Qualitative and quantitative aspects of the epiphytic component in a mixed seagrass meadow from Papua New Guinea

During 1982, structural and functional aspects of the epiphytic component in a tropical mixed seagrass meadow, have been investigated for each seagrass species separately. This meadow consisted of the seagrasses Thalassia hemprichii (Ehrenb.) Aschers., Cymodocea serrulata (R.Br.) Aschers. et Magnus, C. rotundata Ehrenb. et Hempr. ex Aschers., Halodule uninervis (Forssk.) Aschers. and Syringodium isoetifolium (Aschers.) Dandy.No significant differences were observed in floristic composition, number of algal species, abundance and diversity of the epiphytic component. On an area basis, annual mean above-ground biomass (seagrass leaves and epiphytes), amounted to 82 g ADW, of which 18% could be ascribed to the epiphytic component. The contribution of the epiphytic component to the annual mean above-ground production ranged from 16% on leaves of Thalassia hemprichii to 33% on leaves of Cymodocea serrulata. Total annual mean epiphyte production was 4.6 g ADW m⁻² sediment surface day⁻¹ (19%).When including the macroalgal component of this mixed seagrass meadow, total annual mean above-ground plant biomass amounted to 93 g ADW (212 g DW) on an area basis, of which the epiphytes contributed 15.5% (28.5% DW), the macroalgal component 12% (32.5% DW) and the seagrass leaves 72.5% (39.5% DW). Aspects of the epiphytic component (e.g., floristic composition, abundance, biomass and production) in monospecific and mixed seagrass communities are discussed.     

Some structural and functional aspects of the epiphytic component of four seagrass species (Cymodoceoideae) from Papua New Guinea

The epiphytic component of four monospecific seagrass beds from Papua New Guinea was studied structurally and functionally. The floristic composition and abundance of the epiphytes on leaves of four seagrass species (Cymodoceoideae) showed considerable variation, but on all four seagrass species, the same algae were among the five quantitatively most important epiphytes: encrusting coralline algae, Cyanophyta, Ceramium gracillimum (Harv.) Mazoyer, Polysiphonia savatierii Hariot and Audouinella spp. The temporal pattern of the epiphytic algae showed more or less the same features on the four seagrass species.Annual mean biomass of epiphytes and seagrass leaves ranged from 54 g ADW m^?2 in a community of Cymodocea rotundata Ehrenb. and Hempr. ex Aschers. to 169 g ADW m^?2 in a community of Syringodium isoetifolium (Aschers.) Dandy. The contribution of the epiphytic component to the total above-ground biomass ranged from 22 to 24%. Productivity of epiphytes was highest on leaves of Halodule uninervis (Forssk.) Aschers. (2.12 g ADW m^?2 sediment surface day^?1) and the epiphytic community contributed 35–44% of the total above-ground production of these four seagrass communities.     

EPIPHYTE-GRAZER INTERACTIONS ON ZOSTERA MARINA (ANTHOPHYTA: MONOCOTYLEDONES): EFFECTS OF DENSITY ON COMMUNITY FUNCTION1

Interactions between algal epiphytes and their grazers can have a significant impact on the structure and function of eelgrass (Zostera marina L.) meadows. In Puget Sound, the herbivorous gastropod Lacuna variegata Carpenter and its congeners appear to remove large quantities of the epiphytic community from eelgrass blades. When snails at typical field densities were used in microcosms, Lacuna significantly reduced epiphytic biomass and areal productivity. Biomass-specific productivity of the epiphytic community showed an increasing trend with increasing snail density. Epiphytic productivity increased nonlinearly with increasing epiphytic biomass. The commonly used logistic population growth formula adequately described this relationship. Grazing rate also increased nonlinearly with increasing epiphytic biomass. The Hailing equation adequately described the relationship between grazing rate and epiphytic biomass. The proportion of the epiphytic biomass found on the oldest blade of an eelgrass shoot was related linearly to epiphytic biomass, suggesting that a constant fraction of the epiphytic community is lost regardless of epiphytic density. Lacuna clearly removed large quantities of epiphytic material from eelgrass blades, significantly altering community function. Modified Lotka-Volterra equations, incorporating the logistic growth form and Hailing grazing equation, should prove useful in modeling the epiphyte–grazer interaction.     

