Welcome mats? The role of seagrass meadow structure in controlling post-settlement survival in a keystone sea-urchin species

Processes acting on the early-life histories of marine organisms can have important consequences for the structuring of benthic communities. In particular, the degree of coupling between larval supply and adult abundances can wield considerable influence on the strength of trophic interactions in the ecosystem. These processes have been relatively well described in rocky systems and soft-sediment communities, and it is clear that they are governed by very different bottlenecks. Seagrass meadows make interesting study systems because they bear structural affinities to both soft sediments as well as rocky substrates. We examined the early-life history of Paracentrotus lividus, one of the dominant herbivores in Mediterranean seagrass meadows, to identify the drivers of population dynamics in this species. We measured spatial and temporal variability in sea urchin post-settlement in 10 Posidonia oceanica meadows in the North-Western Mediterranean over a period of two years, and compared the numbers with the one-year old cohort a year later (i.e. the new population recruitment) as well as between successive size–age groups. Urchin post-settlers differed substantially between meadows but were present in both years in all meadows surveyed, suggesting that larval supply was not limiting for any of the studied sites. However, in six of the studied meadows, the one-year cohort of urchins was absent in both years, indicating that post-settlement processes strongly affected urchins in these meadows. In contrast, in four of the studied meadows, there was a strong coupling between post-settlers and one-year cohort individuals. These meadows were structurally different from the others in that they were characterised by an exposed matrix of rhizomes forming a dense seagrass mat. This mat apparently strongly mediates post-settlement mortality, and its presence or absence dictates the successful establishment of urchin populations in seagrass meadows. As the population aged, the relationship between size–age groups decreased evidencing the action of other processes. Yet, these results indicate that differences in physical structure are a vital bottleneck for sea urchin populations in seagrass meadows. Exploring the interaction between ecosystem structure and early-life history may provide a broader and more unified framework to understand the dynamics of a range of benthic habitats, including rocky substrates, soft sediments and seagrass meadows.     

In Situ Measurement of Pinna nobilis Shells for Age and Growth Studies: A New Device

Abstract. Pinna nobilis Linnaeus 1758 is an endemic bivalve mollusc in the Mediterranean Sea, where it inhabits seagrass meadows, especially Posidonia oceanica (L.) Delile. It is the largest bivalve in the Mediterranean, reaching lengths up to 120 cm. In its natural habitat, P. nobilis lives with the anterior part of the valve buried in the seabed, attached to Posidonia rhizomes by byssus threads.
This habit makes it impossible to measure its total length directly in situ. As the only way to determine the individual age is the relationship between age and total length, several equations have been proposed to estimate total length by relating it to the unburied parts of the shell. Such measurements are essential to ecological studies that consider age, growth, and population dynamics, and that evaluate the environmental factors that affect this species.
Accurately estimating total length depends on the accuracy and precision of the method employed to measure the unburied shell parts. In this paper, we point out the lack of precision of the instruments and methods used until now; we also demonstrate the reason for this imprecision. A new device to measure unburied parts of Pinna nobilis with a precision comparable to that obtained when measuring extracted valves is described. This device is unaffected by substratum type and reduces measurement time. The latter is a very important feature, because these procedures are usually performed whilst SCUBA diving. Finally, a growth equation has been fitted to the measurements obtained with the new device from a population located in Moraira (Alicante, western Mediterranean).     

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.     

Human influence on seagrass habitat fragmentation in NW Mediterranean Sea

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

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

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

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

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

Low densities of sea urchins influence the structure of algal assemblages in the western Mediterranean

