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.     

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.     

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.     

Variability in the structure of epiphytic assemblages of the Mediterranean seagrass Posidonia oceanica in relation to depth

The aim of the present study was to evaluate whether the variability in the structure of the epiphytic assemblages of leaves and rhizomes of the Mediterranean seagrass Posidonia oceanica differed between depths at a large spatial scale. A hierarchical sampling design was used to compare epiphytic assemblages at two different depths (10 and 20 m) in terms of both species composition and abundance and patterns of spatial variability in the Tuscan Archipelago (North Western Mediterranean Sea, Italy). Results showed significant differences in the structure of assemblages on rhizomes and leaves at different depths. These differences were related to species composition and abundance; differences were not significant for total biomass, total percentage cover and percentage cover of animals and algae. Whereas the higher variability was observed among shoots in all the studied systems, patterns of spatial variability at the other spatial scales investigated differed between the two studied depths. Moreover, in the present study, analogous patterns between depths resulted for both the assemblages of leaves and rhizomes, suggesting that factors that change with depth can be responsible for the spatial variability of both the assemblages (leaves and rhizomes), and operate regardless of the microclimatic conditions and the structure of assemblages.     

Temporal and spatial zonation of macroepiphytes on Posidonia oceanica (L.) Delile leaves in a meadow off Tunisia

The aim of this survey was to study the cover and the composition of macroepiphytic species on the leaves of Posidonia oceanica in the east of Tunisia. Surveys were conducted in December 2009 (winter period), March (spring period) and August 2010 (summer period) in a fringing reef located in Chebba. At each sampling date, 15 adult leaves were randomly collected and divided into basal and apical parts. The inner face of each part was examined with ocular glasses and microscope to estimate species cover by orthogonal projection. PERMANOVA and ANOSIM were used to test for differences of cover between sampling dates and leaf parts. SIMPER, cluster analysis, and PCA were used to ordinate species assemblages. Comparison of epiphytic cover along leaf blades showed significant differences for all groups, except cyanophycea, with high cover of hydrozoans and bryozoans in the basal part and high cover of algae in the apical part. The species composition and cover also vary with sampling date; minimum values were detected in December and the epiphytic community was composed of a few pioneer species, whereas maximum epiphytic cover values were registered in August, with the epiphytic community being composed of a more mature and more diverse community, termed ‘climax’. The main regulatory factors for this distribution are discussed.     

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.     

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.     

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.     

Temporal and depth distribution of microepiphytes on Posidonia oceanica (L.) Delile leaves in a meadow off Tunisia

A survey on the epiphytic microorganisms growing on Posidonia oceanica leaves was conducted along a depth transect along the coast of Eastern Tunisia (Mahdia). Samples were collected by SCUBA diving at depths of 3, 5, 10 and 12 m in July 2008 and January 2009. A total of 58 microepiphyte taxa were identified. Multivariate analyses revealed temporal and spatial variation of the abundance of epiphytic microalgae. Water motion, light availability, temperature and motility of species seem to be responsible of temporal and bathymetric variations of epiphytic microalgae. Unlike diatoms, dinoflagellates were more abundant in deep waters, suggesting that they are more vulnerable to hydrodynamics. The significant correlation between leaf area index and abundance of epiphytic species indicates that the phenological parameters of the host plant influence the abundance of the epiphytic microorganisms. Among the epiphytic dinoflagellates, our data showed a great number of potentially toxic species (Alexandrium minitum, Amphidinium carterae, Karenia selliformis, Coolia monatis, Karlodinium veneficum, Ostreopsis siamensis, Prorocentrum concavum, Prorocentrum minimum, Prorocentrum rathymum and Prorocentrum lima). These species were more abundant in the deep station under more sheltered conditions than found at inshore shallower stations, which are subjected to high water motion.     

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.     

Role of environmental factors in affecting macrophyte dominance in transitional environments: The Italian Lagoons as a study case

One of the main challenges to facilitate the classification of water bodies is to identify direct relationships between anthropogenic pressures and the behavior of biological organisms such as macrophytes in different environments including transitional areas. The investigation of many lagoons and ponds described here shows that macrophyte variables and the community composition have strong and univocal relationships with ecological parameters that are a measure of anthropogenic pressure on the ecological status of water bodies. The areas surveyed represent about 78% of the Italian transitional waters (169 sites sampled both in spring and fall). Anthropogenic impacts affect the availability of nutrients in the water column and surface sediments, causing changes in water transparency and phytoplankton concentration (as chlorophyll‐a [Chl‐a]) that act as the main drivers of variation for macrophyte assemblages, changing species dominance and the conditions that govern their presence or absence. The response of macrophytes to anthropogenic pressure is quite similar in all the examined transitional environments, even when the basin morphology, species richness and composition are different. Some taxa and species assemblages are so sensitive to environmental changes that monitoring them can be considered the most suitable and rapid method for assessing the quality of the environment they inhabit.     

