Understorey benthic microalgae and their consumers depend on habitat complexity and light in a microtidal coastal ecosystem

Presence of habitat-forming macroalgae is supposed to mitigate effects of altered resources on benthic microalgae and their consumers. In a field experiment in a microtidal area of the Western Baltic Sea, we tested the interactive effects of nutrient enrichment, artificial shading, and habitat complexity on microalgal biomass and diversity as well as invertebrate abundance and richness. Habitat complexity comprised three levels, the presence of macroalgal canopy of Fucus vesiculosus, the presence of macroalgal propagules, and the absence of both (=control). Microalgal biomass (and richness) was significantly reduced by canopy presence (−88%, compared to control) and shading (−42%), with the highest biomass in the absence of both canopy and macroalgal propagules at ambient light. Within the microalgal assemblage, higher biomass was related to lower evenness (higher dominance). Density of two main invertebrate groups (snails and amphipods) strongly increased with canopy presence (on average from 53 to 154 individuals m⁻² stone area for snails, and from 234 to 1203 individuals m⁻² for amphipods) and so did invertebrate richness (from 4.3 to 10.3). Additionally, snail density doubled with increasing light availability. The snail responses to light and canopy were independent, the former relating to higher availability of microalgal prey, the latter to more structure. Microalgal taxon richness and biomass decreased with increasing invertebrate richness and with density of snails and amphipods. Our experiment thus showed that the presence of habitat-forming macroalgae alters biomass and diversity across trophic levels in benthic coastal communities as well as their response to resource manipulations.     

The relative importance of nutrient enrichment and herbivory on macroalgal communities near Norman's Pond Cay, Exumas Cays, Bahamas: a “natural” enrichment experiment

The simultaneous effects of grazing and nutrient enrichment on macroalgal communities were experimentally investigated using plastic mesh enclosure/exclosure cages along a natural nutrient (DIN, SRP) gradient from the discharge of a tidal mangrove creek on the west side of Norman's Pond Cay, Exumas Cays, Bahamas. Nutrient enrichment was the only factor affecting total biomass of macroalgae whereas selective herbivory moderated species composition. Biomass ranged from >2 kg dry weight m⁻² at the DIN-enriched Waterfall to <600 g dry weight m⁻² in the DIN-limited Algal Halo. Grazing by the queen conch Strombus gigas and the sea hare Aplysia dactylomela significantly reduced biomass of the epiphyte Laurencia intricata, but not its host Digenea simplex at the DIN-enriched Waterfall. These two rhodophytes dominated the macroalgal community at the DIN-enriched Waterfall and inner Algal Halo, whereas the chlorophytes Microdictyon marinum and Cladophora catenata dominated the relatively DIN-depleted outer Algal Halo. Higher grazing rates by S. gigas, A. dactylomela, and juvenile parrotfish and surgeonfish at the DIN-enriched Waterfall compared to the more oligotrophic Algal Halo suggested selective grazing on higher quality (nitrogen) diets by these herbivores. A pronounced increase in the carbon/nitrogen (C:N) ratios of D. simplex and L. intricata from the Waterfall to the Algal Halo indicated increased DIN-limitation of growth in the more offshore, lower DIN macroalgal communities. In contrast, only modest increases in C:N ratios of M. marinum and C. catenata between the Waterfall and Algal Halo suggested that these species are better adapted to growth in reef environments with lower levels of DIN enrichment. Low coral cover (<2%) co-occurred with high macroalgal cover (>35%) at DIN-enriched reef sites around Norman's Pond Cay (Waterfall, Patch Reef, North Perry Reef) compared to relatively high coral cover (10-20%) and low macroalgal cover (<20%) at the lower DIN reef sites (Rainbow Reef, Lang's Reef). These results support ecological theory that bottom-up control via nutrient enrichment is a primary factor regulating overall biomass and taxonomic assemblages of macroalgae on coral reefs, whereas grazing is more important in controlling relative species composition via dietary preferences.     

