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.     

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.     

The use of surface alkaline phosphatase activity in the seagrass Posidonia oceanica as a biomarker of eutrophication

Eutrophication is one of the most relevant man‐induced changes occurring in coastal waters. The identification and assessment of specific responses to eutrophication in seagrasses can provide a useful tool for the detection of changes in the water quality in coastal zones, given the wide range of distribution of these organisms. In this study, we combine a correlational (across‐sites comparison) and a manipulative (fertilization experiment) approach to evaluate the usefulness and potential of alkaline phosphatase activity (APA) in the endemic Mediterranean seagrass Posidonia oceanica as an eutrophication biomarker. Our results showed that APA decreases promptly following nutrient additions, the response being maintained except during the winter period. APA also varies across natural meadows under different levels of nutrient discharges at scales relevant for monitoring purposes. AP activity seems to be an optimal ‘physiological biomarker’ that responds promptly and reliably to a pulse of eutrophication exposure. However, other considerations, such as the seasonality (the response disappears in winter), suggest its use with some caution and, as far as possible, as a complement of other bio‐indicators.     

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.     

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.     

Study of a Crustacea Decapoda Taxocoenosis of Posidonia oceanica Beds from the Southeast of Spain

Abstract. During 1986 and 1987 a decapod crustacean taxocoenosis of Posidonia oceanica beds from the southeast of Spain has been studied.
Data on abundance-dominance of the 50 species found and other information such as reproduction and the selective use of gastropod shells by the dominant hermit crabs arc given.
Differences in richness and cquitability between shallow (3–4 m) and deeper (5–7 m) samples arc also shown.
The taxocoenosis presents a geometrical distribution in which only 9 species have a relative abundance higher than 1 % and in which the structure is basically controlled by the two dominant hermit crabs, Cestopagitius limidus and Calcium tubularis , and hydrodynamic conditions.     

Systematics and Ecology of the Hydroid Population of two Posidonia oceanica Meadows

Abstract. The ecology and systematics of the hydroid population of Posidonia oceanica meadows have been studied in two different zones of the Western Mediterranean. A new sampling method has been employed and the results obtained have been compared with those of a standard method. The distribution of the species and several morphological variations are pointed out. Depth-related faunistic zones are evidenced. The key-species of the Posidonia hydroid population arc individuated.     

A Map of the Posidonia oceanica Beds of Marina ďElbu (Corsica, Mediterranean)

Abstract. A large-scale map of the Posidonia oceanica seagrass beds as well as other benthic substrata and communities was compiled for a cove of a marine reserve free from pollution and man-made disturbance. An extensive diving survey of the cove and the interpretation of aerial colour photographs were used. The greater part of the cove is occupied by the P. oceanica meadow. Patches of dead "matte" were observed at a depth of between 7 and 14 m. They may be the result of a natural hydrodynamic equilibrium between building and erosion.     

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.     

Structure of the Epiphytic Community of Posidonia oceanica Leaves in a Shallow Meadow

Abstract. The structure of the epiphytic community (hydroids, bryozoans, and the most abundant macrophytes) of Posidonia oceanica leaves in a shallow meadow near Lacco Ameno, Ischia was studied. The structure of the community changed along an age gradient that extended from the inner to the outer leaves and from the basis to the apex of each leaf. This structure was also affected by microclimatic factors. The dynamics of community "maturation" is described.     

Posidonia oceanica (L.) Delile: A Structural Study of the Photosynthetic Apparatus

Abstract. P. oceanica forms broad prairies which completely surround the island of Ischia. This is a histochemical and ultrastructural study of the photosynthetic apparatus of the Posidonia plant. In the leaf, epidermal cells have a porous region underneath the cuticle and contain most of the chloroplasts. Fiber bundles run hypodermally and phenol storing cells are present in the mesophyll. Chloroplast ontogenesis is characterized by juvenile stages with provesicular bodies. Sea depth apparently affects some morphological features of the photosynthetic apparatus, like leaf blade thickness, phenolic cell and hypodermal fiber frequency.     

