The seagrass, Zostera Marina L.: Plant morphology and bed structure in relation to sediment ammonium in izembek lagoon,Alaska

The relationship between the morphology of eelgrass, Zostera marina L., and ammonium in the sediment interstitial water was examined. The size of eelgrass plants collected from an intertidal terrace showed a consistent relationship with the size of the interstitial ammonium pool. Leaf area, length, and width all showed a linear increase in size towards stations having higher ammonium, while within the stations with the highest ammonium the leaf size was relatively unchanged. The size and extent of the root system in eelgrass varied across the environmental gradient. Eelgrass shoot density and flower abundance were inversely correlated with sediment nitrogen across the eelgrass meadow. Shoot density described a strong logarithmic relation with interstitial ammonium, opposite to the relationships for leaf size. Comparison of these results indicates the importance of sediment nitrogen in determining eelgrass bed structure.     

Restoring Eelgrass (Zostera marina L.) Habitats Using a Simple and Effective Transplanting Technique

Eelgrass beds in coastal waters of China have declined substantially over the past 30 years. In this study, a simple new transplanting technique was developed for eelgrass (Zostera marina L.) restoration. To assist in anchoring single shoots, several rhizomes of rooted shoots were bound to a small elongate stone (50–150 g) with biodegradable thread (cotton or hemp), and then the bound packet was buried at an angle in the sediments at a depth of 2–4 cm. This stone anchoring method was used to transplant eelgrass in early November 2009 and late May 2010 in Huiquan Bay, Qingdao. The method led to high success. Three month survivorship of the transplanted shoots at the two transplant sites was >95%. From April 20 to November 19, 2012, the following characteristics of the 2009 and 2010 transplanted eelgrass beds were monitored: morphological changes, shoot density, shoot height, leaf biomass, and sediment particle size. Results showed that the sexual reproduction period of the planted eelgrass was from April to August, and vegetative reproduction reached its peak in autumn. Maximum shoot height and biomass were observed in June and July. After becoming established, the transplanted eelgrass beds were statistically equal to natural eelgrass beds nearby in terms of shoot height, biomass, and seasonal variations. This indicates that the transplant technique is effective for eelgrass restoration in coastal waters.     

Variable responses of native eelgrass Zostera marina to a non-indigenous bivalve Musculista senhousia

The transport and establishment of non-indigenous species in coastal marine environments are increasing worldwide, yet few studies have experimentally addressed the interactions between potentially dominant non-native species and native organisms. We studied the effects of the introduced mussel Musculista senhousia on leaf and rhizome growth and shoot density of eelgrass Zostera marina in San Diego Bay, California. We added M. senhousia over a natural range in biomass (0–1200 g dry mass/m²) to eelgrass in transplanted and established beds. The effects of the non-indigenous mussel varied from facilitation to interference depending on time, the abundance of M. senhousia, and the response variable considered. Consistent results were that mussel additions linearly inhibited eelgrass rhizome elongation rates. With 800 g dry mass/m² of M. senhousia, eelgrass rhizomes grew 40% less than controls in two eelgrass transplantations and in one established eelgrass bed. These results indicate that M. senhousia, could both impair the success of transplantations of eelgrass, which spread vegetatively by rhizomes, and the spread of established Z. marina beds to areas inhabited by M. senhousia. Although effects on leaf growth were not always significant, in August in both eelgrass transplantations and established meadows leaf growth was fertilized by mussels, and showed a saturation-type relationship to sediment ammonium concentrations. Ammonium concentrations and sediment organic content were linear functions of mussel biomass. We found only small, non-consistent effects of M. senhousia on shoot density of eelgrass over 6-month periods. In established eelgrass beds, but not in transplanted eelgrass patches (≈0.8 m in diameter), added mussels suffered large declines. Hence, eelgrass is likely to be affected by M. senhousia primarily where Z. marina beds are patchy and sparse. Our study has management and conservation implications for eelgrass because many beds are already seriously degraded and limited in southern California where the mussel is very abundant. Received: 31 May 1997 / Accepted: 4 September 1997     

Seasonal changes in eelgrass functions: current velocity reduction,prevention of sediment resuspension,and control of sediment–water column nutrient flux in relation to eelgrass dynamics

