The ecology of eelgrass, Zostera marina (L.), fish communities. II. Functional analysis

Consumption, production, and respiration of fish communities utilizing two eelgrass beds in a shallow estuarine system near Beaufort, North Carolina have been estimated for 1971–1972: annual production was 21.7 kcal/m² in each bed with pinfish accounting for 45 and 68% of the production in the Phillips Island and Bogue Sound beds, respectively. Annual community respiration was 57.9 and 69.7 kcal/m² in the two beds with pinfish accounting for 62.6 and 26.7% of the total in the Bogue Sound and Phillips Island beds, respectively. Estimation of the annual food energy consumed by the eelgrass fish community using the Winberg and daily ration methods gave values within 6% of each other.Energy turnover was high (2.8), and the efficiency of energy dissipation low for the two eelgrass fish communities, suggesting that the resident fish populations were adapted to the temperature extremes within the bed. High ecological efficiencies of 0.24 and 0.23 and the high overall efficiency of the eelgrass system (production/solar radiation) of 0.0051 and 0.0086% indicate that the eelgrass beds are efficient systems for converting consumed energy and solar radiation into fish.     

Seasonal occurrence and food habits of larvae and juveniles of two temperate basses in the Shimanto estuary,Japan

Larvae and juveniles ofLateolabrax japonicus andL. latus occurred from January to May 1986 in the shallow waters of the Shimanto estuary.L. japonicus markedly outnumberedL. latus. Distinct ecological differences were recognized in habitats and food habits between the two species:L. japonicus mainly inhabited eelgrass beds composed ofZostera nana, whileL. latus appeared evenly in both eelgrass beds and non-eelgrass habitats; the former fed on copepods and cladocerans, while the latter fed on copepods and fish larvae. From these habitat and food habit analyses, estuaries were considered to be important as a main habitat forL. japonicus, but not forL. latus. The fact that ecological differences have occurred during the early life stages was inferred to be one of the possible keys to speculate on the speciation of the two species.     

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     

Photoadaptation and growth of Zostera marina L. (eelgrass) transplants along a depth gradient

Photosynthetic and growth responses were assessed in Zostera marina L. transplants within and beyond the natural extent of an eelgrass meadow in Great Harbor, Woods Hole, MA. Transplant survival and rapid growth inshore of the shallow edge of the meadow (0.5 and 0.8 m water depth) indicated a periodic disturbance factor maintaining the shallow edge of the meadow. Transplant mortality, reduced growth, and a negative carbon balance of eelgrass transplanted offshore the deep edge of the meadow (7 and 10m) supported the hypothesis of light-limited eelgrass growth in the deep regions of the Great Harbor meadow. Photoadaptive responses occurred along the water depth gradient, and both photosynthesis and growth responses were used to assess the genetic vs. phenotypic components of eelgrass response to the water depth gradient. Reciprocal transplants between shallow (1.3 m) and deep (5.5 m) areas within the eelgrass meadow indicated photosynthetic and growth responses were primarily a result of growth habitat rather than genetic differentiation within the eelgrass meadow.     

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.     

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.     

Eelgrass,Zostera marina,as essential fish habitat for young‐of‐the‐year winter flounder,Pseudopleuronectes americanus (Walbaum, 1792) in Maine estuaries

Distribution and density by habitat for age‐0, young‐of‐the‐year (YOY) winter flounder, Pseudopleuronectes americanus (Walbaum, 1792), were compared for two Maine estuaries to help define essential fish habitat for this life history stage. Two estuaries (Weskeag River and Penobscot Bay) along Mid‐coast Maine were sampled monthly with daytime 1.0 m² fixed‐frame throw traps around neap low tide, May–December over two consecutive years (2003–2004). Both eelgrass and adjacent sand/mud (20–60 cm deep) were randomly sampled with equal effort (4–12 samples per month) at two sites in both the Weskeag River and Penobscot Bay. Significantly higher densities of YOY winter flounder (2–9 cm TL) occurred in eelgrass relative to sand/mud. Density increased significantly in both habitats in 2004, and was higher in Penobscot Bay relative to the Weskeag River. YOY densities compared by eelgrass coverage within throw traps were found to be significantly higher in eelgrass that exceeded 30% coverage when compared with adjacent sand/mud areas and eelgrass coverage of 10–20%. YOY occurred in all months sampled (May–December); no density differences existed by month. These results indicate that very shallow (<0.6 m) eelgrass habitat is of key importance to YOY winter flounder in Maine estuaries and should be viewed as essential fish habitat (EFH) for this species and life stage.     

