Identifying keystone trophic groups in benthic ecosystems: Implications for fisheries management

Many species inhabiting the benthic marine ecosystems of the central and northern Chilean coast have been intensively harvested and this exploitation has increased considerably in recent years. Despite this harvest pressure, few studies have attempted to establish a more holistic, systems-based management plan. On the contrary, research continues to rely on population models in which the species of interest are isolated from their ecological context. This work offers several keystone indices in order to help multispecies fisheries management. The indices used are: (1) functional indices based on steady-state and dynamic trophic models; (2) structural indices based on bottom-up and top-down control mechanisms; and (3) qualitative keystone species indices using loop models (mixed control). The quantitative trophic models were constructed using Ecopath with Ecosim (EwE; v. 5.0) software, and the qualitative model was analysed using Loop Analysis. All models describe the interactions of the most representative species and functional groups inhabiting the benthic ecosystems of Tongoy Bay, La Rinconada Marine Reserve (Antofagasta Bay), and the kelp forest of Mejillones Peninsula (Antofagasta). Even though our results only represent the short-term dynamics of these systems, we have found keystoneness properties of several species and functional groups, including primary producers, herbivores, and top predators. Despite this wide variability of groups, we detected a different core set of species or functional groups, each of which contained prey–predator and plant–herbivore relationships. Because the traditional keystone concept of a single species is difficult to apply, we suggest shifting away from this view towards a more holistic alternative such as that of a keystone species complex. This kind of approach would facilitate the design and assessment of sustainable management strategies for ecological marine ecosystems. Despite the ecological relevance of our results, further experimental studies and modelling using other theoretical frameworks should be performed.     

Assessment of Top-Down and Bottom-Up Controls on the Collapse of Alewives (<Emphasis Type="Italic">Alosa pseudoharengus</Emphasis>) in Lake Huron

Food web models are powerful tools to inform management of lake ecosystems, where top-down (predation) and bottom-up (resource) controls likely propagate through multiple trophic levels because of strong predator–prey links. We used the Ecopath with Ecosim modeling approach to assess these controls on the Lake Huron main basin food web and the 2003 collapse of an invasive pelagic prey fish, alewife (Alosa pseudoharengus). We parameterized two Ecopath models to characterize food web changes occurring between two study periods of 1981–1985 and 1998–2002. We also built an Ecosim model and simulated food web time-dynamics under scenarios representing different levels of top-down control by Chinook salmon (Oncorhynchus tshawytscha) and of bottom-up control by quagga mussels (Dreissena rostriformis bugensis) and nutrients. Ecopath results showed an increase in the relative importance of bottom-up controls between the two periods, as production decreased across all trophic levels. The production of non-dreissenid benthos decreased most, which could cause decreases in production of pelagic prey fishes feeding on them. Ecosim simulation results indicated that the alewife collapse was caused by a combination of top-down and bottom-up controls. Results showed that while controls by Chinook salmon were relatively constant before alewife collapse, controls by quagga mussels and nutrients increased jointly to unsustainable levels. Under current conditions of low nutrients and high quagga mussel biomass, simulation results showed that recovery of alewives is unlikely regardless of Chinook salmon biomass in Lake Huron, which implies that the shrinking prey base cannot support the same level of salmonine predators as that prevailed during the 1980s.     

The relative importance of food and shelter for seagrass-associated invertebrates: a latitudinal comparison of habitat choice by isopod grazers

The generality of mechanisms affecting habitat choice and grazing in seagrass meadows was evaluated in a latitudinal comparison of seagrass grazers from the temperate (60°N) Baltic Sea and the subtropical (30°N) Gulf of Mexico. Using similar habitat choice experiment set-ups in Finland and the USA, the role of food type, habitat complexity and predation hazard on habitat choice of the isopods Idotea baltica (Pallas) and Erichsonella attenuata Harger were tested. When shelter was provided by both living and artificial seagrass, epiphytic food resources on artificial vegetation were clearly preferred by both species, although Idotea was attracted to epiphyte-free seagrass when no alternative food was present. When choosing between food and shelter, both species preferred epiphytic food over shelter. However, under predation hazard of fish, Erichsonella clearly switched to the habitat offering shelter, while the presence of a predatory fish produced no preference for shelter by Idotea. Food type may be considered as an universal mechanism that partly determines the presence of grazers in seagrass habitats and is, in the absence of a predator, more important than shelter. Predation risk affected the behaviour of the grazers, but the response varied between species possibly due to varying importance of fish predation in the areas studied. Received: 16 November 1998 / Accepted: 13 February 1999     

