Vertical distribution and seasonal variation of eelgrass beds in Iwachi Bay,Izu Peninsula,Japan

The vertical distribution and population structure of eelgrass beds were surveyed in Iwachi Bay, along the Pacific coast of central Japan. Samplings were conducted from May through November 1977 by SCUBA. Eelgrass was distributed between 3 and 11 m in depth. The relative light intensity at 12 m depth was 11% at the lower range. The highest population density was 290 shoots/m² in September and the fresh weight of biomass was 888 g/m² in July at 7 m depth. The maximum mean leaf area index was about 3 at 10 m depth in July. The ratio of reproductive shoots to the total shoots was about 36% at 7 m depth in June. Eelgrass showed good growth at 7–10 m depth, which is comparatively deeper than other eelgrass habitations. The high values of water transparency and sunshine duration, as well as solar radiation compared with other localities was believed to contribute to the growth of eelgrass in deeper waters in Iwachi Bay.     

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     

鳗草在荣成天鹅湖不同生境中生长的适应性

鳗草在北半球温带海域广有分布,受到不同程度的人为干扰和环境胁迫,海草床面积急剧退化.为探讨鳗草对不同环境胁迫的适应性,于2014年11月—2015年10月对荣成天鹅湖草床的潮间带和潮下带的斑块区、鳗草草场边缘和鳗草草场内部4个生境的鳗草形态和繁殖能力进行了研究.结果表明: 叶长、叶宽、节间长度/直径、地上生物量/地下生物量和分枝频率在不同月份和生境间差异显著,其中除了分枝频率最大值出现在草场边缘(88.4%)外,其他各项指标的最大值均出现在草场内部(分别为78.54 cm、7.93 mm、7.03和3.88).地上生物量/地下生物量的可塑性指数在各站位均较高(0.77～0.92),叶宽的可塑性(0.41～0.64)略小.不同生境中单株鳗草佛焰苞数差异不显著,而单位面积的佛焰苞数差异显著,在草场内部偏向于克隆生殖,而在人为干扰较大的斑块区倾向于有性生殖.     

Effects of edge/interior and patch structure on reproduction in Zostera marina L. in Chesapeake Bay,USA

We examined the effects of location (patch edge vs. interior) and shoot density (individual, patchy, continuous) on reproduction in three natural and two transplanted Chesapeake Bay (USA) stands of the submerged marine angiosperm Zostera marina L. (eelgrass; Zosteraceae). There were no edge effects on demographic-based reproductive effort or reproductive output (propagule production), and patch structure (individual, patchy, continuous) alone never accounted for the majority of variability in any metric. Transplant site was the most important predictor of eelgrass reproduction response, and relationships among metrics were predictable within sites. Our results suggest that, in Chesapeake Bay eelgrass, environmental factors acting at a regional scale (km) have a stronger impact on reproductive investment than those at a patch scale (1–10 m). Since tradeoffs between clonal and sexual production are mediated primarily by exogenous environmental factors, site selection may be more critical than transplant configuration for producing self sustaining stands, and achieving long-term restoration success.     

Using Acoustics to Determine Eelgrass Bed Distribution and to Assess the Seasonal Variation of Ecosystem Service

Eelgrass beds are an important source of primary production in coastal ecosystems. Understanding seasonal variation in the abundance and distribution of eelgrass is important for conservation, and the objectives of this study were to 1) monitor seasonal variation in eelgrass beds using an acoustic monitoring method (Quantitative echo sounder) and 2) broadly quantify the carbon circulation function. We obtained acoustic data of eelgrass beds in coastal areas north and east of Ikunojima Island. Surveys were conducted nine times over the 3-year period from 2011 to 2013 in order to monitor seasonal variation. Acoustic data were obtained and used to estimate the spatial distribution of eelgrass by geostatistical methods. To determine supporting services, we determined carbon sink and carbon fixation by eelgrass beds using data from the National Research Institute of Fisheries and Environment of Inland Sea (2011). The height and distribution of eelgrass beds were at a maximum in May and at a minimum in November of each year. Distribution trends were different between the north and east areas. Supporting services showed the same patterns throughout the year. The area of distribution was considered to be coincident with the life history of eelgrass. Distribution differed by area and changed yearly due to the effects of bottom characteristics and wind direction. Quantifying the supporting services of eelgrass beds was shown to be useful for managing the conservation of coastal ecosystems.     

