Peculiar growth dynamics of the surfgrass <Emphasis Type="Italic">Phyllospadix japonicus</Emphasis> on the southeastern coast of Korea

The surfgrass Phyllospadix japonicus is endemic to exposed shores of the northeastern Pacific Ocean. Unlike the majority of seagrasses, P. japonicus grows on rocky substrata. The specific physical features of the habitat are likely related to the peculiar ecological characteristics of P. japonicus. However, few studies have been conducted thus far on the growth dynamics of Phyllospadix spp., largely due to the turbulent water conditions in its habitat. P. japonicus is a dominant seagrass species, and it plays critical ecological roles on the eastern coast of Korea. Here, we examined its growth dynamics for the first time on the Korean coast. We measured shoot density, biomass, leaf production, phenology, morphology, tissue nutrient content, as well as environmental factors including underwater photon flux density (PFD), water temperature and water column nutrient concentrations from March 2003 to December 2005. Shoot density, biomass, leaf productivity and morphological characteristics exhibited significant seasonal variations; maximum values of these variables occurred in winter and early spring, and the minima were recorded in late summer and early fall. PFD and water temperature were, respectively, positively and negatively correlated with leaf production. Nutrient availability fluctuated substantially, but there was no evidence of distinct seasonal variation, nor was it correlated with leaf production. Leaf chlorophyll concentrations were correlated strongly with leaf production, whereas tissue nutrient contents were unrelated to leaf production. Maximum potential seed production ranged from 1,200 seeds m⁻² in 2004 to 3,445 seeds m⁻² in 2003; however, seedlings were rarely detected through the observation period. Thus, P. japonicus meadows at the study site appeared to persist through vegetative propagation. Leaf C content varied bimodally, with peaks in spring and fall. Leaf N content was minimal during months in which leaf productivity was lowest. These patterns in tissue nutrient content are clearly different from those of the majority of soft-substratum seagrasses and appear to relate to the reduced physiological tolerance of high temperature in P. japonicus compared to other temperate seagrasses.     

Significance of salinity and silicon levels for growth of a formerly estuarine eelgrass (Zostera marina) population (Lake Grevelingen, The Netherlands)

Since the early 1980s, the eelgrass, Zostera marina L., population in the saline Lake Gevelingen, The Netherlands, is rapidly declining. An earlier study, in which long-term data on eelgrass coverage in this former estuary were correlated with several environmental variables, showed only one significant correlation: coverage was positively related to water column silicon levels. In addition, a negative correlation with salinity was observed, but this was not significant. In the present study, the effect of silicon and the effect of salinity on the development of Z. marina were investigated experimentally. Enhancement of dissolved silicon concentrations in the water did not stimulate Z. marina above-ground production or an increase in final above- and below-ground biomass. The highly significant correlation between eelgrass coverage and water column silicon levels, thus, remains to be explained. The results of the growth experiments did, however, demonstrate a clear effect of salinity on Z. marina growth. Plants cultured at 22 psu showed a higher production of shoots and leaves, resulting in more above-ground biomass, than plants grown at 32 psu. In addition, below-ground biomass was also higher at 22 psu. Measurements of chlorophyll a fluorescence, performed with a PAM-fluorometer, indicated a reduction of photosynthesis in the high-salinity treatments. Thus, low salinity stimulates development of Z. marina from Lake Grevelingen. Eelgrass from such a historically estuarine area may be more sensitive to high salinities than other, more marine populations. Recovery of the autochthonous eelgrass population is expected to be favoured when the estuarine conditions of the seagrass area are re-established, or when restoration programmes are carried out with allochthonous ecotypes that are less sensitive to high salinities. Received: 23 June 1998 / Accepted: 19 November 1998     

Development of epiphytic communities on eelgrass (Zostera marina) along a nutrient gradient in a Danish estuary