Numerous studies of interactions between urchins and algae in temperate areas have shown an important structuring effect of sea urchin populations. These studies focused almost wholly on the effect of high urchin densities on laminarian forests. In contrast, algal communities below 5–6 m depth in the northwestern Mediterranean are characterised by low sea urchin densities (<5 ind m⁻²) and the absence of laminarian forests. No previous research has addressed sea urchin/algal interactions in this type of community. To determine the effect of the most abundant echinoid species, Paracentrotus lividus, on well-established algal communities in this area, we performed a removal–reintroduction experiment in rocky patches located between 13 and 16 m depth in the northwestern Mediterranean, where sea urchin densities ranged between 0.9 and 3.4 ind m⁻². After 6 months, the cover of non-crustose algae was significantly higher in the plots from which sea urchins had been removed than in control plots (84 vs 67% cover). These removal plots reverted to their original state upon reintroduction of sea urchins. The non-crustose algae consisted of turfing and frondose forms, with the former representing some 70% of the non-crustose algal cover. Change in the cover of turfing algae was responsible for the significant increase in algal development in the sea urchin removal plots. The response of frondose algae to the treatment varied between algal species. It is concluded that grazing by P. lividus exerts a significant effect on habitat structure, even in communities with low sea urchin densities, such as those found in vast areas of the Mediterranean sublittoral.     

Relationships among predatory fish, sea urchins and barrens in Mediterranean rocky reefs across a latitudinal gradient

Previous studies conducted on a local scale emphasised the potential of trophic cascades in Mediterranean rocky reefs (involving predatory fish, sea urchins and macroalgae) in affecting the transition between benthic communities dominated by erected macroalgae and barrens (i.e., bare rock with partial cover of encrusting algae). Distribution patterns of fish predators of sea urchins (Diplodus sargus sargus, Diplodus vulgaris, Coris julis and Thalassoma pavo), sea urchins (Paracentrotus lividus and Arbacia lixula) and barrens, and fish predation rates upon sea urchins, were assessed in shallow (3-6m depth) sublittoral rocky reefs in the northern, central and southern sectors of the eastern Adriatic Sea, i.e., on a large spatial scale of hundreds of kilometres. No dramatic differences were observed in predatory fish density across latitude, except for a lower density of small D. sargus sargus in the northern Adriatic and an increasing density of T. pavo from north to south. P. lividus did not show any significant difference across latitude, whereas A. lixula was more abundant in the southern than in the central Adriatic. Barrens were more extended in the southern than in the central and northern sectors, and were related with sea urchin density. Fish predation upon adult sea urchins did not change on a large scale, whereas it was slightly higher in the southern sector for juveniles when predation rates of both urchins were pooled. Results show that: (1) assemblages of predatory fish and sea urchins, and barren extent change across latitude in the eastern Adriatic Sea, (2) the weak relations between predatory fish density and predation rates on urchins reveal that factors other than top-down control can be important over large scale (with the caveat that the study was conducted in fished areas) and (3) patterns of interaction among strongly interacting taxa could change on large spatial scales and the number of species involved.     

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.     

Sedimentary iron inputs stimulate seagrass (Posidonia oceanica) population growth in carbonate sediments

The relationship between sedimentary Fe inputs and net seagrass population growth across a range of Posidonia oceanica meadows growing in carbonate Mediterranean sediments (Balearic Islands, Spain; SE Iberian Peninsula, Spain; Limassol, Cyprus; Sounion, Greece) was examined using comparative analysis. Sedimentary Fe inputs were measured using benthic sediment traps and the net population growth of P. oceanica meadows was assessed using direct census of tagged plants. The meadows examined ranged from meadows undergoing a severe decline to expanding meadows (specific net population growth, from −0.14 yr⁻¹ to 0.05 yr⁻¹). Similarly, Fe inputs to the meadows ranged almost an order of magnitude across meadows (8.6–69.1 mg Fe m⁻² d⁻¹). There was a significant, positive relationship between sedimentary iron inputs and seagrass net population growth, accounting for 36% of the variability in population growth across meadows. The relationship obtained suggested that seagrass meadows receiving Fe inputs below 43 mg Fe m⁻² d⁻¹ are vulnerable and in risk of decline, confirming the pivotal role of Fe in the control of growth and the stability of seagrass meadows in carbonate sediments.     

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.     

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.     

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.     

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.     

