Seasonal Changes in Nitrogen, Free Amino Acids, and C/N Ratio in Mediterranean Seagrasses

Abstract. Seasonal changes in nitrogen, free amino acids, and carbon were investigated in the three Mediterranean seagrasses Posidonia oceanica, Cymodocea nodosa , and Zostera noltii. Leaves, rhizomes, roots, as well as dead plant material were analysed separately. Highest N-concentrations were obtained in the winter months, regardless of species or plant part. In contrast to the other two species, the N-content in Posidonia was higher in the rhizomes than in the leaves.
In the investigated species, marked differences in the free amino acid (FAA) composition were detected between species: Proline, lacking in Posidonia , was the main component in Cymodocea. In Posidonia , FAA decreased from 320umol g^-1 (dry wt) in leaf sheaths to 1.5 umol g^-1 (dry wt) in the leaf tips. The function of proline as a possible storage and/or stress metabolite is discussed.
High C/N values in dead P. oceanica and C. nodosa rhizomes as well as in P. oceanica wrack seem to be related to considerable resistance to decomposition. Low C/N ratios did not increase in detached C. nodosa and Z. noltii leaves, which began to decompose shortly after detachment from the plants.     

Seasonal Changes in Soluble Carbohydrates, Starch, and Energy Content in Mediterranean Seagrasses

Abstract. In all scagrass species investigated, rhizomes were the main storage organs for soluble carbohydrate reserves, and the highest levels were found in Cymodocea nodosa (U cria ) A scherson . Sucrose was the primary soluble carbohydrate identified in Posidonia oceanica (L.) D elile , and Zostera noltii H ornem . This disaccharide was dominant in Cymodocea nodosa roots, as well as in its rhizome in winter. Myo-inositol was the main cyclitol in Zostera noltii. In Cymodocea nodosa leaves 1-chiro-inositol was the main sugar component; it was also present to a lesser extent in the roots and rhizomes. The amounts of glucose and fructose were correlated with growth to a certain extent in all species. Higher amounts of starch were stored in the rhizomes of Posidonia oceanica and Zostera noltii. The energy content of Mediterranean scagrasses was significantly higher than in tropical species. No distinct seasonal trend in energy content could be observed.     

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.     

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.     

Growth Dynamics in Posidonia oceanica (L.) Delile

Abstract. Amounts of photosynthate in the rhizomes, in photosynthetically inactive basal parts of the leaves and different old leaf blades were examined. Winter leaf growth was supported by mobilisation of starch in the rhizome. This winter growth enables Posidonia oceanica to utilize the increased energy influx in early spring via the substantial leaf area already developed and to approach highest productivity in spring. During summer and autumn considerable concentrations of soluble carbohydrates were found in the leaves and rhizomes. Starch was stored in the rhizomes in concentrations of up to 6.8 % of dry weight. Levels of nitrogen and free amino acids were correlated with growth rates. The percentage of total nitrogen present as free amino acid-nitrogen decreased from November (35 %) to summer (less than 1 %), by which time leaf growth had stopped. In contrast to this, amounts of organic anions were low in winter and spring and reached their maximum in summer.
Within a shoot, sites of leaf growth were characterized by high amounts of total nitrogen and free amino acids (innermost leaves), while carbon Fixation was highest in the leaves #2–#4. Particularly high concentrations of soluble carbohydrates were found midway along leaf blades.
It is evident that this unusual growth rhythm of Posidonia oceanica was only possible because of the ability to store considerable amounts of carbon and nitrogen in the rhizomes.
The results of this investigation demonstrate on the one hand that the chemical composition of the plant is strongly correlated with growth and production, and on the other hand that it is dependent on environmental factors, such as energy influx and temperature, which change with season.     

Respiratory Rate and Assessment of Secondary Production in the Holothuroid Holothuria tubulosa (Echinodermata) from Mediterranean Seagrass Beds

Abstract. Respiration in Holothuria tubulosa was investigated in individuals from the Posidonia oceanica meadow off Lacco Ameno (Ischia Island, Italy). Respiratory rates increase with increasing body weight and increasing sea water temperature. Oxygen consumption of an average individual (7g dw body wall) ranges from 0.409 (14 °C) to 1.300 (26 °C) mg O₂· h^-1. Data on population density, mean size of individuals, and annual sea water temperature variations allow an assessment of holothuroid production. Values of 45.65 and 13.75 kJ · m^-2· y^-1 were calculated for shallow (3 to 10 m) and deep (25 to 33 m) areas of the Posidonia meadow, respectively. Holothuroid production shows a bathymetric pattern similar to primary production of the Posidonia -epiphytes complex and the production of Posidonia litter.     

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.     

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.     

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.     

