Effects of in situ CO2 enrichment on the structural and chemical characteristics of the seagrass Thalassia testudinum

Seagrasses commonly display carbon-limited photosynthetic rates. Thus, increases in atmospheric pCO₂, and consequentially oceanic CO_2(aq) concentrations, may prove beneficial. While addressed in mesocosms, these hypotheses have not been tested in the field with manipulative experimentation. This study examines the effects of in situ CO_2(aq) enrichment on the structural and chemical characteristics of the tropical seagrass, Thalassia testudinum. CO_2(aq) availability was manipulated for 6 months in clear, open-top chambers within a shallow seagrass meadow in the Florida Keys (USA), reproducing forecasts for the year 2100. Structural characteristics (leaf area, leaf growth, shoot mass, and shoot density) were unresponsive to CO_2(aq) enrichment. However, leaf nitrogen and phosphorus content declined on average by 11 and 21 %, respectively. Belowground, non-structural carbohydrates increased by 29 %. These results indicate that increased CO_2(aq) availability may primarily alter the chemical composition of seagrasses, influencing both the nutrient status and resilience of these systems.     

Risk assessment of heavy metal contamination in shrimp farming in Mai Po Nature Reserve, Hong Kong

An ecological survey was carried out to determine the sediment concentrations of nutrients and heavy metals and bioaccumulation of heavy metals in fish and shrimp including tilapia (Oreochromis mossambicus×O. nilotica), grey mullet (Mugil cephalus), gei wai shrimp (Metapenaeus ensis) and caridean shrimp (Macrobrachium nipponensis) in the traditional tidal shrimp ponds (gei wais) of Mai Po Nature Reserve, Hong Kong. The sediments collected from the landward sites contained higher nutrient contents, as well as zinc (Zn), chromium (Cr), copper (Cu), nickel (Ni) and cadmium (Cd) than those collected from the seaward sites, but vice versa for lead (Pb) and mercury (Hg). However, the concentrations of all metals were exceptionally high in the two sites located outside the reserve, suggesting that waters from Deep Bay might be the possible source of metal contamination affecting the reserve. All metals studied seemed to accumulate in the viscera of fish. Body size was the determining factor for the accumulation of heavy metals in caridean shrimp and gei wai shrimp but not fish. Concentrations of the metals studied in tissues of grey mullet and gei wai shrimp were found to be safe for human consumption. Concentrations of Cr in tilapia whole body (0.68–1.10 mg kg⁻¹ wet weight) were close to or over the guideline value of 1 mg kg⁻¹ set by the Food Adulteration (Metallic Contamination) Regulations of Hong Kong. Tilapia flesh and small caridean shrimp collected from gei wais were contaminated by Cr and Pb but still fit for human consumption. Caution is required if large caridean shrimp is to be consumed in large amounts continuously because the concentration of Pb exceeded the maximum permitted concentration (6 mg kg⁻¹). The rather high Cr concentrations in tilapia whole body should not be overlooked as the fish will serve as a food source for migratory birds visiting the site.     

Effects of nutrients, herbivory, and depth on the macroalgal community in the rocky sublittoral

We studied the interacting roles of nutrient availability and herbivory in determining the macroalgal community in a rocky littoral environment. We conducted a factorial field experiment where we manipulated nutrient levels and herbivory at two sublittoral depths and measured macroalgal colonization and the following young assemblage during the growing season. At the community level, grazing reduced algal colonization, though the effect varied with depth and its interaction with nutrient availability varied in time. In shallow water, the total density of macroalgae increased in response to nutrient enrichment, but the ability of grazers to reduce macroalgal density also increased with the nutrient enrichment, and thus, the community could not escape from the top-down control. In deep water, the algal density was lower, except in July when nutrient enrichment caused a very dense algal growth. Grazing at the greater depth, though effective, was generally of smaller magnitude, and in July it could not limit algal recruitment and growth. Species richness peaked at the intermediate nutrient level in deep but not in shallow water during most of the growing season. Grazing had no effect on diversity of the algal community at either depth and only a minor effect on species richness at the greater depth. Opportunistic and ephemeral algae benefited from the nutrient enrichment but were also grazed to very low densities. Slowly growing and/or perennial species colonized poorly in the nutrient enriched treatments, and depending on the species, either suffered or indirectly benefited from herbivory. For all species, effects of nutrients on colonization depended on depth; usually both nutrient and herbivory effects were more pronounced at the shallow depth. We conclude that grazers are able to reduce macroalgae over a large range of nutrient availabilities, up to 12-fold nutrient enrichment in the current experiment, and that the sublittoral depth gradient generates variation in the algal community control exerted by both herbivory and nutrient availability. Thus temporal and spatial variability in both top-down and bottom-up control and in their interaction, especially along the depth gradient, may be crucially important for producer diversity and for the successional dynamic in a rocky sublittoral environment.     