The effect of nutrient enrichment on epiphyte development was examined by following the seasonal development of epiphyte biomass on eelgrass (Zostera marina L.) at four localities along a nutrient gradient in Roskilde Fjord, Denmark between March and December 1982. In the most nutrient-poor area, epiphyte biomass followed a distinct bimodal seasonal pattern with maxima in spring and early fall. Low nutrient availability and a high rate of eelgrass leaf renewal kept epiphyte biomass at a low level throughout the summer period. Unlike phytoplankton, the epiphytic community was not stimulated by nutrient enrichment during spring, however, from May through August, the biomass of both components increased exponentially with increasing concentrations of total N in the water. Along the nutrient gradient, phytoplankton biomass increased 5- to 10-fold, while epiphyte biomass increased 50- to 100-fold. Thus differences in nutrient conditions among study sites were more clearly reflected by epiphytes than phytoplankton.Contribution No. 419 from the Freshwater-Biological Laboratory, University of Copenhagen     

The role of the amphipod <Emphasis Type="Italic">Gammarus locusta</Emphasis> as a grazer on macroalgae in Swedish seagrass meadows

Mesograzers are thought to play a critical role in seagrass beds by preventing overgrowth of ephemeral algae. On the Swedish west coast, eelgrass Zostera marina has decreased in recent decades as a result of eutrophication and increased growth of macroalgal mats (mainly filamentous Ulva spp. and Ectocarpales), with no indication of grazer control of the algae. The aim of this study was to investigate the ability of the amphipod Gammarus locusta to control algal blooms during nutrient-enriched and ambient conditions, using a combination of laboratory, field and model studies. Laboratory experiments demonstrated that juvenile and adult G. locusta could consume both Ulva spp. and Ectocarpales, but that consumption of Ulva spp. was significantly higher. Cannibalism was common in individual treatments involving multiple size-classes of G. locusta, but only large, male gammarids consumed smaller juveniles in the presence of Ulva spp. as an alternative food source. However, no negative effects of cannibalism were found on total grazing impact. A model using size-specific grazing rates and growth rates of G. locusta and of Ulva spp. suggests that approximately 62 young juvenile, or 27 adult G. locusta are needed per gram DW of Ulva spp. to control the algal growth during ambient nutrient conditions, and approximately 2.6 times as many gammarids during enhanced nutrient conditions. On the Swedish west coast, densities and mean sizes of G. locusta in eelgrass beds are below these critical values, suggesting that the gammarids will not be able to control the growth of the filamentous macroalgae. However, in the field cage experiment, immigration of juveniles and reproduction of encaged adult G. locusta resulted in unexpectedly high densities of G. locusta (>4,000 individual m⁻²), and very low biomass of Ulva spp. in both ambient and nutrient-enriched treatments. Although the high numbers of juveniles in all cages precluded any significant treatment effects, this suggests that in the absent of predators, the population of G. locusta can grow significantly and control the biomass of Ulva spp. Furthermore, low grazing of Ectocarpales in the laboratory and high biomass of these filamentous brown algae in the field indicate a preference for the more palatable green algae Ulva spp. This study indicates that the high grazing capacity of G. locusta, in combination with high reproduction and growth rates, would allow the amphipod to play a key role in Z. marina ecosystems by controlling destructive blooms of filamentous green algae. However, high predation pressure appears to prevent large populations of G. locusta in eelgrass beds on the Swedish west coast today.     

Drastic decline of an extensive eelgrass bed in Nova Scotia due to the activity of the invasive green crab (Carcinus maenas)

In 2001–2002, there were severe declines of eelgrass (Zostera marina) in estuaries along the Nova Scotia coast of the southern Gulf of St. Lawrence. We examined the relationship between the recent invasion of European green crabs (Carcinus maenas) and the eelgrass decline in Benoit Cove in Tracadie Harbour. The abnormally abundant eelgrass wrack consisted mainly of entire shoots, not the usual blades. Three separate methods yielded similar estimates of the rate of shoot removal from the eelgrass beds (direct quadrat counts, “mark–recapture” of tagged shoots in the eelgrass bed, and rate of shoot appearance in the shore wrack). From 14 July to 8 September 2002, the shoot density in the bed fell by about 75 %. Similar rates of decline occurred in crab enclosures (4.4 crabs m^?2) placed in the eelgrass bed. Green crab foraging, involving the tearing of shoots and the digging of large pits, was the reason for the drastic decline of the eelgrass bed in Benoit Cove. It is clear that an invasion of green crabs to a region where they had not previously existed can both destroy eelgrass beds and restrict their recovery.     