Effect of nutrient availability, grazer assemblage and seagrass source population on the interaction between Thalassia testudinum (turtle grass) and its algal epiphytes

Seagrass leaves are often densely covered by epiphytic algae which can suppress seagrass productivity and has been implicated in declines of seagrass meadows worldwide. The net effect of epiphytes on seagrass growth and morphology depends on the independent and interactive effects of a variety of factors, including nutrient availability and the intensity of grazing on epiphytes. Here I report the results of a mesocosm experiment designed to test the effects of nutrient addition and within-functional group variation (grazer species composition and the source population of seagrass) on the strength of the interactions among grazers, epiphytes, and turtle grass (Thalassia testudinum). Turtle grass ramets from two sites in the northern Gulf of Mexico were cleared of epiphytes and transplanted into common-garden mesocosms. Replicate ramets were grown in a split-split plot design with two levels of dissolved nutrients and four different grazer species combinations (Tozeuma carolinense alone, Pagurus maclaughlinae alone, both species together, and no grazers present). As expected, grazers had a significant negative effect on epiphyte biomass/leaf area and a significant positive effect on turtle grass growth in the mesocosms. The two species were more similar in their direct effects on epiphyte biomass than in their indirect effects on turtle grass growth; this may reflect differences in epiphyte community composition under different grazer treatments. The effect of nutrient addition on turtle grass growth depended critically on the intensity of grazing: in the presence of grazers, turtle grass tended to produce a greater biomass of new leaf tissue in the tanks with nutrients added than in the control tanks. However, when grazers were absent, the direction of the effect was reversed, and plants with nutrients added grew less than the control plants. The two source populations of turtle grass differed significantly in epiphyte biomass/leaf area accrued in the mesocosms as well as in the strength of the effect of grazers on turtle grass growth. This suggests that population differentiation in seagrass interactions with epiphytes, as well as spatial and temporal variation in resources and grazer community composition, can greatly effect the role of epiphytes in limiting seagrass productivity.     

Importance of seagrass vegetation for habitat partitioning between closely related species,mobile macrofauna <Emphasis Type="Italic">Neomysis</Emphasis> (Misidacea)

Seagrass meadows provide both habitats and a range of food sources for macrofaunal communities. These functions facilitate the coexistence of less mobile invertebrates (in comparison with mysids, such as amphipods) that are associated with seagrass leaves, and may also enhance the coexistence of highly mobile invertebrates such as mysid. We investigated the function of seagrass in supporting the coexistence of two mysid species, Neomysis awatschensis and N. mirabilis. These taxa are dominant in seagrass ecosystems of temperate coastal areas. We compared patterns of habitat use between the two species at mesoscales (among seagrass patches) and microscales (among seagrass leaves) by performing field surveys and laboratory experiments. The field survey results showed positive correlations in the abundance of the two mysid species, indicating that both species select similar habitats at the mesoscale level. In the laboratory experiments, the pattern of microhabitat selection (fundamental habitat) was similar for both species, even at increased densities and with the presence of an immobile habitat-competitor (the gastropod Barleeia angustata) on the leaves. However, this pattern changed significantly when a food source (epiphytic microalgae) was present on the leaves. This result indicates that (i) inter- and intraspecific interference competition does not affect microhabitat selection in these two mysids and (ii) both Neomysis species use similar habitats at the feeding stage. Although these two closely related mysids species may have similar requirements for microhabitat and food, the evidence that they did not act as competitors is attributable to unrestricted microhabitat and food (e.g., epiphytic algae) in the presence of seagrass vegetation.     

