion of hydrozoans, rhodophytes and chlorophytes in relatively deep meadows seem promising monitoring tools for detecting coastal environmental deterioration.     

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.     

Spatial pattern of coastal fish assemblages in different habitats in North‐western Mediterranean

Variability of fish assemblages across habitat structures can depend on spatial scales. A hierarchical sampling design was used to assess the spatial variability of temperate fish assemblages in different habitats and at multiple scales. Underwater visual censuses were carried out along the coasts of Elba Island (NW Mediterranean) on Posidonia oceanica beds, rocky algal reefs and sandy habitat at three spatial scales, namely tens of metres (individual replicates), hundreds of metres (sites) and tens of kilometres (locations). At the assemblage level, there was a clear relationship between fish and habitat type and the observed habitat‐related differences were largely dependent on species identity. Fish assemblages on P. oceanica beds and rocky reefs shared a high number of species, whereas overlap with sandy assemblages was negligible. Multivariate analyses revealed significant differences in fish assemblages among habitats, although there was also a significant habitat × site interaction. These differences relied mainly upon assemblage composition and species richness. Assemblages on rocky reefs and P. oceanica meadows usually harboured a higher number of species and individuals compared with sandy assemblages. Nevertheless, the patterns of habitat‐related differences in species richness and, especially, in the total number of fish, changed significantly from site to site. Eight species showed significant differences over habitats, but they were not consistent due to the interaction of habitat with site. Predictability of fish at both assemblage and population levels decreased with the scale of observation, and the spatial pattern of fish observed at the smallest scale was likely dependent on factors other than habitat type.     

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.     

Spatial variability of Posidonia oceanica (L.) Delile epiphytes around the mainland and the islands of Sicily (Mediterranean Sea)

This paper investigates patterns of variability in epiphytes of Posidonia oceanica leaves at various spatial scales around Sicily, including geographical differences among the Mediterranean basins, differences between the small islands and mainland, and the variability among and within replicated meadows in each of the previous conditions. Data on percentage cover of the most common epiphytic organisms were analysed by univariate techniques. Encrusting red algae, encrusting brown algae, filamentous algae, encrusting bryozoans, erect bryozoans, hydroids and Foraminifera were the most abundant taxa. Significant differences in the abundance of taxa were detected among geographical regions, while no significant differences were found between the islands and mainland. At smaller scales, variability was concentrated mostly among leaves 100s of centimetres apart and among meadows a few kilometres apart. These results suggest that both geographical and local processes are important in structuring the epiphytes of P. oceanica leaves in this area of the Mediterranean.     

Diversity and temporal fluctuations of epiphytes and sessile invertebrates on the rhizomes Posidonia oceanica in a seagrass meadow off Tunisia

The aim of this work is to study the temporal dynamics of rhizome epiphytes and sessile animals living on the rhizomes of the seagrass Posidonia oceanica in the east of Tunisia. Surveys were conducted in October 2009, and in January, April and August 2010 on a fringing reef located in Chebba. Rhizomes were sampled by SCUBA diving at three stations. Samples were examined with a microscope to estimate the cover of macroinvertebrate and macroalgal organisms on the top 10 cm of each rhizome. Results revealed a high diversity of epiphytes on P. oceanica rhizomes with a dominance of red and brown algae, ascidians, and bryozoans. Distinct temporal changes were observed in Oued Lafrann, with a high January cover (winter period) for all groups. These winter increases can be attributed to: (i) the low phenological parameters of P. oceanica in winter that reduce the effects of shading, (ii) life cycles of the epiphytes and invertebrates, (iii) water motion and (iv) grazing.     