Spatial patterns and environmental correlates in leaf-associated epifaunal assemblages of temperate seagrass (<Emphasis Type="Italic">Zostera marina</Emphasis>) meadows

We estimated and tested variability of seagrass leaf-associated epifaunal assemblages at a range of scales. Sampling was performed in 36 seagrass (Zostera marina) meadows within three regions along the Swedish west coast following a hierarchical design (samples separated by 10 s m, km or 100 km). Results showed strongest variability (43–81%) at the intermediate amongst-meadow (km) scale using biomass of functional categories, while considering taxa composition the within-meadow (10 s m) scale contributed most to variability (60%). Using functional categories, we found that embayment exposure and seagrass shoot density were the most important predictor variables explaining part of the variability in biomass of suspension feeders (bivalves and barnacles) and grazers. In contrast, variability in epifaunal taxa composition was predicted mainly by sediment chemistry, substratum coverage and geographical positioning. Our findings suggest that models to develop predictive power and mechanistic understanding should focus on variables and processes varying at small and intermediate scales rather than those varying at larger scales.     

Eelgrass fairy rings: sulfide as inhibiting agent

Distinct ‘fairy rings’ consisting of narrow fringes of eelgrass (Zostera marina L.) expand radially over a bottom of chalk plates outside the calcium carbonate cliffs of the island of Møn, Denmark. We conducted a survey to evaluate possible explanations for the formation of the rings and, more specifically, for the apparent die-off of eelgrass shoots on the inner side of the rings. The fairy rings were up to 15 m in diameter consisting of 0.3- to 1-m-wide zones of sea grass shoots at densities of up to 1,200 shoots m^?2 and rooted in an up to 10-cm-thick sediment layer. On the outer side, shoots expanded over the bare chalk plates. On the inner side, shoots were smaller, had lower absolute and specific leaf growth, shoot density was lower and the sediment eroded leaving the bare chalk with scattered boulders behind. Sediment organic matter and nutrients and tissue nutrient contents were not different among positions. Sediment pools of acid volatile sulfides and chromium-reducible sulfur increased from outer to the middle positions of the rings, and so did total sulfur content of eelgrass tissues, while tissue δ³⁴S isotope ratios, regardless of position in the fringes, were low reflecting substantial invasion of sulfide from the sediment. Neither the clonal growth pattern of eelgrass, sediment burial of shoots, hydrodynamic forcing nor nutrient limitation could explain the ring-shaped pattern. We conclude that the most likely explanation must be found in invasion of eelgrass shoots by toxic sulfide accumulating in the sediment due to low iron availability in the carbonate-rich environment.     

Growth dynamics of the seagrass <Emphasis Type="Italic">Halophila nipponica</Emphasis>, recently discovered in temperate coastal waters of the Korean peninsula

Seagrass species in the genus Halophila are usually distributed in tropical or subtropical areas, but a Halophila species identified as H. nipponica was first observed in temperate coastal regions of Korea in 2007. Since this species mainly occurs in warm temperate regions influenced by warm currents, we hypothesized that H. nipponica may exhibit different growth patterns from those of other temperate seagrass species in Korea, instead showing similar growth dynamics to tropical/subtropical species. The growth and morphology of H. nipponica in relation to coincident measurements of environmental factors were investigated from July 2008 to September 2009 to examine the growth dynamics of this species. Water temperature at the study site ranged from 9.7°C in January to 25.1°C in August. Shoot density, biomass, and productivity exhibited significant seasonal variation, increasing during summer and decreasing during winter. Productivity was severely restricted to nearly ceasing at water temperatures less than 15°C, and winter minimum growth lasted until May. The optimal temperature for H. nipponica growth was approximately 25°C, which was the maximum water temperature at the study site, and no growth reduction in high summer water temperature was observed. Thus, H. nipponica on the temperate coast of Korea exhibited a distinctly different growth pattern from those of temperate seagrass species in Korea, which have shown great reductions in growth at water temperatures higher than 20°C. Higher below- to above-ground ratio and leaf burial into sediments with shorter leaf petioles during winter might be overwintering strategies in this species. The growth patterns of H. nipponica at the study site imply that this species still possess the tropical characteristics of the genus Halophila.     

How do nutrient conditions and species identity influence the impact of mesograzers in eelgrass-epiphyte systems?