Preliminary data on the impact of fish farming facilities on Posidonia oceanica meadows in the Mediterranean

While the extension of aquaculture along the coasts of the Mediterranean islands is today an economic priority, no data is available on the impact of these facilities on the littoral environment in general, and in particular on the Posidonia oceanica meadows that are responsible for the richness and diversity of these biotopes. The impact of fish farming facilities in the marine littoral environment is assessed at three Mediterranean sites in Corsica and Sardinia. Various parameters are considered: (i) the amount of light available, (ii) the nature of the sediment, and (iii) the vitality of the Posidonia oceanica beds. The findings confirm the impact of the aquaculture facilities within the water column (increase in turbidity, enrichment of the sediment in organic matter and nutrients), and it would appear that the Posidonia oceanica beds also undergo significant alterations. The decline in density of the meadows in the area influenced by aquaculture farming and their total disappearance beneath the facilities is particularly disturbing, even if the areas affected are not very extensive, at least for the smaller farms (a few hundred square metres), and if certain beneficial effects may be recorded (increase in benthic primary production). Because of its ability to record environmental alterations caused by these facilities (light, nutrients, trace metals), the Posidonia oceanica meadow is a good bioindicator for use in monitoring studies.     

Changes in the Composition of the Diatom Flora During the Last Century Indicate Increased Eutrophication of the Oder Estuary, South-western Baltic Sea

Four short sediment cores from the Oder estuary, south-western Baltic Sea, were studied with respect to their siliceous microfossil and organic carbon content. The sediments were dated by²¹⁰Pb. The objective was to detect and date changes in the composition of the diatom flora and link these changes to increased human impact in the drainage area during recent centuries. Two of the cores showed an unperturbed sedimentary sequence representing a complete historical record. A change in the composition of the diatom assemblages attributable to anthropogenic factors was recorded. This was dated to about 1900 in the Oderhaff. The change consisted of an increase in species that thrive in eutrophic waters and those indicating increased salinity or the availability of inorganic nutrients.     

A Study in Ultra-Ecology: Microorganisms on the Seagrass Posidonia oceanica (L.) DELILE

Abstract. Microbial colonization on the leaves of a shoot of the mediterranean seagrass Posidonia oceanica (L.) DELILE was studied using Scanning Electron Microscopy. Methods of field ecology such as transect, random plot and stratified sampling survey were applied to the microbial niveau to gain both qualitative and quantitative information on the microbial assemblage. While macro-epiphytic growth was significantly greater on the outer leaf sides, microbial colonization density was significantly higher on the inner leaf sides, both on leaf surface and epiphyte surface. Diatoms colonized the surface of incrusting algae and epiphytic animals in significantly lower numbers than the Posidonia leaf surface and were absent on erect epiphytic algae. Bacterial densities on epiphyte surfaces even exceeded values of the corresponding leaf surfaces on algal thalli near the leaf tips and on old leaves. Diatoms reach highest mean density on mature leaves and close to the leaf tips, while bacteria reach their greatest density on the oldest leaf and closer to the leaf base. Diatom density in general increases with exposure time of plant surface, while greatest bacterial density was observed at 7–10 weeks exposure. Basal leaf parts on younger leaves were dominated by rod-shaped bacteria, while distal leaf parts and old leaves were dominated by small coccoid bacteria. Surfaces of epiphytic algae were always distinctly dominated by small coccoid bacteria, and edges of thalli attracted high microbial densities. Microbial biomass (calculated from cell volumes using standard conversion factors) amounts to 2.3 g dry weight m-² in the Posidonia stand where the shoot was sampled. The observed patterns of epiphytic colonization are interpreted as the result of a complex, dynamically changing system of interactions both within the epiphytic community and between the epiphytic community, the host plant, and it's environment. A model of the organization of the epiphytic community on Posidonia leaves is presented. “Ultra-ecology” is a term introduced to denote a type of SEM research in the micro-environment which is analogous to in situ investigation in “macroscopic” ecological work.     

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.     

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.     

Leaf partitioning of the seagrass Posidonia oceanica between two herbivores: Is Sarpa salpa herbivory underestimated because of Paracentrotus lividus grazing?

In the Mediterranean region, herbivory appears to be a factor controlling the production of the endemic species Posidonia oceanica, which is mainly due to two main macroherbivores: the sea urchin Paracentrotus lividus, and the sparid fish Sarpa salpa. In this context the present study is a contribution in clarifying herbivory on P. oceanica by testing: 1) whether the abundance of grazing marks of the two herbivores is variable across different spatial scales, 2) whether spatial variation of P. lividus grazing marks and its density is consistent, 3) whether there is a dependence of the number of P. lividus grazing marks on its density, 4) whether the grazing of both macroherbivores, P. lividus and S. salpa, involves the entire leaf length, and 5) whether there is a dependence of the number of P. lividus grazing marks on the number of S. salpa grazing bites.     

Comparison between amphipod assemblages associated with Caulerpa racemosa var. cylindracea and those of other Mediterranean habitats on soft substrate

The spread of the invasive alga Caulerpa racemosa var. cylindracea in shallow-water habitats can present different faunal assemblage composition. We compared the amphipod assemblages associated with C. racemosa and natural habitats found on shallow-water Mediterranean soft substrata. Four vegetated habitats were compared: C. racemosa, Caulerpa prolifera, Cymodocea nodosa and Posidonia oceanica with unvegetated substrata. Samples were collected during two sampling periods (September 2004 and March 2005). A total of 63 amphipod species were recorded. The results showed that the vegetated habitats sampled, including C. racemosa stands, supported a higher abundance and species richness of amphipods. Furthermore, the assemblage structure differed between the different habitats, while the abundance of some species was significantly different, depending on habitat. For example, Microdeutopus obtusatus was favoured by C. racemosa habitat; Ampelisca diadema was associated with C. prolifera beds; and Hyale schmidti was more abundant in P. oceanica meadows. Habitat invasion by C. racemosa can exert an important influence on biotic assemblages, modifying habitat structure and associated fauna.