Seasonal growth and reproduction of Ruppia maritima L. s.l. in Lake Pontchartrain,Louisiana, USA

Ruppia maritima L. s.l. is an important submerged aquatic plant in Lake Pontchartrain, a water quality impaired oligohaline estuary located north of New Orleans, Louisiana, USA. Efforts to restore Lake Pontchartrain are in progress, and a major restoration goal is the re-establishment of historic submerged macrophyte beds that have declined since first studied in 1953. This study was part of a long-term monitoring program to distinguish natural from anthropogenic causes of macrophyte population changes. Ruppia populations were monitored monthly for a 2-year period to obtain information on seasonal growth, peak biomass periods, flowering, seed production, and germination. We found that Lake Pontchartrain R. maritima had relatively high peak biomass and two seasonal peaks of biomass and flowering due to high water temperature and absence of competition from other macrophytes. However, biomass peaks were often disrupted by storms. Shoot biomass was high from May through November, and low from December through April. Spring shoot growth from seeds, roots, and rhizomes was rapid. Plants matured and produced abundant inflorescences and seeds between May and October. Reproductive shoots bearing seeds were detached in May and June. Remaining shoots formed dense beds through vegetative propagation during the summer. Turion production occurred during the winter.     

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.     

Quantifying the Effects of Green Crab Damage to Eelgrass Transplants

Mesocosm experiments were conducted in the summer of 1996 to quantify the effect of bioturbation by Carcinus maenas (the introduced European green crab) on survival of transplanted Zostera marina (eelgrass). The research grew out of a successful 2.52 ha eelgrass transplant project in the Great Bay Estuary of New Hampshire. At several subtidal sites, green crabs were found to damage transplanted eelgrass by cutting the shoots to the extent that some sites demonstrated poor survival. In three separate experiments, eight replicate mesocosm tanks were transplanted with 36 shoots of eelgrass, and different crab densities were introduced into the tanks. The number of shoots damaged by crabs was significantly higher in tanks with moderate (4.0 crabs/m²), high (7.0 crabs/m²), or very high (15.0 crabs/m²) crab densities than in tanks with low (1.0 crabs/m²) crab densities. Up to 39% of viable shoots were lost within one week of exposure to green crab activities. The mesocosm results demonstrated that green crabs were not directly attracted to eelgrass but that they significantly decreased transplant survival through their activity. Field densities of green crabs were found to exceed the density at which most damage occurred in the experiments, suggesting that this introduced species can be a major determinant of eelgrass transplant survival. The results underscore the major influence that biological components of transplant sites can have on transplant survival, and the need for their consideration in the site selection process.     

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.     

Population Genetic Analyses of Transplanted Eelgrass (Zostera marina) Beds Reveal Reduced Genetic Diversity in Southern California

Seagrass ecosystems fulfill ecologically and economically valuable functions in coastal marine environments. Unfortunately, seagrass beds are susceptible to natural and human disturbances, and their distrubution is declining worldwide. Although intentional disturbance of seagrass beds must be mitigated pursuant to U.S. law, to date mitigation of seagrass beds has not prevented a net loss of habitat. Transplantation of vegetative material from small areas of nearby beds is the primary method of seagrass mitigation. Restoration research on seagrasses has focused primarily on establishment of the plants and secondarily on the functional equivalency of the habitats. We questioned whether transplanted seagrass beds were comparable to “natural” beds in terms of genetic diversity and structure. We sampled Zostera marina L. (eel-grass) from 12 sites in the highly urbanized area of San Diego County and from pristine sites in Baja California. Using allozyme electrophoresis, we determined that genetic diversity (percentage of polymorphic loci, allele richness, expected and observed heterozygosities, and proportion of genetically unique individuals) was significantly reduced in transplanted eelgrass beds. Eelgrass from Baja California exhibited the highest genetic diversity. Based on Wright's F statistics, most of the genetic variation was distributed within rather than among sites (F_ST= 0.139), and the degree of genetic structure was only moderate at the greatest geographical scale (San Diego—Baja). Using a spatial statistical analysis (second-order analysis), we found virtually no evidence for nonrandom distribution of alleles or genotypes at scales of 3–50 m within beds. We discuss several hypotheses for reduced genetic diversity in transplanted eelgrass beds, including transplantation protocol, small size of transplantations, and reduced or failed sexual reproduction.     

A simulation model of production,seasonal changes in biomass and distribution of eelgrass (Zostera marina) in Lake Grevelingen