Multiple stressors threaten the imperiled coastal foundation species eelgrass (Zostera marina) in Chesapeake Bay,USA

Interactions among global change stressors and their effects at large scales are often proposed, but seldom evaluated. This situation is primarily due to lack of comprehensive, sufficiently long‐term, and spatially extensive datasets. Seagrasses, which provide nursery habitat, improve water quality, and constitute a globally important carbon sink, are among the most vulnerable habitats on the planet. Here, we unite 31 years of high‐resolution aerial monitoring and water quality data to elucidate the patterns and drivers of eelgrass (Zostera marina) abundance in Chesapeake Bay, USA, one of the largest and most valuable estuaries in the world, with an unparalleled history of regulatory efforts. We show that eelgrass area has declined 29% in total since 1991, with wide‐ranging and severe ecological and economic consequences. We go on to identify an interaction between decreasing water clarity and warming temperatures as the primary drivers of this trend. Declining clarity has gradually reduced eelgrass cover the past two decades, primarily in deeper beds where light is already limiting. In shallow beds, however, reduced visibility exacerbates the physiological stress of acute warming, leading to recent instances of decline approaching 80%. While degraded water quality has long been known to influence underwater grasses worldwide, we demonstrate a clear and rapidly emerging interaction with climate change. We highlight the urgent need to integrate a broader perspective into local water quality management, in the Chesapeake Bay and in the many other coastal systems facing similar stressors.     

The ecology of eelgrass, Zostera marina (L.), fish communities. I. Structural analysis

Fish populations utilizing eelgrass, Zostera marina (L.), beds in two different estuarine areas near Beaufort, North Carolina were analysed and compared to determine aspects of their community structure. The fish community of the eelgrass beds was characterized by low diversity and high standing crops of biomass and energy, both of which showed seasonal variation. Wide temperature fluctuations related to the overall shallowness of the beds probably regulated the diversity of fishes utilizing the beds. This community was dominated by pinfish, Lagodon rhombiodes (L.), which comprised 45% and 67% of the fish biomass in the Phillips Island and Bogue Sound beds, respectively.Changes in total body caloric content were probably related to developmental stages and changes in diet. Adult fish often had significantly higher weight-specific caloric contents than juvenile fish. Monthly or seasonal variations in caloric content of the organic matter of pinfish had little influence on the caloric content within the various sizes of pinfish.There was a significant correlation between fish biomass, temperature, and Zostera biomass. Fish biomass was highest when temperature and grass biomass were at a maximum. In general, water depth over the beds had little effect on the standing crop of fish within the bed, but cooler waters which occurred at night, darkness, or both, had a large effect.     

Evidence of Eelgrass (Zostera marina) Seed Dispersal by Northern Diamondback Terrapin (Malaclemys terrapin terrapin) in Lower Chesapeake Bay

The initial discovery in May 2009 of eelgrass (Zostera marina) seeds in fecal samples of wild-caught northern diamondback terrapins (Malaclemys terrapin terrapin) was the first field evidence of eelgrass seed ingestion in this species. This finding suggested the potential of terrapins as seed dispersers in eelgrass beds, which we sampled for two additional years (2010 and 2011). Seeds were only found in feces of terrapins captured prior to June 8 in all three years, coinciding with eelgrass seed maturation and release. Numbers of seeds in terrapin feces varied annually and decreased greatly in 2011 after an eelgrass die off in late 2010. The condition of seeds in terrapin feces was viable-mature, germinated, damaged, or immature. Of terrapins captured during time of seed release, 97% were males and juvenile females, both of which had head widths <30 mm. The fraction of individuals with ingested seeds was 33% for males, 35% for small females, and only 6% for large (mature) females. Probability of seed ingestion decreased exponentially with increasing terrapin head width; only males and small females (head width <30 mm) were likely to be vectors of seed dispersal. The characteristic that diamondback terrapins have well-defined home ranges allowed us to estimate the number of terrapins potentially dispersing eelgrass seeds annually. In seagrass beds of the Goodwin Islands region (lower York River, Virginia), there were 559 to 799 terrapins, which could disperse between 1,341 and 1,677 eelgrass seeds annually. These would represent a small proportion of total seed production within a single seagrass bed. However, based on probable home range distances, terrapins can easily traverse eelgrass meadow boundaries, thereby dispersing seeds beyond the bed of origin. Given the relatively short dispersion distance of eelgrass seeds, the diamondback terrapin may be a major source of inter-bed seed dispersal and genetic diversity.     

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.     

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.     

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.     