The short-term influence of herbivory near patch reefs varies between seagrass species

The coexistence of multiple species within a trophic level can be regulated by consumer preferences and nutrient supply, but the influence of these factors on the co-occurrence of seagrass species is not well understood. We examined the biomass and density responses of two seagrass species in the Florida Keys Reef Tract to grazing pressure near patch reefs, and evaluated how nutrient enrichment impacted herbivory dynamics. We transplanted Halodule wrightii (shoalgrass) sprigs into caged and uncaged plots in a Thalassia testudinum (turtlegrass) bed near a patch reef. Nutrients (N and P) were added to half of the experimental plots. We recorded changes in seagrass shoot density, and after three months, we measured above- and belowground biomass and tissue nutrient content of both species. Herbivory immediately and strongly impacted H. wrightii. Within six days of transplantation, herbivory reduced the density of uncaged H. wrightii by over 80%, resulting in a decrease in above- and belowground biomass of nearly an order of magnitude. T. testudinum shoot density and belowground biomass were not affected by herbivory, but aboveground biomass and leaf surface area were higher within cages, suggesting that although herbivory influenced both seagrass species, T. testudinum was more resistant to herbivory pressure than H. wrightii. Nutrient addition did not alter herbivory rates or the biomass of either species over the short-term duration of this study. In both species, nutrient addition had little effect on the tissue nutrient content of seagrass leaves, and N:P was near the 30:1 threshold that suggested a balance between N and P. The different impacts of grazing on these two seagrass species suggest that herbivory may be an important regulator of the distribution of multiple seagrass species near herbivore refuges like patch reefs in the Caribbean.     

Epiphyte grazing enhances productivity of remnant seagrass patches

Anthropogenic nutrient enrichment is increasingly modifying community structure and ecosystem functioning in terrestrial and aquatic ecosystems. In marine ecosystems, the paradigm is that nutrient enrichment leads to a decline of seagrasses by stimulating epiphytic algal growth, which shades and overgrows seagrasses. This ignores the potential for herbivores, which graze upon epiphytic algae, to partially or wholly counter such nutrient effects. We conducted a field experiment to assess the role that the trochid gastropod Calthalotia fragum plays in reducing nutrient impacts on the seagrass, Posidonia australis, in an urbanized Australian estuary, Botany Bay, Sydney. In a field experiment, where nutrient loading and grazer density were orthogonally manipulated, nutrient enrichment failed to promote epiphyte biomass or diminish growth and primary productivity of P. australis. To the contrary, nutrient enrichment enhanced photosynthesis of the seagrass in plots where the grazer was present at higher density. Epiphytic growth was negatively affected by increased C. fragum density, while P. australis shoot growth was positively influenced. Thus, in this study system, grazing appears to play a much greater role in determining seagrass primary productivity and above‐ground growth than moderate nutrient loading, suggesting that the interaction between grazers and nutrients depends on the relative levels of each. Our study contributes to a growing body of literature suggesting that effects of nutrient loading on benthic assemblages are not universally negative, but are dependent on the biotic and abiotic setting.     

Gastropod grazing of epiphytic lichen-dominated communities depends on tree species

