Local Regime Shifts Prevent Natural Recovery and Restoration of Lost Eelgrass Beds Along the Swedish West Coast

Along the Swedish northwest coast, over 60% of the eelgrass meadows have been lost since the 1980s. Despite improved water quality, no recovery has occurred, and restoration is presently considered to mitigate historical losses. However, the factors preventing natural recovery of eelgrass are not known, and it is not clear if conditions would allow restoration. Here, we present the results from 5 years of field studies with the aim of identifying the key processes affecting eelgrass growth and survival at historical eelgrass areas. Continuous light measurements and comparison with historic eelgrass distribution indicate that maximum depth distribution has decreased locally with 1.5–2.3 m in areas that have lost large eelgrass beds in the last 10–30 years. Field studies suggest that wind-driven local resuspension of sediments that are no longer stabilized by eelgrass beds is the main cause behind the deteriorated light conditions. Field experiments show that a combination of low light condition and disturbance from drifting algal mats prevents eelgrass recovery in these areas, whereas the sulfide intrusion from the sediment and dislodgement of shoots by waves had little effect on growth and survival. These results suggest that local regime shifts acting on a scale of 40–200 ha have occurred after the loss of eelgrass beds, where increased sediment resuspension and proliferation of drifting algal mats act as feedback mechanisms that prevent both natural recovery and restoration of eelgrass. The feedbacks appear to be interacting and causing an accelerating loss of eelgrass that is presently spreading to neighboring areas.     

A Dynamic Model of an Estuarine Invasion by a Non-Native Seagrass

Mathematical and simulation models provide an excellent tool for examining and predicting biological invasions in time and space; however, traditional models do not incorporate dynamic rates of population growth, which limits their realism. We developed a spatially explicit simulation model that allows patch or population growth rate to change with population size through the incorporation of field data. We used the model to evaluate the invasion of a west coast estuary by the non-indigenous Japanese eelgrass, Zostera japonica (Zosteraceae). Specifically, we tested the relative importance of stochastic, abiotic disturbance, interspecific competition, and vegetative and seedling survival. Our model predicted that vegetative shoot and seedling survival limited by competition are the most important limiting factors for Z. japonica growth, although stochastic disturbance was also a limiting factor. Population cycles and patchy distribution were also predicted, with the eelgrass apparently coexisting with the competitor. The model should be applicable to a variety of invasive species, with various types of disturbance and limiting factors.     

Morphological variation and habitat modification are strongly correlated for the autogenic ecosystem engineer<Emphasis Type="Italic">Spartina anglica</Emphasis> (common cordgrass)

We explored to what extent morphological variation and habitat modification are correlated for an autogenic ecosystem engineer, which is an organism that modifies its habitat via its own physical structures. The intertidal salt marsh speciesSpartina anglica is well known for its capacity to enhance sediment accretion within its canopy by reducing hydrodynamic energy. Sediment accretion is favorable toSpartina, as it reduces inundation stress, enhances soil drainage, and enhances nutrient availability. Shoot density and clonal architecture showed a large variation that was strongly correlated with the marsh elevation and sediment type. This correlation showed that at the lowest elevations at the muddy site,Spartina tussocks had the highest shoot density, which is known to be favorable for sediment accretion by reduction of hydrodynamic energy. There was also a strong positive correlation between the amount of sediment that accumulated within a tussock and gully formation around thatSpartina tussock. The tussocks at the lowest elevations at the muddy site had the lowest lateral tussock growth. At the highest elevations at the sandy site, stem densities were lower and there was a relatively high rate of clonal expansion and marsh formation. At this location, we also observed tussock mortality due to erosion of several of these openly structured tussocks. Based on the observed correlations, we hypothesize that the morphology ofSpartina represents trade-offs between the capacity to maximize habitat modification through sediment trapping by having a high shoot density versus the capacity to maximize clonal expansion by spreading shoots widely and the capacity of maximal clonal expansion by spreading shoots widely versus the risk of tussock mortality due to insufficient modification of the habitat that makes the tussock vulnerable to erosion. Our results indicated that morphological variation and habitat modification are strongly correlated for the autogenic ecosystem engineerS. anglica.     