Changes in the trophic interactions and the community structure of Lake Taihu (China) ecosystem from the 1960s to 1990s

As the third largest freshwater lake in China, Lake Taihu has suffered from overfishing, eutrophication, and physical disturbance over the last several decades. Evaluating and quantifying changes in the ecosystem can help us better understand and develop hypotheses to explain the dynamics of the ecosystem. In this study, trophic interactions and community structure of commercial fisheries species of Lake Taihu ecosystem were analyzed and compared for three time periods (1961–1965, 1981–1987, and 1991–1995) using the Ecopath with Ecosim model with the aim of evaluating the changes in the population dynamics and ecosystem development mechanism spanning the period from the 1960s to 1990s. The results show that the biomass of large predators decreased over the three decades, while the biomass of small species increased. Increases in the P/B ratios and fishing mortality levels observed for species groups reveal rapidly intensifying fishery stress over the three decades. The fisheries operated at the highest trophic level during the 1980s, and there are some indications of “fishing down the food web” in this ecosystem between the 1980s and the 1990s. Drawing upon Odum’s theory of ecosystem maturity, the structured, web-like ecosystem of the 1960s developed into a highly mature system during the 1980s; yet, in the 1990s, this structure became less complex and the system’s maturity fell to its lowest observed level. During this period, the successional development of the system occurred in reverse.     

Importance of epiphytic cyanobacteria as food sources for heterotrophs in a tropical seagrass bed

 The natural carbon and nitrogen stable isotope ratios (δ¹³C, δ¹⁵N) of various autotrophs and heterotrophs were measured in a Syringodium isoetifolium-dominated seagrass bed at Dravuni Island, Fiji to define carbon and nitrogen sources for heterotrophic organisms in a system where few animals graze directly on seagrass leaves. The organic carbon, nitrogen, and phosphorus content of organisms was also determined. The δ¹³C and δ¹⁵N data suggest that herbivorous heterotrophs in this seagrass bed depend significantly on epiphytic cyanobacteria rather than seagrass leaves and its detritus. This can be attributed to relative differences in nitrogen content of those organic materials. The cyanobacteria nitrogen content (3.6–4.8% of DW) is nearly half that of heterotrophs (7.0–8.6% N of DW) while that of S. isoetifolium origin (0.6–1.1% N of DW) is less than one third of the cyanobacteria nitrogen content. Phosphorus content was similar among cyanobacteria (0.8–1.1 mg g^-1) and S. isoetifolium (0.4–1.4 mg g^-1). These results suggest that cyanobacteria are important food sources for heterotrophs at the study site, and that inorganic nitrogen released through breakdown of cyanobacteria by heterotrophs may support the continued production of S. isoetifolium. Accepted: 24 May 1999     

Habitat preference of three common fishes for seagrass, <Emphasis Type="Italic">Caulerpa taxifolia</Emphasis>, and unvegetated substrate in Moreton Bay,Australia

A decrease in seagrass cover and a commensurate increase in Caulerpa taxifolia distribution in Moreton Bay have prompted concern for the impact that habitat change may have on faunal communities. Therefore, it is important to understand the patterns of habitat use. We examined habitat selection of three common seagrass species: double-ended pipefish (Syngnathoides biaculeatus), eastern trumpeter (Pelates quadrilineatus), and fan-bellied leatherjacket (Monacanthus chinensis) using a mesocosm experiment. Fish were given three possible habitat pairings (1) seagrass and C. taxifolia, (2) seagrass and unvegetated, and (3) C. taxifolia and unvegetated. Observation trials were conducted during the day and night over two days. In all trials, fish preferred vegetated habitat (seagrass or C. taxifolia) over unvegetated habitat (sand). In seagrass and C. taxifolia trials, all species preferred seagrass significantly over C. taxifolia. Habitat use patterns did not differ between day and night trials. Caulerpa taxifolia provides a valuable structured habitat in the absence of seagrass; however, it is unclear if C. taxifolia meadows provide other resource benefits to fishes beyond that of shelter.     