Lionfish alter benthic invertebrate assemblages in patch habitats of a subtropical estuary

Invasive lionfish (primarily Pterois volitans) have spread throughout the Caribbean region, Gulf of Mexico, and the Southeast US coast, and resulting impacts on reef fish populations have been well documented. We examined whether lionfish can likewise affect benthic invertebrate communities, using an in situ caging experiment, in the Loxahatchee Estuary, Florida, USA. We found that lionfish caused significant declines in the three most abundant benthic invertebrate species, driving an overall shift in assemblage composition. For example, grass shrimp (Palaemonidae) abundance was reduced by nearly 90 % in the presence of lionfish. Species richness of benthic organisms was significantly higher when lionfish were present, suggesting potentially complex emergent effects of lionfish predation on benthic assemblages. Despite the fact that this experiment was conducted in just a single location using relatively small experimental units, we show altered benthic invertebrate communities could well be an additional outcome of the lionfish invasion.     

Anthropogenic subsidies alter stream consumer-resource stoichiometry, biodiversity, and food chains.

Urbanization is dramatically changing nutrient and organic matter regimes in streams, yet the community and ecosystem implications often remain obscure. We assessed the consequences of sewage-derived particulate organic matter (SDPOM) for invertebrate community structure and function in a headwater stream. Using stable isotope analyses, we found assimilation of organic SDPOM to double community secondary production, and stoichiometric analyses revealed SDPOM enriched in phosphorus (P) to foster putatively fast-growing, P-rich consumers in the subsidized reach. This altered consumer-resource stoichiometry impacted both community structure and nutrient fluxes through the invertebrate community. Community structure shifted toward significantly reduced diversity and evenness in the subsidized reach and consequently toward shorter food chains. Our integration of ecological stoichiometry with stable isotope analyses and food web ecology expands the previous focus of traditional ecotoxicology and ecophysiology to an ecosystem-level appreciation of pollutant ecology.     

Combining gut fluorescence technique and spatial analysis to determine <Emphasis Type="Italic">Littorina littorea</Emphasis> grazing dynamics in nutrient-enriched and nutrient-unenriched littoral mesocosms

Spatiotemporal distribution patterns in relation to feeding behavior of herbivorous gastropods have been studied extensively, but still knowledge about small-scale patterns is limited in relation to eutrophication. This experimental study aimed to describe the small-scale distribution of Littorina littorea in nutrient-enriched and nutrient-unenriched mesocosms in a merely atidal region and relate the distribution to food abundance and possible competing organisms, while checking simultaneously for feeding activities. The latter part was accomplished through the “gut fluorescence technique” GFT (which, to our knowledge, has not previously been used for benthic grazers) to estimate per capita grazing rates and the former part through monitoring of spatial heterogeneity of L. littorea and co-variation with sessile organisms (using semivariograms and cross-semivariograms, respectively). After 5 months of nutrient addition, the abundance and biomass of L. littorea had increased in enriched systems, which also had significantly higher total biomass of green algae. Gut pigment content was higher in L. littorea from enriched mesocosms, and gut depletion rate was higher in L. littorea from unenriched mesocosms. Spatial analysis showed that L. littorea exhibited generally random patterns (suggesting feeding activities) but sometimes (often in the morning) spatial patchiness (clumped distribution) in both enriched and unenriched conditions. There was mainly positive co-variation between L. littorea and biofilm, while different nutrient conditions exhibited contrasting co-variation between L. littorea and barnacles (positive co-variation in enriched and negative co-variation in unenriched mesocosms). The study offered insights into how feeding behavior and spatial distribution of a species may interact with community components differently under different nutrient regimes. The applied methodology can be useful for purposes of faster examination of grazing effects among different regions and also to compare grazing intensities and interactions between grazers and the benthic communities in disturbed (including pollution and nutrient enrichment) and non-disturbed systems, as well as in up-welling versus non-upwelling areas.     