Organic Matter Derived from a Seagrass Meadow: Origin, Properties, and Quality of Particles

Abstract. An investigation of the properties of particles found in wrack beds and suspended in the water body of a Posidonia oceanica system led to the following conclusion: except for the period of the leaf fall and at times of strong water movement (e. g. winter storms), the majority of the particles in the water show no similarity with seagrass particles. This is based on the comparison of C: N ratios and SEM inspection. Information on the magnitude of DOC fluxes in this system led to the formulation of the phase shift hypothesis from DOM to POM. Because wrack particles were shown to have relatively fast settling rates, it is expected that processes like adsorption onto inorganic particles, micellation and aggregate formation have more importance in regulating particle density in the water body than the resuspension of directly fractionated Posidonia debris. The implication of these findings are discussed and the importance of the Mediterranean seagrass P. oceanica as an energy source supporting secondary production in adjacent ecosystems is questioned.     

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.     

Modelling formation of complex topography by the seagrass Posidonia oceanica

Posidonia oceanica is a slow growing seagrass species that extends via growing rhizomes that grow only centimetres both horizontally and vertically each year. Posidonia oceanica forms topographically complex biogenic reefs of dead rhizome and sediments that are up to 4 m in height that are called “matte”. This study investigates the role of slow horizontal and vertical growth of rhizomes in the formation of topographic complexity in P. oceanica matte using agent-based modelling. The simulated infilling of landscapes by P. oceanica was run over 600 iterations (years) for 10 random starts of 150 agents each. Initial infilling rates were very slow and P. oceanica had limited cover after a century of growth. Growth accelerated after 100 years but plateaued after 400 years such that after 600 years only two-thirds of the landscape was occupied by P. oceanica. The pattern of spread of agents was initially random in direction but after larger patches were formed spread was radial from these patches. The seagrass landscape was initially highly fragmented with many small separate patches made up of a few agents each, with a Landscape Division index close to 1. Between 300 and 600 years Landscape Division declined sharply to 0.42, indicating patches had coalesced into larger more continuous meadows forming a less fragmented landscape. Perimeter to area ratio of seagrass patches declined exponentially from >1 to approximately 0.2 over 600 years of simulation. The matte developed from growth of patches and its greatest height occurred in more continuously occupied cells of the grid. The topography of the reef that occupied two-thirds of the landscape after six centuries of growth could be described as a pattern of channels between reef plateaus elevated 1–2 m above channels. These results demonstrate that development in P. oceanica meadows of three-dimensional structure, in the formation of biogenic reefs, can be explained by, and is an emergent property of, slow horizontal and vertical rhizome growth rates combined with the time it takes for the accumulation of rhizomes in any region of the landscape. As such, the model provides a parsimonious explanation for the development of complex matte topography.     

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.     

Biometry, Bathymetric Distribution, and Reproductive Cycle of the Holothuroid Holothuria tubulosa (Echinodermata) from Mediterranean Sea grass Beds

Abstract. Holothuria tubulosa individuals were sampled regularly at three different depths (6, 19, and 33m) over a 20-month period in a Posidonia oceanica meadow off Ischia Island (Italy). Biometrical analysis demonstrated that holothuroids have a well-marked size-related distribution according to depth, with the smallest individuals in the shallowest part of the meadow and the large ones in its deepest part. The observed distributions reflect the ability of H. tubulosa to grow more efficiently in areas of low water turbulence where Posidonia shoots are scattered and detrital food is more easily accessible. Recruitment takes place in the shallow meadow, and holothuroids with high reproductive potential only occur in the deep meadow. Small holothuroids (shallow individuals of similar sizes) may be either juveniles or sexually mature individuals; environmental constraints apparently restrict the growth of individuals and their gonads. There is a progressive downward migration of growing individuals when the whole population is considered, although at the level of the individual the migration occurs randomly. Accordingly, the size of a sexually mature individual is more a function of its past history in the meadow than of its real age.     

Heavy Metal Contents in Soft-Bottom Marine Macrophytes and Sediments Along the Mediterranean Coast of Spain

Abstract. Hg, Cd, Pb and Zn concentrations were determined in sediment and in tissues of five species of soft-bottom marine macrophytes ( Posidonia oceanica, Cymodocea nodosa, Zostera noltii, Ruppia cirrhosa and Caulerpa prolifera ) along the Spanish Mediterranean coast. Levels of metals were low in most of the sampling stations and similar to those found by other authors in uncontaminated zones. Certain locations, however, showed some degree of contamination (Cambrils, Almassora, Alacant, Mar Menor and El Portús). In Santa Pola we found high contents of metals in one sample of sediment due to the high proportion of the fine fraction (particules < 63µm) and organic matter, but not in the seagrass species. Mercury and zinc concentrations in the sediment are correlated to those in at least some anatomic fractions of Posidonia oceanica, Cymodocea nodosa and Caulerpa prolifera , suggesting that these species reflect the levels of these metals in the environment. Lead contents in the alga Caulerpa prolifera are also correlated to those in the sediment, while no significant correlations were obtained for cadmium. Among the species studied, Posidonia oceanica and Cymodocea nodosa seem to be better biomonitors than Caulerpa prolifera .     

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 restudy of ostracode assemblages in the surface sediments of the Huanghai Sea

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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.