Posidonia oceanica (Linnaeus) Delile

Posidonia oceanica (L.) Delile is a seagrass species endemic to the Mediterranean Sea that forms dense and extensive underwater meadows with leaves that can attain 1 metre in height. These meadows provide important ecological functions and services and support a highly diverse community, including species of economic interest.

Since the 1970´s, a worldwide decline of seagrass distribution and abundance has been detected and causes are mainly attributed to the negative influence of anthropogenic impacts (Orth et al., 2006). P. oceanica is very sensitive to specific impacts such as bottom trawling (Sánchez and Ramos, 1996), anchoring (Francour et al., 1999), coastal constructions (Ruiz and Romero, 2003), chemical wastes (Pergent-Martini and Pergent, 1995), fish farm effluents (Delgado et al., 1999; Ruiz et al., 2001; Pergent-Martini et al., 2006) desalination plants (Gacia et al in press), geodynamic alterations (Badalamenti et al., 2006, biological invasions (Villèle and Verlaque, 1995) and many others. The effect of these impacts, alone or combined; cause either a loss of vegetated areas, a reduction in seagrass abundance (cover and/or shoot density) or a deterioration of plant health.

P. oceanica beds are identified as a priority habitat for conservation under the European Union’s Habitats Directive (Dir 92/43/CEE). Conservation management is mainly focused on protection from physical damage through the installation of artificial reefs and seagrass-friendly moorings for boats, which reduce the erosive pressure of otter-trawling and free anchoring in shallow meadows. The control of invasive species has also been performed recurrently in some P. oceanica beds.

Regressed meadows are prone to invasion by one or more of the potential substitutes for P. oceanica (Bianchi and Peirano, 1995; Montefalcone et al., 2006) such as the other common Mediterranean seagrass Cymodocea nodosa (Ucria) Ascherson, the native Mediterranean green alga Caulerpa prolifera (Forsskal) Lamouroux and the two alien green algae Caulerpa taxifolia (Vahl) C. Agardh and Caulerpa racemosa (Forskal) J. Agardh.

There is a need to further develop regulations for activities that have a negative impact on P. oceanica beds (e.g. pollutants level limits and allowed minimum distances of impact sources to meadows) and to implement them through a vigilance system that is coordinated with the existing seagrass monitoring networks.

Once the cause of habitat perturbation is eliminated, the slow growth of P. oceanica beds means that recovery can take centuries. Measures like remediation of seagrass sediments enriched with organic matter, or transplanting of P. oceanica, are at an experimental stage.

Table of Contents

1. Introduction

2. Morphology

3. Propagation

4. Ecological Benefits

5. Species that Depend on the Posidonia oceanica Habitat - Key Fauna:

• Sea urchins
• Fish
• Molluscs

6. Threats:

• Sedimentation/Erosion Balance
• Eutrophication
• Direct Erosion by Boat-trawling and Boat Anchoring
• Expansion of Invasive Algal Species
• Salinity Increase in the Vicinity of Water Desalination Facilities
• Fish Farm Activity
• Climate Change

7. Trends

8. Conservation Measures

9. Restoration Initiatives:

• Dredging Recovery
• Transplantation of Posidonia oceanica
• Transplantation of Seedlings
• Success of Posidonia Oceanica Transplantation

10. See Also

Introduction

Posidoniaceae is one of the 5 families of seagrasses, descendants of terrestrial plants that re-colonised the ocean between 100 and 65 million years ago. Seagrasses are monocotyledons that are not true grasses (family Poaceae) but are closely related to the lily family, Magnolyophyta.

The Posidonia genus has 9 species: P. angustifolia, P. australis, P. sinuosa, P. coriacea, P. denhartogii, P. kirkmanii, P. ostenfeldii, P. robertsonae and P. oceanica. Whilst the other species are found around southern Australia, P. oceanica is unique to the Mediterranean Sea and grows within a temperature range of 10ºC to about 30ºC. Temperature is therefore considered the central parameter controlling the geographical distribution of this species. P. oceanica beds cover between 25,000 and 50,000 km2 of the coastal areas of the Mediterranean, corresponding to 25% of the sea bottom at depths between 0 and 40 m.

