Benthic primary producers--a neglected environmental problem in Mediterranean maricultures?

Marine fish farming is increasing rapidly in the Mediterranean and in contrast to the Atlantic the coastal zone in the Mediterranean is characterized by clear waters with high transparency. This allows benthic primary producers such as the slow-growing seagrass Posidonia oceanica to grow at large depths at locations suitable for fish farming and generating a conflict between the conservation of these meadows and the growth of aquaculture operations in the Mediterranean. In this paper we review the current knowledge on environmental interactions between fish farming and benthic primary producers with particular focus on P. oceanica, as this seagrass is a key component along Mediterranean coasts. The recovery times of P. oceanica are very long, in the order of centuries, and losses of this species are thus considered to be irreversible at managerial time scales.     

Benthic input rates predict seagrass (Posidonia oceanica) fish farm-induced decline

Fish farms represent a growing source of anthropogenic disturbance to benthic communities, and efficient predictors of such impacts are urgently needed. We explored the effects of fish farm benthic organic and nutrient inputs on the population dynamics of a key seagrass species (Posidonia oceanica) in four Mediterranean deep meadows adjacent to sea bream and sea bass farms. We performed two annual plant censuses on permanent plots at increasing distance from farms and measured benthic sedimentation rates around plots. High shoot mortality rates were recorded near the cages, up to 20 times greater than at control sites. Recruitment rates increased in variability but could not compensate mortality, leading to rapid seagrass decline within the first 100 m from cages. Seagrass mortality increased with total sedimentation rates (K=0.55, p<0.0002), and with organic matter (K=0.50, p=0.001), total nitrogen (K=0.46, p=0.002) and total phosphorus (K=0.56, p<3.10(-5)) inputs. P. oceanica decline accelerated above a phosphorus loading threshold of 50mg m(-2)day(-1). Phosphorus benthic sedimentation rate seems a powerful predictor of seagrass mortality from fish farming. Coupling direct measurements of benthic sedimentation rates with dynamics of key benthic species is proposed as an efficient strategy to predict fish farm impacts to benthic communities.     

Urban seagrass: status of Posidonia oceanica facing the Genoa city waterfront (Italy) and implications for management

A system of five adjacent Posidonia oceanica meadows facing the waterfront of Genoa city (Ligurian Sea, NW Mediterranean) was investigated over different spatial scales (meters-kilometers) using three environmental indices: conservation index (CI), substitution index (SI) and phase-shift index (PSI). CI revealed differences mostly at large spatial scale, distinguishing the poor condition of the meadows closest to Genoa centre and harbour from the comparatively healthy condition of the farthest meadows. SI showed differences mostly at small spatial scale (i.e., within meadows), suggesting the influence of local factors in the re-colonisation of regressed meadows by the seagrass Cymodocea nodosa and/or the invasive alga Caulerpa racemosa. Mapping of PSI showed that the meadows closest to Genoa centre and harbour have undergone a nearly total phase shift and have no real potential for recovery: attempts to re-establish P. oceanica there might be a waste of time and money. On the contrary, the meadows farthest from Genoa centre and harbour showed a comparatively low level of phase shift and could still fully recover given specific management actions.     

Effects of fish farm loadings on seagrass (Posidonia oceanica) distribution, growth and photosynthesis