Oxygen and ammonium dynamics during a farming cycle of the bivalve <Emphasis Type="Italic">Tapes philippinarum</Emphasis>

Fluxes of dissolved oxygen and ammonium across the water sediment interface were measured in a control and in an experimental area farmed with the clam Tapes philippinarum. Young clams were seeded in March 2003 at mean (~500 ind m⁻²) and high (~1500 ind m⁻²) densities in a sandy area (2100 m²) of the Sacca di Goro Lagoon, Italy. Approximately every two months, until October 2003, intact sediment cores were collected and incubated in the light and in the dark and surface sediments (0–2 cm) were analysed for organic matter and nitrogen content. Clams farming induced pronounced changes in sediment characteristics and metabolism. Oxygen consumption and ammonium production at the high density area were, on average, 3 to 4 and 1.9 to 4.9 folds higher than those measured in the control field respectively; rates were positively correlated with clams biomass. Experimental fields resulted “Net and Total Heterotrophyc” in 3 out of 4 sampling dates and clams were the major factor shifting the benthic system towards this status. In only one occasion the appearance of the macroalgae Ulva spp. pushed the system rapidly towards hyperautotrophic conditions. Our results indicated that clams have the potential to drive benthic metabolism in farmed areas and to sustain macroalgal growth through regeneration of a limiting nutrient for seawater as inorganic N.     

Macroalgal-mediated transfers of water column nitrogen to intertidal sediments and salt marsh plants

In many temperate estuaries, mats of opportunistic macroalgae accumulate on intertidal flats and in lower elevations of salt marshes, perhaps playing a role in linking water column nitrogen (N) supply to these benthic habitats. Using a flow-through seawater system and tidal simulator, we varied densities (equivalent to 0, 1, 2, or 3 kg m⁻² wet mass) of ¹⁵N-labelled macroalgae (Enteromorpha intestinalis) on estuarine sediments in microcosms with/without pickleweed (Salicornia virginica) to assess N transfers from algae. In the 6-week experiment, macroalgal biomass increased from initial levels in the lower density treatments but all algae lost N mass, probably through both leakage and decomposition. With all densities of algae added, sediments and pickleweed became enriched in ¹⁵N. With increasing mat density, losses of algal N mass increased, resulting in stepwise increases in ¹⁵N labeling of the deeper sediments and pickleweed. While we did not detect a growth response in pickleweed with macroalgal addition during the experiment, N losses from algal mats that persist over many months and/or recur each year could be important to the mineral nutrition of N-limited marsh plants. We conclude that N dynamics of intertidal sediments and lower salt marsh vegetation are linked to the N pools of co-occurring macroalgae and that further study is needed to assess the magnitude and importance of N transfers.     

Macroalgal detrital systems: an overlooked ecological niche for heterotrophic nitrogen fixation

Diazotrophic macroalgal associations (DMAs) can contribute fixed nitrogen (N) to the host macroalgae. Biological nitrogen fixation (BNF) rates investigated using acetylene reduction assays with living macroalgae surrounding Santa Catalina Island were low (maximum: 36 nmol N × g⁻¹ (dw) × h⁻¹) and probably insufficient towards helping meet macroalgal N demand. However, DMAs were observed during periods of low nitrate availability in Southern California coastal waters, highlighting the potential importance of diazotrophs during N depleted conditions. Eleven long-term (16–32 days) litter bag decomposition experiments with various macroalgae, especially those with high (> 10) C:N ratios, resulted in much higher BNF rates (maximum: 693 nmol N × g⁻¹ (dw) × h⁻¹) than observed with living macroalgae. BNF rates were lower at the beginning of macroalgal decomposition but rapidly increased during the second phase before declining towards the end of decomposition. Labile carbon availability is likely influencing BNF rates throughout macroalgal degradation and limits BNF in the final decomposition stage. Comparable dark and light BNF rates with most macroalgae surveyed suggest macroalgal detrital systems are an overlooked, potentially global, niche for heterotrophic N₂ fixation. Lastly, suppressed BNF rates with sodium molybdate additions highlight the prevalence of sulfate reducing diazotrophs.     