Antioxidant response of the seagrass Posidonia oceanica when epiphytized by the invasive macroalgae Lophocladia lallemandii

The aim was to evaluate the antioxidant defences and the occurrence of oxidative damage in Posidonia oceanica under a stress situation due to the epiphytism of the invasive macroalgae Lophocladia lallemandii. P. oceanica leaves were collected in the absence (control station) and in the presence of the epiphytic algae L. lallemandii and the antioxidant enzyme activities, markers of oxidative damage, and hydrogen peroxide production were determined. Antioxidant enzyme - catalase, glutathione peroxidase and superoxide dismutase - activities were significantly higher in Posidonia epiphytized by L. lallemandii. Malondialdehyde, protein carbonyl derivates, and glutathione levels were also higher in L. lallemandii epiphytized P. oceanica leaves compared to control samples. The production of hydrogen peroxide was also significantly increased when Posidonia was epiphytized by L. lallemandii. The invasion of P. oceanica meadows by L. lallemandii appeared to induce oxidative stress in the seagrass as evidenced by increased levels of oxidative stress markers and antioxidant defences.     

Physiological response of the sea urchin Paracentrotus lividus fed with the seagrass Posidonia oceanica and the alien algae Caulerpa racemosa and Lophocladia lallemandii

The aim was to determine the effects of alien algae feeding on biomarkers of oxidative stress in the sea urchin Paracentrotus lividus. Sea urchins were fed during three months with the native seagrass Posidonia oceanica, and the alien macroalgae Caulerpa racemosa and Lophocladia lallemandii and biochemical analysis were performed in the gonads. A control group was immediately processed after sampling from the sea. Antioxidant enzyme and glutathione S-transferase activities and GSH concentration were significantly higher in sea urchins fed with alien algae when compared with the control group and the one fed with P. oceanica group. This response was more intense in the group fed with L. lallemandii respect to the C. racemosa group. The concentration of MDA, protein carbonyl derivates and 8-OHdG reported no significant differences between treatments. In conclusion, the invasive algae C. racemosa and L. lallemandii induced an antioxidant response in P. lividus without evident oxidative damage.     

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.     

Carbonate sedimentation and hydrodynamic pattern on a modern temperate shelf: The strait of Bonifacio (western Mediterranean)

The sedimentary features of the inner-middle shelf of the strait of Bonifacio (western Mediterranean) were analyzed to evaluate the relationship between the production and transport of biogenic carbonate sediments and the basin morphology and hydrodynamics. A three-dimensional hydrodynamic modeling was performed in order to simulate the influence of waves and currents at seabed level. Superficial sediments were collected at depths ranging from 5 to 80 m and were analyzed for grain size, mineralogical composition and skeletal carbonate composition. Posidonia oceanica seagrass meadows border the coasts in a narrow strip on both sides of the strait down to a depth of 40 m. At greater depths, the seabed is characterized by the presence of plateaus and ridges which are controlled by outcropping bedrock morphology.     

Habitat modification in tidepools by bioeroding sea urchins and implications for fine‐scale community structure

By creating novel habitats, habitat‐modifying species can alter patterns of diversity and abundance in marine communities. Many sea urchins are important habitat modifiers in tropical and temperate systems. By eroding rocky substrata, urchins can create a mosaic of urchin‐sized cavities or pits separated by exposed, often flat surfaces. These microhabitats appear to harbor distinct assemblages of species. We investigated how a temperate rocky intertidal community uses three small‐scale (<100 cm²) microhabitats created by or adjacent to populations of the purple sea urchin (Strongylocentrotus purpuratus): pits occupied by urchins, unoccupied pits, and adjacent flat spaces. In tidepools, flat spaces harbored the highest percent cover of algae and sessile fauna, followed by empty pits and then occupied pits. The Shannon diversity and richness of these sessile taxa were significantly higher in flat spaces and empty pits than in occupied pits. The composition of these primary space holders in the microhabitats also varied. Unlike primary space holders, mobile fauna exhibited higher diversity and richness in empty pits than in flat spaces and occupied pits, although results were not significant. The protective empty pit microhabitat harbored the highest densities of most trophic functional groups. Herbivores, however, were densest in flat spaces, concordant with high algal coverage. These results suggest the habitats created by S. purpuratus in addition to its biological activities alter community structure at spatial scales finer than those typically considered for sea urchins.