The spatial extent and timing of the impact of fish farms on the distribution and performance of a Posidonia oceanica meadow were examined in an embayment of the south-eastern coast of Spain (Hornillo Bay, Murcia). Changes in seagrass distribution were determined using available seagrass mapping (from 1988, i.e., before the onset of aquaculture activities and 1998) and by successive sampling in 1994 and 1998. Environmental variables (light attenuation coefficient, water-column dissolved nutrients and organic content of sediments) together with plant performance (shoot biomass, leaf growth rate, photosynthetic activity, carbohydrate reserves, the number of leaves per shoot, epiphyte loads and herbivore pressure) were measured in plants affected by organic discharges, and were compared with those found in reference healthy plants over an annual growth cycle. Since the onset of fish farm activity, 11.29 ha of P. oceanica meadow has been completely lost and 9.86 ha significantly degraded, thus resulting in a total affected area which accounts for about 53% of the former meadow, or 7-fold the fish farming area. Unequal propagation of seagrass die-off or degradation reflects the relevance of local factors such as depth and hydrodynamism on the true extent of fish farm impact. Water transparency decreases and dissolved nutrient and organic content of sediments increases in the vicinity of cages compared to distant reference stations, thus supporting the notion of environmental gradients caused by the organic release from cages, which spreads outwards. Shoot size, leaf growth rate and the number of leaves per shoot in plants close to the fish farm decreased. Moreover, low leaf growth and low rhizome carbohydrate concentration (always relative to that found in an undisturbed area) indicated carbon budget imbalances. Since light reduction in the affected area was only modest (31% of light reaching the sea surface, while at the same depth this figure was 39% at the reference site), and light availability was well above the minimum requirement estimated for this species, neither this factor nor epiphyte overgrowth (epiphyte load was lower in the affected area) seem to explain such carbon imbalances or the observed meadow regression. Alternatively, the high herbivore pressure found in the affected zone suggests that overgrazing is one of the main causes of decreasing shoot sizes and hence of carbon imbalance, reduced growth and shoot mortality. The impact of fish farms on seagrasses, therefore, seems to be highly variable and depends on complex interactions between a large number of processes.     

Impacts of a fuel oil spill on seagrass meadows in a subtropical port, Gladstone, Australia--the value of long-term marine habitat monitoring in high risk areas

We used an established seagrass monitoring programme to examine the short and longer-term impacts of an oil spill event on intertidal seagrass meadows. Results for potentially impacted seagrass areas were compared with existing monitoring data and with control seagrass meadows located outside of the oil spill area. Seagrass meadows were not significantly affected by the oil spill. Declines in seagrass biomass and area 1month post-spill were consistent between control and impact meadows. Eight months post-spill, seagrass density and area increased to be within historical ranges. The declines in seagrass meadows were likely attributable to natural seasonal variation and a combination of climatic and anthropogenic impacts. The lack of impact from the oil spill was due to several mitigating factors rather than a lack of toxic effects to seagrasses. The study demonstrates the value of long-term monitoring of critical habitats in high risk areas to effectively assess impacts.     

Effects of climate, invasive species and anthropogenic impacts on the growth of the seagrass Posidonia oceanica (L.) Delile in Liguria (NW Mediterranean Sea)

Eight shallow water Posidonia oceanica meadows were sampled in June 1999 along 300 km of the Ligurian coast and were compared through shoot density and lepidochronology. The growth of the seagrass was examined in the light of climate fluctuations and local stresses, colonisation by alien, invasive alga Caulerpa taxifolia, and effects of the oil spill from the tanker "Haven", and other anthropogenic impacts. Both shoot density and lepidochronology pointed to a generalised state of regression of all the meadows. The analysis of long-term growth curves of the rhizomes showed a positive trend parallel to the increase of air temperature. Two main groups of meadows were individuated on the basis of growth curve similarity. The first included four meadows, namely Ventimiglia, Imperia, Noli and Prelo, that were characterised by average values of rhizome growth of 8-9.1 mmyear(-1) and shoot density greater than 200 shootsm(-2). Although the Imperia meadow was the only one where the alien invasive alga C. taxifolia was found, it did not show differences for rhizome growth in comparison to the other meadows. The second group was formed by meadows that had suffered past anthropogenic impacts: Arenzano and Monterosso al Mare. They showed higher rhizome growth rates (9.4-10.6 mmyear(-1)) and shoot densities between 200 and 100 shootsm(-2). At Arenzano, where "Haven" oil was stranded in April 1991, no rhizome older than 8 years was found, thus confirming the shoot mortality induced by the oil spill event. The two last meadows exhibited growth curves very different from all the others: Portovenere, is a shallow meadow where P. oceanica merely survives in an extremely degraded situation with highest rhizome growth rate (12 mmyear(-1)), the other, Riva Trigoso, is the only meadow implanted on rock and had the lowest growth rates (7.1 mm year(-1)).     