We assessed seasonal changes in eelgrass (Zostera marina) functions, i.e., reduction of current velocity, buffering of sediment resuspension, and control of dissolved inorganic nitrogen flux between the sediments and the water column, using field observations and experiments in the Akkeshi-ko estuary, Hokkaido, Japan. We also analyzed the relationships between eelgrass traits and functions. The efficiency of the reduction in current velocity increased with the development of the eelgrass canopy. Sediment resuspension was inhibited from May to August, during which time the eelgrass canopy developed. Eelgrass controlled the NH₄ ⁺ concentration of sediment porewater through root nutrient uptake, affecting NH₄ ⁺ flux between the sediments and the water column. Fluctuations in eelgrass functions and coincident changes in dynamics resulted in seasonal changes in the eelgrass environment, which may in turn affect the dynamics of organisms inhabiting eelgrass beds, e.g., mysids and epiphytic algae. Moreover, the developed eelgrass canopy trapped a large amount of material during spring and summer, which was resuspended into the surrounding ecosystem in autumn when the canopy and its functions (i.e., reduction of current and sediment resuspension) diminished. These results suggest that seasonal changes in eelgrass functions also affect communities within marginal coastal ecosystems through the control of allochthonous resources. Handling editor: P. Viaroli     

Effects of sediment nutrients on seagrasses: Literature review and mesocosm experiment

A review of the literature shows that seagrass growth, abundance and morphology are strongly linked to available nutrient resources. In north temperate climates and in habitats with terrigenous sediments, nitrogen limitation occurs in both intertidal and subtidal beds. Typically, seagrasses growing in terrigenous sediments have ample phosphorus but lack nitrogen, and the plants' chemical composition is depleted in nitrogen. However, seagrasses occurring in tropical environments and carbonate sediments appear to experience phosphorus limitation due to binding of phosphate in the sediments. Thus, it is the sediment geochemistry in seagrass beds that is important in determining the limiting nutrient to seagrass growth.Examination of the literature indicates that field research on seagrass relationships involves too many interactive factors to be able to say certainly that any one plant characteristics is caused by any one environmental factor. Using mesocosms (partially enclosed outdoor experimental set-ups) one environmental factor can be changed between the treatment tank and the control. Therefore, we can determine experimentally that a plant characteristic is affected by a particular altered environmental factor. Experimental mesocosms used to grow eelgrass, Zostera marina L., in substrata of varied nitrogen composition showed the dramatic effect of insufficient nitrogen on eelgrass growth, abundance and leaf morphology. Additionally, eelgrass leaf tissue from low ammonium sand substratum is significantly depleted in nitrogen, demonstrating the supposition that nitrogen is limiting in terrigenous environments.     

The role of seedlings and seed bank viability in the recovery of Chesapeake Bay,USA, <Emphasis Type="Italic">Zostera marina</Emphasis> populations following a large-scale decline

The objective of this study was to quantify the spatial and temporal recolonization characteristics of Zostera marina beds in the lower Chesapeake Bay following large scale declines in the late summer of 2005. Transects were established and monitored monthly for changes in eelgrass abundance at three sites (two downriver, one upriver) in the York River from April–October 2006 and 2007. Measurements included percent bottom cover, above ground biomass, shoot density, shoot origin (seedling or vegetative), seed bank abundance and seed viability. During 2006, the eelgrass beds at all sites recovered with seedlings providing the largest proportion of the total shoot abundance. This trend shifted in 2007 and surviving vegetative shoots were the dominant component of shoot standing crop. A second consecutive decline related to low light conditions occurred during the summer of 2006 in the upriver site and recovery there was minimal in 2007. These results highlight that after a single die off event, seed germination with subsequent seedling growth is the principal method for revegetation in lower Chesapeake Bay Z. marina beds. However, no viable seeds remain in the seed bank during this first year of recovery and shoots produced by the seedling growth do not flower and produce seeds until their second year of growth. Therefore the seed-bank density is low and is not immediately replenished. This suggests that the resiliency of perennial Chesapeake Bay Z. marina populations to repeated disturbances is restricted and repeated annual stress may result in much longer term bed loss.     

