Transplantation as a method for restoring the seagrass Posidonia australis

Transplant trials of the seagrass Posidonia australis were carried out after loss of seagrasses following eutrophication and increased turbidity in two marine inlets on the south coast of Western Australia. A pilot study in Oyster Harbour measured survival and growth in situ for 4 years. Long-term survival rates were high (96–98%), providing plants were anchored into the sediment. All unanchored plants were lost in the first winter. Following the success of the pilot study, a more comprehensive program began 3 years later with over 500 transplant units collected from either actively growing edges of nearby patches (plagiotropic growth form) or within established meadows (orthotropic growth form). Transplant units from edges expanded at a faster rate compared to units from mid-meadow but increases in shoot numbers were similar. Growth rates in the first 2.5 years averaged 10–20 cm yr⁻¹ horizontal rhizome extension, depending on the source of the transplant units, and 4–12 shoots per initial shoot yr⁻¹, depending on the initial shoot number of the transplant unit. After 5 years, shoot numbers of individual transplants were similar to shoot densities recorded for natural meadows, >500 shoots m⁻². Approximately, 10% of transplants from mid-meadow flowered in the first year, whereas transplants from edges flowered only after 5 years.     

Architectural characteristics of two bed types of the seagrass Posidonia oceanica over different spatial scales

Seagrass beds occur in various morphological forms, ranging from small patches to continuous meadows. The endemic Mediterranean seagrass Posidonia oceanica forms dense and extensive stands that occur in several different morphotypes, including reticulate (seagrass interspersed with a different habitat type, such as bare sand) and continuous beds. This study, undertaken in the Maltese Islands, examined whether reticulate and continuous P. oceanica beds, located adjacent to each other and at similar depths, had different within-bed architectural characteristics. Five commonly used architectural measures (shoot density, number of leaves per shoot, mean leaf length, mean leaf width and shoot biomass) were measured from P. oceanica shoots collected from the two bed types at three different spatial scales: (1) tens of metres (‘small’ scale); (2) hundreds of metres (‘medium’ scale); and (3) kilometres (‘large’ scale). Results of 2-factor ANOVA (factor 1=bed type; factor 2=sampling locality) carried out at the three spatial scales indicated significant differences between the two bed types in shoot density (P<0.01) and leaf length (P<0.05) at the small scale, and in leaf number (P<0.05) at the large scale. Significant interactions were also apparent for shoot density (at the large scale) and for shoot biomass (at the medium scale). However, the results obtained did not indicate consistent architectural differences between the two P. oceanica bed types over the spatial scales considered. Spatial variations in within-bed architectural characteristics observed were therefore thought to be attributable mainly to the influence of local environmental factors. The findings are discussed with reference to the conservation and management of P. oceanica habitat.     

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.     

Photosynthesis,growth and survival of the Mediterranean seagrass Posidonia oceanica in response to simulated salinity increases in a laboratory mesocosm system

This study aims to examine the effect of increased salinity on the photosynthetic activity of the Mediterranean seagrass Posidonia oceanica in a laboratory mesocosm system. To do this, large rhizome fragments were transplanted in a mesocosm laboratory system and maintained at 37 (ambient salinity, control treatment), 39, 41 and 43 (hypersaline treatments) for 47 days. Pigment content, light absorption, photosynthetic characteristics (derived from P vs. E curves and fluorescence parameters), and shoot size, growth rates and net shoot change were determined at the end of the experimental period. Both net and gross photosynthetic rates of plants under hypersaline conditions were significantly reduced, with rates some 25–33% and 13–20% lower than in control plants. The pigment content (Chla, Chlb, Chlb:Chla molar ratio, total carotenoids and carotenoids:Chla ratio), leaf absorptance and maximum quantum yield of PSII (F_v/F_m) of control plants showed little or no changes under hypersaline conditions, which suggests that alterations to the capacity of the photosynthetic apparatus to capture and process light were not responsible for the reduced photosynthetic rates. In contrast, dark respiration rates increased substantially, with mean values up to 98% higher than in control leaves. These results suggest that the respiratory demands of the osmoregulatory process are likely to be responsible for the observed decrease in photosynthetic rates, although alterations to photosynthetic carbon assimilation and reduction could also be involved. As a consequence, leaf carbon balance was considerably impaired and leaf growth rates decreased as salinity increased above the ambient (control) salinity. No significant differences were found in the percentage of net shoot change, but mean values were clearly negative at salinity levels of 41 and 43. Results presented here indicate that photosynthesis of P. oceanica is highly sensitive to hypersaline stress and that it likely account for the decline in leaf growth and shoot survival reported in this and previous studies in response to even small increments of the ambient salinity.     