P. oceanica is a marine flowering plant (angiosperm) with a millenary life span, a need for light and clear water, and a very slow growth (a few centimetres per year) and poor reproduction rate. P. oceanica propagates slowly, through the elongation of horizontally growing rhizomes, which eventually forms tightly knit mattes of rhizomes that hold the sandy seabed in place. Thus the meadow rises, over decades, producing reefs up to 3 m high that can be thousands of years old. These meadows accumulate sediment and mediate wave motion, minimising the effect of wave action and therefore helping to stabilise the coastline. This process also reduces the amount of sediment suspended in the water, helping to maintain the clear water conditions P. oceanica requires for growth.

Morphology

Like other angiosperms P. oceanica has roots, stems, leaves, flowers and fruits. At the base of each plant is a rhizome, which is actually a modification of the stem. The rhizomes of P. oceanica can easily be distinguished from those of the other three European seagrass species by the dense, hairy remains of old, degrading leaf sheaths found around the rhizomes. These remains can also be found as conspicuous balls of fibres washed onto the beaches, known as egagropili.

Vertical rhizomes are attached to horizontal rhizomes that branch and expand by terminal apices. Rhizome internodes are short (0.5 to 2 mm) reflecting the slow horizontal growth of the plant, and the thickness of the rhizomes vary between 5 and 10 mm. The roots are 3-4 mm thick, up to 40 cm long and richly branched, attaching the plant to the substratum and allowing the absorption of nutrients from the sediment. Nutrients are taken up from the sediments by the roots and transported to the meristems and leaves for growth. Leaves themselves can also absorb nutrients, and are the main structures for absorbing carbon dioxide CO2 from the water column. Leaf life span in P. oceanica is almost a year with shoots living for decades.

P. oceanica has leaf bundles consisting of 5 to 10 leaves attached to a vertical rhizome. The leaves are broad (5 to 12 mm) and the length usually varies from 20 to 40 cm in length, but may be up to 1 m. A section of the petiole of a leaf shows a true network of lacunae throughout the plant from the tip of the leaf to the end of the roots, called the aerarium, and all the tissues are steeped in gas. This is the main difference between the marine phanerogams and other marine vegetation, which never left the sea.

The rate of formation of seagrass leaves, rhizomes and roots depends on the activity of meristems, where active cell division takes place. The horizontal growth and vertical extension of P. oceanica rhizomes is at a rate of only a few centimetres per year, producing, on average, a branch every 30 years. Shoots produce new leaves every 50 days on average.

The vascular and lacunal systems of the roots and rhizomes facilitate the transport and exchange of fluids and gasses respectively. A proportion of the oxygen O2 that is produced in the leaves during photosynthesis is diverted to the lacunae in the leaves, and then diffuses through the rhizomes to the roots. Some of the O2 diffuses out of the roots to maintain less hypoxic conditions around the rhizosphere. Seagrasses growing in normally hypoxic or anoxic sediments are dependent on transporting sufficient O2 down to their roots to maintain aerobic respiration and to reduce sulphide formation around the roots.

When factors that negatively inhibit O2 production, such as low light, occur simultaneously with factors that increase the O2 demand, such as increased organic loading of the sediments, the risk of sudden, dramatic loss of seagrass beds is increased, accelerated by the further increase in O2 demand created when the dead plant material is degraded.

Propagation

P. oceanica flowers between August and November. The number of shoots flowering in meadows is generally lower than 3 % per year. However, massive flowering events (more than 10% shoots flowering) have been observed associated with extremely warm summers. Flowering intensity is negatively correlated with water depth.

P. oceanica flowers are yellow and can produce half a dozen seeds per shoot. Fruit are large (10 mm) and known as sea olives. Many female flowers do not develop viable fruits due to abortion and predation, and actual seed production is less than 1% of potential. Among the European seagrasses, only P. oceanica has buoyant seeds capable of long-range (10’s of km) dispersal. Nonetheless, young individuals originating from seedlings are rarely found and P. oceanica primarily propagates vegetatively by elongating the rhizomes; a whole meadow may be one single clone resulting from one ancient seedling.

The little investment and low success of sexual reproduction, combined with the extremely slow clonal spread explains the extremely slow colonisation rate of P. oceanica plants. Numerical models simulating the occupation of space by a P. oceanica meadow indicate that it would need 600 years to cover 66 % of the available space around the Mediterranean coastal strip at depths in which it is able to grow. Similar colonisation time scales have been retrospectively calculated based on patch size and patch growth rate in patchy P. oceanica meadows. The very long time scales for colonisation of this species indicate that recovery of disturbed meadows, where important plant losses have occurred, would involve several centuries.