A metabolism-based trophic index for comparing the ecological values of shallow-water sediment habitats

We determined fluxes of oxygen and nutrients between water and sediments at 21 sites primarily in Virginia and North Carolina estuaries, over the past 15 yr. These sites represented broad ranges in salinity, tidal amplitude, hydrology, nutrient availability, turbidity, light availability, depth, sediment grain size, and anthropogenic disturbance. In general, we found that heterotrophically dominated sediments had the potential to degrade water quality, whereas photoautotrophy in the sediments ameliorated this impact. We propose a benthic trophic state index as a management tool to make general assessments of the degree to which sediments support ecological processes related to photoautotrophy. The index can be based on simple measurements of metabolic parameters. We also evaluated the relative significance of variability in the index across a number of spatial and temporal scales. Reduced photoautotrophy and/or enhanced heterotrophy tended to be associated with finer-grained, organic-rich sediments. This sediment type was common in oligohaline areas at water depths exceeding 2 m. Temporally, autotrophy declined from winter to spring particularly at sandy sites, while interannual variability was more pronounced for mud sites. *** DIRECT SUPPORT *** A01BY074 00011     

Sediment-water exchange of dissolved nutrients at an intertidal site in the upper reaches of the Bay of Fundy

Concentrations of dissolved nitrate, silicate, and phosphate in water flooding intertidal sediments at Pecks Cove and along the axis of Cumberland Basin, Bay of Fundy were measured throughout the year. Exchanges of dissolved nutrients between intertidal sediments and overlying water were measured by enclosing water in chambers over undisturbed sediment. Nitrate concentrations in the water usually decreased during incubations while silicate was released by sediments during summer and consumed during fall. Particles which settled in sediment traps exposed during periods of high tide were stirred in filtered seawater to measure nutrient exchange. The flux of nutrients between the intertidal sediments and settled particles and seawater was estimated from incubation experiments and the observed nitrogen content in surface sediments and suspended particulate material. There was a net import of dissolved nitrate and silicate into Cumberland Basin from Chignecto Bay during early summer, at all other times there was a net export. Despite the low primary productivity and rigorous physical environment, biological activity has a measurable impact on dissolved nutrient concentrations in the waters of Cumberland Basin.     

Factors influencing spatial and annual variability in eelgrass (Zostera marina L.) meadows in Willapa Bay, Washington, and Coos Bay, Oregon, estuaries

Environmental factors that influence annual variability and spatial differences (within and between estuaries) in eelgrass meadows (Zostera marine L.) were examined within Willapa Bay, Washington, and Coos Bay, Oregon, over a period of 4 years (1998–2001). A suite of eelgrass metrics were recorded annually at field sites that spanned the estuarine gradient from the marine-dominated to mesohaline region of each estuary. Plant density (shoots m^?2) of eelgrass was positively correlated with summer estuarine salinity and inversely correlated with water temperature gradients in the estuaries. Eelgrass density, biomass, and the incidence of flowering plants all increased substantially in Willapa Bay, and less so in Coos Bay, over the duration of the study. Warmer winters and cooler summers associated with the transition from El Niño to La Niña ocean conditions during the study period corresponded with this increase in eelgrass abundance and flowering. Large-scale changes in climate and nearshore ocean conditions may exert a strong regional influence on eelgrass abundance that can vary annually by as much as 700% in Willapa Bay. Lower levels of annual variability observed in Coos Bay may be due to the stronger and more direct influence of the nearshore Pacific Ocean on the Coos Bay study sites. The results suggest profound effects of climate variation on the abundance and flowering of eelgrass in Pacific Northwest coastal estuaries.     