Tree species differ in longevity, canopy structure, and bark texture, chemistry and water storage. Tree species-specific traits play a role in shaping epiphytic vegetation and likely influence the community assembly of organisms feeding on epiphytes. Lichenivorous gastropods, species with calcium-rich shells in particular, need calcium and likely occur more abundantly in and around tree species with high available calcium. We quantified gastropod grazing on the epiphytic lichens Lobaria pulmonaria and Lobaria scrobiculata transplanted to blocks of adjacent trunks of Acer platanoides, Quercus robur and Tilia cordata. We tested the hypothesis that tree species known to have more available Ca, exhibit more grazing damage on transplanted lichens than trees with lower Ca-availability. The grazing pressure was 1.6–1.8 times higher for lichen transplants on Acer and Tilia known to produce litter with easily soluble Ca than on Quercus, which binds Ca as oxalate. Trees with a high grazing pressure on transplants had greater natural abundance of Lobaria virens than of L. pulmonaria. Gastropods preferred L. scrobiculata to L. pulmonaria, evidenced by more observed grazing marks and greater measured biomass loss. We attribute this difference to the lower concentration of carbon-based secondary compounds in L. scrobiculata. However, the strength of the preference varied between the three tree species receiving lichen transplants and was strongest on A. platanoides, while gastropods on T. cordata grazed equal amounts of each transplanted lichen. In conclusion, tree species influenced grazing patterns of gastropods on epiphytic lichens. In addition to bark pH and other factors, we have shown that tree species-specific differences in grazing pressure play a role in shaping the epiphytic macrolichen community.     

Plant-herbivore interactions in seagrass meadows

During the past two decades we have gained much insight into the factors that regulate the productivity of seagrass dominated ecosystems, especially those at low latitudes. Here, we review and reassess the importance of plant-herbivore interactions in seagrass meadows, focusing on recent studies that have examined: 1) grazing on live seagrass leaves; 2) consumption of epiphytic algae growing on seagrass leaves; and 3) consumption of planktonic algae from the waters surrounding seagrass meadows. The major conclusion is that, in contrast to what has been reported in much of the literature on food webs in seagrass meadows, a diverse grazing pathway continues to represent an important conduit for the transfer of energy from the primary producers to higher order consumers. This remains true, although in many areas consumption of seagrasses is reduced in an historical context, owing to the overharvesting of many large species of herbivorous waterfowl, turtles and mammals.We also summarize our view of the important gaps in understanding the broadly defined topic of herbivory in seagrass-dominated ecosystems. We suggest that future studies should focus on: understanding the foraging strategies of seagrass herbivores; quantifying the impact of herbivory on seagrass demography, including effects on sexual reproduction, the fate of flowers, and the production of fruits and seeds; and documenting the commonness of compensatory responses to grazing. In addition, the role of chemical defenses in seagrass species remains inadequately investigated. Studies of the roles of nutritional content (as measured by C/N/P ratios) and chemical defenses are also fertile grounds for future studies of epiphytes and their grazers, as are additional experiments to quantify the relative roles of top-down and bottom-up factors as they determine algal growth and abundance. There is also a need to expand the geographical scope of studies of epiphyte-grazer interactions from cold temperate to sub-tropical and tropical waters. Suspension feeders also need to be studied more broadly, with additional experiments required to quantify their effects on water clarity and their ability to fertilize pore waters, and whether benefits from these activities balances the costs of shading and competition for space that can result from both epifaunal and infaunal suspension feeders.     

Specificity in Mesograzer-Induced Defences in Seagrasses

Grazing-induced plant defences that reduce palatability to herbivores are widespread in terrestrial plants and seaweeds, but they have not yet been reported in seagrasses. We investigated the ability of two seagrass species to induce defences in response to direct grazing by three associated mesograzers. Specifically, we conducted feeding-assayed induction experiments to examine how mesograzer-specific grazing impact affects seagrass induction of defences within the context of the optimal defence theory. We found that the amphipod Gammarus insensibilis and the isopod Idotea chelipes exerted a low-intensity grazing on older blades of the seagrass Cymodocea nodosa, which reflects a weak grazing impact that may explain the lack of inducible defences. The isopod Synischia hectica exerted the strongest grazing impact on C. nodosa via high-intensity feeding on young blades with a higher fitness value. This isopod grazing induced defences in C. nodosa as indicated by a consistently lower consumption of blades previously grazed for 5, 12 and 16 days. The lower consumption was maintained when offered tissues with no plant structure (agar-reconstituted food), but showing a reduced size of the previous grazing effect. This indicates that structural traits act in combination with chemical traits to reduce seagrass palatability to the isopod. Increase in total phenolics but not in C:N ratio and total nitrogen of grazed C. nodosa suggests chemical defences rather than a modified nutritional quality as primarily induced chemical traits. We detected no induction of defences in Zostera noltei, which showed the ability to replace moderate losses of young biomass to mesograzers via compensatory growth. Our study provides the first experimental evidence of induction of defences against meso-herbivory that reduce further consumption in seagrasses. It also emphasizes the relevance of grazer identity in determining the level of grazing impact triggering resistance and compensatory responses of different seagrass species.     