Estimating the carrying capacity of French Frigate Shoals for the endangered Hawaiian monk seal using Ecopath with Ecosim

The carrying capacity of the French Frigate Shoals (FFS) region for the endangered Hawaiian monk seal was appraised using an updated version of the original FFS Ecopath model ( Polovina 1984 ). Model parameters were updated using recent literature, and data from surveys of the seal population and its bottom‐associated prey. Together they produced a static mass balance model for 1998 when the prey surveys began. The Ecopath‐estimated monk seal biomass was 0.0045 t/km², which was in close agreement with the biomass calculated from monk seal field beach counts (0.0046 t/km²). Model simulations through time were done in Ecosim using the Ecopath balanced model and included fisheries data time series from 1998 to 2008. Monk seal biomass declined concurrently with decreases in benthic bottomfish biomass, which were influenced by large‐scale changes in the environment of the North Pacific. This model scenario was extended from 2010, when the last permitted fishery in the Northwestern Hawaiian Islands was closed, through to 2040, assuming a constant environmental signal. Model results for this time period did not show a recovery of monk seals that exceeded the initial 1998 model biomass levels, highlighting the importance of including environmental variability in estimates of monk seals recovery at FFS.     

Wasting disease regulates long-term population dynamics in a threatened seagrass

The role of disease in the long-term dynamics of threatened species is poorly quantified, as well as being under-represented in ecology and conservation management. To understand persistent host–pathogen interaction operating in a vulnerable habitat, we quantified dynamics driving patterns of seagrass density using a longitudinal study in a relatively pristine site (Isles of Scilly, UK). Replicated samples of eelgrass (Zostera marina) density and wasting disease prevalence, presumably caused by Labyrinthula zosterae, were taken from five meadows at the height of the growing season, over the years 1997–2010. Data were used to parameterise a population dynamic model, incorporating density-dependent factors and sea temperature records. We found that direct density and disease-mediated feedback operate within a network of local populations. Furthermore, our results indicate that the strength of limitation to seagrass growth by disease was increased at higher temperatures. This modification of the coupled host–pathogen dynamics forms a novel hypothesis to account for dramatic die-backs of Z. marina widely reported elsewhere. Our findings highlight the importance of disease in structuring distributions of vulnerable species, as well as the application of population modelling in order to reveal ecological processes and prioritise future mechanistic investigation.     

Simple measures of channel habitat complexity predict transient hydraulic storage in streams

Stream thalweg depth profiles (along path of greatest channel depth) and woody debris tallies have recently become components of routine field procedures for quantifying physical habitat in national stream monitoring efforts. Mean residual depth, standard deviation of thalweg depth, and large woody debris (LWD) volumes are potential metrics of habitat complexity calculated from these survey data. We used 42 intensive dye-transit studies to demonstrate the relevance of these easily measured channel habitat complexity metrics to transient hydraulic (“dead zone”) storage, a channel process important for biotic habitat as well as retention and “spiraling” of dissolved and particulate nutrients. We examined transient storage and channel morphology in small gravel and cobble-bedded upland streams (wetted width 2–5 m; slopes 2.6–8.3%) representing a wide range of flow stages, LWD loading, and channel complexity, including measurements before and after LWD was added to enhance fish habitat. While transient storage volume fraction decreased as flow stage increased in simple channels, those with complex morphology and well-developed riparian vegetation maintained high transient storage fractions even during storm flows. LWD additions increased transient storage and channel complexity over the 2 years of post-treatment measurements. We predict with considerable precision two different formulations of transient hydraulic storage fraction using single-variable linear regressions on residual depth (R ² = 0.61–0.89), thalweg depth variance (R ² = 0.64–0.91), or large woody debris volume (R ² = 0.48–0.74). Demonstration of these likely causal associations contributes to understanding the process of transient storage and redefines the use of thalweg profile metrics as a new approach to quantifying morphologic and hydraulic complexity in streams.     