THE EFFECT OF A HORN SNAIL ON ULVA EXPANSA (CHLOROPHYTA): CONSUMER OR FACILITATOR OF GROWTH?1

To determine whether California horn snails are more likely to be consumers or facilitators of Ulva expansa (Setch) S. & G. growth in estuaries, we conducted manipulative experiments that evaluated algal growth and the movement of N between the water column, algal tissue, and, in the second experiment, sediments. Algae grew poorly in the absence of sediments, drawing on their own sequestered N supplies (3.5% of dry weight reduced to <2%) and N released by snails and by depleting inorganic N in the water column. There was no evidence of consumption when snail densities ranged from 0 to 900.m^?2 (0, 3, 6, and 9 per aquarium), as algal growth was similar for all snail densities, and snail lengths did not increase during the 21–d experiment. when sediment was provided, N was depleted in the sediment and enhanced in the algal tissue. As in the first experiment, the water column was depleted of inorganic N and enriched with organic N, mostly in the dissolved form. Because both snails and macroalgae often dominate the shallow waters of southern California's lagoons and estuaries, our evidence that the snails are primarily facilitators of algal growth (via transfer of N from sediments to the water column) suggests that snails may play an important role in both food web and N dynamics.     

UPTAKE AND RELEASE OF NITROGEN BY THE MACROALGAE GRACILARIA VERMICULOPHYLLA (RHODOPHYTA)1

Macroalgae, often the dominant primary producers in shallow estuaries, can be important regulators of nitrogen (N) cycling. Like phytoplankton, actively growing macroalgae release N to the water column; yet little is known about the quantity or nature of this release. Using ¹⁵N labeling in laboratory and field experiments, we estimated the quantity of N released relative to assimilation and gross uptake by Gracilaria vermiculophylla (Ohmi) Papenfuss (Rhodophyta, Gracilariales), a non‐native macroalgae. Field experiments were carried out in Hog Island Bay, a shallow back‐barrier lagoon on the Virginia coast where G. vermiculophylla makes up 85%–90% of the biomass. There was good agreement between laboratory and field measurements of N uptake and release. Daily N assimilation in field experiments (32.3±7.2 μ mol N·g dw^?1·d^?1) was correlated with seasonal and local N availability. The average rate of N release across all sites and dates (65.8±11.6 μ mol N·g dw^?1·d^?1) was 67% of gross daily uptake, and also varied among sites and seasons (range=33%–99%). Release was highest when growth rates and nutrient availability were low, possibly due to senescence during these periods. During summer biomass peaks, estimated N release from macroalgal mats was as high as 17 mmol N·m^?2·d^?1. Our results suggest that most estimates of macroalgal N uptake severely underestimate gross N uptake and that N is taken up, transformed, and released to the water column on short time scales (minutes–hours).     

Effect of grazing by isopods and amphipods on growth of Ulva spp. (Chlorophyta)

Eutrophication of shallow coastal waters often leads to blooms of macroalgae. Grazing by crustaceans, such as amphipods and isopods, can reduce macroalgal biomass accumulation. At the same time, growth of the macroalgae can be stimulated by epiphyte removal. The role of grazing by isopods and amphipods on Ulva spp. biomass development was investigated in the Veerse Meer, a brackish lagoon situated in the southwest Netherlands. Exclusion of grazing in the field did not stimulate Ulva spp. growth. In fact, growth rates were higher in exclosures that allowed grazers to enter. Edibility tests identified the amphipod Gammarus locusta, and the isopods Idotea chelipes and Sphaeroma hookeri as potential grazers on Ulva spp. However, when epiphytic diatoms were present on the Ulva spp. thalli, Gammarus and Sphaeroma grazed on ephiphytes and not on Ulva tissue. Only Idotea continued to graze on Ulva spp. A laboratory growth experiment revealed a positive effect of Gammarus presence on Ulva spp. growth, probably caused by preferential removal of epiphytic diatoms from the Ulva spp. thalli. The growth stimulation by epiphyte removing grazers such as Gammarus may explain the higher growth rates in the presence of grazers observed in the field. When determining the potential role of invertebrate grazers in controlling macroalgal biomass accumulation, it is important to include an assessment of the epiphyte abundance on the macroalgae, as preferential removal of epiphytes may stimulate growth and thus have the opposite effect.     