Summary A simulation model has been described, based on data from Lake Grevelingen, The Netherlands, as a predictive tool for lake management. The model has been developed as part of a large-scale aquatic modelling effort in Lake Grevelingen, carried out in close cooperation with the Delft Hydraulics Laboratory and the Delta Department, Environmental Research Division of the Ministry of Transport and Public Works. Available data on growth rates per unit eelgrass biomass, obtained with the leaf-marking technique, and on above- en below ground biomass and shoot density changes per unit area have been used. A space limitation depending on density of the above ground biomass and a growth limitation due to shortage in below ground biomass have been introduced. The seasonal changes in eelgrass production, both above and below ground, have been simulated as functions of the external forcing variables light, water temperature, wind generated water movements and of the internal control variables due to aging of the plant material. The vertical distribution of eelgrass can be partly explained from the modelling results on space, light and below ground biomass limitations. From the shore down to about 1 m waterdepth the above ground eelgrass biomass suffers from space limitation. Between 1 and 2 m production and biomass reach maximum values. Between 2 and 3 m waterdepth the above ground eelgrass growth is limited by the availability of below ground biomass. Between 3 and 5 m waterdepth both below ground biomass and light are the growth-limiting factors. Below 5 m waterdepth light is not sufficient to sustain net growth of eelgrass from rhizomes. Together with additional data — not used in the model — on seed production and growth of eelgrass shoots from seeds the vertical and horizontal distribution of the dominant macrophyte in the lake can be explained.     

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.     

Growth dynamics of Zostera in Sevastopol Bay (Crimea,Black Sea)

Density, biomass and primary productivity of eelgrass (Zostera marina L.) and density and biomass of Zostera noltii Hornem. were studied at three locations in Sevastopol Bay, Crimean Peninsula, Ukraine, in April 2002, late September 2002, and late January 2003 at 3 m and 5 m depths. All three growth parameters varied by station, depth, and month. The aboveground biomass (AGB) was greater in September than in April or January at 3 m. It was greater at 5 m than at 3 m in April at one station. For Z. noltii, results were more variable as to month. For belowground biomass (BGB), month and station relationships were not clear cut. It was generally true that total biomass was greatest in September. Significant differences in density, AGB and total biomass were observed for station and month in eelgrass, while in BGB, these differences were significant for depth and for month. Primary production in eelgrass was greatest at both depths in April as compared to September or January. Reproductive shoots were found only during April. Overall, seedlings were found in all collection periods and depths, but were most abundant at the three stations in April.     

Effects of hyposalinity in Thalassia testudinum (Hydrocharitaceae) from Parque Nacional Morrocoy, Venezuela

The year 1996 had a high pluviosity in Morrocoy National Park (western coastal zone, Venezuela) and low salinity in December 1996 affected the seagrass beds, dominated by Thalassia testudinum. Patches without T. testudinum were observed in localities of the park that used to have very dense populations of this plant. Sampling was done at Las Luisas to determine leaf productivity, turnover rate, short shoot density and relative biomass of plant sections, in order to compare with data obtained in September 1996, previous to the event. Green leaves, roots and rhizomes were the most affected parts. Mean green leaf biomass decreased in January and February 1997 to 5% of the plant's total biomass; mean root biomass decreased in March to 40% and mean rhi zome biomass decreased in February to 30%. The density of the active short shoots decreased to a minimum in February, but in April it reached a value similar to that of September 1996. The density of the inactive short shoots decreased to a minimum in March, and they dissappeared in April, matching the increasing density of the active short shoots between these two months. In February 1997, 56% of the inactive short shoots showed evidence of leaf initiation. In January 1997 the leaf productivity and turnover rate values (2.72 +/- 0.35 g/m2/d and 2.15% leaf DW/d) were similar to the annual mean previously determined from Las Luisas (2.35 +/- 0.72 g/m2/d and 1.96% leaf DW/d). Nevertheless, the values of productivity and turnover rate detected at Las Luisas in April 1997 (4.88 +/- 2.14 g/m2/d and 4.66% leaf DW/d) were higher than those values previously reported for this location. In response to the mortality episode, the leaf productivity and turnover rate of T. testudinum increased and the leaf initiation was activated in the inactive short shoots.     

Growing Zostera marina (eelgrass) from Seeds in Land‐Based Culture Systems for Use in Restoration Projects

The use of aquaculture systems to grow the seagrass Zostera marina (eelgrass) from seeds for restoration projects was evaluated through laboratory and mesocosm studies. Along the mid‐Atlantic coast of North America Z. marina seeds are shed from late spring through early summer, but seeds typically do not begin to germinate until the late fall. Fall is the optimal season to plant both seeds and shoots in this region. We conducted studies to determine if Z. marina seeds can be induced to germinate in the summer and seedlings grown in mesocosms to a size sufficiently large enough for out‐planting in the fall. Seeds in soil‐less culture germinated in the summer when held at 14°C, with percent germination increasing with lower salinities. Cold storage (4°C) of seeds prior to planting in sediments enhanced germination and seedling survival. Growth rates of seedlings were significantly higher in nutrient enriched estuarine sediments. Results from preliminary studies were used in designing a large‐scale culture project in which 15,000 shoots were grown and out‐planted into the Potomac River estuary in the Chesapeake Bay and compared with an equal number of transplanted shoots. These studies demonstrate that growing Z. marina from seeds is an alternative approach to harvesting plants from donor beds when vegetative shoots are required for restoration projects.     