The response of interstitial ammonium in eelgrass (Zostera marina L.) beds to environmental perturbations

Natural and human-induced perturbations of eelgrass (Zostera marina L.) beds were used to examine the interaction between the sediment interstitial ammonium pool and nitrogen uptake by the plants. Eelgrass colonization of unvegetated areas was accompanied by a substantial decrease in the interstitial ammonium pool over a 4-yr period. The changes in interstitial ammonium and shoot density during colonization support an already determined relationship between shoot density and ammonium pool measurements. In field perturbation experiments, removing eelgrass leaves and sealing the sediment surface altered the flux of ammonium from the interstitial ammonium pool, and resulted in a rapid increase in interstitial ammonium concentrations. Measurements of ammonium accumulation under the various perturbation conditions and a control permitted calculation of the sediment ammonium flux. These estimates include uptake by eelgrass roots, regeneration in the root zone, and diffusion from the sediments. Nitrogen limitation was observed in some eelgrass beds.     

Effects of salinity and water temperature on the ecological performance of Zostera marina

We tested the effects of salinity and water temperature on the ecological performance of eelgrass (Zostera marina L.) in culture-experiments to identify levels that could potentially limit survival and growth and, thus, the spatial distribution of eelgrass in temperate estuaries. The experiments included eight levels of salinity (2.5, 5, 10, 15, 20, 25, 30 and 35‰) and seven water temperatures (5, 10, 15, 20, 25, 27.5 and 30 °C). Low salinity (i.e. 5 and 2.5‰) increased mortality (3–6-fold) and had a strong negative effect on shoot morphology (number of leaves per shoot reduced by 40% and shoot biomass reduced by 30–40%), photosynthetic capacity (P_max—reduced by 30–80%) and growth (production of new leaves reduced by 50–60%, leaf elongation rate reduced by 60–70% and production of side-shoots reduced by 40–60%), whereas eelgrass performed almost equally well at salinities between 10 and 35‰. The optimum salinity for eelgrass was between 10 and 25‰ depending on the response parameter in question. Extreme water temperatures had an overall negative impact on eelgrass, although via different mechanisms. Low water temperatures (5 °C) slowed down photosynthetic rate (by 75%) and growth (production of new leaves by 30% and leaf elongation rate by 80%), but did not affect mortality, whereas high temperatures (25–30 °C) increased mortality (12-fold) and lowered both photosynthetic rate (by 50%) and growth (production of new leaves by 50% and leaf elongation rate by 75%). The optimum water temperature for eelgrass appeared to lie between 10 and 20 °C. These results show that extreme conditions may affect the fitness of eelgrass and, thus, may potentially limit its distribution in coastal and estuarine waters.     

The Ichthyofauna Associated with an Intertidal Creek and Adjacent Eelgrass Beds in the Kariega Estuary,South Africa

The structure and functioning of salt marsh fish communities in the overall ecology of southern African estuaries is poorly understood. This study compares the ichthyofauna associated with a salt marsh creek and eelgrass bed in an attempt to evaluate the relative importance of these habitats to fishes. Taylor's salt marsh creek and adjacent eelgrass bed in the Kariega Estuary were sampled twice per season between the winter of 1992 and the autumn of 1994. The average density and standing stock of fishes were found to be considerably higher in the eelgrass bed than in the intertidal creek. Both habitats had similar fish diversities but were dominated by different taxa, the most notable of which was the dominance of mugilids in the creek and their scarcity in the eelgrass. Taylor's intertidal creek and adjacent eelgrass beds were dominated by juvenile fish, with both habitats functioning as nursery areas for juvenile fish, albeit for totally different ichthyofaunal communities. The similar fish diversities but lower abundances in the intertidal creek compared to the eelgrass beds are in contrast to similar North American studies, and refute the hypothesis that intertidal salt marsh creeks have higher fish densities but lower diversities than eelgrass beds.The first author is also the senior author     

Light control on phytoplankton production in a shallow and turbid estuarine system

Tagus estuary is one of the largest estuaries of Western Europe. With the aim of unravelling the drivers of primary production in this shallow and turbid nutrient replete estuary, we tested the hypothesis that light availability is a major factor controlling phytoplankton production. Environmental parameters, phytoplankton biomass, community composition, and photosynthetic parameters were monitored at two sites in the estuary during a complete annual cycle. Despite the fact that nutrient concentrations were always above growth-limiting values, Chl a concentrations were relatively low throughout the study period. High water column turbidity, due to riverine inputs, promoted a rapid attenuation of light and created a compressed profile with optimal photosynthetic conditions. Therefore, the phytoplankton community, dominated by small cells, such as diatoms and cryptophycean flagellates, displayed highly photosynthetic efficiency and low light-saturated photosynthetic rates as a photo-acclimation response to low light conditions year-round. Primary production rate was unimodal, peaking in the summer months. In such estuarine system, gross primary production could thus be predicted by an existing robust empirical model based on pigment standing crop (Chl a), surface irradiance (E ₀) and optical depth (Z _eup). Compared to other shallow estuaries, the Tagus can be classified as a low- to moderately productive estuary, being the turbidity-induced low light conditions the principal factor limiting phytoplankton growth.     