Ecological Benefits

P. oceanica meadows play an important biological and ecological role, providing benefits to humans that rank among the highest of all ecosystems on earth. They are vital within their ecosystem for the production of oxygen and organic material and they support numerous animal species that utilise them as a site for breeding, feeding and shelter. The leaves and rhizomes increase the surface available to sessile species and offer shelter to mobile species, thereby sustaining a diverse community (Templado 1984). Posidonia beds are especially valuable as nursery grounds for several commercial species (Francour 1997).

A moderately wide (1 km) belt of P. oceanica meadow may produce litter in excess of 125 kg of dry seagrass material per metre of coastline each year (mostly during autumn). This material accumulates on the beach, developing cushions up to 4 metres high, which can in turn sustain a complex invertebrate food web, protect the shoreline from erosion, deliver inorganic material in the form of carbonate and silica shells and, when transported further inland by the wind, may act as seed material for dune formation (Borum et al. 2004).

Dried P. oceanica leaves were traditionally used in Mediterranean countries as packing material to transport fragile items of glassware and pottery, and also to ship fresh fish from the coast to cities. As parasites are less successful in P. oceanica leaves than in straw, they were utilised in stables, as roof insulation, and as a filling material for mattresses to prevent respiratory infections. Further medicinal uses included the alleviation of skin diseases and leg pain caused by varicose veins.

The seasonality of P. oceanica allows other ecosystems to be enriched by the swathe of organic material that is carried by the currents and waves. The biomass of P. oceanica decomposes slowly and stores a significant amount of carbon in the sediment over long periods. Seagrasses are responsible for 12% of the carbon stored in ocean sediments and play a significant role in the regulation of the global carbon cycle. In daylight, P. oceanica meadows oxygenate coastal waters (Bay 1984), and have been called the lungs of the Mediterranean Sea.

Left undisturbed over a very long period P. oceanica seagrass beds form into reefs that slow down wave movement and protect the shore from erosion. The leaves trap larger grains of sand, providing a natural filter that ensures water reaching the shore is clearer and cleaner. P. oceanica meadows are excellent indicators of environmental quality as they can only grow in clean unpolluted waters. Moreover, their rhizomes concentrate radioactive, synthetic chemicals and heavy metals, recording the environmental levels of such persistent contaminants.

Species that Depend on the Posidonia oceanica Habitat

P. oceanica meadows serve as sanctuaries for numerous species during breeding and as a year round refuge and food source for others. Calcifying organisms such as coralline algae, molluscs and foraminifera, some of which grow between the seagrass shoots and some as epiphytes are important components of the meadows. Epiphytic communities, growing on the leaves and rhizomes of the plant, provide a food source for sea slugs, sea hares (e.g. Aplysia fasciata and Aplysia depilans) and several species of nudibranch, which also deposit their eggs on the Posidonia leaves.

In healthy meadows, the red algae Fosliella spp. and Hydrolithon spp., and brown algae, like the complex Giraudio-Myrionemetum orbicularis, cover the tips of the leaves. Sessile animals, such as hydroids (over 44 species identified including the obligate species Sertularia perpusilla and Plumularia obliqua posidoniae), bryozoans (more than 90 species, like the obligate species Electra posidoniae and Lichenopora radiata) and encrusting ascidia (e.g. Botryllus schlosseri) are also a common component of the leaf epiphytic community.

Algae adapted to low levels of light intensity, mostly red algae, colonize the rhizomes (e.g. Peyssonnelia squamaria and Udotea petiolata) and light dependent algae like Jania rubens may appear on meadow borders. In fact, the primary production of P. oceanica meadows is a combination of seagrass leaf growth and that of micro- and macro-epiphytic and benthic seaweeds, with the latter groups occasionally contributing as much to the ecosystem production as the seagrass itself (Hemminga and Duarte 2000).

The sediment that accumulates within seagrass beds is much richer in organic matter and nourishing salts than adjacent sediment of the bare sandy areas, and an army of different suspension-feeding animals such as feather stars, ascidia, sponges, hydrozoa, and tube worms take advantage of this, together with sediment-feeding animals such as sea cucumbers and brittle stars. These in turn serve as food for carnivorous animals such as crabs, fish, octopus, cuttlefish and starfish.

Key Fauna

Information on key fauna associated with P. oceanica beds can be directly relevant for the interpretation of seagrass monitoring results, particularly in cases where the fauna grazes the seagrasses. Moreover, the associated species assemblages often reflect plant health and their monitoring adds to the general understanding of the importance of seagrass beds for coastal biodiversity.