Coastal eutrophication is thought to cause excessive growth of epiphytes in eelgrass beds, threatening the health and survival of these ecologically and economically valuable ecosystems worldwide. Mesograzers, small crustacean and gastropod grazers, have the potential to prevent seagrass loss by grazing preferentially and efficiently on epiphytes. We tested the impact of three mesograzers on epiphyte biomass and eelgrass productivity under threefold enriched nutrient concentrations in experimental indoor mesocosm systems under summer conditions. We compared the results with earlier identical experiments that were performed under ambient nutrient supply. The isopod Idotea baltica, the periwinkle Littorina littorea, and the small gastropod Rissoa membranacea significantly reduced epiphyte load under high nutrient supply with Rissoa being the most efficient grazer, but only high densities of Littorina and Rissoa had a significant positive effect on eelgrass productivity. Although all mesograzers increased epiphyte ingestion with higher nutrient load, most likely as a functional response to the quantitatively and qualitatively better food supply, the promotion of eelgrass growth by Idotea and Rissoa was diminished compared to the study performed under ambient nutrient supply. Littorina maintained the level of its positive impact on eelgrass productivity regardless of nutrient concentrations.     

Feeding preferences,seasonal gut repletion indices,and diel feeding patterns of the sea urchin<Emphasis Type="Italic"> Tripneustes gratilla</Emphasis> (Echinodermata: Echinoidea) on a coastal habitat off Toliara (Madagascar)

The feeding biology of the echinoid Tripneustes gratilla from a seagrass bed off Toliara, on the south-west coast of Madagascar, was investigated during the summer and winter seasons. Repletion indices of the gut of T. gratilla varied seasonally, in such a way that higher amounts of food occurred in the gut in winter than in other seasons. Conversely, a more detailed analysis of one summer and one winter sample of the gut tissue dry weights revealed higher weights in summer (February) than in winter (August). This was interpreted as a varying capability of nutrient storage and closely related to the feeding activity of the echinoid. This inverse relationship between feeding activity and nutrient storage was suggested to be dependent on gonadal growth. Indeed, higher food consumption was observed during the post-spawning period (August), that is, when most of the energy accumulated was used up for gonadal growth. Low feeding activity occurred when a sufficient amount of energy was accumulated in gut tissues (February), the time corresponding to the initiation of gonadal growth. Investigations on the feeding habits, using Ivlev's electivity index, revealed preferential feeding of T. gratilla on Syringodium isoetifolium, the dominant seagrass species found in the gut. This was partly related to the higher availability of that particular seagrass species in the field. However, its soft and terete leaf blade's morphology also makes it easier to manipulate and ingest. Studies on the feeding behaviour indicated the presence of a diel feeding cycle in T. gratilla individuals in both summer and winter seasons. Yet, its pattern varied between the two studied months (one summer and one winter) as the cycle was less contrasted during period of active feeding, i.e. in winter compared to summer. This study provides novel information on how intrinsic factors, such as physiological state, can govern the feeding activity of T. gratilla living in a seagrass habitat.     

The contribution of seagrass beds (<Emphasis Type="Italic">Zostera noltii</Emphasis>) to the function of tidal flats as a juvenile habitat for dominant,mobile epibenthos in the Wadden Sea

Since the substantial loss of subtidal eelgrass (Zostera marina L.) in the 1930s, seagrass beds in the Wadden Sea are limited to the intertidal zone and dominated by Z. noltii Hornem. This study deals with the effect of vegetated tidal flats on quantities of mobile epifauna and proves empirically the function of seagrass canopies as a refuge for marine animals remaining in the intertidal zone at ebb tide. Drop-trap samples were taken in the Sylt-Rømø Bight, a shallow tidal basin in the northern Wadden Sea, on vegetated and unvegetated tidal flats during July and August 2002, and during the entire growth period of Z. noltii from May to September in 2003. The species composition in Z. noltii and bare sand flats showed minor differences since only two isopod species (Idotea baltica and I. chelipes) occurred on Z. noltii flats exclusively. Juvenile shore crabs (Carcinus maenas L.), brown shrimps (Crangon crangon L.) and common gobies (Pomatoschistus microps Krøyer) were also found abundantly on bare sand flats. However, the results showed significantly higher abundances and production of these dominant species on vegetated tidal flats. Additionally, the analyses of faunal size classes indicated higher percentages of small individuals in the seagrass bed during the entire sampling period. Despite drastic diurnal fluctuations of dissolved oxygen at low tide, faunal density in the residual water layer remaining in seagrass canopies at ebb tide was found to be consistently higher than that found in artificially created tide-pool units. Although species composition of mobile epifauna did not basically differ between vegetated and unvegetated tidal flats, Z. noltii beds are considered to contribute quantitatively to the function of tidal flats, as an extended juvenile habitat for some of the most important species of the Wadden Sea food web.Communicated by O. Kinne, Oldendorf/Luhe     