Trophic models of four benthic communities in Tongoy Bay (Chile): comparative analysis and preliminary assessment of management strategies

Steady-state trophic flow models of four benthic communities (seagrass, sand-gravel, sand and mud habitats) were constructed for a subtidal area in Tongoy Bay (Chile). Information of biomass, catches, food spectrum and dynamics of the commercial and non-commercial populations was used and the ECOPATH II software of Christensen and Pauly [Ecol. Modell. 61 (1992a) 169] was applied. The sea star Meyenaster gelatinosus and the crabs Cancer polyodon, C. porteri and Paraxanthus barbiger were found to be the most prominent predators in the benthic system. The scallop Argopecten purpuratus as well as other bivalves represented the principal secondary producers in the seagrass, sand-gravel and sand habitats, while the Infauna dominated the mud habitat. The highest total biomass and system throughput (33579.3 t/km²/year) was calculated for the sand-gravel habitat. The sand habitat had a negative net system production due to the amount of primary production imported from deeper waters to satisfy the food requirements of the large beach clam (Mulinia sp.) populations. The mean trophic level of the fishery varied between 2.06 (sand-gravel) and 3.92 (sand) reflecting the fact that the fishery concentrates on primary producers (i.e. algae and filter feeding), and on top predators (i.e. snails and crabs). Fishery is strongest in sand-gravel habitat, where annual catches amount to 122.05 g/m². Low values of the relative Ascendency (A/C) (from 27.4 to 32.7%) suggest that the systems analysed are immature and highly resistant to external perturbations. Manipulations of the input data for the exploited species suggest that seagrass and sand-gravel habitats have a potential for a ∼3 times higher than the present production of scallops and the red algae Chondrocanthus chamissoi. Preliminary results of Mixed Trophic Impacts (MTI) analysis suggest that any management policy aimed at a man-made increase in the standing stocks of A. purpuratus and Ch. chamissoi in seagrass and sand-gravel habitats, and a removal of the seastar M. gelatinosus in the seagrass habitat appears justified.     

Comparison between community structures of fishes in <Emphasis Type="Italic">Enhalus acoroides</Emphasis>- and <Emphasis Type="Italic">Thalassia hemprichii</Emphasis>-dominated seagrass beds on fringing coral reefs in the Ryukyu Islands,Japan

To clarify differences in community structures and habitat utilization patterns of fishes in Enhalus acoroides- and Thalassia hemprichii-dominated seagrass beds on fringing coral reefs, visual censuses were conducted at Iriomote and Ishigaki islands, southern Japan. The numbers of fish species and individuals were significantly higher in the E. acoroides bed than in the T. hemprichii bed, although the 15 most dominant fishes in each seagrass bed were similar. Cluster and ordination analyses based on the number of individuals of each fish species also demonstrated that fish community structures were similar in the two seagrass beds. Species and individual numbers of coral reef fishes which utilized the seagrass beds numbered less than about 15% of whole coral reef fish numbers, although they comprised about half of the seagrass bed fishes. Of the 15 most dominant species, 5 occurred only in the two seagrass beds, including seagrass feeders. Ten other species were reef species, their habitat utilization patterns not differing greatly between the two seagrass beds. Some reef species, such as Lethrinus atkinsoni and L. obsoletus, showed ontogenetic habitat shifts with growth, from the seagrass beds to the coral areas. These results indicate that community structures and habitat utilization patterns of fishes were similar between E. acoroides- and T. hemprichii-dominated seagrass beds, whereas many coral reef fishes hardly utilized the seagrass beds.     