Compression behaviour of minerogenic low energy intertidal sediments

Existing geotechnical approaches that describe volumetric changes in intertidal sediments in response to applied vertical effective stresses are limited by a lack of empirical research into their one-dimensional compression behaviour. In this paper we address this deficiency by presenting the results of an investigation into the compression behaviour of minerogenic low marsh and tidal flat sediments. We have tested samples of these sediment types obtained from Greatham Creek (Cowpen Marsh, Tees Estuary, UK). Analysis of physical properties and oedometer compression tests demonstrates that, contrary to the implicit assumptions of existing models, the surface sediments studied are overconsolidated. Structural variability between samples arises due to sedimentological factors, notably variations in organic content. We attribute overconsolidation to tidal exposure and falls in groundwater level that permit desiccation and cause capillary suction stresses. Greater rates of compression with respect to effective stress occur in sediments with higher initial voids ratios and more open, unstable initial structures. Variability in structure decreases with application of higher effective stresses due to the destructuration of the sediments, which also creates increased homogeneity of compression behaviour under higher effective stresses. We subsequently develop a new conceptual framework to describe compression behaviour in minerogenic intertidal sediments that incorporates overconsolidation. We advocate a statistical approach that accounts for structural variability and variations in compression behaviour at effective stresses less than and greater than the yield stress. We argue that our conceptual framework is broadly applicable to minerogenic intertidal sediments at different locations and burial depths within Holocene stratigraphic sequences providing site-specific compression data are collected. Inter-site transfer and application of measured material properties should not be undertaken due to local variations in compression behaviour resulting from varying ecological, sedimentological, geochemical, climatic, geomorphic and hydrographic conditions. The individual characteristics of different field locations should be carefully considered before the suggested framework is routinely applied.     

江苏王港潮间带表层沉积物特征及输运趋势

潮滩沉积物粒度参数可以用来反映沉积物沉积的水动力条件和沉积环境。根据对江苏王港潮间带采集的15 4个表层样品的粒度资料进行分析,结果表明,沉积物类型以粉砂为主;随着水动力环境由海向陆的减弱,沉积物粒度参数表现出有规律的变化趋势,即由岸向海粒径具有粗化趋势,分选变好,逐渐正偏并达到近对称。研究区与世界其他地区的潮滩在沉积特征上有较大差异,主要是由于沉积物来源、水动力环境及受生物作用的强弱程度不同所致。沉积物输运趋势分析显示,该地区沉积物输运主要受当地潮波特征、大型人工建筑物、地貌形态及生物作用的影响;在低潮线附近主要向西北方向输运,随着向陆地方向的延伸,沉积物由向西输运逐渐转变为向南输运。     

Vertical disturbance of sediments by horseshoe crabs (Limulus polyphemus) during their spawning season

Biological and physical reworking of sediments can profoundly affect the structure and functioning of benthic communities. The depth of the disturbance is an important factor that controls the types of organisms that can exist within the sediments. Large numbers of horseshoe crabs,Limulus polyphemus, spawn each spring on the sandy shores of Delaware Bay beaches. We have used this abundance peak to provide an estimate of the depth of sediment disruption caused by this species on an intertidal flat adjoining a major spawning beach. Vertical columns of marked sediment were placed in three locations of an intertidal flat. Some columns were protected with cages while others remained unprotected. Analysis of variance of the depth of disruption of the marked sediment indicated that different areas of the intertidal zone were disrupted to different depths. Caged marked sediment was disturbed to an average depth of 3.2 cm while unprotected sediment was disturbed to a mean depth of 11.1 cm. Deepest mixing occurred in a trough between sandwave crests and averaged 17.7 cm deep. These mixing depths are greater than the 1.2 cm deep disturbance produced by nonstorm wave action in the study area.     