Whole-system nutrient enrichment increases secondary production in a detritus-based ecosystem

Although the effects of nutrient enrichment on consumer-resource dynamics are relatively well studied in ecosystems based on living plants, little is known about the manner in which enrichment influences the dynamics and productivity of consumers and resources in detritus-based ecosystems. Because nutrients can stimulate loss of carbon at the base of detrital food webs, effects on higher consumers may be fundamentally different than what is expected for living-plant-based food webs in which nutrients typically increase basal carbon. We experimentally enriched a detritus-based headwater stream for two years to examine the effects of nutrient-induced changes at the base of the food web on higher metazoan (predominantly invertebrate) consumers. Our paired-catchment design was aimed at quantifying organic matter and invertebrate dynamics in the enriched stream and an adjacent reference stream for two years prior to enrichment and two years during enrichment. Enrichment had a strong negative effect on standing crop of leaf litter, but no apparent effect on that of fine benthic organic matter. Despite large nutrient-induced reductions in the quantity of leaf litter, invertebrate secondary production during the enrichment was the highest ever reported for headwater streams at this Long Term Ecological Research site and was 1.2-3.3 times higher than predicted based on 15 years of data from these streams. Abundance, biomass, and secondary production of invertebrate consumers increased significantly in response to enrichment, and the response was greater among taxa with larval life spans < or = 1 yr than among those with larval life spans >1 yr. Production of invertebrate predators closely tracked the increased production of their prey. The response of invertebrates was largely habitat-specific with little effect of enrichment on food webs inhabiting bedrock outcrops. Our results demonstrate that positive nutrient-induced changes to food quality likely override negative changes to food quantity for consumers during the initial years of enrichment of detritus-based stream ecosystems. Longer-term enrichment may impact consumers through eventual reductions in the quantity of detritus.     

Transient dynamics of pelagic producer-grazer systems in a gradient of nutrients and mixing depths

Phytoplankton-grazer dynamics are often characterized by long transients relative to the length of the growing season. Using a phytoplankton-grazer model parameterized for Daphnia pulex with either flexible or fixed algal carbon:nutrient stoichiometry, we explored how nutrient and light supply (the latter by varying depth of the mixed water column) affect the transient dynamics of the system starting from low densities. The system goes through an initial oscillation across nearly the entire light-nutrient supply space. With flexible (but not with fixed) algal stoichiometry, duration of the initial algal peak, timing and duration of the subsequent grazer peak, and timing of the algal minimum are consistently accelerated by nutrient enrichment but decelerated by light enrichment (decreasing mixing depth) over the range of intermediate to shallow mixing depths. These contrasting effects of nutrient vs. light enrichment are consequences of their opposing influences on food quality (algal nutrient content): algal productivity and food quality are positively related along a nutrient gradient but inversely related along a light gradient. Light enrichment therefore slows down grazer growth relative to algal growth, decelerating oscillatory dynamics; nutrient enrichment has opposite effects. We manipulated nutrient supply and mixing depth in a field enclosure experiment. The experimental results were qualitatively much more consistent with the flexible than with the fixed stoichiometry model. Nutrient enrichment increased Daphnia peak biomass, decreased algal minimum biomass, decreased the seston C:P ratio, and accelerated transient oscillatory dynamics. Light enrichment (decreasing mixing depth) produced the opposite patterns, except that Daphnia peak biomass increased monotonously with light enrichment, too. Thus, while the model predicts the possibility of the "paradox of energy enrichment" (a decrease in grazer biomass with light enrichment) at high light and low nutrient supply, this phenomenon did not occur in our experiment.     

Grass shrimp (Palaemonetes spp.) play a pivotal trophic role in enhancing Ruppia maritima

Coupled trophic-engineer interactions are potentially important for maintaining habitat function and ecosystem services. As ephemeral submerged aquatic vegetation (SAV), Ruppia maritima has a short well-defined growth-senescence cycle and should benefit from any ecological interaction that enhances its physical condition and longevity. Grass shrimp (Palaemonetes spp.) are abundant facultative grazers of epiphytic algae and conveyors of nutrients in tidal marsh and SAV habitats. Grass shrimp addition consistently enhanced Ruppia biomass and shoot density in a series of three field experiments conducted in Grand Bay National Estuarine Research Reserve, Mississippi, USA. In two experiments, epiphyte grazing by grass shrimp enhanced Ruppia by inhibiting die-back during the mid- and latter stages of the Ruppia life cycle. Despite a nonsignificant epiphyte grazing effect, grass shrimp also enhanced Ruppia during its early growth stage in a third experiment. In that experiment, nutrient addition also significantly increased epiphyte biomass. Grass shrimp may have fostered the early growth of Ruppia through direct deposition of feces to the sediment in the third experiment. Grass shrimp play a pivotal trophic role in the maintenance of Ruppia through context-dependent interactions involving stage of the SAV life cycle, season, and nutrient limitation.     