EFFECTS OF IRRADIANCE AND GRAZING ON LOTIC ALGAL ASSEMBLAGES1

A laboratory experiment was conducted for 75 days to examine how irradiance levels and grazing influence algal biomass and community structure. Twelve laboratory streams were used for experimental analyses, with four channels exposed to one of three irradiance levels (15, 100, or 400 μE·m^?2·s^?1). Three of the four stream at each light level were stocked with the snail Juga silicula (250·m^?2), leaving one stream at each light level without snails. Grazed stream exposed to low light levels developed low amounts of algal biomass (<2 g AFDW·m^?2) and were dominated by adnately attached diatoms. Mean algal biomass increased over time in the grazed streams exposed to intermediate light; by day 75, these streams were characterized by moderate algal biomasses (30-40 g AFDW·m^?2) and filamentous chlorophytes. Algal assemblages in high light, grazed channels had high levels of biomass at day 43 (70 g AFDW·m^?2) that declined to 30 g AFDW·m^?2at day 75 and were dominated by chlorophytes. Among ungrazed streams, algal biomass at day 75 was relatively low in the low light streams (<7g AFDW·m^?2) and was dominated by adnately attached diatoms. Ungrazed streams exposed to intermediate and high light levels had moderate biomasses (23 and 19 g AFDW·m^?2, respectively) and were dominated by chlorophytes and large diatoms. Grazing appeared both to delay and alter successional trajectories of algal assemblages, with alterations most noticeable during early seral stages at intermediate and high light levels. Grazing had the least effect on successional trajectories at low light.     

Impact of invasive apple snails in Hong Kong on wetland macrophytes,nutrients, phytoplankton and filamentous algae

1. Grazing by invasive species can affect many aspects of an aquatic system, but most studies have focused on the direct effects on plants. We conducted mesocosm and laboratory experiments to examine the impact of the invasive apple snail Pomacea canaliculata on macrophytes, filamentous algae, nutrients and phytoplankton. 2. In a freshwater pond, we confined 500 g of Myriophyllum aquaticum or Eichhornia crassipes with 0, 2, 4 or 8 apple snails in 1 m × 1 m × 1 m enclosures for approximately 1 month. Apple snails grazed heavily on both species of macrophytes, with higher overall weight losses at higher snail densities. The damage patterns differed between the two macrophytes. In M. aquaticum, both leaves and stems suffered from substantial herbivory, whereas in E. crassipes, only the roots suffered significant weight reduction. 3. In addition to grazing on macrophytes, apple snails appeared to have controlled the growth of filamentous algae, as these did not develop in the snail treatments. The ability of P. canaliculata to control filamentous algae was supported by a laboratory experiment where the consumption was as high as 0.25 g g⁻¹ snail DW d⁻¹. Because of a lack of native herbivorous snails in the pond, the growth of filamentous algae (mainly Spirogyra sp.) reached 80.3 g m⁻², forming a spongy pond scum in the no‐apple snail control. Together with previous reports that apple snails could eat the juveniles and eggs of other freshwater snails, our results indicated that P. canaliculata could have out‐competed native herbivorous snails from the pond by predation on their juveniles or eggs. Alternatively, P. canaliculata might have out‐competed them by monopolisation of food resources. 4. Nitrogen and phosphorous concentrations remained low throughout both experiments and were not correlated with apple snail density. The treatment effects on chlorophyll a (Chl a) and phytoplankton composition varied in the two experiments. In the M. aquaticum experiment, with increasing snail density, Chl a increased, and the phytoplankton community became dominated by Cryptophyceae. In the E. crassipes experiment, Chl a level was independent of snail density, but with increasing snail density, the phytoplankton community became co‐dominated by Cryptophyceae, Chlorophyceae and Bacillariophyceae. 5. Given the multiple effects of P. canaliculata on wetland biodiversity and function, management strategies should be developed to prevent its further spread. In invaded wetlands, strategies should be developed to eradicate the apple snail and re‐introduce native snails which can control the development of filamentous algae.     

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.     

THE PERFORMANCE OF SPRAY‐IRRIGATED ULVA LACTUCA (ULVOPHYCEAE,CHLOROPHYTA) AS A CROP AND AS A BIOFILTER OF FISHPOND EFFLUENTS1

The seaweed Ulva lactuca L. was spray cultured by mariculture effluents in a mattress‐like layer, held in air on slanted boards by plastic netting. Air‐agitated seaweed suspension tanks were the reference. Growth rate, yield, and ammonia‐N removal rate were 11.8% · d^?1, 171 g fresh weight (fwt) · m^?2 · d^?1, and 5 g N · m^?2 · d^?1, respectively, by the spray‐cultured U. lactuca, and 16.9% · d^?1, 283 g fwt · m^?2 · d^?1, and 7 g N · m^?2 · d^?1, respectively, by the tank U. lactuca. Biomass protein content was similar in both treatments. Dissolved oxygen in the fishpond effluent water was raised by >3 mg · L^?1 and pH by up to half a unit, upon passage through both culture systems. The data suggest that spray‐irrigation culture of U. lactuca in this simple green‐mattress‐like system supplies the seaweed all it needs to grow and biofilter at rates close to those in standard air‐agitated tank culture.     