Experimental test of biodeposition and ammonium excretion from blue mussels (Mytilus edulis) on eelgrass (Zostera marina) performance

Blue mussels and eelgrass have been found to coexist in many locations. However, knowledge of the interactions between these species is limited. Two experiments were conducted in the laboratory, a “Deposit” and an “Epiphyte” experiment. The Deposit experiment examined possible effects of increasing load of blue mussel (Mytilus edulis) biodeposits on sediment biogeochemistry and eelgrass (Zostera marina) performance. Z. marina mesocosms received normal or high loads of mussel biodeposits (Normal and High), while no biodeposits were added to the Control. High dosage had overall negative effects on Z. marina, which was reflected as lower leaf numbers and biomass and accumulation of elemental sulphur in rhizomes. The sediment biogeochemical conditions were altered, as the mussel biodeposits enhanced sulphate reduction rates and increased sulphide concentrations in the porewater, which may result in sulphide invasion and reduced growth of Z. marina.In the Epiphyte experiment effects of mussel excretion, with particular emphasis on ammonium, on the growth of Z. marina and their epiphytes were examined. A thick cover of epiphytes developed on Z. marina growing together with M. edulis, and the relative growth rate was reduced with 20% compared to plants from control without mussels. Overall the experiments showed negative effects on Z. marina growing together with M. edulis, thereby supporting a preceding field study by Vinther et al. [Vinther, H.F., Laursen, J.S., Holmer, M. 2008. Negative effects of blue mussel (Mytilus edulis) presence in eelgrass (Zostera marina) beds in Flensborg fjord, Denmark. Est. Coast Shelf. Sci. 77, 91-103.].     

Bay-scale assessment of eelgrass beds using sidescan and video

The assessment of the status of eelgrass (Zostera marina) beds at the bay-scale in turbid, shallow estuaries is problematic. The bay-scale assessment (i.e., tens of km) of eelgrass beds usually involves remote sensing methods such as aerial photography or satellite imagery. These methods can fail if the water column is turbid, as is the case for many shallow estuaries on Canada’s eastern seaboard. A novel towfish package was developed for the bay-scale assessment of eelgrass beds irrespective of water column turbidity. The towfish consisted of an underwater video camera with scaling lasers, sidescan sonar and a transponder-based positioning system. The towfish was deployed along predetermined transects in three northern New Brunswick estuaries. Maps were created of eelgrass cover and health (epiphyte load) and ancillary bottom features such as benthic algal growth, bacterial mats (Beggiatoa) and oysters. All three estuaries had accumulations of material reminiscent of the oomycete Leptomitus, although it was not positively identified in our study. Tabusintac held the most extensive eelgrass beds of the best health. Cocagne had the lowest scores for eelgrass health, while Bouctouche was slightly better. The towfish method proved to be cost effective and useful for the bay-scale assessment of eelgrass beds to sub-meter precision in real time.     

The role of the blue mussel Mytilus edulis in the cycling of nutrients in the Oosterschelde estuary (The Netherlands)

The fluxes of particulate and dissolved material between bivalve beds and the water column in the Oosterschelde estuary have been measured in situ with a Benthic Ecosystem Tunnel. On mussel beds uptake of POC, PON and POP was observed. POC and PON fluxes showed a significant positive correlation, and the average C:N ratio of the fluxes was 9.4. There was a high release of phosphate, nitrate, ammonium and silicate from the mussel bed into the water column. The effluxes of dissolved inorganic nitrogen and phosphate showed a significant correlation, with an average N:P ratio of 16.5. A comparison of the in situ measurements with individual nutrient excretion rates showed that excretion by the mussels contributed 31–85% to the total phosphate flux from the mussel bed. Ammonium excretion by the mussels accounted for 17–94% of the ammonium flux from the mussel bed. The mussels did not excrete silicate or nitrate. Mineralization of biodeposition on the mussel bed was probably the main source of the regenerated nutrients.From the in situ observations net budgets of N, P and Si for the mussel bed were calculated. A comparison between the uptake of particulate organic N and the release of dissolved inorganic N (ammonium + nitrate) showed that little N is retained by the mussel bed, and suggested that denitrification is a minor process in the mussel bed sediment. On average, only 2/3 of the particulate organic P, taken up by the mussel bed, was recycled as phosphate. A net Si uptake was observed during phytoplankton blooms, and a net release dominated during autumn. It is concluded that mussel beds increase the mineralization rate of phytoplankton and affect nutrient ratios in the water column. A comparison of N regeneration by mussels in the central part of the Oosterschelde estuary with model estimates of total N remineralization showed that mussels play a major role in the recycling of nitrogen.     