Spatial scale variability in shoot density and epiphytic leaves of Posidonia oceanica on Kerkennah Island (Tunisia) in relation to current tide effects

A study was undertaken of the patterns of spatial variability, epiphytic biomass and distribution of epiphytic fauna and flora of Posidonia oceanica. Samples were taken at four stations located approximately 4 km apart, exposed to different current conditions. Stations A and B, situated near the Oued Mimoun tidal channel with its relatively strong bi‐directional flows, were affected by high current tide. The other two stations, North Oued Mimoun (L1) and South Oued Mimoun (L2), were located further from the channel, in low current tide conditions. Sampling was conducted in the Attaya area of Kerkennah Island (Tunisia) in August 2009 at depths between 2 and 3 m, with the results indicating differences among the stations. Shoot density decreased when exposed to high levels of hydrodynamic activity generated by current tides whereas the epiphytic biomass of P. oceanica leaves decreased at sheltered stations located far from the channel. Epiphytic algae such as Heterokontophyta, Rhodophyta and Chlorophyta, and epiphytic fauna represented by Bryozoa, Hydrozoa, Annelida, Porifera and Tunicata, dominated the epiphytic assemblages and were abundant at the station most exposed to high current tide hydrodynamics. Cyanobacteria, however, were dominant in stations exposed to low current tide.     

Long-term effect of beach replenishment on natural recovery of shallow Posidonia oceanica meadows

The recovery capacity of shallow Posidonia oceanica meadows degraded by beach replenishment eighteen years before was assessed in two impacted meadows and compared with other two undisturbed localities. Inside each locality, we selected randomly three sites separated by 500–1000 m. At site level we study the vitality of P. oceanica meadow assessing the vegetative growth, leaf characteristics, and non-structural carbohydrates of the plants. Additionally, at locality level, silt-clay fraction, organic matter, pH and light intensity incident on the sea bottom were measured to evaluate the environmental conditions. Covering of P. oceanica was significantly lower at the impacted localities while amount of dead “matte” was higher. Leaf production of horizontal rhizomes (14.6 ± 1.11 vs 19.47 ± 1.45 leaves y⁻¹), net total rhizomes recruitment (2.33 ± 0.17 vs 4.3 ± 0.33 branches y⁻¹) and starch concentration (43.625 ± 0.67 vs 54.45 ± 0.74 mg per g of rhizome) at impacted meadows were significantly lower than controls. Leaf features, epiphytes biomass, colonization, elongation and horizontal and vertical rhizome production did not show significant differences. Sediments at impacted localities contained higher silt-clay fraction and higher organic matter load while pH was lower. Light intensity on the sea bottom measured at all localities was over the minimum light requirements estimated for P. oceanica. Our results show that the press impact produced by beach replenishment was enduring in the time slowing natural recovery by 45%. This impact may be related with changes in the sediment features.     

Photosynthetic performance,epiphyte biomass and nutrient content of two seagrass species in two areas with different level of nutrients along the Dar es Salaam coast

Heavy nutrient loads in coastal waters often lead to excessive growth of microalgal and macroalgal epiphytes on seagrass leaves, with varying effects on the underlying seagrasses. This study evaluates the photosynthetic performance, epiphytic biomass and tissue nutrient content of two tropical seagrasses, Cymodocea serrulata and Thalassia hemprichii, in two intertidal areas along the Dar es Salaam coast in the Indian Ocean, a nutrient-rich region at Ocean Road (near the city centre), and a nutrient-poor region at Mjimwema (south of the city centre). Epiphyte biomass was significantly higher at the nutrient-rich site, and epiphytes were associated with reduced photosynthetic performance in both seagrass species at both sites. Likewise, nitrogen and phosphorus tissue content was higher in both species at the nutrient-rich site than at the nutrient-poor site. Epiphytic species composition on the seagrass leaves varied between seagrass species and between sites. Cymodocea serrulata had a higher number of epiphytic species at Mjimwema than at Ocean Road, whereas Thalassia hemprichii had more epiphytic species at Ocean Road than at Mjimwema. Seagrass photosynthetic performance, epiphytic biomass and nutrient content of the seagrasses were shown to be affected by nutrient concentration in the water column. Thus, for the future monitoring of the seagrass meadow, we recommend the use of combined measures such as seagrass performance, epiphytic biomass, nutrient contents and nutrient concentration levels in the water column.     