Effects of an invasive reef-building polychaete on the biomass and composition of estuarine macroalgal assemblages

Invasive species can transform ecological communities. Their profound effects may alter the sources and pathways of primary production. We investigated the effects of the reef forming polychaete Ficopomatus enigmaticus invasion on the biomass and distribution of estuarine macroalgae in a SW Atlantic coastal lagoon (Mar Chiquita, 37° 40′S, 57° 23′W, Argentina). Reefs built by this species serve as substrates for macroalgal development and furnish structures that modify physical and biological conditions for the surrounding benthos. We showed that (1) the red macroalga Polysiphonia subtilissima settles and grows almost exclusively on the surface of the reef, (2) the green macroalgae Cladophora sp. and Enteromorpha intestinalis are found almost exclusively in areas without reefs attached to mollusk shells and, (3) no macroalgae occur in the sediment between reefs. Manipulative experiments show that reefs provide a complex substrate for settlement and survival and therefore benefit red macroalga. These experiments also show that the invasive reef builder has negative indirect effects on green macroalgae by increasing grazing and probably by increased sedimentation between reefs. Via these direct and indirect effects, reefs change the relative biomass contribution of each macroalgal species to the overall production in the lagoon. Knowledge of these processes is important not only for predicting net effects on primary production but also because changes in macroalgal species composition may produce effects that cascade through the food web.     

High Nitrogen and Phosphorous Acquisition by Belowground Parts of Caulerpa prolifera (Chlorophyta) Contribute to the Species' Rapid Spread in Ria Formosa Lagoon,Southern Portugal

Despite worldwide proliferation of the genus Caulerpa and subsequent effects on benthic communities, little is known about the nutritional physiology of the Caulerpales. Here, we investigated the uptake rates of ammonium, nitrate, amino acids, and phosphate through the fronds and rhizoids + stolon, the internal translocation of nitrogen, and developed a nitrogen budget for the rapidly spreading Caulerpa prolifera in Ria Formosa lagoon, southern Portugal. Caulerpa prolifera acquired nutrients by both aboveground and belowground parts at similar rates, except nitrate, for which fronds showed 2-fold higher uptake rates. Ammonium was the preferential nitrogen source (81% of the total nitrogen acquisition), and amino acids, which accounted for a significant fraction of total N acquisition (19%), were taken up at faster rates than nitrate. Basipetal translocation of ¹⁵N incorporated as ammonium was nearly 3-fold higher than acropetal translocation, whereas ¹⁵N translocation as nitrate and amino acids was smaller but equal in either direction. The estimated total nitrogen acquisition by C. prolifera was 689 μmol · m⁻² · h⁻¹, whereas the total nitrogen requirement for growth was 672 μmol · m⁻² · h⁻¹. The uptake of ammonium and amino acids by belowground parts accounted for the larger fraction of the total nitrogen acquisition of C. prolifera and is sufficient to satisfy the species nitrogen requirements for growth. This may be one reason explaining the fast spreading of the seaweed in the bare sediments of Ria Formosa where it does not have any macrophyte competitors and the concentration of nutrients is high.     

Selective grazing by the mud snail Ilyanassa obsoleta

Summary Mud snails (Ilyanassa obsoleta) starved for 48 h were allowed to feed on sediments in laboratory microcosms. Sediment cores sliced at 2 mm intervals were compared to snail stomach contents for per cent carbon and nitrogen, plant pigment contents and species composition of benthic diatoms. Concentrations of carbon, nitrogen, phaeopigments, phycocyanin and chlorophyll were enriched in the top 2 mm of the sediments compared to 7–10 mm depth by a factor of 2–10. In turn, these materials were 20–40 times more concentrated in snail guts than in the surface sediments. Snail feces were enriched for carbon and nitrogen by 5–7 times over the surface sediments. Bacterial chlorophyll peaked at about 3–4 mm in the sediments and was not detectable in the snail stomach contents. The C/N ratio of the snail stomach contents was only 6 compared to a ratio of 8.5 for their feces and 12 for the surface sediments.The percentage of migratory diatoms (e.g. Nitzschia and Navicula) decreased with depth where non-migratory species, such as Fragilaria pinnata, dominated. These migratory species were more common in the snails than in the sediments on which they were feeding.A comparison of daily ingestion rates to the animal's energy budget shows that this selective ingestion is sufficient to meet Ilyanassa's energy needs.     