The role of the sea urchin,Tripneustes gratilla (Linnaeus), in decomposition and nutrient cycling in a tropical seagrass bed

The sea urchin,Tripneustes gratilla, which feeds mainly on living leaves of the seagrass,Thalassia hemprichii, was studied in its habitat on the southern coast of Papua New Guinea, and its roles in decomposition and nutrient cycling in a seagrass bed were assessed through the excretion of ammonium and metabolism of feces produced by the sea urchin. Carbon content of the fresh feces (21% of dry weight) was similar to that of intact dead leaves of the same species (22–23%). Carbon/nitrogen and carbon/phosphorus ratios of the feces (21.7 and 466, respectively), however, were significantly lower than those of the dead leaves (25.9–27.7 and 656–804, respectively), indicating that the feces retain more nitrogen and phosphorus in comparison with carbon. Net consumption of ammonium and orthophosphate typically concurred with oxygen consumption during dark incubation of both the dead leaves and the sea urchin feces. Compared with the same oxygen consumption rate, however, the dead leaves consumed more orthophosphate than the feces. Under sunlight, dead leaves showed a net accumulation of carbon by epiphytic algae, while the feces showed a carbon loss. Ammonium excretion by this sea urchin (1.7–5.4 mg nitrogen/individual/day) would thus appear to make a significant contribution to nitrogen recycling since biological communities associated with dead leaves and sea urchin feces tend to demand an external supply of nitrogen, such as ammonium.     

Macrozoobenthic assemblages in littoral sediments in the enclosed Rhine-Meuse Delta

Littoral macrozoobenthos in the enclosed Rhine-Meuse Delta was investigated by taking 95 sediment samples from 17 sites between 1984 and 1990. In addition, a set of environmental parameters was determined. The aim was to identify the main assemblages and the environmental conditions under which they occur. By the use of TWINSPAN, three main littoral zoobenthic assemblages were distinguished, which were related to geographical zones and differences in sediment grain size distribution. The ‘litoral river sand’ assemblage was found in the most upstream part; it mainly consisted of ‘interstitial’ invertebrates, including the indicator speciesVejdovskiella comata, Propappus sp. andKloosia pusilla. The ‘littoral sedimentation area silt’ assemblage was dominated byGammarus tigrinus, Einfeldia dissidens andPisidium sp. It was found in several river sections and contained the indicator speciesEinfeldia dissidens, Potamopyrgus antipodarum andValvata piscinalis. The ‘littoral sandy basin’ assemblage was concentrated in the littoral fine sands of the Haringvliet and contained the indicator speciesPisidium henslowanum, P. moitessierianum, Cladotanytarsus sp. andLipiniella arenicola. These three assemblages are the reflection of an interaction between habitat, food and disturbance. Palaeoecological analysis of insect remains revealed that 14 out of the 24 insect taxa, that were formerly common in the river sand habitat, are now extinct from the Rhine. The river silt habitat seems less impoverished: two out of the 19 insect taxa found in palaeoecological analysis are now extinct from the Rhine and seven are rate. Exotic species (Corbicula fluminea, C. fluminalis andCorophium curvispinum) have recently colonized the Rhine-Meuse Delta, but their impact on the macrozoobenthos seems limited.Corbicula spp. have become abundant in the ‘littoral river sand’ assemblage only.     

Selection and preference of benthic habitat by small and large ammocoetes of the least brook lamprey (<Emphasis Type="Italic">Lampetra aepyptera</Emphasis>)

In this laboratory study, we quantified substrate selection by small (<50 mm) and large (100–150 mm) ammocoetes of the least brook lamprey (Lampetra aepyptera). In aquaria, ammocoetes were given a choice to burrow into six equally-available substrate types: small gravel (2.360–4.750 mm), coarse sand (0.500–1.400 mm), fine sand (0.125–0.500 mm), organic substrate (approximately 70% decomposing leaves/stems and organic sediment particles, and 30% silt and fine sand), an even mixture of silt, clay, and fine sand, and silt/clay (<0.063 mm). Fine sand was selected with a significantly higher probability than any other substrate. Fine sand habitat is limited in many streams, in part owing to geology, but also as a result of channelization and excessive silt/clay sedimentation, which is a conservation concern. Our results indicate that ammocoetes of least brook lampreys are habitat specialists that prefer fine sand habitat. Hence, availability of fine sand habitat may limit distributions and population sizes.     