Detecting the impacts of notorious invaders: experiments versus observations in the invasion of eelgrass meadows by the green seaweed <Emphasis Type="Italic">Codium fragile</Emphasis>

Biological invasions can vary in the extent of their effects on indigenous communities but predicting impacts for particular systems remains difficult. In coastal marine ecosystems, the green seaweed Codium fragile ssp. fragile is a notorious invader with its reputation based on studies conducted largely on rocky shores. The green seaweed has recently invaded soft-bottom eelgrass communities by attaching epiphytically to eelgrass (Zostera marina) rhizomes, thereby creating the potential for disruption of these coastal habitats through competition or disturbance. We investigated the effect of this invader on various aspects of eelgrass performance (shoot density and length, shoot growth, above- and below-ground biomass, carbohydrate storage) using both small-scale manipulative and large-scale observational experiments. Manipulative experiments that varied Codium abundance demonstrated clear negative effects over a 4-month period on shoot density and carbohydrate reserves, but only for high, but realistic, Codium biomass levels. Light levels were much lower under canopies for high and medium density Codium treatments relative to low and control Codium cover treatments, suggesting that shading may influence eelgrass growing under the algal cover. In contrast, these effects were either not detectable or very weak when examined correlatively with field surveys conducted at larger spatial scales, even for sites that had been invaded for over 4 years. It is premature to extend generalizations of Codium’s impact derived from studies in other systems to eelgrass communities; further efforts are required to assess the long-term threats that the alga poses to this ecosystem. This study demonstrates the need to investigate impacts of invasions over multiple scales, especially those that incorporate the temporal and spatial heterogeneity of the invader’s abundance.     

Rapid recovery of the intertidal seagrass Zostera japonica following intense Manila clam (Ruditapes philippinarum) harvesting activity in Korea

Although the Manila clam (Ruditapes philippinarum) culture grounds are occasionally located in Zostera japonica beds along the coasts of Korea, plant responses to the clamming activity have not been reported for this seagrass species. Intense Manila clam harvesting activity took place in the intertidal Z. japonica bed during April 2004. The Z. japonica bed at the study site has been monitored since January 2003. Thus, this study provided a unique opportunity to compare the structure of the Z. japonica population before and after the clamming activity, which was conducted for approximately 1 week in April 2004. All Z. japonica shoots were removed and buried in the sediment immediately after the clamming activity. However, a few shoots were found at the disturbed area in July 2004, 3 months after the clamming activity. By September 2004, 5 months after the disturbance, shoot density and biomass were almost recovered to the levels reported before the clamming activity. No Z. japonica seedlings were observed when the shoot density rapidly increased in August and September 2004, 4-5 months after the disturbance, because revegetation of the disturbed seagrass bed has occurred before the seed germination time which is typically winter or early spring in this area. Thus, the initial rapid revegetation of the disturbed area occurred via asexual reproduction through new shoot formation from the buried below-ground tissues. The reproductive shoot density and reproductive efforts of Z. japonica were significantly higher after the disturbance relative to the levels recorded before the disturbance, and the duration of the fertile period was approximately three times longer following the clamming activity. The belowground biomass after the disturbance was also significantly higher than that before the disturbance. These results suggest that Z. japonica allocated more energy to sexual reproduction, as well as the maintenance of belowground tissues, to persist their population under unstable environmental conditions.     

微齿眼子菜(Potamogeton maackianus A.Benn.)茎段的无性繁殖特点研究

匍匐茎型的沉水植物微齿眼子菜(Potam ogeton maackianusA.Benn.)在自然生境中的繁殖方式主要是无性繁殖,彼此相连的分株通常受到水流、草食性鱼类和人为干扰的影响导致分离或者枝条片断化,片断化的枝条在新的生境中再生进行繁殖。实验模拟微齿眼子菜枝条的片断化来验证假设:微齿眼子菜直立茎段与匍匐茎段的无性繁殖能力是否一致,茎段可能有一个最小长度来保证它的存活,并且茎段着泥的方向也会影响它的成活率。实验结果表明:直立茎段与匍匐茎段的成活率、生物量有显著差异,而且茎段的长度显著影响它们的成活率、生物量,而直立茎段与匍匐茎段克隆株的每株分枝数没有显著差异;直立茎段比匍匐茎段更易成株成活,茎段的长度越长,成活率越高,具有3个节间的茎段成活率接近100%;同时茎段着泥的方向显著影响茎段成活率、所产生抽条的株高以及可产生抽条的节数,对总生物量没有显著影响,而对根的生物量有显著影响;茎段正向着泥有利于它们的成活率、定植能力提高。     

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.     

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.