Spatio-Temporal Distribution of Algal and Animal Communities in a Posidonia oceanica Meadow

Abstract. Investigations were carried out in a Posidonia oceanica meadow at Ischia (Gulf of Naples) along a depth gradient from 1 to 32 m in November 1979 and May 1981.
In these two months, different discontinuities were found at the sampling stations (1, 5, 10, 20, 30 m) at 10 m for leaf features, between 1 and 5 m for the algal community and the vagile fauna. The leaf features (length, surface, and biomass) along the transects show a delay in leaf production toward the deeper stations, below the thermocline.
The algal community shows in both seasons and in all stations a persistence of an encrusting layer, mostly represented by Corallinaceae and the brown alga Myrionema orbiculare , while at the shallowest station (1 m) the community is characterized by a more developed upright layer.
The vagile fauna of the leaf stratum, mostly represented by Polychaeta, Mollusca , and Crustacea (Peracarida and Decapoda) , shows a consistent zonation in both seasons along the transect. A superficial community at I m, characterized by a low number of species and individuals and specialized for high environmental energy levels, and a deeper community, persistent in time and more strictly related to the Posidonia oceanica meadows, are identified.
The influence of environmental factors and the importance of meadow structure for the zonation of the algal and animal communities are discussed.     

Long-term change in the structure of a Posidonia oceanica landscape and its reference for a monitoring plan

Abstract On the basis of a detailed cartographic survey carried out by Side Scan Sonar and a towed underwater video camera during 2005, and from a series of historical maps (1959, 1980, 1990), an extensive regression of Posidonia oceanica (L.) Delile beds was evidenced for a vast area of the central Tyrrhenian Sea (Latium coast, Italy). The total loss of P. oceanica surface was assessed through GIS estimate. In 1959, the Posidonia beds extended over 7290 ha, while in the 2005 survey they had regressed to 2899 ha, a loss of about 60% of their coverage. Also the seagrass lower limit showed a general depth decrease in time. Total seagrass coverage loss and lower limit regression were not uniform along the whole investigated areas and three main sub-areas have been identified with different degrees of regression somehow related with coastal potential human-mediated impacts. From different coverage estimates of the present survey and of the previous maps, minimum sampling areas were calculated through bootstrapping simulation routines from small sampling areas (Landscape Units) to reach the nearest estimate of the observed condition in the different periods.     

The Detritic Compartment in a Posidonia oceanica Meadow: Litter Features, Decomposition Rates, and Mineral Stocks

Abstract. The ecosystem associated to the Mediterranean seagrass Posidonia oceanica shows a clear distinction in two subcompartments regarding turnover time: aboveground and belowground. Aboveground parts (leaves) are highly dynamic, and most of the leaf material is decomposed or exported in less than one year, representing a net loss of nutrients. In contrast, belowground biomass (roots and rhizomes) has a turnover time of the order of centuries, with a consequent accumulation of organic matter in the sediment. The accumulation rates for the single elements rank in the order C > N > P. This ecosystem may be considered as a sink for biogenic elements.     

Seagrass meadows at the extreme of environmental tolerance: the case of Posidonia oceanica in a semi-enclosed coastal lagoon

Atoll‐like structures of the endemic Mediterranean seagrass Posidonia oceanica were encountered in the innermost area of the Stagnone di Marsala, a semi‐enclosed coastal lagoon along the western coasts of Sicily. The area is characterized by limited water exchange with the open sea and by a marked seasonal variation of water salinity and temperature, reaching beyond the theoretical tolerance limits of the species. In the present study we determined the genetic composition of the atoll‐like structures, as well as the growth performance and flowering rate of these stands. We also assessed whether and to what extent the atoll‐like structures are genetically isolated from plants growing in meadows outside the lagoon. For this purpose we utilized 13 microsatellite markers to genotype single shoots sampled inside and outside the lagoon. Lepidochronological analyses were performed on the same shoots to determine the annual rhizome growth rate, the number of leaves and the inflorescences formed as an estimate of growth‐ and reproductive performance over the years. The innermost area of the lagoon showed a lower number of alleles, a lower percentage of polymorphic loci, a lower clonal diversity, but higher heterozygosis excess with respect to the other areas analysed. Spatial autocorrelation was here significant, up to slightly below 300 m. Shoots collected in the atolls exhibited a 25% lower vertical growth rate and 16% lower leaf formation in comparison to those in open‐sea meadows. No flowering events were recorded during 24 years of investigation, whereas inflorescences were observed frequently in meadows outside the lagoon. Results from F_st and factorial correspondence analysis confirmed the expected genetic isolation of the confined atolls with respect to the meadows outside the lagoon and revealed limited gene flow within the lagoon itself. Apparently, the enclosed system of the Stagnone lagoon is genetically isolated, with a possible selection of genotypes adapted to persistent stressful conditions, consistent with reduced growth and lack of flowering events.     