Periphyton density and shading in relation to tidal depth and fiddler crab activity in intertidal seagrass beds of the Banc d'Arguin (Mauritania)

Periphyton development was studied on microscopic glass slides and leaves of Zostera noltii Hornem. in an intertidal area in the Banc d'Arguin (Mauritania). The effects of shading, tidal depth and grazing activities by the fiddler crab Uca tangeri Eydoux were evaluated. For all experiments, periphyton ash content was high (52–93%) and ash-free dry weight ranged between 0.10–0.63 mg cm^–2. Slides accumulated more periphyton than leaves.Artificial shading (62–99%) for 13 days had no effect on periphyton densities on leaves. Increased tidal depth resulted in higher ash-free dry weight on slides, but in lower ash-free dry weight on leaves. Significant variation along the coastline also existed. The effect of fiddler crabs was studied using exclosures. Presence of fiddler crabs reduced periphyton density on slides, whereas light transmittance was increased. On leaves, no significant fiddler crab effect was found. This difference between leaves and slides was probably caused by a storm at the day before the end of the experiment, and by the higher periphyton density on slides as compared with leaves. As visual inspection during the experiment showed clear differences in appearance of leaves inside and outside the exclosures, the storm probably sloughed off mainly the older leaves, i.e. those on which the differences in periphyton cover were the highest.It is hypothesized that periphyton accumulation is higher with increased tidal depth, whereas fiddler crab grazing pressure also increases in this direction. The result is a decreased periphyton density with increased tidal depth.The presently found light extinction coefficients (mean 0.8 m^–1) and periphyton light attenuance (up to 25%) in Banc d'Arguin are not likely to affect seagrass leaf growth.     

Plant–animal associations in two species of seagrasses in Western Australia

Relationships between algal epiphytes and epifaunal invertebrates (amphipods, molluscs and polychaetes) occurring within meadows of the seagrasses Posidonia sinuosa and Amphibolis griffithii were compared along the south west coast of Western Australia. Although the seagrasses are very different structurally, many species of algal epiphytes and epifaunal grazers were common to both. However, meadows of Amphibolis supported a greater number of both algal epiphyte and epifaunal species. The long-lived stems of Amphibolis supported a larger biomass of algal epiphytes and grazers than did the leaves of either Posidonia or Amphibolis. The densities and biomass of epifauna were variable but on a comparison adjusted to the biomass of seagrass, both the density and biomass of the taxonomic groups were similar between seagrass species except that the density of grazing gastropods and the biomass of polychaetes were greater in Amphibolis (by 238% and 252%, respectively). Nested analyses of variance (ANOVA) indicated that variations in plant and animal biomass differed at all spatial scales (sites, meadows within sites and replicates) and the pattern was inconsistent amongst biota. However, a significant proportion of the variability occurred between replicate samples. Canonical correlation and multiple regression analyses indicated that associations between algal epiphytes and epifauna were also inconsistent and differed between seagrass species. These patterns highlight the importance of seagrass species and structural complexity in affecting both the epiphytic and grazer community. The importance of spatial scales at which seagrasses and their associated communities are sampled are equally important because of the differing levels of spatial patchiness.     