A model for measuring leaf growth rate of vegetative shoot in the seagrass, Zostera marina

A new model for determining leaf growth in vegetative shoots of the seagrass Zostera marina (eelgrass) is described. This model requires the weights of individual mature and immature whole leaves and leaf plastochrone interval (P_L) as parameters, differing from the conventional leaf marking technique (CLM) that requires cutting and separation between new and old tissue of leaves. The techniques required for the model are the same as for the plastochrone method, but the parameters differ between both methods in use of the weight of individual immature leaves. In a mesocosm study, eelgrass growth was examined, and parameters for the new model and plastochrone method (the weights of individual mature and immature leaves and P_L) were measured. Leaf growth rate was measured using the CLM and determined by the new method and the plastochrone method. The results were then compared between the CLM, the new model, and the plastochrone method. The results obtained with the new model were similar to those obtained with the CLM. However, the results of the plastochrone method differed from those of the CLM, while the weight of immature leaves varied seasonally. The new model was also used to determine leaf growth in a natural eelgrass bed in Mikawa Bay, Japan, and revealed the growth rates in all shoots and those of different ages. This method would be advantageous as an accurate means of direct measurement in fieldwork, and should therefore be a useful tool for monitoring seagrass growth.     

Effects of anaerobiosis on root metabolism of Zostera marina (eelgrass): implications for survival in reducing sediments

The temperate seagrass Zostera marina L. typically grows in highly reducing sediments. Photosynthesis-mediated O₂ supplied to below-ground tissues sustains aerobic respiration during photosynthetic periods. Roots, however, experience daily periods of anoxia and/or hypoxia at night and under conditions that reduce photosynthesis. Rhizosphere cores of Z. marina were collected in August 1984 from Great Harbor, Massachusetts, USA. We examined short-term anaerobic metabolism of [U-¹⁴C]sucrose in excised roots and roots of intact plants. Under anaerobic conditions roots showed appreciable labeling of CO₂, ethanol and lactate, and slight labeling of alanine and other metabolites. Over 95% of the ¹⁴C-ethanol was recovered in the root exudate. Release of other metabolites from the roots was minimal. Ethanol was also released from hypoxic/anoxic roots of intact plants and none of this ethanol was transported to the shoot under any experimental conditions. Loss of ethanol from roots prevented tissue levels of this phytotoxin from increasing during anaerobiosis despite increased synthesis of ethanol. Anaerobic metabolism of [U-¹⁴C]glutamate in excised roots led to appreciable labelling of -aminobutyrate, which was known to accumulate in eelgrass roots. Roots recovered to fully aerobic metabolism within 4 h after re-establishment of aerobic conditions. The contributions of these root metabolic responses to the ability of Z. marina to grow in reducing marine sediments are related to light-regulated interactions of shoots and roots that likely dictate depth penetration, distribution and ecological success of eelgrass.     

Animals on marine flowers: does the presence of flowering shoots affect mobile epifaunal assemblage in an eelgrass meadow?

Eelgrass, Zostera marina, produces two types of shoots: morphologically simple vegetative shoots and highly branched flowering (reproductive) shoots, the latter found only in summer months. We examined whether the abundance and diversity of mobile epifaunal assemblage are affected by the presence of flowering shoots in an eelgrass meadow of Otsuchi Bay, northeastern Japan. Comparisons of epifauna in natural vegetation revealed that density and species richness did not differ significantly between sites consisting of both flowering and vegetative shoots, and those only of vegetative shoots. A transplant experiment, conducted to examine the colonization rates of epifauna to defaunated eelgrass planted with different combination of vegetative and flowering shoots, showed no obvious variation in abundance and species richness. At species level, the density of some species such as a tanaid Zeuxo sp. and a polychaete Platynereis sp. was higher at sites and/or treatments with flowering shoots, whereas that of some gastropods, such as Lirularia iridescens and Siphonacmea oblongata was higher at sites without flowering shoots. The species-specific response led to dissimilarity of epifaunal assemblage between sites and among treatments with different densities of vegetative and flowering shoots. Similar patterns observed for natural vegetation and the transplant experiment suggest that the variation in assemblage structure is caused by habitat selection of each species, for example, the utilization of flowering shoots as feeding ground and nursery by Zeuxo sp.     