Spatio-temporal variation in the structure of a deep water Posidonia oceanica meadow assessed using non-destructive techniques

The Malta‐Comino Channel (Maltese islands, central Mediterranean), supports extensive meadows of the seagrass Posidonia oceanica that in some places extend to a depth of around 43 m, which is rare for this seagrass. To assess spatial and temporal variation in the state of the deeper parts of the P. oceanica meadow with time, data on the structural characteristics of the seagrass meadow at its lower bathymetric limit were collected during the summers of 2001, 2003 and 2004 from four stations (two stations within each of two sites) located at a similar depth, over a spatial extent of 500 m. Shoot density was estimated in situ, while data on plant architecture (number of leaves, mean leaf length, and epiphyte load) were successfully obtained using an underwater photographic technique that was specifically designed to avoid destructive sampling of the seagrass. Results indicated that P. oceanica shoot density was lower than that recorded from the same meadow during a study undertaken in 1995; the observed decrease was attributed to the activities of an offshore aquaculture farm that operated during the period 1995–2000 in the vicinity of the meadow. ANOVA indicated significant spatial and temporal variations in meadow structural attributes at both sites during the 3‐year study; for example, shoot density values increased overall with time at site A; a indication of potential recovery of the meadow following cessation of the aquaculture operations. Lower shoot density values recorded from site B (compared with site A) were attributed to higher epiphyte loads on the seagrass, relative to those at site A. The findings, which include new data on the structural characteristics of P. oceanica occurring at depths >40 m, are discussed with reference to the use of the non‐destructive photographic technique to monitor the state of health of deep water seagrass meadows.     

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.     

Polychaete Depth Distribution in a Posidonia oceanica Bed (Rhizome and Matte Strata) and Neighbouring Soft and Hard Bottoms

Abstract. The study was carried out in a Posidonia oceanica bed and in the neighbouring sandy and rocky bottoms, at Ponza Island (Central Tyrrhenian Sea), in order to investigate the distributional pattern of the polychaete community at different depths. A total of 15232 individuals of polychaetes belonging to 218 species was collected and three new species of Syllidae were found. The polychaete community of the matte can be clearly distinguished from the neighbouring communities; this is due to the high species richness in Posidonia , which is related to the heterogeneity of the biotope. In the Posidonia bed the shallow stations (1 m and 5 m depth) were the richest in terms of species numbers and individuals due to the high shoot density of the seagrass bed. In the shallow waters polychaete community structure changed according to the different exposure to water movement. Rocky bottom species, which preferentially inhabit crevices, were more abundant in the matte exposed to water movement (5–10 m depth). In contrast, soft-bottom species were more abundant in the sheltered zone (1 m depth) as well as in the deeper zone due to the reduction of the Posidonia bed.     

Fish Assemblages of Posidonia oceanica Beds at Port-Cros (France, NW Mediterranean): Assessment of Composition and Long-Term Fluctuations by Visual Census

Abstract. Observations of the fish fauna of Posidonia oceanica (L.) DELILE seagrass beds were undertaken in two shallow stations and one deeper station in Port-Cros National Park, between November 1984 and September 1987. Demographic structure, density and biomass of the 19 most abundant species or taxa were assessed by several non-destructive sampling methods. The fish assemblages studied were dominated in number and biomass by a small number of species (Labridae: Symphodus ocellatus and S. tinca ; Sparidae: Diplodus annularis and Sarpa salpa ). Large individuals were infrequent and small size class fish were dominant especially in sheltered shallow areas. Recruitment and migrations (shortor long-term) have been identified as the most important causes of abundance variations over the year. The four trophic categories were sampled only in the sheltered shallow zone, where the herbivore Sarpa salpa represents a strong proportion of the fish fauna (40–70%) during summer. The fish assemblages differ between the three seagrass beds, and the depth appears as the most important factor affecting the fish assemblages in P. oceanica . The functional difference between shallow and deeper seagrass beds is therefore discussed.     

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.