Eelgrass Detritus as a Food Source for the Sea Cucumber Apostichopus japonicus Selenka (Echinidermata: Holothuroidea) in Coastal Waters of North China: An Experimental Study in Flow-Through Systems

Eelgrass ecosystems have a wide variety of ecological functions in which living tissues and detritus may be a food source for many marine animals. In this study, we conducted a laboratory simulating experiment to understand the trophic relationship between the eelgrass Zostera marina L and the sea cucumber Apostichopus japonicus. A mixture of decaying eelgrass debris and seafloor surface muddy sediments was used as food to feed A. japonicus, and then specific growth rates (SGR) and fecal production rates (FPR) were measured. According to the proportion of eelgrass debris, we designed five treatment diets, i.e., ES0, ES10, ES20, ES40, and ES100, with eelgrass debris accounting for 0%, 10%, 20%, 40%, and 100% in dry weight, respectively. Results showed that diet composition had a great influence on the growth of A. japonicus. Sea cucumbers could use decaying eelgrass debris as their food source; and when the organic content of a mixture of eelgrass debris and sediment was 19.6% (ES40), a relatively high SGR (1.54%·d⁻¹) and FPR (1.31 g·ind.⁻¹ d⁻¹) of A. japonicus were obtained. It is suggested that eelgrass beds can not only provide habitat for the sea cucumber A. japonicus but can also provide an indirect food source for the deposit feeder. This means that the restoration and reconstruction of eelgrass beds, especially in coastal waters of China, would be a potential and effective measure for sea-cucumber fisheries, in respect to both resource restoration and aquaculture of this valuable species.     

Ecosystem engineers: Interactions between eelgrass Zostera capensis and the sandprawn Callianassa kraussi and their indirect effects on the mudprawn Upogebia africana

Increasing recognition is being given to the concept of ‘ecosystem engineers’, which alter the physical nature of the environment and thereby influence other species by means that are neither competitive nor trophic. This paper examines a case study of such effects, in which negative correlations between the abundances of the sandprawn Callianassa kraussi and the eelgrass Zostera capensis imply mutual exclusion because Z. capensis stabilises sediment and inhibits burrowing by C. kraussi, whereas sandprawn bioturbation smothers eelgrass. To test this, we undertook manipulative experiments in which Z. capensis was transplanted into sandflats where C. kraussi was either left undisturbed or eliminated by defaunation. This demonstrated that (1) Z. capensis thrived and expanded in sandflats in the absence of bioturbation but deteriorated and disappeared if C. kraussi was present. (2) In the short term, introduction of Z. capensis reduced densities of C. kraussi, and in established beds of eelgrass, C. kraussi was rare. (3) The mudprawn Upogebia africana was largely restricted to treatments that contained Z. capensis, and its densities were greater in the absence of C. kraussi than in its presence. The presence of eelgrass and the exclusion of C. kraussi also diminished sediment penetrability, suspension of particles and surface burial rates compared to the situation in undisturbed Callianassa-dominated sandflats. These results confirm a mutually negative interaction between C. kraussi and Z. capensis and strengthen the belief that the underlying mechanisms are the antagonistic effects of bioturbation by C. kraussi versus sediment stabilisation by Z. capensis. U. africana seems to benefit directly from Z. capensis but possibly also benefits indirectly from the exclusion of C. kraussi by Z. capensis, since its filter-feeding mode of life requires sufficient sediment stability to maintain semi-permanent U-tubes.     

Factors controlling spatial variability in ammonium release within an estuarine bay (Alfacs Bay,Ebro Delta,NW Mediterranean)

Sediment-water ammonium fluxes, oxygen uptake and sediment characteristics were studied in an estuarine bay influenced by temporal freshwater discharges. Sediment at seven stations representing a gradient imposed by freshwater inputs was sampled for sediment-water ammonium and oxygen fluxes, chlorophyll a derivative pigments, organic content, porosity and elemental composition (Fe, Mn, Si, Al). Oxygen uptake decreased along the gradient and correlated with total chlorophyll a derivatives indicating the close coupling between aerobic metabolism and short-time sedimentation events. Ammonium release showed a discontinuous pattern of decrease along the gradient and only correlated with the Fe:Mn atomic ratio. Correlation between the structural properties of the sediment (Si:Al atomic ratio, porosity and organic content) and ammonium release was also found (excluding data from the station with the highest ammonium flux). The extent of the influence of metabolism and sediment structure on ammonium release is discussed.     