Productivity,carbon assimilation and intra-annual change in tropical reef platform seagrass communities of the Torres Strait,north-eastern Australia

Detailed data on seagrass distribution, abundance, growth rates and community structure information were collected at Orman Reefs in March 2004 to estimate the above-ground productivity and carbon assimilated by seagrass meadows. Seagrass meadows were re-examined in November 2004 for comparison at the seasonal extremes of seagrass abundance. Ten seagrass species were identified in the meadows on Orman Reefs. Extensive seagrass coverage was found in March (18,700 ha) and November (21,600 ha), with seagrass covering the majority of the intertidal reef-top areas and a large proportion of the subtidal areas examined. There were marked differences in seagrass above-ground biomass, distribution and species composition between the two surveys. Major changes between March and November included a substantial decline in biomass for intertidal meadows and an expansion in area of subtidal meadows. Changes were most likely a result of greater tidal exposure of intertidal meadows prior to November leading to desiccation and temperature-related stress.     

Impacts of a fuel oil spill on seagrass meadows in a subtropical port,Gladstone, Australia – The value of long-term marine habitat monitoring in high risk areas

We used an established seagrass monitoring programme to examine the short and longer-term impacts of an oil spill event on intertidal seagrass meadows. Results for potentially impacted seagrass areas were compared with existing monitoring data and with control seagrass meadows located outside of the oil spill area.Seagrass meadows were not significantly affected by the oil spill. Declines in seagrass biomass and area 1 month post-spill were consistent between control and impact meadows. Eight months post-spill, seagrass density and area increased to be within historical ranges. The declines in seagrass meadows were likely attributable to natural seasonal variation and a combination of climatic and anthropogenic impacts. The lack of impact from the oil spill was due to several mitigating factors rather than a lack of toxic effects to seagrasses. The study demonstrates the value of long-term monitoring of critical habitats in high risk areas to effectively assess impacts.     

Protecting the hand that feeds us: Seagrass (Zostera marina) serves as commercial juvenile fish habitat

Although fisheries are of major economic and food security importance we still know little about specific juvenile habitats that support such production. This is a major issue given the degradation to and lack of protection afforded to potential juvenile habitats such as seagrass meadows. In the present study we investigate the role of seagrass in supporting juvenile fish of commercial value. By assessing seagrass relative to adjacent sand we determined the presence of abundant juvenile fish. Nine commercial species were recorded and the most abundant of these were Plaice, Pollock and Herring. We provide the first quantitative evidence of the presence of juvenile fish of commercial value in seagrass surrounding Great Britain. Although the species that we found in seagrass as juveniles are not obligate seagrass users the resources that seagrass meadows offer to these fish provide significant long-term fitness benefits, potentially enhancing the whole population.     

Courage under fire: Seagrass persistence adjacent to a highly urbanised city–state

Due to increasing development Southeast Asia’s coastlines are undergoing massive changes, but the associated impacts on marine habitats are poorly known. Singapore, a densely populated island city–state, is a quintessential example of coastal modification that has resulted in the (hitherto undocumented) loss of seagrass. We reconstructed the historic extent and diversity of local seagrass meadows through herbarium records and backwards extrapolation from contemporary seagrass locations. We also determined the current status of seagrass meadows using long-term monitoring data and identified the main threats to their presence in Singapore. Results show that, even though ∼45% of seagrass has been lost during the last five decades, species diversity remains stable. The main cause of seagrass loss was, and continues to be, land reclamation. We conclude that strict controls on terrestrial runoff and pollution have made it possible for seagrass to persist adjacent to this highly urbanised city–state.     