Grazing effects of a gammaridean Amphipoda,Ampithoe sp., on the seagrass,Syringodium isoetifolium,and epiphytes in a tropical seagrass bed of Fiji

In the pure stand of tropical seagrass,Syringodium isoetifolium, in a small oceanic island, Fiji, grazing effects of the seagrass-associated gammarid,Ampithoe sp., on seagrass and epiphytes were assessed in October 1989, November 1991, November 1992. Density of the gammarid was estimated with two methods, mesh bag method and tuft method. During the three years surveyed the density of the gammarid increased remarkably from 1989 to 1991, with heavy epiphytism. Gut contents of the gammarid were examined. Grazing rates on seagrass leaf with and without epiphytic blue-green algae were measured in a bottle experiment. Litter bag experiments were conducted using different mesh sizes each containing seagrass only and seagrass and gammarids. The seagrass leaf biomass in the litter bag reduced abruptly in both bags. After one week, 78–86% of seagrass biomass disappeared from the bags. Enhancement of decomposition of seagrass leaf by the gammarid grazing was observed. Oxygen consumption and ammonium excretion rates were measured simultaneously in bottle experiments. Carbon budget in the seagrass bed was estimated as follows: 0.9 gC m⁻² day⁻¹ in seagrass growth, gammarid grazing was about a half of it and further assimilated a half of it, about 0.1 gC m⁻² day⁻¹, and more than half of it become CO₂ by respiration. Grazing effects on epiphyte and seagrass growth and production were discussed through the carbon budget and indirect interactions between seagrass, epiphytes and associated gammarids to explain the temporal change of seagrass and epiphyte dynamics.     

Variation in multiple traits of vegetative and reproductive seagrass tissues influences plant–herbivore interactions

Plant–herbivore interactions have strong ecological and evolutionary consequences, but have been traditionally overlooked in marine higher plants. Despite recent advances in seagrass ecology that highlight the importance of herbivory, the mechanisms that regulate the feeding behaviour of seagrass consumers remain largely unknown. Herbivores have been shown to reduce the sexual reproductive success of seagrasses through direct consumption of inflorescences and seeds, but we know little about intraspecific variation in susceptibility to grazing of different seagrass tissues. We contrasted the relative palatability of reproductive and vegetative tissues of the temperate seagrass Posidonia oceanica in the field, and we assessed the feeding preferences among these tissues of the main consumers of the plant, the fish Sarpa salpa and the urchin Paracentrotus lividus. Moreover, we identified the plant traits that explained the observed feeding behaviour. We provide strong evidence for herbivore selectivity among seagrass tissues. In the field, 70–90% of inflorescences were damaged by herbivores compared to 3–60% of leaves of similar age. In feeding assays, the urchin P. lividus showed over a twofold preference for reproductive tissue at various stages of development. By contrast, we detected no feeding activity on either leaves or inflorescences from the fish S. salpa, which is known to migrate to deeper waters soon after flowering starts and during the period of fruit maturation. Despite being the preferred food of urchins, inflorescences were chemically defended, had higher levels of phenolics and lower nutrient and calorific content than leaves. We experimentally demonstrated that leaf structural defences are the primary factor in determining urchin feeding preferences. Removal of plant structure results in a drastic shift in urchin selectivity towards the most nutritious and less chemically defended leaf tissue, indicating that multiple mechanisms of defence to herbivory may coexist in seagrasses.     