Causes of rarity and range restriction of an endangered, endemic limpet, Siphonaria compressa

The pulmonate limpet Siphonaria compressa is South Africa's most endangered marine mollusc. It is endemic to just two localities: Langebaan Lagoon on the west coast, and Knysna Estuary on the south coast, and occurs only on the eelgrass Zostera capensis. In Langebaan Lagoon, eelgrass has fluctuated substantially over the last 34 years, and S. compressa has twice approached extinction. S. compressa is largely confined to the lower edge of the eelgrass beds there, being replaced higher up by another small gastropod, Assiminea globulus. We explored the physical and biological factors underlying the limpet's narrow habitat, using field observations, translocations, caging and transplant experiments. Abundance of S. compressa was positively correlated with Z. capensis cover and negatively correlated with shore height. When moved to the upper portions of the eelgrass bed, S. compressa had lower rates of persistence and survival than in the lowest zone. The lower limit of zonation for S. compressa was set indirectly by bioturbation by the sandprawn Callianassa kraussi, which excluded eelgrass from intertidal sandbanks. Transplants of eelgrass into the sandbanks proliferated provided C. kraussi was experimentally eliminated, and supported densities of S. compressa 20-fold greater than in control eelgrass beds, suggesting that high-shore eelgrass beds to which S. compressa is normally confined are suboptimal for the limpet. A. globulus showed patterns opposite to those of S. compressa: its persistence and survival were greatest in the upper zone and it actively avoided the lower sections of these beds and never colonised eelgrass transplanted into sandflats lower on the shore. There was no evidence that competition between S. compressa and A. globulus influenced the zonation or abundance of either species. Rarity of S. compressa and its endangered status seem dictated by its extremely narrow and temporally changeable habitat-range, which is defined by physical stress in the high-shore and bioturbation by C. kraussi in the low-shore. Fluctuations in eelgrass abundance and limitation of S. compressa to just two localities add substantially to the risks of extinction for this embattled stenotypic limpet.     

Photosynthetic and morphological responses of eelgrass (Zostera marina L.) to a gradient of light conditions

Zostera marina L. (eelgrass) from Great Bay Estuary, New Hampshire and Maine (USA), was transplanted in outdoor mesocosms and subjected to four light treatments (100, 58, 34 and 11% surface irradiance, SI) between May and September 2003 to investigate the relationship between light availability and the growth and survival of eelgrass. Evaluating eelgrass seedlings and adult mature plants demonstrated no differences in photosynthetic response after 22 days of acclimation. During at least the first 19 days of shading, maximum electron transport rate (ETR_max) rate of eelgrass did not differ significantly between light treatments. After 40 days, a significant reduction in ETR_max and minimum saturating light was observed in plants growing at 34% SI and below. Morphological responses exhibited a linear increasing trend with greater light. 34% SI exhibited drastic reductions (to less than 25% of control) in rhizome growth, shoot density, shoot production, number of nodes per plant and plant weight at the end of the study (81 days). Shoot to root ratio at 34% SI increased by > 50%. Plants shaded to 58% SI showed no significant difference from the control in plant parameters except an increased rate of rhizome elongation. Our results link the lower shoot densities with shading to the slow growth rate of horizontal rhizomes and a total lack of lateral expansion at 11% SI. ETR_max declined over time in plants at 11% SI resulting in 81% mortality, no lateral branching and no morphological development, indicating that the minimum light required for long-term eelgrass growth and survival is greater than the previously suggested 11% SI. We demonstrate that eelgrass plants at these latitudes can persist at light levels of 58% SI and above, and are light-limited at 34% SI and below.     

Influences of biological interactions on community structure within seagrass beds and sandprawn-dominated sandflats