Seagrass population dynamics and water quality in the Great Barrier Reef region: a review and future research directions

Seagrasses in the Great Barrier Reef region, particularly in coastal habitats, act as a buffer between catchment inputs and reef communities and are important habitat for fisheries and a food source for dugong and green turtle. Within the Great Barrier Reef region there are four different seagrass habitat types now recognised. The spatial and temporal dynamics of the different types of seagrass habitat is poorly understood. In general seagrass growth is limited by light, disturbance and nutrient supply, and changes to any or all of these limiting factors may cause seagrass decline. The capacity of seagrasses to recover requires either recruitment via seeds or through vegetative growth. The ability of seagrass meadows to recover from large scale loss of seagrass cover observed during major events such as cyclones or due to anthropogenic disturbances such as dredging will usually require regeneration from seed bank. Limited research into the role of pollutants on seagrass survival suggests there may be ongoing impacts due to herbicides, pesticides and other chemical contaminants. Further research and monitoring of seagrass meadow dynamics and the influence of changing water quality on these is needed to enhance our ability to manage seagrasses on the Great Barrier Reef.     

Early warning indicators for river nutrient and sediment loads in tropical seagrass beds: a benchmark from a near-pristine archipelago in Indonesia

In remote, tropical areas human influences increase, potentially threatening pristine seagrass systems. We aim (i) to provide a bench-mark for a near-pristine seagrass system in an archipelago in East Kalimantan, by quantifying a large spectrum of abiotic and biotic properties in seagrass meadows and (ii) to identify early warning indicators for river sediment and nutrient loading, by comparing the seagrass meadow properties over a gradient with varying river influence. Abiotic properties of water column, pore water and sediment were less suitable indicators for increased sediment and nutrient loading than seagrass properties. Seagrass meadows strongly responded to higher sediment and nutrient loads and proximity to the coast by decreasing seagrass cover, standing stock, number of seagrass species, changing species composition and shifts in tissue contents. Our study confirms that nutrient loads are more important than water nutrient concentrations. We identify seagrass system variables that are suitable indicators for sediment and nutrient loading, also in rapid survey scenarios with once-only measurements.     

Sediment characteristics and macrofauna distribution along a human-modified inlet in the Gulf of Oristano (Sardinia, Italy)

We studied the spatial variability and within-year temporal changes in hydrological features, grain size composition and chemical characteristics of sediments, as well as macrofaunal assemblages, along a heavily modified inlet in the Gulf of Oristano (western Sardinia, Italy). The inlet connects the Cabras lagoon to the gulf through a series of convoluted creeks and man-made structures, including a dam and fish barriers built in the last three decades. Sediments were muddy and mainly composed of the "non-sortable" fraction (i.e., <8 microm particle size) in all four areas investigated: Lagoon, Creeks, Channel and Seaward. Along the inlet, however, the ratio between the <8 microm and the 8-64 microm fractions was highest in Creeks and Channel, between the fish barriers and the dam, suggesting impaired hydrodynamics. Consistently, steep gradients in water salinity, temperature and dissolved oxygen concentrations were found in proximity to the fish barriers. The whole inlet was characterized by a major organic enrichment of sediments, with up to an annual mean of 33.6% of organic matter and 11.7% of total organic carbon in Seaward due to the presence of seagrass leaf litter. Acid-volatile sulphide and chromium-reduced sulphur concentrations were highest throughout the year in Seaward and Lagoon, respectively, with a peak in summer. Consistently, the whole inlet supported low structured macrofaunal assemblages dominated by few opportunist species, with a relatively lower diversity in Lagoon throughout the year and the highest abundances in Seaward in summer. We infer that the presence of artificial structures along the inlet, such as fish barriers and the dam, impair the lagoon-gulf hydrodynamics, sediment exchange and animal recruitment and colonization. We suggest that the removal of these structures would favour water renewal in the Cabras lagoon, but would also increase the outflow of organic C-bonding fine particles into the gulf with serious consequences for Posidonia oceanica and Cymodocea nodosa seagrass meadows. We conclude that all possible consequences of such initiatives should be carefully considered before any action is taken.     