Microbial activity and carbon, nitrogen, and phosphorus content in a subtropical seagrass estuary (Florida Bay): evidence for limited bacterial use of seagrass production

Bacterial abundance, production, and extracellular enzyme activity were determined in the shallow water column, in the epiphytic community of Thalassia testudinum, and at the sediment surface along with total carbon, nitrogen, and phosphorus in Florida Bay, a subtropical seagrass estuary. Data were statistically reduced by principle components analysis (PCA) and multidimensional scaling and related to T. testudinum leaf total phosphorus content and phytoplankton biomass. Each zone (i.e., pelagic, epiphytic, and surface sediment community) was significantly dissimilar to each other (Global R = 0.65). Pelagic aminopeptidase and sum of carbon hydrolytic enzyme (esterase, peptidase, and α- and β-glucosidase) activities ranged from 8 to 284 mg N m⁻² day⁻¹ and 113–1,671 mg C m⁻² day⁻¹, respectively, and were 1–3 orders of magnitude higher than epiphytic and sediment surface activities. Due to the phosphorus-limited nature of Florida Bay, alkaline phosphatase activity was similar between pelagic (51–710 mg P m⁻² day⁻¹) and sediment (77–224 mg P m⁻² day⁻¹) zones but lower in the epiphytes (1.1–5.2 mg P m⁻² day⁻¹). Total (and/or organic) C (111–311 g C m⁻²), N (9.4–27.2 g N m⁻²), and P (212–1,623 mg P m⁻²) content were the highest in the sediment surface and typically the lowest in the seagrass epiphytes, ranging from 0.6 to 8.7 g C m⁻², 0.02–0.99 g N m⁻², and 0.5–43.5 mg P m⁻². Unlike nutrient content and enzyme activities, bacterial production was highest in the epiphytes (8.0–235.1 mg C m⁻² day⁻¹) and sediment surface (11.5–233.2 mg C m⁻² day⁻¹) and low in the water column (1.6–85.6 mg C m⁻² day⁻¹). At an assumed 50% bacterial growth efficiency, for example, extracellular enzyme hydrolysis could supply 1.8 and 69% of epiphytic and sediment bacteria carbon demand, respectively, while pelagic bacteria could fulfill their carbon demand completely by enzyme-hydrolyzable organic matter. Similarly, previously measured T. testudinum extracellular photosynthetic carbon exudation rates could not satisfy epiphytic and sediment surface bacterial carbon demand, suggesting that epiphytic algae and microphytobenthos might provide usable substrates to support high benthic bacterial production rates. PCA revealed that T. testudinum nutrient content was related positively to epiphytic nutrient content and carbon hydrolase activity in the sediment, but unrelated to pelagic variables. Phytoplankton biomass correlated positively with all pelagic components and sediment aminopeptidase activity but negatively with epiphytic alkaline phosphatase activity. In conclusion, seagrass production and nutrient content was unrelated to pelagic bacteria activity, but did influence extracellular enzyme hydrolysis at the sediment surface and in the epiphytes. This study suggests that seagrass-derived organic matter is of secondary importance in Florida Bay and that bacteria rely primarily on algal/cyanobacteria production. Pelagic bacteria seem coupled to phytoplankton, while the benthic community appears supported by epiphytic and/or microphytobenthos production.     

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.     

Crustacean epifauna of seagrass and macroalgae in Apalachee Bay,Florida, USA

Epifaunal crustaceans on turtlegrass (Thalassia testudinum) and five dominant macroalgae (Anadyomene stellata, Digenia simplex, Halimeda incrassata, Laurencia poitei and Penicillus lamourouxii) were quantitatively sampled bimonthly over a one-year period from September 1979 to September 1980 in a subtropical seagrass meadow in Apalachee Bay, Florida (northeastern Gulf of Mexico). These plant species exhibited a wide range of morphologies, with surface area-to-biomass ratios differing by over 2.5 times. A similar suite of crustaceans occurred on all macrophytes despite differences in shape or architecture among plant species. Relative abundances of many crustaceans, however, varied among plant hosts. Similarity analysis indicated that the epifaunal associates of T. testudinum were distinct from those of the macroalgae. Species richness was generally higher on turtlegrass than on any of the macroalgae. Abundances of total crustaceans per plant biomass or per plant surface area, on the other hand, were greater on all macroalgal species compared to the seagrass. Abundances (per plant biomass or plant surface area) of 14 of the 16 numerically dominant epifaunal species differed significantly among macrophytes. Twelve of the 16 species had greater abundance on one or more macroalgae, while only two species were more abundant on T. testudinum. Almost half of the dominant species had greatest abundances on the branching red alga L. poitei. Although abundances per plant biomass and plant surface area were greater on macroalgae relative to turtlegrass, densities (individuals per meter square of bottom) of animals associated with T. testudinum were significantly greater than those associated with macroalgae, primarily because of the greater abundance of turtlegrass in the grass bed. Both surface area-to-biomass ratios and degree of branching were poorly correlated with epifaunal abundance and number of species. Neither structural feature is an adequate predictor of faunal abundance and species richness among plant species, especially when macrophytes with very different morphologies are compared.     