The faunal composition of high-shore beds of eelgrass Zostera capensis versus unvegetated sandflats dominated by the sandprawn Callianassa kraussi was investigated by experimentally transplanting plots of eelgrass into sandflats where the sandprawn was either left unmanipulated or eliminated by prior faunal suffocation. After 12 months, multidimensional scaling defined five faunal clusters that were dictated primarily by the presence or absence of eelgrass rather than sandprawns. By 18 months, eelgrass and sandprawns played equal roles in defining faunal clusters. After 30 months, sandprawns had displaced the eelgrass transplants and faunal composition converged on that of sandflat control plots.Most species were either ‘eelgrass-associated’ or ‘sandflat-associated’, but three had unique responses to the treatments: two high-shore species (Assiminea globulus and Hydrobia sp.) that were abundant in eelgrass beds never appeared in the eelgrass transplants, whereas another eelgrass-occupant, Siphonaria compressa, became superabundant on the transplants for 18 months.Unexpectedly, the eelgrass fauna was never more rich or diverse than that of unvegetated sandflats, although abundance was greater in the eelgrass. As hypothesised, the sandflats were disproportionally populated by burrowers and eelgrass by non-burrowers, but a second hypothesis that these habitats should support mainly hard-bodied and soft-bodied animals respectively was rejected.The distinctively different faunas of eelgrass and unvegetated sandflats are seemingly respectively maintained by sediment stabilisation by eelgrass versus bioturbation by sandprawns. The transplants of eelgrass into sandflats developed an eelgrass-associated fauna provided sandprawns were excluded, but ultimately reverted to a sandflat fauna once sandprawns reinvaded.     

Light dependence of Zostera marina annual growth dynamics in estuaries subject to different degrees of eutrophication

We examined the coupling between eelgrass growth dynamics and surface irradiance over an annual cycle in four shallow estuaries of the Waquoit Bay system (MA, USA) that have similar physical characteristics, but are subject to different land-derived nitrogen loading rates and eutrophication. Contrary to our hypothesis, the results show that most measures of eelgrass demographics were positively correlated with surface irradiance in all four estuaries. Of the 45 regression models adjusted between irradiance and demographic variables (density, plastochrone intervals, and above- or belowground biomass, growth, and production, on both a per shoot and areal basis), only nine were non-significant, and only six of those corresponded to the eutrophic estuaries. There was a lack of correlation between shoot density and irradiance in the eutrophic estuaries, in contrast to the strong coupling in estuaries with the lowest nitrogen loads. Severe light limitation and other deleterious impacts imposed by macroalgal canopies on newly recruiting shoots in the eutrophic estuaries likely contributed to the lack of correlation between shoot density and irradiance at the water's surface. Because the range in eutrophication included the range of conditions at which eelgrass can survive, the relatively consistent temporal coupling between surface irradiance and most eelgrass demographic variables found here may also be a feature of other shallow temperate systems undergoing eutrophication, and indicates a measure of plant recruitment (density) to be one of the first parameters to become uncoupled from light reaching the water's surface.     

In situ phytoremediation of PAH- and PCB-contaminated marine sediments with eelgrass (Zostera marina)

In view of the fact that there are presently no cost-effective in situ treatment technologies for contaminated sediments, a 60-week-long phytoremediation feasibility study was conducted in seawater-supplied outdoor ponds to determine whether eelgrass (Zostera marina) is capable of removing polynuclear aromatic hydrocarbons (PAHs) and polychlorinated biphenyls (PCBs) from submerged marine sediments. It was determined that all PAHs and PCBs, independent of the number of aromatic rings and degree of chlorination, respectively, were removed to a much larger extent in planted sediments compared to unplanted controls. After 60 weeks of treatment, the concentration of total PAHs decreased by 73% in planted sediments but only 25% in unplanted controls. Similarly, total PCBs declined by 60% in the presence of plants while none were removed in the unplanted sediment. Overall, the apparent PAH and PCB biodegradation was greatest in the sediment layer that contained most of the eelgrass roots. Abiotic desorption tests conducted at week 32 confirmed that the phytoremediation process was not controlled by mass-transfer or bioavailability limitations since all PAHs and PCBs desorbed rapidly and to a large extent from the sediment. PAHs were detected in both roots and shoots, with root and shoot bioaccumulation factors for total PAHs amounting to approximately 3 and 1, respectively, after 60 weeks of phytoremediation treatment. Similarly, the root bioaccumulation factor for total PCBs was around 4, while no PCBs were detected in the eelgrass leaves at the end of the experiment. The total mass fraction of PAHs and PCBs absorbed and translocated by plant biomass during the 60-week period was insignificant, amounting to less than 0.5% of the total mass of PAHs and PCBs which was initially present in the sediment. Finally, the number of total heterotrophic bacteria and hydrocarbon degraders was slightly but not statistically significantly greater in planted sediments than in unplanted controls. After ruling out contaminant loss to the water column or absorption and transformation within plant cells, it is most likely that the presence of eelgrass stimulated the microbial biodegradation of PAHs and PCBs in the rhizosphere by releasing root exudates, plant enzymes, or even oxygen. Additional research is needed to further elucidate these potential phytoremediation mechanisms.     