Short-term morphological change and sediment dynamics in the intertidal zone of a macrotidal beach

This paper describes the morphological and sedimentological evolution of a macrotidal beach over a 20 day period under varying hydrodynamic conditions (significant breaker heights of 0·3–2 m and tidal ranges of 2–5 m). During the field campaign, an intertidal bar developed around the mid‐tide level, migrated onshore, welded to the upper beach and was then flattened under energetic wave conditions. The bar had a wave breakpoint origin and its formation was triggered by a reduction in tidal range, causing more stationary water‐level conditions, rather than an increase in wave height. Most of the onshore bar migration took place while the bar was positioned in the inner to mid‐surf zone position, such that the bar moved away from the breakpoint and exhibited ‘divergent’ behaviour. The depth of disturbance over individual tidal cycles was 10–20% of the breaker height. Such values are more typical of steep reflective beaches, than gently sloping, dissipative beaches, and are considered to reflect the maximum height of wave‐generated ripples. The grain size distribution of surficial sediments did not vary consistently across the beach profile and temporal changes in the sedimentology were mostly unrelated to the morphological response. The lack of clear links between beach morphology and sedimentology may be in part due to shortcomings in the sampling methodology, which ignored the vertical variability in the sediment size characteristics across the active layer.     

The effects of semi-lunar spring and neap tidal change on nutrients cycling in the intertidal sediments of the Yangtze estuary

During a semi-lunar tidal cycle from full moon till new moon, tide is characterized by the periodic change in spring and neap tide. Under the in situ conditions of light and temperature, the influence of a semi-lunar spring and neap tidal cycle on nutrient cycling in intertidal flat of the Yangtze estuary was simulated in the laboratory in July 2002. Lab experiments showed that NH₄⁺ and PO₄^3– were always released into overlying waters, while NO₃^– was directed into sediments in the permanently waterlogged systems, suggesting that the long-term waterlogged sediment acts as a significant source for NH₄⁺ and PO₄^3–, and a sink for NO₃^– in water columns. In contrast, reflooding of intertidal sediments after long-term desiccation promoted the considerable effluxes of NH₄⁺, NO₃^– and PO₄^3– into overlying waters, reflecting that the long-term exposed sediment is an important source for nutrients in overlying waters. In addition, the semi-lunar tidal cycle led to the intricate depth distribution patterns of nutrients in intertidal sediments. During long-term exposure, NH₄⁺ in sediments was quickly transformed into NO₂^– and NO₃^–, while organic P pool might be converted to slightly adsorbed and iron-bound P. Therefore, it is considered that the semi-lunar tidal circulation has the significant influence on the biogeochemical cycle of nutrients in intertidal systems.     

Effects of the intertidal burrowing crab<Emphasis Type="Italic">Chasmagnathus granulatus</Emphasis> on infaunal zonation patterns,tidal behavior,and risk of mortality

Rhythmic movements in response to tidal cycles are characteristic of infaunal inhabitant of intertidal soft-bottoms, allowing them to remain in the area with best living conditions. The effect of bioturbators as modifier of local environmental conditions and thus of gradients in intertidal habitats, has not been investigated yet. The Atlantic estuarine intertidal areas are dominated by the burrowing crabChasmagnathus granulatus that generates strong environmental heterogeneity by affecting the physical-chemical characteristics of the sediment. The comparison between intertidal areas with and without crab shows that sediments in the crab beds remain more humid, softer, and homogeneous across the intertidal and along the tidal cycle than areas without crabs. The densities of infauna were higher at high intertidal zones in crab beds than in similar areas without crabs. Infaunal organisms performed vertical movements into the sediment following the tidal cycle that were always of higher magnitude in habitats without crabs. Infaunal species tend to spend most of the time buried into the sediment in the crab bed. Migratory shorebirds use the Atlantic estuarine environments as stopover or wintering sites. They feed (mainly on polychaetes) in the low intertidal zones of both habitats (with and without crabs), but they also feed in the upper intertidal of the crab bed; polychaete per capita mortality rate is higher in the upper part of the crab bed. Environmental heterogeneity produced by crab disturbance has an effect on the infaunal behavior, risk of mortality, and the zonation pattern. This is another example of the ecosystem engineering ability of a burrowing intertidal species.     