Metabolic adaptation of Zostera marina (eelgrass) to diurnal periods of root anoxia

The temperate seagrass Zostera marina L. is common in coastal marine habitats characterized by the presence of reducing sediments. The roots of this seagrass grow in these anoxic sediments, yet eelgrass is highly productive. Through photosynthesis-dependent oxygen transport from leaves to roots, aerobic respiration is supported in eelgrass roots only during daylight; consequently, roots are subjected to diurnal periods of anoxia. Under anoxic root conditions, the amino acids alanine and -amino butyric acid accumulate within a few hours to account for 70% of the total amino acid pool, while glutamate and glutamine decline. Little ethanol is produced, and the pool size of the organic acid malate changes little or declines slowly. Upon the resumption of shoot photosynthesis and oxygen transport to the roots, the accumulated -amino butyric acid declines rapidly, glutamate and glutamine pools increase, and alanine declines over a 16-h period. These adaptive metabolic responses by eelgrass to diurnal root anoxia must contribute to the successful exploitation of shallow-water marine sediments that have excluded nearly all vascular plant groups. A metabolic scheme is presented that accounts for the observed changes in organic and amino acid pool sizes in response to anoxia.     

Nutrient enrichment of seagrass beds in a rhode island coastal lagoon

Seagrass and algal beds showed a variety of reponses when the water column was treated with low level additions of ammonium, nitrate and phosphate. The nutrients were added separately to 3 uniform seagrass beds of a temperature coastal lagoon during 1979 and 1980. (1) Ammonium caused the production of dense mats of free-floating green algae Enteromorpha plumosa and Ulva lactuca. It also stimulated growth in both the leaf and root-rhizome fractions of Zostera marina. This growth response in Z. marina was greater in the area where current reached 12 cm · s^-1 than in the area with little or no current. The concentration of nitrogen in the tissue did not change. In contrast, where current was lacking, Z. marina growth increase with ammonium was small, but the concentration of nitrogen in the tissue doubled over that in control plots. The growth of Ruppia maritima was inversely related to the growth of green algae in the same plots. The red alga Gracilaria tikvahiae did not grow better in ammonium, but its tissue reddened. (2) Nitrate additions enhanced the growth of the green seaweeds Enteromorpha spp. and U. lactuca, but not Z. marina or R. maritima. G. tikvahiae, when fertilized in isolation from other plants, showed a marginal response to this nutrient, and the tissue always reddened. (3) Phosphate enhanced growth in Z. marina and R. maritima exposed to moderate current. G. tikvahiae growing alone showed a small growth response to phosphate. The phosphate made no difference in the growth of the green seaweeds. (4) None of the nutrient supplements noticeably altered the species composition of either epiphytic or planktonic algae associated with the beds, although we did detect small increases in their numbers. The rapid and dense growth of green algae in nitrogen-enriched water probably limited growth of adjacent seagrasses and red algae. Because these seaweeds did not use the phosphate, it became available to other plant components. The overall floral response to nutrient addition in seagrass communities depends, therefore, upon the particular nutrient supplied, the ability of alternate species in the area to compete for that nutrient and the velocity of current in the specific area.     

Analysis of annual fluctuations of C. nodosa in the Venice lagoon: Modeling approach

A simple simulation model was developed to describe the growth trends of Cymodocea nodosa (Ucria) Ascherson based on data sets from the Venice lagoon. The model reproduces the seasonal fluctuations in the above and belowground biomass and in shoot density. The modeling results are in good agreement with data on net production, growth rates and chemical–physical parameters of water. It was assumed that light and temperature are the most important factors controlling C. nodosa development, and that the growth was not limited by nutrient availability. The aim was to simulate biomass production as a function of external forcing variables (light, water temperature) and internal control (plant density). A series of simulation experiments were performed with the basic model showing that among the most important phenomena affecting C. nodosa growth are: (1) inhibition of production and recruitment of new shoots by high temperature and (2) light attenuation due to seasonal fluctuation.     