Relevant key fauna to measure in connection with P. oceanica monitoring programmes are listed below:

Sea urchins – are often important grazers of seagrasses. Grazing by the sea urchin Paracentrotus lividus occasionally (overgrazing events) can be so intense that it may even result in the elimination of extensive seagrass patches. The density of sea urchins increases with increasing nutrient in the environment, and, hence, concentration in plant tissues. An increased grazing activity by sea urchin has, for example, been observed in P. oceanica meadows situated under fish cages. However, at more typical or natural nutrient levels, sea urchins have a relatively minor impact on the seagrass whilst grazing by the fish species Sarpa salpa can outstrip the plants' leaf production.

The most abundant Echinoderms within P. oceanica meadows are sea cucumbers (16 species described) which play an important ecological role as sediment filterers. Among them, Holothuria tubulosa predominates in dense, sandy meadows, while H. polii is more prevalent in sparse or degraded meadows, although it is very difficult to distinguish these two species. At night, many mobile species living within the rhizomes migrate to feed in the canopy.

Fish – As indicated above, Sarpa salpa, or cow bream, form large schools in Posidonia oceanica meadows during summer causing mowed patches in which the biomass can be reduced by as much as 50% (Tomas et al 2005). Many fish species utilise P. oceanica meadows as nurseries during their juvenile stage. There are also resident species, the most common of which are Gobius spp. (living on rhizomes), as well as Labrus merula, L. viridis, Symphodus spp., Diplodus spp, Sarpa salpa, Coris julis and Chromis chromis. There are also some obligate species living within the leaf canopy, like the cryptic species Opeatogenys gracilis and Syngnathus typhle. The endangered species Hippocampus hippocampus is also found within the canopy.

Molluscs – some large species, like the Mediterranean bivalve Pinna nobilis, are exclusively dependent on seagrasses, and are therefore inherently affected by physical impacts on the meadows, e.g. boat anchoring. Presence of P. nobilis is a characteristic of healthy seagrass meadows. Some species of snail (e.g. the genus Rissoa) are also frequent on seagrasses, whereas predatory molluscs such as the cuttlefish Sepia officinalis and octopus Octopus vulgaris are frequently seen around the edges of P. oceanica beds. Mollusca (more than 185 species described) and Crustacea (more than 120 species of Copepoda, Decapoda and Amphipoda) are the most abundant faunal groups in P. oceanica meadows.

Threats:

As for most submerged plants, light is the key abiotic factor controlling P. oceanica productivity and spatial distribution. The levels of solar irradiance reaching a meadow controls both daily growth and seasonal growth, with levels being reduced by environmental factors such as water depth, turbidity, the state of surface ripples and waves. Temperature and salinity are also important abiotic factors controlling P. oceanica production.

Sedimentation/Erosion Balance

While very intense sedimentation processes may kill Posidonia plants, it is also particularly sensitive to an increase in the turbidity of sea water caused by waste and spillages, and is similarly affected by the persistent erosion of the sea bed which can rip out its roots and rhizomes. Increased sedimentation caused by storms, flooding, coastal soil erosion, dredging near seagrass meadows or coastal construction smother the plants irreversibly or create high concentrations of suspended sediments that reduce the light reaching the seagrasses.

In the Ligurian Sea it is estimated that nearly 30% of the original surface area of P.oceanica meadows was lost during the 1960s, a period of rapid urban and industrial coastal development. However, this decline also corresponds with a cold phase in global temperature trends within this decade making it difficult to distinguish between the effects of climate change and those of direct human pressure. The first results of a long-term monitoring programme in the Ligurian Sea reported that at three stations (5, 10 and 17 m depth, sampled each March, June, September and December by SCUBA diving) showed a similar pluriannual trend, consisting of a decrease in density and an increase in mean leaf length, suggesting a continued decline in the vitality of the meadow (Peirano et al 2001).

A study of P. oceanica meadows near a coastal construction site in the Ligurian Sea suggested that light reduction accounted for the reduced seagrass productivity and abundance, although there were also other complex interactions such as nutrient-epiphytes-grazers and water quality-siltation involved in seagrass mortality (Ruiz et al 2003).

Coastline transformation, with the proliferation of roads and houses and the regulation of continental river-flow, sharply reduces sediment inputs to the submersed coastal habitats, thereby promoting meadow erosion in their area of influence. Dredging and sand reclamation activities close to meadows have a high risk of direct meadow removal and may produce bed siltation or erosion.