The influence of periphyton biomass and density on grazing in Physella virgata

We studied how differences in periphyton colonization interval and snail density affected grazing rates in Physella virgata, and whether snails controlled periphyton biomass. Both egestion rates and incorporation rates of ¹⁴C labeled periphyton were estimated in laboratory experiments. Periphyton biomass increased with field colonization interval in all experiments, but did not consistently influence estimates of grazing rate. However, increased periphyton abundance in one of the experiments could still explain higher grazer rates in that year, although larger snail body size is a confounding explanation. Increased snail density also resulted in decreased grazing rates, as observed in earlier studies with this snail species, as well as in studies with other snail grazers. Our results suggest grazing rates and resulting impacts may change seasonally with variation in either periphyton biomass, grazer life-history stage or population density.     

The effect of horse, cattle and sheep grazing on the diversity and abundance of land snails in nutrient-poor calcareous grasslands

Livestock grazing is a common management practise in semi-natural grasslands in Central Europe. Different types of livestock (horses, cattle, sheep) and grazing intensity are known to affect the richness and composition of plant species. However, knowledge of grazing-dependent effects on invertebrates is limited. We examined the influence of horse, cattle and sheep grazing on the richness, abundance and composition of land snail species in 21 calcareous nutrient-poor grassland areas in the northwestern Jura Mountains, Switzerland. Grazing by different livestock species did not affect the species richness, abundance and species composition of land snails. Furthermore, the number of open-land species and the ratio of large- to small-sized snail species or individuals did not differ among the three pasture types. However, independent of livestock species, grazing intensity negatively influenced the snail fauna. Snail species richness, abundance and number of Red list species decreased with increasing grazing intensity. Grazing intensity also affected the occurrence of individual snail species (Truncatellina cylindrica, Cecilioides acicula, Candidula unifasciata and Trichia plebeia). To preserve the snail fauna in nutrient-poor grasslands, pastures can be stocked with horses, cattle or sheep. However, both maximum stocking rate (number of livestock units per hectare) and grazing duration (number of grazing days per year) must be carefully defined for the proper management of the pastures.     

Effects of phosphorus enrichment and grazing snails on modern stromatolitic microbial communities

1. The effects of phosphorus enrichment and grazing snails on a benthic microbial community that builds stromatolic oncolites were examined in an experiment at Rio Mesquites, Cuatro Ciénegas, Mexico. Chemical analyses of stream water samples indicated that overall atomic ratios of total nitrogen (N) to total phosphorus (P) were approximately 110, indicating a strong potential for P‐limitation of microbial growth. 2. Phosphorus enrichment involved addition of 5 μmol Na₂HPO₄ L^?1 to streamside microcosms receiving intermittent inputs of stream water while grazer manipulation involved removal of the dominant grazer, the snail Mexithauma quadripaludium. After 7 weeks, we examined responses in organic matter content, C : N : P ratios, metabolism (P removal, primary production, dark respiration, and calcification), and microbial community structure using molecular fingerprinting of 16S rRNA genes. 3. Manipulation of snails did not affect response variables measured in these treatments (organic matter, C : P ratio, P removal rate). However, P enrichment significantly decreased the C : P and N : P ratios of surficial materials in the oncolites (organic matter content was unchanged), increased net and gross photosynthesis (oxygen consumption in the dark was unchanged), increased rates of calcification, and increased diatoms relative to cyanobacteria. Heterotrophic Eubacteria and Archaea were only modestly affected. Thus, our results indicate weak grazing effects but strong impacts of P in this benthic system. 4. We hypothesise that a state of severe P‐limitation is imposed on autotrophic production in this food web due, at least in part, to co‐precipitation of phosphate during calcite deposition. This produces severe P‐limitation of the benthic algae and cyanobacteria, resulting in high C : P ratio of microbial mats relative to the biomass of photoautotrophs (phytoplankton, terrestrial foliage) in other ecosystems. In turn, this high C : P ratio is likely to generate severe stoichiometric constraints on the herbivores, thus limiting their populations and resulting in weak overall grazing impacts.     