波浪导致黄河口海床沉积物超孔压响应现场试验研究

黄河口海床特殊的工程地质性质与复杂的工程动力稳定性问题,均与海床沉积物在波浪荷载作用下的孔压动力响应密切相关。在现代黄河水下三角洲潮间带岸滩选择4个典型研究点,现场模拟波浪作用对原状海床沉积物实施循环加载,利用孔隙水压力观测、沉积物强度测试、样品采集与实验室土工测试等方法手段,测定黄河口原状海床沉积物在循环荷载作用不同阶段的孔压响应与强度变化。研究发现,黄河口原状海床沉积物在经历循环加载过程中,典型的超孔压响应可分为逐渐累积、部分消散、快速累积、累积液化和完全消散5个阶段,分别对应沉积物强度的衰减、增大、衰减、丧失和恢复过程,沉积物的粒度组成与结构性强弱决定了超孔压的具体响应模式。波浪导致原状海床液化深度受沉积物的干密度、孔隙比、饱和度等初始物理性质影响显著,细颗粒组分的相对含量高低也在很大程度上控制着沉积物的液化特性。     

Small-scale heterogeneity in the geochemistry of seagrass vegetated and non-vegetated estuarine sediments: causes and consequences

In addition to nutrient and light availability, sedimentary biogeochemical processes can play an essential role in seagrass productivity. Previous investigations of the interactions between seagrasses and their underlying sediments have failed to clearly identify the spatio-temporal variability of the major geochemical parameters involved. Dissolved and solid phase chemical parameters in eelgrass vegetated and nearby non-vegetated sediments were investigated in this study to determine their vertical, lateral, and temporal distributions. Solid-state microelectrodes were used to investigate dissolved O₂, ΣH₂S, Fe²⁺, and Mn²⁺ on mm space scales. In this study, spatial heterogeneity was assessed and diurnal “ventilation” by seagrass productivity (i.e., the translocation of photosynthetically produced oxygen to the anoxic sedimentary environment) was not observed probably because benthic infaunal activity (bioturabation and bioirrigation) and microzones established by microbial processes led to highly heterogeneous sediment geochemistry where temporal variability was obscured by small-scale spatial variability. Non-vegetated sediments were less geochemically variable laterally than vegetated sediments, however, in some cases, they had similar vertical variability, possibly because they had been vegetated at an earlier time. This study demonstrates that in vegetated sediments where there is also substantial benthic macrofaunal activity it is difficult to separate the impacts of the two types of biota on sediment geochemistry and their spatial patterns, and it also raises the question of the applicability of traditional one-dimensional diagenetic models for such spatially–temporally complex sediments.     

Regulation of eelgrass (Zostera marina) cover along depth gradients in Danish coastal waters

A large data set, collected under the national Danish monitoring program, was used to evaluate the importance of photon flux density (PFD), relative wave exposure (REI), littoral slope, and salinity in regulating eelgrass cover at different depth intervals in Danish coastal waters. Average eelgrass cover exhibited a bell-shaped pattern with depth, reflecting that different factors regulate eelgrass cover at shallow- and deep-water sites. The multiple logistic regression analysis was used to identify regulating factors and determine their role in relation to eelgrass cover at different depth intervals. PFD, REI, and salinity were main factors affecting eelgrass cover while littoral slope had no significant effect. Eelgrass cover increased with increasing PFD at water depths of more than 2 m, while cover was in versely related to REI in shallow water. This pattern favored eelgrass cover at intermediate depths where levels of PFD and REI were moderate. Salinity had a minor, but significant, effect on eelgrass cover that is most likely related to the varying costs of osmoregulation with changing salinity. The analysis provided a useful conceptual framework for understanding the factors that regulate eelgrass abundance with depth. Although the regression model was statistically significant and included the factors generally considered most important in regulating eelgrass cover, its explanatory power was low, especially in shallow water. The largest discrepancies between predicted and observed values of cover appeared in cases where no eelgrass occurred despite sufficient light and moderate levels of exposure (almost 50% of all observations). These discrepancies suggest that population losses due to stochastic phenomena, such as extreme wind events, played an important regulating role that is not adequately described by average exposure levels. A more thorough knowledge of the importance of such loss processes and the time scales involved in recovery of seagrass populations after a severe disturbance are necessary if we are to understand the regulation of seagrass distribution in shallow coastal areas more fully.     