Size‐frequency distributions of Joania cordata and Argyrotheca cuneata (Brachiopoda: Megathyrididae) from the Central Tyrrhenian Sea

Size‐frequency distributions can support reliable inferences concerning population dynamics of brachiopods, but only a few data are available so far. In this study, length and width frequency distributions of dead specimens of the Recent brachiopods Joania cordata and Argyrotheca cuneata from the Marine Protected Area ‘Secche di Tor Paterno’, Central Tyrrhenian Sea, Italy (41°35′ N, 12°20′ E), are reported in order to add new data about size‐frequency distributions of brachiopods. The studied specimens came from death assemblages in the coralligenous substrate, in the Posidonia oceanica meadows, and in the sand channels. The observed patterns vary from left‐skewed (J. cordata) to right‐skewed (A. cuneata), indicating respectively a low and high mortality of smaller individuals. Significant differences between the coralligenous substrate and the P. oceanica meadow were observed for both species, revealing a variation among different habitats. All length and width distributions are clearly polymodal, but the biological meaning of the peaks is difficult to interpret, as the two species seem to have a 2‐year life span. A biometric analysis of shell sizes revealed that length and width are the most variable parameters during the growth of the animal.     

Functional changes due to invasive species: Food web shifts at shallow Posidonia oceanica seagrass beds colonized by the alien macroalga Caulerpa racemosa

Multiple stable isotope analyses were used to examine the trophic shifts at faunal assemblages within the invading macroalga Caulerpa racemosa in comparison to established communities of Posidonia oceanica seagrass meadows. Sampling of macrobenthic invertebrates and their potential food sources of algal mats and seagrass meadows in Mallorca (NW Mediterranean) showed differences in species composition of faunal and primary producers among seagrass and C. racemosa. Accordingly, changes in food web structure and trophic guilds were observed, not only at species level but also at community level. The carbon and nitrogen isotope signatures of herbivores, detritivores and deposit feeders confirmed that the seagrass provided a small contribution to the macrofaunal organisms. δ¹³C at the P. oceanica seagrass and at the C. racemosa assemblages differed, ranging from −6.19 to −21.20‰ and −2.67 to −31.41‰, respectively. δ¹⁵N at the Caulerpa mats was lower (ranging from 2.64 to 10.45‰) than that at the seagrass meadows (3.51–12. 94‰). Significant differences in isotopic signatures and trophic level among trophic guilds at P. oceanica and C. racemosa were found. N fractionation at trophic guild level considerable differed between seagrass and macroalgae mats, especially for detritivores, deposit feeders, and herbivores. Filter feeders slightly differed with a relatively lower N signal at the seagrass and CR values at community level and at trophic guild level were higher in the C. racemosa invaded habitats indicating an increase in diversity of basal resource pools. C. racemosa did seem to broaden the niche diversity of the P. oceanica meadows it colonised at the base of the food web, may be due to the establishment of a new basal resource. The extent of the effects of invasive species on ecosystem functioning is a fundamental issue in conservation ecology. The observed changes in invertebrate and macrophytic composition, stable isotope signatures of concomitant species and consequent trophic guild and niche breadth shifts at invaded Caulerpa beds increase our understanding of the seagrass systems.     

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.     

Coccolith assemblages and their response to climate and surface hydrography in the Yellow Sea,Northwest Pacific,AD 1780–2011