GROWTH DYNAMICS OF ULVA ROTUNDATA (CHLOROPHYTA) IN A FISH FARM: IMPLICATIONS FOR BIOMITIGATION AT A LARGE SCALE1

Changes in biomass of several macroalgae [Ulva rotundata Bliding; Gracilariopsis longissima (S. G. Gmel.) Steentoft, L. M. Irvine et Farnham; Ulva intestinalis L.; and Cladophora sp.] and marine plants (Zostera noltii and Ruppia cirrhosa) growing naturally in earthen ponds of a fish farm (Acuinova, San Fernando, Southern Spain) were recorded during a year. The farm is mainly devoted to the culture of gilthered seabream (Sparus aurata). The most conspicuous algal species thriving in the ponds was U. rotundata, which reached densities up to 600 g dry mass · m⁻² and produced up to 20.45 g C · m⁻² · d⁻¹. Dissolved nutrients (phosphate and ammonium), tissue nutrient content, and growth rates of this species were estimated during 2001 and 2002. Evidence of natural biomitigation by U. rotundata when water circulates throughout the fish farm is presented. Due to the fish cultivation, both phosphate and ammonium increased as water circulated from the preculture ponds to the postculture ponds. As a consequence, U. rotundata tissue nitrogen (N) and phosphorus (P) increased from algae growing in preculture ponds to algae growing in the outflow channel, so that mean C:N:P ratio varied from 773:57:1 in preculture ponds to 567:64:1 in the outflow channel. Phosphorus limited growth of U. rotundata during the spring. As growth rates increased as a function of tissue P, data were fitted to the Droop equation. From this equation, the estimated maximal growth rate was 0.295 ± 0.041 d⁻¹, the subsistence quota was 0.05 ± 0.01% P of dry mass, and the critical quota was 0.215% P of dry mass. The results suggest that management of the fish farm based on a large-scale integrated mariculture system of fish and macroalgae may increase the total ecological and economic benefits, both for the farm and for the environment.     

The effect of natural populations of the burrowing and grazing soldier crab (Mictyris longicarpus) on sediment irrigation, benthic metabolism and nitrogen fluxes

The aim of this study was to determine the effect of sediment grazing and burrowing activities of natural populations of Mictyris longicarpus on benthic metabolism, nitrogen flux and irrigation rates by comparing sediments taken from minimum disturbance exclusion cages and adjacent sediments subject to M. longicarpus activities. M. longicarpus reduced sediment surface chlorophyll a (approximately 77%), organic carbon (approximately 95%) and total nitrogen concentrations (approximately 99%) in comparison to ungrazed sediments. Consequently, they significantly reduced gross benthic O₂ production (about 71%) and sediment O₂ consumption (approximately 46%). Mean N₂ fluxes showed net effluxes (276-430 μmol m⁻² day⁻¹) in the presences of M. longicarpus and net uptakes (194.09-449.21 μmol m⁻² day⁻¹) where they were excluded. The net uptake of N₂ was most likely due to cyanobacteria fixing of N₂, as dense microbial mats became established over the sediment surface in the absence of M. longicarpus grazing activity. Sediment irrigation/transport rates calculated from CsCl tracer dilution indicated greater irrigation rates in the exclusions (12.12-16.22 l m⁻² h⁻¹) compared to inhabited sediments (6.33-11.73 l m⁻² h⁻¹) and this was again was most likely due to the lack of grazing pressure which allowed large populations of small burrowing polychaetes to inhabit the organic matter rich exclusion sediments. As such, the main influence of M. longicarpus was the interception and consumption of transported organic material, benthic microalgae and other small infaunal organisms resulting in the removal of approximately 0.06 g m⁻² day⁻¹ of nitrogen and 12.12 g m⁻² day⁻¹ of organic carbon. This “cleansing” of the sediments reduced sediment metabolism and the flux of solutes across the sediment water interface and ultimately the heavy predation of M. longicarpus by transient species such as stingrays, results in a net loss of carbon and nitrogen from the system.