Factors affecting the structure of epiphytic gastropod communities in the St. Lawrence River (Quebec,Canada)

A study of epiphytic gastropods associated with two submerged macrophytes (Myriophyllum spicatum and Vallisneria americana) was conducted at a soft and at a hard water site in the St. Lawrence River during two non-consecutive years in order to compare effects of macrophyte species and biomass, site and year in gastropod community structure. There was no effect of macrophyte species on total gastropod abundance nor on gastropod diversity, and few gastropod species showed a marked preference for either macrophyte species. Inter-site and inter-annual variations in diversity, total gastropod abundance and gastropod community structure were greater than variations among macrophytes. However, analysis of variations of epiphytic gastropod communities in one of the two sites shows that abiotic factors are important in explaining epiphytic gastropod distribution. Our results and results of other studies on gastropod population dynamics in the St. Lawrence River indicate that intra- and interspecific competition between gastropods is important on macrophytes and that they must be carefully considered in order to understand epiphytic community structure and population dynamics.     

The effects of sea urchin grazing and drift algal blooms on a subtropical seagrass bed community

Subtropical seagrass beds can be subject to relatively high levels of direct herbivory and large blooms of drift algae, both of which can have important effects on the floral and faunal components of the community. Caging experiments were used to investigate these factors in a Thalassia testudinum bed in Biscayne Bay, Florida. Abundance of sea urchins, Lytechinus variegatus, and drift algae was manipulated within the cages. Naturally occurring levels of urchin grazing do not appear to affect the T. testudinum population. With experimentally increased urchin densities in the winter, seagrass shoot density and aboveground biomass decreased significantly. Similar effects were not detected in the summer, indicating that the impact of grazing on T. testudinum is lessened during this time of year. Shoot density was more vulnerable to grazing than aboveground biomass. This may be a result of grazing-induced increases in seagrass productivity, in which the remaining shoots produce more or longer leaves. In the winter, drift algal blooms form large mats that cover the seagrass canopy. Under the normal grazing regime these algal blooms do not have significant negative effects on the seagrass. With increased grazing pressure, however, there is a synergistic effect of grazing and drift algae on seagrass shoot density. At intermediate urchin density (10 per m(-2)), cages without algae did not undergo significant decreases in shoot density, while those with algae did. At the high density of urchins, the number of seagrass shoots in cages both with and without algae decreased, but the effect was more pronounced for cages with algae. Invertebrate abundance at the field site was low relative to other seagrass beds. There were no discernible effects, either positive or negative, of urchin and algae manipulations on the sampled invertebrate community.     

Quantitative and qualitative variability of epiphytic algae on one Apiaceae (<Emphasis Type="Italic">Apium nodiflorum</Emphasis> L.) in a karstic river (Southeast of France)

Spatial distribution of the firmly attached epiphytic algae was investigated directly on the leaves of one Apiaceae (Apium nodiflorum L.) commonly found in chalky Mediterranean rivers, at two sampling dates (summer and autumn). The composition and structure of the epiphytic communities of the samples were determined at different levels: along the leaves, on both their sides and within each leaflet (n = 13). Forty-eight leaflets from four randomly selected plants were cut along the longitudinal and vertical axes so as to define 16 microhabitats located on the edge, the centre and the vein. Only a few species of algae were found to be constantly present whatever the sampling date, and dominated all microzones (n = 768). Their presence seemed to depend, rather, on the architectural structure of the plant. The other algae represented the major part of the epiphytic community and were only occasionally present. Those were more subject to temporal and environmental variations and less specific of the substrate. The epiphytes showed considerable spatial heterogeneity within their microhabitats, with an evident preference for the centre and the edge of the leaves, whereas two inhospitable spaces were the vein and apex. On the other hand, when the study was performed at a larger scale (along the leaf), the epiphytic populations of the youngest leaves (near the surface) and the eldest ones (close to the sediments) were homogenous, hence suggesting that other factors, independent from the physiology of the plant, may have an impact on epiphytic distribution.     

Are epiphytes a significant component of intertidal Zostera noltii beds?