Legal protection is not enough: Posidoniaoceanica meadows in marine protected areas are not healthier than those in unprotected areas of the northwest Mediterranean Sea

Using the Conservation Index, which measures the proportional amount of dead matte relative to live Posidonia oceanica, we assessed the health of 15 P. oceanica meadows at a regional scale along the coast of Liguria (NW Mediterranean). These areas were characterized by different degrees of anthropization, from highly urbanized sites to marine protected areas. Two different scenarios were identified according to depth: in shallow zones, the health of P. oceanica meadows was related to the degree of anthropization along the coastline. In contrast, in deep zones, most meadows exhibited poor health, independent of both the degree of disturbance and the legal measures protecting the area. Working synergistically with the regional impact of increased water turbidity, local impacts from the coast were recognized as the main causes of the severe regression of most Ligurian P. oceanica meadows. We conclude that marine protected areas alone are not sufficient to guarantee the protection of P. oceanica meadows. We emphasize the need for a management network involving the Sites of Community Interest (SCIs) containing P. oceanica meadows.     

Biogeomorphology of the Mediterranean Posidonia oceanica seagrass meadows

Here we review the multiple interactions between the endemic Mediterranean seagrass, Posidonia oceanica, and coastal geomorphologic processes as an outstanding example of biogeomorphology, taking into account recent advances in the field. Seagrass meadows are among the most important elements for the functioning of marine coastal ecosystems, and represent a major focus for research and conservation. Being considered a priority habitat, P. oceanica meadows are protected by several European Union directives and national laws. In this paper we examine: the role of sedimentary features in controlling the development of the meadows; the interplay between P. oceanica leaf litter (i.e. beached necromass) cast ashore and erosional‐depositional processes on the beaches; the interactions between meadows and nearshore hydrodynamics, and; possible linkages between geomorphological features of the seafloor and the architecture of meadows. Finally, we provide perspectives for future research on P. oceanica and other Mediterranean seagrass meadows in a biogeomorphological context with specific reference to climate change. Copyright © 2016 John Wiley & Sons, Ltd.     

Water quality in the Great Barrier Reef region: responses of mangrove, seagrass and macroalgal communities

Marine plants colonise several interconnected ecosystems in the Great Barrier Reef region including tidal wetlands, seagrass meadows and coral reefs. Water quality in some coastal areas is declining from human activities. Losses of mangrove and other tidal wetland communities are mostly the result of reclamation for coastal development of estuaries, e.g. for residential use, port infrastructure or marina development, and result in river bank destabilisation, deterioration of water clarity and loss of key coastal marine habitat. Coastal seagrass meadows are characterized by small ephemeral species. They are disturbed by increased turbidity after extreme flood events, but generally recover. There is no evidence of an overall seagrass decline or expansion. High nutrient and substrate availability and low grazing pressure on nearshore reefs have lead to changed benthic communities with high macroalgal abundance. Conservation and management of GBR macrophytes and their ecosystems is hampered by scarce ecological knowledge across macrophyte community types.     

Contrasting recovery of shallow and deep water seagrass communities following climate associated losses in tropical north Queensland,Australia

Tropical seagrass decline and recovery from severe storm impacts was assessed via quarterly measurements of seagrass biomass, species composition and experimental investigations of recovery in north Queensland. Shallow and deep seagrass meadows suffered major declines. Significant recovery in the two years following loss only occurred at deeper sites. Halophila spp. in deep water areas had a high capacity for recovery through the availability of seed banks. In contrast, the shallow species did not recover quickly from experimental disturbance, had poor seed reserves and relied on asexual propagation. The potential for shallow species to recover rapidly from widespread losses was limited as seed banks were limited or non-existent. Understanding inter- and intra-specific differences in seagrass recovery and how this interacts with location is critical to predict the consequences of climate events to tropical seagrasses. This is especially important as more frequent severe storms are predicted as a consequence of climate change.