Composition of epiphytic leaf community of Posidonia oceanica as a tool for environmental biomonitoring

The demand for sensitive biological tools to assess the environmental quality of coastal waters at broad spatial scales is increasing. Many of the tools used are based on the taxonomic composition of biotic assemblages. They usually require a valuable taxonomic expertise while are unique reflecting the overall ecosystem integrity. Here, we evaluate the potential indicator value of several features of the epiphytic community (overall assemblage composition, species richness, and proportion of the main taxonomic groups) growing on the seagrass Posidonia oceanica leaves. We do so by empirically examining their changes along a disturbance gradient where multiple human activities have interactive and cumulative impacts, sampling at different spatial scales and at two different depths (5 and 15 m). Our results show that the specific composition of the epiphytic assemblages (i.e. species composition) closely reflects, in the deep meadows, the combined effects of different anthropogenic stressors along the gradient, showing an integrative and non-specific response. Similarly, an increase in the proportion of hydrozoans, and a decrease in the proportion of rhodophytes and chlorophytes are observed in deep meadows along the gradient. In shallow meadows, grazing and biotic features of the seagrass seem the main forcing factors determining species composition, and therefore masking the response of epiphytes to the deterioration gradient. After address the effect of natural sources of variability (water depth, within- and between-meadow heterogeneity), changes in epiphyte assemblages and in the proportion of hydrozoans, rhodophytes and chlorophytes in relatively deep meadows seem promising monitoring tools for detecting coastal environmental deterioration.     

Use of a high-resolution profiling sonar and a towed video camera to map a Zostera marina bed,Solent, UK

Seagrasses are flowering plants that develop into extensive underwater meadows and play a key role in the coastal ecosystem. In the last few years, several techniques have been developed to map and monitor seagrass beds in order to protect them. Here, we present the results of a survey using a profiling sonar, the Sediment Imager Sonar (SIS) and a towed video sledge to study a Zostera marina bed in the Solent, southern UK. The survey aimed to test the instruments for seagrass detection and to describe the area for the first time.     

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.     

Much damage for little advantage: Field studies and morphodynamic modelling highlight the environmental impact of an apparently minor coastal mismanagement

While coastal management activities have long been known to exert a strong influence on the health of marine ecosystems, neither scientists nor administrators have realized that small interventions may lead to disproportionately larger impacts. This study investigated the broad and long-lasting environmental consequences of the construction of an ill-planned, although small (only 12 m long) jetty for pleasure crafts on the hydrodynamic conditions and on the meadow of the seagrass Posidonia oceanica of an embayed cove in the Ligurian Sea (NW Mediterranean). There, P. oceanica used to develop on a high (>1.5 m) matte (a lignified terrace causing seafloor elevation) in which the leaves reach the surface and form a compact natural barrier to waves in front of the beach. Such a so-called ‘fringing reef’ of P. oceanica is today recognized of high ecological value and specific conservation efforts are required. The construction of the jetty implied the cutting of the matte, which directly destroyed part of the fringing reef. In addition, meadow mapping and sedimentological analyses coupled with morphodynamic modelling showed that the ecosystem of the whole cove had been greatly altered by the jetty. We used the geometric planform approach, a proper tool in the study of headland-controlled embayment, both to characterise the present situation of Prelo cove and to simulate the original one, before the jetty was built. In the long term, such a small jetty completely altered the configuration and the hydrodynamic conditions of the whole cove, splitting the original pocket beach into two smaller ones and creating strong rip-currents flowing seaward along the jetty. These rip-currents enhanced erosion of residual shallow portions of the meadow and further modified the sedimentary fluxes in shallow waters. A century after the construction of the jetty, an irreversible environmental damage has occurred, as the slow growing rate of P. oceanica implies that the high matte terrace and the fringing reef will hardly form again, even after the removal of the jetty. The lesson learnt from this study is that even such small, and therefore reputed intrinsically ‘innocent’, interventions on the coastal zone require accurate planning based on interdisciplinary studies to understand and respect the delicate interplay among morphological, hydrodynamic and ecological components.     

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.     

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.     

SeagrassNet monitoring across the Americas: case studies of seagrass decline

Seagrasses are an important coastal habitat worldwide and are indicative of environmental health at the critical land–sea interface. In many parts of the world, seagrasses are not well known, although they provide crucial functions and values to the world's oceans and to human populations dwelling along the coast. Established in 2001, SeagrassNet, a monitoring program for seagrasses worldwide, uses a standardized protocol for detecting change in seagrass habitat to capture both seagrass parameters and environmental variables. SeagrassNet is designed to statistically detect change over a relatively short time frame (1–2 years) through quarterly monitoring of permanent plots. Currently, SeagrassNet operates in 18 countries at 48 sites; at each site, a permanent transect is established and a team of people from the area collects data which is sent to the SeagrassNet database for analysis. We present five case studies based on SeagrassNet data from across the Americas (two sites in the USA, one in Belize, and two in Brazil) which have a common theme of seagrass decline; the study represents a first latitudinal comparison across a hemisphere using a common methodology. In two cases, rapid loss of seagrass was related to eutrophication, in two cases losses related to climate change, and in one case, the loss is attributed to a complex trophic interaction resulting from the presence of a marine protected area. SeagrassNet results provide documentation of seagrass change over time and allow us to make scientifically supported statements about the status of seagrass habitat and the extent of need for management action.     

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.     

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.     