Scale-dependent patterns of variability of a grazing parrotfish (<Emphasis Type="Italic">Leptoscarus vaigiensis</Emphasis>) in a tropical seagrass-dominated seascape

Although herbivorous fish form critical linkages between primary producers and higher trophic levels, the major factors regulating their spatial structure in seagrass systems remain poorly understood. The present study examined the parrotfish Leptoscarus vaigiensis in seagrass meadows of a tropical embayment in the western Indian Ocean. Stomach content analysis and direct field observations showed that L. vaigiensis is an efficient grazer, feeding almost exclusively on seagrass leaves. Seagrass shoot density was highly correlated to all density variables (total, juvenile and subadult) and juvenile biomass of L. vaigiensis, while subadult biomass was predicted by distance to neighbouring coral habitat. Moreover, density and biomass of predatory fish (piscivores) were predicted by seagrass canopy height and the distribution patterns of predators followed those of L. vaigiensis. Hence, factors at local (seagrass structural complexity and feeding mode) and landscape scale levels (seascape context and distribution of piscivores) likely mutually structure herbivorous fish communities. The findings underscore the importance of incorporating multiple scale-dependent factors when managing coastal seagrass ecosystems and their associated key species.     

Sediment ammonium availability and eelgrass (Zostera marina) growth

The interaction of sediment ammonium (NH ₄ ⁺ ) availability and eelgrass (Zostera marina L.) growth, biomass and photosynthesis was investigated using controlled environment and in-situ manipulations of pore water ammonium concentrations. Sediment diffusers were used to create pore water diffusion gradients to fertilize and deplete ammonium levels in sediments with intact eelgrass rhizospheres. Between October, 1982 and September, 1983 controlled environment experiments using plants from shallow (1.3 m) and deep (5.5 m) stations in a Great Harbor, Woods Hole, Massachusetts, USA eelgrass meadow along with in-situ experiments at these stations provided a range of sediment ammonium concentrations between 0.1 and 10 mM (adsorbed+interstitial NH ₄ ⁺ ). The results of the in-situ experiments indicate that nitrogen limitation of eelgrass growth does not occur in the Great Harbor eelgrass meadow. A comparison of NH ₄ ⁺ regeneration rates and eelgrass nitrogen requirements indicates an excess of nitrogen supply over demand and provides an explanation for the lack of response to the manipulations. Results of controlled environment experiments combined with in-situ results suggest that sediment ammonium pool concentrations above approximately 100 mol NH ₄ ⁺ per liter of sediment (interstitial only) saturate the growth response of Zostera marina.     

Disturbance by ice and life-history adaptations of the seagrassZostera marina

The formation of winter ice, and its movement with the tides, has had a major influence on the life-history parameters of shallow-water populations of the rhizomatous marine angiosperm eelgrass (Zostera marina) in Nova Scotia, Canada. In this region, and annual form of eelgrass inhabits intertidal and shallow subtidal mudflats and a perennial form is common in subtidal areas. Where lowtide water depth was greater than winter ice thickness (25 cm), ice movement removed much of the aboveground biomass from perennial ramets, but did not influence the density of ramets. Measurements of primary production showed that perennial plants allocated a greater proportion of their total production to below-ground structures than measured in all previous studies on eelgrass. Thus perennials are well anchored in the sediment, are more able to withstand removal by ice, and have reserves available for production of new leaves when ice melts. In the spring, seed germination was greatest in areas where ice had removed whole plants (or their aboveground parts) than among mature ramets which had survived winter intact. Survival of seedlings (new genets) was not affected by shading from the adult canopy, but a shading experiment showed that competition for light with mature ramets had a significant negative effect on morphology, growth and the allocation of seedling biomass to below-ground parts, thus reducing the ability of new genets to survive ice disturbance in the next winter. The annual form ofZ. marina was restricted to areas where low tide water depth was much less than winter ice thickness. Annual plants did not survive winter, had small investment in below-ground parts and high reproductive effort, and overwintered as seeds. The genetic status of the two forms and the restriction of the annual to very shallow water are discussed in the light of previous work on eelgrass