自然和人类活动作用下月湖沉积物有机质来源及其演变

潟湖海湾可以保存环境演化和人类活动的良好记录,碳氮元素含量及同位素特征可用来有效地反演这些演化。对位于山东半岛的月湖柱状样和沿岸表层样总有机碳、总氮、稳定碳氮同位素的分析表明,陆源物质对月湖沉积物和有机质的贡献很小,其沉积物中有机质的主要来源是维管植物大叶藻和各种海洋藻类。在2002年前大约140 a的沉积历史中,大叶藻对沉积物有机质的贡献比例呈上升趋势,其中在约1930年和1980年有两次明显的上升,前者可能因为种群的快速繁育,后者则可能是1979年口门筑坝后由于湾内环境恶化,大叶藻开始大量死亡的表现。月湖海洋藻类的有机碳埋藏通量在沉积记录中保持相对稳定,而大叶藻则增加了一倍,但两者都有一定程度的波动。研究结果表明自然环境演变（如沙嘴生长、口门缩小）和人类活动（如筑坝、修筑虾池）对月湖生态环境及有机碳埋藏的影响在沉积物中有清晰的记录。     

Effect of Sediment Nutrient Enrichment and Grazing on Turtle Grass <Emphasis Type="Italic">Thalassia testudinum</Emphasis> in Jobos Bay,Puerto Rico

We examined the effect of nutrients and grazers on Thalassia testudinum in Jobos Bay, Puerto Rico by fertilizing sediment and manipulating grazer abundances. Bottom-up effects were variable: Added nutrients did not increase seagrass aboveground biomass, but decreased belowground biomass—perhaps as a result of less biomass being allocated to belowground structures in response to greater nutrient supply in porewater. Experimental fencing of 1.5 × 1.5 m plots provided shelter that attracted large aggregations of fish, including seagrass herbivores. Seagrass biomass and shoot density decreased with increasing abundance of herbivorous fish, indicating a significant top-down effect. There were interactions between nutrient supply, provision of shelter, and grazing pressure. Fertilization enhanced seagrass %N; however, %N also increased in unfertilized plots that were fenced, most likely due to uptake of N excreted from the large numbers of fish associated with the fences. Only plots where shelter was provided and fertilizer was applied to sediments exhibited evidence of heavy grazing, reducing both seagrass cover and aboveground biomass. In the unfertilized fenced plots, signs of grazing were fewer despite large abundances of fish and enhanced nutritional quality of seagrass leaves. This suggests the possibility that high nutrient availability in sediments lowered concentrations of chemical defense compounds in the seagrass and that cues other than %N may have been involved in stimulating grazing. This study highlights the complexity of bottom-up and top-down interactions in seagrass systems and the important role of refuge availability in shaping the relative strengths of these controls.     