The role of epiphytes in an intertidal Zostera noltii seagrass bed in Marennes-Oléron Bay was assessed in comparison with the other main benthic primary producers (Z. noltii, microphytobenthos) at two bathymetric levels and on a seasonal basis. Assemblage and biomass of epiphytes were studied using scanning electron microscopy (SEM). Z. noltii and its detrital matter followed a typical seasonal pattern: microphytobenthos was present in large quantities throughout the year representing 21% of the total biomass while detrital matter, above-ground parts and below-ground parts accounted for 65, 9 and 5%, respectively. Only two species of epiphytic diatoms, Cocconeis scutellum and Cocconeis placentula, were observed on seagrass leaves. Epiphyte biomass was very low, representing on average less than 0.001% of that of microphytobenthos or leaves. This low epiphyte biomass is linked with the absence of macroalgae and also with the low biovolume of Cocconeis, which formed a monolayer of cells on leaves. This can be explained by the severe conditions of the intertidal and the high leaf turn-over of Z. noltii leaves.     

Grazing effects of a gammaridean Amphipoda,Ampithoe sp., on the seagrass,Syringodium isoetifolium,and epiphytes in a tropical seagrass bed of Fiji

In the pure stand of tropical seagrass,Syringodium isoetifolium, in a small oceanic island, Fiji, grazing effects of the seagrass-associated gammarid,Ampithoe sp., on seagrass and epiphytes were assessed in October 1989, November 1991, November 1992. Density of the gammarid was estimated with two methods, mesh bag method and tuft method. During the three years surveyed the density of the gammarid increased remarkably from 1989 to 1991, with heavy epiphytism. Gut contents of the gammarid were examined. Grazing rates on seagrass leaf with and without epiphytic blue-green algae were measured in a bottle experiment. Litter bag experiments were conducted using different mesh sizes each containing seagrass only and seagrass and gammarids. The seagrass leaf biomass in the litter bag reduced abruptly in both bags. After one week, 78–86% of seagrass biomass disappeared from the bags. Enhancement of decomposition of seagrass leaf by the gammarid grazing was observed. Oxygen consumption and ammonium excretion rates were measured simultaneously in bottle experiments. Carbon budget in the seagrass bed was estimated as follows: 0.9 gC m⁻² day⁻¹ in seagrass growth, gammarid grazing was about a half of it and further assimilated a half of it, about 0.1 gC m⁻² day⁻¹, and more than half of it become CO₂ by respiration. Grazing effects on epiphyte and seagrass growth and production were discussed through the carbon budget and indirect interactions between seagrass, epiphytes and associated gammarids to explain the temporal change of seagrass and epiphyte dynamics.     

Seasonal shifts in seagrass bed primary producers in a cold-temperate estuary: Dynamics of eelgrass Zostera marina and associated epiphytic algae

The seasonal dynamics of seagrass and epiphytic algal primary production were measured in an eelgrass (Zostera marina) bed in the Akkeshi-ko estuary, Hokkaido, Japan (43°02′N, 144°52′E). During spring and early summer, eelgrass biomass increased, with a high production (maximum: 2.89 g C m⁻² day⁻¹), but the production and biomass of epiphytic algae remained low. In contrast, epiphytic algae bloomed in August, with a high production (5.21 g C m⁻² day⁻¹), but eelgrass production ceased and its biomass subsequently decreased. Therefore, the major primary producers in this eelgrass bed switched seasonally from eelgrass in spring and early summer to epiphytic algae in late summer and autumn. Epiphytic algae maintained similar productivity because of the change of photosynthetic kinetics and the dominant epiphytic diatom changed from highly adhesive species to less adhesive or filamentous small species during the bloom. This suggests that the change of epiphyte density and biomass was due to change of its loss rate, possibly due to herbivorous grazing rate. Moreover, competition between epiphytic algae and eelgrass for nutrients and light may also affect the dramatic seasonal changes in the major primary producers.