A study of coccolith assemblages from a box core from the central South Yellow Sea(SYS) was performed revealing fluctuations on their relative abundance(%) that can be related to climatic and hydrographic changes over the last 230 years(1780–2011). Total coccolith abundances ranged from 7.0 to 55.1×10~6 coccoliths·g~(-1)sediment. Although the abundance of different species varied widely throughout the core, seven taxa dominated the assemblage. Among these species, Gephyrocapsa oceanica was the most dominant species, and it showed an average percentage of 50.1%. The pattern of G. oceanica(eutrophic species) was opposite to that of the combined percentage of Braarudosphaera bigelowii and Umbilicosphaera sibogae(both oligotrophic species), indicating that in the Yellow Sea(YS), the distribution pattern of G. oceanica might be characteristic of nutrient availability.Similar patterns between G. oceanica and the Siberian High were observed on an inter-decadal time scale,indicating that the East Asian Winter Monsoon(EAWM) may be an important driver of ecological changes in the YS. When the EAWM prevails, both the Yellow Sea Coastal Current(YSCC) and Yellow Sea Warm Current(YSWC)strengthen, and the increasing nutrient availability and warmer water brought by the strengthened YSWC favor eutrophic and warm-water coccolithophore species, such as G. oceanica. This likely mechanism demonstrates that coccolith assemblages can be used as benign and reliable proxy for climate change and surface oceanography.     

Impacts of urban wastewater discharge on seagrass meadows (Zostera noltii)

The abiotic disturbance of urban wastewater discharge and its effects in the population structure, plant morphology, leaf nutrient content, epiphyte load and macroalgae abundance of Zostera noltii meadows were investigated in Ria Formosa coastal lagoon, southern Portugal using both univariate and multivariate analysis. Four sites were assessed, on a seasonal basis, along a gradient from a major Waste Water Treatment Works (WWTW) discharge to a main navigation channel. The wastewater discharge caused an evident environmental disturbance through the nutrient enrichment of the water and sediment, particularly of ammonium. Zostera noltii of the sites closest to the nutrient source showed higher leaf N content, clearly reflecting the nitrogen load. The anthropogenic nutrient enrichment resulted in higher biomass, and higher leaf and internode length, except for the meadow closest to the wastewater discharge (270 m). The high ammonium concentration (158–663 μM) in the water at this site resulted in the decrease of biomass, and both the leaf and internode length, suggesting a toxic effect on Z. noltii. The higher abundance of macroalgae and epiphytes found in the meadow closest to the nutrient source may also affect the species negatively. Shoot density was higher at the nutrient-undisturbed site. Two of the three abiotic processes revealed by Principal Component Analysis were clearly related to the WWTW discharge, a contrast between water column salinity and nutrient concentration and a sediment contrast between both porewater nutrients and temperature and redox potential. A multiple regression analysis showed that these abiotic processes had a significant effect on the biomass-density dynamics of meadows and on the overall size of Z. noltii plants, respectively. Results show that the wastewater discharge is an important source of environmental disturbance and nutrients availability in Ria Formosa lagoon affecting the population structure, morphology and N content of Z. noltii. This impact is spatially restricted to areas up to 600 m distant from the WWTW discharge, probably due to the high water renewal of the lagoon.     

The introduction of Sargassum muticum modifies epifaunal patterns in a Moroccan seagrass meadow

Marine macrophytes sustain valuable epiphytic biodiversity. Nonindigenous macroalgae may induce changes in composition and structure of epifaunal assemblages and therefore support different assemblages from those associated with native species. In this study, differences in faunal community structure between the introduced fucoid Sargassum muticum and the native seagrass Cymodocea nodosa were tested over a year on an intertidal shallow sandy bottom at the southern introduction front, the El Jadida coastline (NW Morocco). Epifaunal community structure consistently differed between macrophytes through seasons, with more species‐rich assemblages associated with S. muticum than C. nodosa despite comparable abundances. The significantly greater epifauna diversity on S. muticum may be related to its structural complexity. However, the species contributing most to differences in assemblages between both macrophytes, such as Steromphala umbilicalis and S. pennanti, were found on both habitats with temporally varying abundances. Some species‐specific affinities were detected (Stenosoma cf. acuminatum, Elasmopus vachoni, Chauvetia brunnea). Nitrogen, dissolved oxygen, suspended matter and temperature were identified as the best explanatory variables contributing to the observed macroepifaunal patterns. This study provides evidence that S. muticum acts as a favourable and additional habitat for epifaunal species and supports a more diverse epifaunal assemblage in this Moroccan seagrass meadow.