Phenology and reproductive biology of eelgrass (Zostera marina L.) at Bahia Kino,sea of Cortez,Mexico

At Bahia Kino, Mexico, seeds of Zostera marina L. (eelgrass) germinate in October and early November. By January seedlings are widespread throughout the area. A very rapid vegetative growth ensues. By March all eelgrass at the depth limit (7 m) is flowering; and most is flowering in shallow water (low tide to 4 m depth). In April all plants are in flower; a few plants display mature seeds. Windrows of detached plants occur on the beaches. In May all reproductive plants have mature seeds, and few attached plants remain. In July all eelgrass in the area has disappeared.All reproductive activities and plant detachment are completed before the lethal upper limit for the species is reached (30°C). Thus, eelgrass at Bahia Kino is a true annual and represents an ultimate response to high water temperatures.     

Relationships between bed age,bed size,and genetic structure in Chesapeake Bay (Virginia,USA) eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.)

Genetic structure and diversity can reveal the demographic and selective forces to which populations have been exposed, elucidate genetic connections among populations, and inform conservation strategies. Beds of the clonal marine angiosperm Zostera marinaL. (eelgrass) in Chesapeake Bay (Virginia, USA) display significant morphological and genetic variation; abundance has fluctuated widely in recent decades, and eelgrass conservation is a major concern, raising questions about how genetic diversity is distributed and structured within this metapopulation. This study examined the influence of bed age (<65years versus<6years) and size (>100ha versus<10ha) on morphological and genetic (allozyme) structure and diversity within Chesapeake Bay eelgrass beds. Although both morphology and genetic diversity varied significantly among individual beds (F _ST=0.198), neither varied consistently with bed age or size. The Chesapeake eelgrass beds studied were significantly inbred (mean F _IS=0.680 over all beds), with inbreeding in old, small beds significantly lower than in other bed types. Genetic and geographic distances within and among beds were uncorrelated, providing no clear evidence of isolation by distance at the scale of 10's of km. These results suggest that local environmental conditions have a greater influence on plant morphology than do bed age or size. They support the hypotheses that eelgrass beds are established by multiple founder genotypes but experience little gene flow thereafter, and that beds are maintained with little loss of genetic diversity for up to 65 years. Since phenotypic and genotypic variation is partitioned among beds of multiple ages and sizes, eelgrass conservation efforts should maximize preservation of diversity by minimizing losses of all beds.     

Responses of Very Shallow Marine Ecosystems to Nutrient Enrichment

The success of simple predictive relationships such as the Vollenweider plot in limnology has encouraged marine ecologists to attempt to develop similar models relating pollutant inputs to ecological conditions in estuaries. Most of these efforts have focused on relatively deep (>5?m) river mouth estuaries and embayments where primary production is dominated by phytoplankton. Experimental nutrient enrichment studies of phytoplankton-based mesocosms at the Marine Ecosystems Research Laboratory (MERL) have confirmed that simple Vollenweider type relationships can be found between the rate of input of inorganic nutrients and annual mean chlorophyll concentrations and primary production. However, much of the coastline of the U.S. is characterized by estuarine ecosystems that are very shallow, and where most of the primary production is carried out by angiosperms, such as eelgrass, Zostera marina, epiphytic algae, drift and attached macroalgae, and epibenthic microalgae, rather than by phytoplankton. We have not been able to find useful relationships between nutrient input and the type of plant providing most of the primary production or between nutrient input and the amount of primary production in such shallow lagoon systems. Attempting to adjust nutrient loading for varying hydraulic residence time did not improve the models. Experimental studies using shallow lagoon mesocosms have shown that there is a large variation in the abundance of the various plant forms in these very shallow systems, and that simple Vollenweider models are not likely to emerge for this type of environment. However, it does seem that total system production increases with nutrient enrichment at very low rates of input, and that eelgrass does not persist when exposed to even moderate levels of fertilization. Zostera responds to inorganic nitrogen enrichment and to shading by increasing the rate of leaf elongation and decreasing the allocation of resources to below ground roots and rhizomes. This reduces or eliminates lateral branching of the rhizomes and causes a decline in the density of shoots. Based on mesocosm studies, we propose several indicators of eelgrass health, including the rate of leaf elongation, plant density, and the shoot: root biomass ratio that all deserve further study and field testing.