Mussels matter: postlarval dispersal dynamics altered by a spatially complex ecosystem engineer

This study investigated postlarval dispersal of soft-bottom macrofauna at a spatially complex intertidal mudflat comprising patches of bare sediment and an ecosystem engineer, the mussel Mytilus edulis. At each of four sites in Guard Point Cove, Maine, USA, we took core samples and deployed bedload traps in bare sediment and mussel bed habitats to estimate ambient densities, rates of sediment flux, and several measures of postlarval dispersal. Univariate and multivariate nonmetric multidimensional scaling (nMDS) results showed few significant site effects and no habitat×site interactions. In contrast, there were numerous significant habitat effects. Compared to the bare sediment, the mussel bed habitat had: fewer species; higher ambient density and proportional abundance of the oligochaete Tubificoides benedeni (the dominant species in both habitats); lower ambient densities and proportional abundances of major taxa and the nonoligochaetes as a group; and higher sediment flux and relative (i.e., per capita) dispersal of nonoligochaetes. Macrofauna species dispersed in relative proportions that were different from those in the ambient assemblage. Per capita T. benedeni transport rates were low in mussel beds compared to those for nonoligochaetes, consistent with the view that beds represent favorable habitat for oligochaetes. The number of total macrofauna individuals trap⁻¹ day⁻¹ was negatively correlated with ambient density and positively correlated with sediment flux in both habitats, but these relationships were significant only in the mussel bed. The results indicate that altered transport rates of sediment and postlarvae are important mechanisms by which mussels act as ecosystem engineers to modify soft-bottom habitats. Differential transport rates caused by aggregations of mussels and other foundation species must be considered in explanations of spatial pattern in soft-bottom communities.     

Eelgrass, Zostera marina, growth along depth gradients: upper boundaries of the variation as a powerful predictive tool

1200 measurements of eelgrass ( Zostera marina ) biomass, shoot density and cover along 19 depth gradients in Øresund, located between Denmark and Sweden, were analysed to characterise growth of eelgrass in relation to depth. The large data set allowed analyses of boundaries of distribution as well as of average trends. Natural variability is large in shallow water where populations are disturbed by wave action and other physical parameters. Models based on average values, therefore, did not adequately describe growth regulation by resources, and only explained a minor part (up to 30%) of the overall variation in data. In contrast, boundary functions, which describe the upper bounds of distributions, focus on the variation produced by the ultimately growth-regulating resource, and therefore provide models with high predictive power. An exponential model explained up to 90% of the variation in upper bounds of eelgrass shoot density as a function of depth and indicated that shoot density was ultimately regulated by light availability. The boundary functions demonstrated that eelgrass shoot density, biomass and cover followed markedly different patterns as functions of depth and were affected differently by the factors governing their distribution. In addition, boundary functions revealed informative spatial structures in data and illustrated whether a given general trend was caused by changes in maximum values, minimum values or both. For example, upper and lower boundaries of biomass-shoot density relations changed markedly with depth, demonstrating depth-related changes in intraspecific succession and competition patterns. Boundary functions are therefore suggested as a promising tool for analysing ultimate regulating factors of distribution and growth of organisms when large data sets are available.     

Nitrate and nitrite in interstitial waters of eelgrass beds in relation to the rhizosphere

The distribution of nitrate and nitrite in the interstitial water of the sediment of eelgrass (Zostera marina) bed of Izembek Lagoon, Alaska, were investigated. Their concentrations were relatively high (0 to 9.8 μg-at.N·1^?1, average 4.8 for nitrate; 0 to 4.0 μ-at.N·1^?1, average 1.9 for nitrite) although the sediments were anoxic and contained hydrogen sulphide. The rates of bacterial denitrification measured by ¹⁵N tracer technique ranged from 0.49×10^?10 to 1.2 × 10^?9 g-atN·g^?1·h^?1. When a steady state is maintained, the loss of nitrate and nitrite must be balanced by their production by bacterial nitrification. Experimentally determined rate of nitrification in the sediment was of the same order. A model experiment demonstrated that oxygen is transported from leaves to rhizomes and roots of eelgrass and released into the sediment. The oxygen is used for nitrification in the rhizosphere in anoxic sediments.