Eutrophication

Eutrophication (nutrient loading) has been found to be responsible for the deterioration of seagrasses on a national and regional scale. Excess nutrients find their way into the sea from urban sewage outlets, industrial outlets, run off from agricultural areas, atmospheric deposition from agriculture and the burning of fossil fuels. Nitrogen and phosphorus are the most important nutrients for regulating the growth of planktonic algae, and hence water transparency and light conditions for seagrasses. Nutrients also stimulate the growth of opportunistic, fast-growing algae living on seagrass leaves, causing additional shading and further diminishing the plants' ability to photosynthesise. Studies over the past 10 years measuring P. oceanica cover and shoot density along the Mediterranean coasts have shown an overall decline. A further effect of eutrophication is the increased loading of organic materials such as dead algae to the sediment, which increases its O2 demand and may cause excessive hypoxia, which kills the seagrass roots.

Direct Erosion by Boat-trawling and Boat Anchoring

Otter trawling is one of the most important causes of large-scale degradation of P. oceanica meadows, particularly in deep meadows (e.g. Ardizzone and Pelusi 1984, Erftemeijer and Robin Lewis 2006). The repeated use of trawl gear over the seabed pulls up P. oceanica leaves and rhizomes (100,000 to 360,000 shoots per hour, Martín et al. 1997)), greatly reducing plant density and cover. As the trawl passes over the seabed, it also re-suspends the sediment and alters the substrate structure, increasing turbidity and nutrient concentrations in the water column. Reduced plant cover and the altered sediment interact to maintain silted conditions. The slow regrowth of P. oceanica further extends the impact of trawling which can sometimes run into decades (González-Correa et al. 2005).

In sites frequently visited by pleasure boats, there is significant removal of seagrasses by boat anchors (Francour et al. 1999). Also moorings consisting of a dead weight lowered to the seabed, attached to a partially crawling chain, form characteristic bare circles in P. oceanica meadows. These clearings persist for many years after the removal of the moorings. If the anchoring density and frequency are too high, the subsequent erosion may be accelerated by increased water movement caused by boating activities.

Expansion of Invasive Algal Species

The sustained increase in global marine transport favours the rapid expansion of exotic species that can harm existing communities (Galil 2007). In the Mediterranean Sea, around 100 exotic macrophytes have been introduced in the last few decades, of which at least 10 have an invasive nature (Ballesteros 2007). Those that most affect P. oceanica meadows are the green algae Caulerpa taxifolia and C. racemosa. Although these species do not apparently penetrate into dense healthy meadows, they may, when associated with other perturbations (e.g. eutrophication, bottom trawling), accelerate meadow decline, since they compete for space and light and increase the quantity of labile organic matter in the sediment.

Recently, the invasive red alga Lophocladia lallemandii has been shown to induce P. oceanica shoot mortality (Ballesteros 2007). L. lallemandii settles on rhizomes and old leaves along the edges of meadows and in low density patches where it grows rapidly, produce disc-like holdfasts along the thalli that enable the formation of a mat of red algal filaments intermingled with P. oceanica leaves. This mat can become so thick that leaves confined within may display chlorosis, and shoots eventually die.

Cosmopolitan filamentous algae may also behave as an invasive species, forming dense mucous layers on P. oceanica canopies in calm periods usually of 1 to 3 months, and, in so doing, reducing light availability to the seagrass (Lorenti et al. 2005). Some studies indicate that such episodes affect P. oceanica growth and survival, while others show no evident impacts. Their effect probably depends on their persistence and frequency as P. oceanica can resist shading for several months (Ruiz and Romero 2001). Finally, Acrothamnion preissii, a new exotic red algae that invades P. oceanica rhizomes has no apparent effects on seagrass integrity but it does displace most of the autochthonous rhizome epiphytes which reduces the meadow’s faunal species diversity and habitat complexity.

Salinity Increase in the Vicinity of Water Desalination Facilities

P. oceanica is especially sensitive to increases in salinity levels (Fernández-Torquemada and Sánchez-Lizaso 2005). Salt concentrations above 39 p.s.u. induce rapid plant death (Sabah et al. 2003). Thus, the brine (40-80 p.s.u.) from water desalination facilities, when poured directly onto P. oceanica meadows, can produce diebacks across large areas. Moreover, pipelines constructed to divert the brine to offshore areas destroy considerable meadow surfaces. The present and projected increase in coastal desalination facilities is therefore an emergent threat to P. oceanica meadows.