Genetic diversity and structure of natural and transplanted eelgrass populations in the Chesapeake and Chincoteague Bays

The objective of this study was to gain baseline population data on the genetic diversity and differentiation of eelgrass (Zostera marïna L.) populations in the Chesapeake and Chincoteague bays. Natural and transplanted eelgrass beds were compared using starch gel electrophoresis of allozymes. Transplanted eelgrass beds were not reduced in genetic diversity compared with natural beds. Inbreeding coefficients (F_IS) indicated that transplanted eelgrass beds had theoretically higher levels of outcrossing than natural beds, suggesting the significance of use of seeds as donor material and of seedling recruitment following transplantation diebacks. Natural populations exhibited very great genetic structure (F_ST=0.335), but transplanted beds were genetically similar to the donor bed and each other. Genetic diversity was lowest in Chincoteague Bay, reflecting recent restoration history since the 1930s wasting disease and geographical isolation from other east coast populations. These data provide a basis for developing a management plan for conserving eelgrass genetic diversity in the Chesapeake Bay and for guiding estuary-wide restoration efforts. It will be important to recognize that the natural genetic diversity of eelgrass in the estuary is distributed among various populations and is not well represented by single populations.     

Restored Eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) as a Refuge for Epifaunal Biodiversity in Mid-Western Atlantic Coastal Bays

As nearshore ecosystems are increasingly degraded by human activities, active restoration is a critical strategy in ensuring the continued provision of goods and services by coastal habitats. After being absent for nearly six decades, over 1800 ha of the foundational species eelgrass (Zostera marina L.) has been successfully re-established in the coastal bays of the mid-western Atlantic, USA, but nothing is known about the recovery of associated animal communities in this region. Here, we determine the patterns and drivers of functional recovery in epifaunal invertebrates associated with the restored eelgrass habitat from 2001 to 2013. After less than a decade, the invertebrate community in the restored bed was richer, more even, and exhibited greater variation in functional traits than a nearby reference bed. Analysis of a suite of environmental and physical variables using random forests revealed these differences were primarily due to the increasing area and density of eelgrass, a direct consequence of ongoing restoration efforts. Based on analysis of functional traits, we propose that the rapid life histories of constituent organisms may have played a key role in their successful recovery. We also speculate that diverse epifaunal communities may have contributed to the restoration success through a well-described mutualism with eelgrass. Given that restored eelgrass now make up 32 % of total seagrass cover in the mid-Atlantic coastal bays, this restoration may conserve regional biodiversity by providing new and pristine habitat, particularly given the general decline of existing eelgrass in this region.     

The autecology and production dynamics of eelgrass (Zostera marina L.) in Netarts Bay,Oregon

Changes in biomass, growth form and shoot net primary production in an eelgrass, Zostera marina L., bed were monitored along transects at three tidal heights in Netarts Bay, Oregon, from May 1979 through June 1981. During the growing season, April through October, the mean plastochrone interval was 16.5 d along the low intertidal transect and 11.6 d along the high intertidal transect. The mean export interval was 13.3 d along the low intertidal transect and 11.6 d along the high intertidal transect. The life span of a leaf averaged 48 d along the low intertidal transect and 36 d along the high intertidal transect. Shoot density was positively correlated with mean leaf area index (LAI) until the LAI reached 3.8 to 5.5, above which LAI was negatively correlated with density. The maximum Zostera biomass ranged from 143 (high intertidal transect) to 463 (low intertidal transect) g dry wt m^?2. Maximum values of shoot net production ranged from 4.7 (high intertidal transect) to 13.6 (low intertidal transect) g dry wt m^?2d^?1. Zostera shoot net production was related to light and to the physical damage to the shoots associated with a rapid accumulation of Enteromorpha biomass in the bay. In addition, patterns of biomass accumulation were related to the duration of water coverage, as determined by both tidal height and local impoundments of water. At all transects, biomass sloughed was equal to at least 50% of the shoot net primary production in that area during that time period; sloughed leaves accounted for 25 to 97% of these losses. An estimate of the total annual net primary production of aboveground Zostera in the bed was 17,500 kg, dry wt (SE=3,080 kg dry wt), which was equivalent to a mean annual rate of 383 g C m^?2 (SE=67 g C m^?2)