Impact of the brine from a desalination plant on a shallow seagrass (Posidonia oceanica) meadow

Although seawater desalination has increased significantly over recent decades, little attention has been paid to the impact of the main by-product (hypersaline water: brine) on ecosystems. In the Mediterranean, potentially the most affected ecosystems are meadows of the endemic seagrass Posidonia oceanica. We studied the effect of brine on a shallow P. oceanica meadow exposed to reverse osmosis brine discharge for more than 6 years. P. oceanica proved to be very sensitive to both eutrophication and high salinities derived from the brine discharge. Affected plants showed high epiphyte load and nitrogen content in the leaves, high frequencies of necrosis marks, low total non-structural carbohydrates and low glutamine synthetase activity, compared to control plants. However, there was no indication of extensive decline of the affected meadow. This is probably due to its very shallow situation, which results in high incident radiation as well as fast dilution and dispersion of the brine plume.     

Evaluating the effects of protection on Paracentrotus lividus distribution in two contrasting habitats

The sea urchin Paracentrotus lividus is common in the Mediterranean in shallow subtidal rocky habitats and in Posidonia oceanica beds. The aim of this study is to investigate whether protection has the same effect on the population structure of P. lividus occurring in rocky reef habitats and in P. oceanica beds. These results are important to generate hypotheses about the influence of human harvesting, predatory pressure and migration processes on P. lividus in the two habitats.     

First evidence of a massive recruitment event in Posidonia oceanica: Spatial variation in first-year seedling abundance on a heterogeneous substrate

A recruitment event of Posidonia oceanica was recorded in the Gulf of Ventilegne (Corsica, Western Mediterranean) in June 2005. One-year seedlings (alive and dead) were found at shallow depth (2–3 m) on a mix of rock and sand substrates. Pattern in the distribution of seedlings was examined over spatial scales ranging from centimetres to hundreds of metres. We also examined (a) whether establishment was preferentially associated with a specific substrate type; (b) the relation between proportional mortality and the density of established seedlings; and (c) the growth characteristics of seedlings. The average density of seedlings varied from 19 to 189 seedlings m⁻², with a maximum of 480 seedlings m⁻². While significant variation was detected at the scale of area (i.e., tens of metres), no difference was found in abundance of seedlings at the scale of sites (i.e., hundreds of metres). Most of the overall variability was, however, at on the smallest scale examined (i.e., among sampling units tens of centimetres apart). No differences were also observed in the establishment probability and seedling abundance between sand and rock substrates. The fine scale analysis of spatial patterns on rock and sand substrates showed a clumped distribution of seedlings. These results indicated that the spatial distribution of seedlings was not random, but organized at small spatial scales within the study sites. Hydrodynamics and micro-scale bottom features appeared to account for such patterning. Proportional seedling mortality was highly variable (ranging from 0.3 to 0.8), and tended to increase with increasing settlement density. A total of 128,000 seedlings were recoded over the 1600 m² surface area investigated. Such event reflected conspicuous seed supply and particularly favourable local conditions. This finding contradicts the current statement of low recruitment ability of P. oceanica and indicates that the species can recruit both on consolidated and unconsolidated substrates, at least in the study area.     

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.     

Fatty acid profiles indicate the habitat of mud snails Hydrobia ulvae within the same estuary: Mudflats vs. seagrass meadows

Mud snails Hydrobia ulvae occupy different habitats in complex estuarine ecosystems. In order to determine if fatty acid profiles displayed by mud snails can be used to identify the habitat that they occupy within the same estuary, fatty acids of H. ulvae from one mudflat and one seagrass meadow in the Ria de Aveiro (Portugal) were analyzed and compared to those displayed by microphytobenthos (MPB), the green leaves (epiphyte-free) of Zostera noltii, as well as those exhibited by the epiphytic community colonizing this seagrass. MPB and epiphytic diatom-dominated samples displayed characteristic fatty acids, such as 16:1n-7 and 20:5n-3, while 18:2n-6 and 18:3n-3 were the dominant fatty acids in the green leaves of Z. noltii. Significant differences between the fatty acid profiles of H. ulvae specimens sampled in the mudflat and the seagrass meadow could be identified, with those from the mudflat displaying higher levels of fatty acids known to be characteristic of MPB. This result points towards the well known existence of grazing activity on MPB by mud snails. The fatty acid profiles displayed by H. ulvae inhabiting the seagrass meadows show no evidence of direct bioaccumulation of the two most abundant polyunsaturated fatty acids of Z. noltii (18:2n-6 and 18:3n-3) in the mud snails, which probably indicates that either these compounds can be metabolized to produce energy, used as precursors for the synthesis of essential fatty acids, or that the snails do not consume seagrass leaves at all. Moreover, the fatty acid profiles of mud snails inhabiting the seagrass meadows revealed the existence of substantial inputs from microalgae, suggesting that the epiphytic community colonizing the leaves of Z. noltii displays an important role on the diet of these organisms. This assumption is supported by the high levels of 20:5n-3 and 22:6n-3 recorded in mud snails sampled from seagrass meadows. In conclusion, fatty acid analyses of H. ulvae can be successfully used to identify the habitat occupied by these organisms within the same estuary (e.g. mudflats and seagrass meadows) and reveal the existence of contrasting dietary regimes.