Fish Farm Activity

Since the onset of fish farm activity on the south-eastern coast of Spain, 11.29 ha of P. oceanica meadow has been completely lost and 9.86 ha significantly degraded, affecting about 53% of the former meadow area (Ruiz et al. 2001). The impact of fish farms on seagrasses seems to be highly variable and depends on complex interactions between a large number of processes such as decreased water transparency, increased dissolved nutrient and organic content of sediments in the vicinity of cages, and overgrazing by herbivores.

Climate Change

The potential effects of global climate change include increasing seawater CO2 levels and decreasing pH (ocean acidification). While this could affect P. oceanica photosynthesis and growth, insufficient data is available to predict the effects of CO2 and pH on seagrass meadows. The effects of global change are likely to accelerate the decline in already stressed seagrass meadows.

Trends

Most information available on the distribution of seagrass meadows is focused on the European Mediterranean coasts, while information along the Mediterranean African coasts is sparse. In a bibliographic review of 46 local or regional studies on 135 surveyed meadows, 20 meadows showed extreme regression (more than 50% of the area lost), 62 showed some degree of decline, 30 appeared stable and 23 experienced some degree of progression.

The following data has been extracted from the Natura 2000 Network database, elaborated by the European Commission with data updated on December 2006. The surface was estimated on the basis of the habitat cover indicated for each protected site and should be considered only as indicative of the habitat surface included in Natura 2000 (approximately 6 to 12% of the total habitat surface).

The slow recovery rates of P. oceanica necessitate the detection of trends in meadow dynamics before declines in seagrass cover and density become noticeably evident. Along the Spanish coasts, this was achieved by studying shoot demography, i.e. variation in shoot recruitment and mortality in meadows. The studies showed declining trends (mortality>recruitment) in 21 of 37 meadows and increasing trends (recruitment>mortality) in 7 meadows, while the remaining 9 meadows were in steady state. Among the meadows showing a declining trend, several are located far away from direct human influences. This finding suggests the existence of a background level of generalised P. oceanica declines possibly caused by global environmental factors, such as general deterioration of water transparency, or seawater temperature increase (both documented changes in Mediterranean waters).

In the National Park of Cabrera Island (Spain), an analysis was carried out by reconstruction of past and present growth, quantification of the demographic status of the established meadows, and quantification of patch formation and growth rates in areas where recolonization was occurring. Whilst regulation of mooring activities has improved the status of the P. oceanica meadows at Cabrera National Park, the process of recolonization of bare patches is still so slow that coalescence of patches and adjacent beds into a homogeneous meadow has been estimated to take more than 6 centuries. At a larger spatial scale in coastal areas of Mallorca (Spain) comparison of aerial photos from 1956 and 2001 shows that the 569 ha of meadow initially present had produced 28 ha of new meadow in the 45 years, representing a gain of 5%. However, P. oceanica losses in the area were higher than the gains, as 81 ha were lost in the same period (Marbà et al 2002).

Within the European seagrass flora, P. oceanica is the species with the lowest recolonization capacity. The slow rhizome elongation rate (a few cm per year) and the sparse flowering of this species, are conducive to a low recolonization capacity. Natural recovery of P. oceanica meadows is an extremely slow process, even following small-scale disturbances and small (m2) gaps within P. oceanica meadows can remain visible over several years.

For example, during the Second World War, in 1943, a bomb dropped and exploded within a dense meadow in the Rade de Villefranche (France): a circular area 80 m in diameter was completely destroyed, with a total area of 170m diameter altered. Forty years later the crater was still perfectly distinguishable, although surrounded by dense and apparently healthy meadows. Many small patches have colonised the zone at an average rate of 3 new patches per ha per year, and the surrounding meadow has migrated slightly from the borders into the centre of the crater. The estimated average linear growth was only 3.4 cm per year, however, which is half the potential horizontal growth of this species. The time necessary to completely recover this small area is estimated at 120 to 150 years (Meinesz and Lefèvre 1984), and large-scale recovery requires time scales of centuries (500 – 800 years, depending on patch formation rate).

Large-scale decline appears to be widespread, involving a number of factors, such as constructions along the shoreline (e.g. ports, wave breakers, etc.), enhanced organic and nutrient inputs from land and from aquaculture activities, coastal erosion and mechanical damage by trawling boats and anchors. The deterioration of sediment conditions due to enhanced organic inputs may slow recolonization even further, as rhizomes cannot extend into anoxic sediments.