Comparative Evidence that Salt Marshes and Mangroves May Protect Seagrass Meadows from Land-derived Nitrogen Loads

Seagrass meadows within estuaries are highly sensitive to increased supplies of nitrogen (N). The urbanization of coastal watersheds increases the delivery of N to estuaries, threatening seagrass habitats; both seagrass production per unit area and the area of seagrass meadows diminish as land-derived N loads increase. The damaging effects of land-derived N loads may be lessened where there are fringes of coastal wetlands interposed between land and seagrass meadows. Data compiled from the literature showed that production per unit area by seagrasses increased and losses of seagrass habitat were lower in estuaries with relatively larger areas of fringing wetlands. Denitrification and the burial of land-derived N within fringe wetlands may be sufficient to protect N-sensitive seagrass habitats from the detrimental effects of land-derived N. The protection furnished by fringing wetlands may be overwhelmed by increases in anthropogenic N loads in excess of 20–100 kg N ha⁻¹ y⁻¹. The relationships of land-derived N loadings, fringing coastal wetlands, and seagrass meadows demonstrate that different units of the landscape mosaic found in coastal zones do not exist as separate units, but instead are coupled and uncoupled by biogeochemical transformations and transport among environments. Received 12 December 2000; accepted 15 August 2001.     

Stimulation of sulfate reduction rates in Mediterranean fish farm sediments inhabited by the seagrass <Emphasis Type="Italic">Posidonia oceanica</Emphasis>

Sulfate reduction rates and biogeochemical parameters of fish farm sediments across the Mediterranean were investigated in the order to evaluate the potential effects of organic matter inputs on habitat quality for the common seagrass Posidonia oceanica. Four study sites were selected in Spain, Italy, Greece and Cyprus to represent the Mediterranean basin. P. oceanica was found in immediate vicinity of all the farms, which were located at physically exposed sites about 1 km from the shore lines. Organic matter accumulation, sulfate reduction rates and sulfur pools were measured in depth profiles along transects from the farms in both bare and vegetated sediments. Results show that although the organic matter accumulation was minor at the sites (POC < 2.8% DW), the sulfate reduction rates were high, in particular at the largest farm in Italy (up to 212 mmol m⁻² d⁻¹), similar to rates found at shallower, temperate fish farm sites, where higher sedimentation rates can be expected. Sulfate reducing bacteria in these low-organic, carbonate-rich Mediterranean sediments respond strongly to organic matter loadings and cause habitat degradation. Sulfate reduction rates measured in the P. oceanica sediments were among the highest recorded (7.8–42.0 mmol m⁻² d⁻¹) similar to rates found in degrading meadows impacted by organic matter loadings. As sulfate reduction rates were correlated with the sedimentation rates along the transects rather than organic matter pools this suggests mineralization processes were controlled by organic matter loading in fish farm sediments. The vegetated sediments near the net cages were more reduced due to accumulation of sulfides compared to control sites, which is a possible contributing factor to the observed seagrass decline in the farm surroundings. It is recommended that Mediterranean fish farms are placed in areas with rapid dispersal of particulate waste products to minimize organic matter loading of the sediments and thereby preserve habitat quality for benthic fauna and flora.     

Resource translocation within seagrass clones: allometric scaling to plant size and productivity

The allometric scaling of resource demand and translocation within seagrass clones to plant size (i.e. shoot mass and rhizome diameter), shoot production and leaf turnover was examined in situ in eight seagrass species (Cymodocea nodosa, Cymodocea serrulata, Halophila stipulacea, Halodule uninervis, Posidonia oceanica, Thalassodendron ciliatum, Thalassia hemprichii and Zostera noltii), encompassing most of the size range present in seagrass flora. One fully developed shoot on each experimental rhizome was incubated for 2–3 h with a pulse of NaH¹³CO₃ (235 μmol) and ¹⁵NH₄Cl (40 μmol). The mobilisation of incorporated tracers across the clone was examined 4 days later. Carbon and nitrogen demand for shoot production across seagrass species scaled at half of the shoot mass, whereas seagrass leaves incorporated tracers (¹³C and ¹⁵N) at rates proportional to the shoot mass. The shoots of all seagrass species shared resources with neighbours, particularly with younger ones. The time scales of physiological integration and the absolute amount of resources shared by seagrass ramets scaled at 2.5 power of the rhizome diameter. Hence, the ramets of larger species were physiologically connected for longer time scales and share larger absolute amounts of resources with neighbours than those of smaller species. The different pattern of resource translocation exhibited by seagrasses helps explain the ecological role displayed by these species and the success of large seagrasses colonising nutrient-poor coastal areas, where they often dominate.