Omnivory and grazer functional composition moderate cascading trophic effects in experimental <Emphasis Type="Italic">Fucus vesiculosus</Emphasis> habitats

We tested the relative strength of direct versus indirect effects of an aquatic omnivore depending on the functional composition of grazers by manipulating the presence of gastropod and amphipod grazers and omnivorous shrimp in outdoor mesocosms. By selectively preying upon amphipods and reducing their abundance by 70–80%, omnivorous shrimp favoured the dominance of gastropods. While gastropods were the main microalgal grazers, amphipods controlled macroalgal biomass in the experiment. However, strong predation on the amphipod by the shrimp had no significant indirect effects on macroalgal biomass, indicating that when amphipod abundances declined, complementary feeding by the omnivore on macroalgae may have suppressed a trophic cascade. Accordingly, in the absence of amphipods, the shrimp grazed significantly on green algae and thereby suppressed the diversity of the macroalgal community. Our experiment demonstrates direct consumer effects by an omnivore on both the grazer and producer trophic levels in an aquatic food web, regulated by prey availability.     

Grazers, producer stoichiometry, and the light : nutrient hypothesis revisited

The stoichiometric light:nutrient hypothesis (LNH) links the relative supplies of key resources with the nutrient content of tissues of producers. This resource-driven variation in producer stoichiometry, in turn, can mediate the efficiency of grazing. Typically, discussions of the LNH attribute this resource-stoichiometry link to bottom-up effects of light and phosphorus, which are mediated through producer physiology. Emphasis on bottom-up effects implies that grazers must consume food of quality solely determined by resource supply to ecosystems (i.e., they eat what they are served). Here, we expand upon this largely bottom-up interpretation with evidence from pond surveys, a mesocosm experiment, and a model. Data from shallow ponds showed the "LNH pattern" (positive correlation of an index of light : phosphorus supply with algal carbon : phosphorus content). However, algal carbon : phosphorus content also declined as zooplankton biomass increased in the ponds. The experiment and model confirmed that this latter correlation was partially caused by the various bottom-up and top-down roles of grazers: the LNH pattern emerged only in treatments with crustacean grazers, not those without them. Furthermore, model and experiment clarified that another bottom-up factor, natural covariation of nitrogen : phosphorus ratios with light : phosphorus supply (as seen in ponds), does not likely contribute to the LNH pattern. Finally, the experiment produced correlations between shifts in species composition of algae, partially driven by grazing effects of crustaceans, and algal stoichiometry. These shifts in species composition might shape stoichiometric response of producer assemblages to resource supply and grazing, but their consequences remain largely unexplored. Thus, this study accentuated the importance of grazing for the LNH; de-emphasized a potentially confounding, bottom-up factor (covarying nitrogen : phosphorus supply); and highlighted an avenue for future research for the LNH (grazer-mediated shifts in producer composition).     

Leaf reddening in the seagrass Thalassia testudinum in relation to anthocyanins, seagrass physiology and morphology, and plant protection

Numerous seagrass species growing in high-light environments develop red coloration in otherwise green leaves, yet the ecophysiology of leaf reddening in seagrasses is poorly understood. To increase our understanding of the process of leaf reddening in Thalassia testudinum found in the lower Florida Keys (USA), we identified the molecules responsible for red coloration in leaves and compared physiological, morphological, and growth attributes of entirely red-leafed shoots to entirely green-leafed shoots. We determined that four anthocyanin molecules are responsible for red coloration in leaves. In addition, we found that red leaves had higher concentrations of photoprotective pigments (anthocyanins and UV-absorbing compounds), higher effective quantum yields (ΔF/F _m′) at midday, and were shorter, narrower, and weighed less than green leaves. No significant difference in growth rates was observed between red- and green-leafed shoots, but patches of red-leafed shoots had shorter canopy heights and smaller LAI compared to patches of green-leafed shoots. Our results demonstrate that leaf reddening in T. testudinum is caused by high concentrations of anthocyanins, is associated with physiological and morphological attributes, and acts as a sunscreen since red leaves were able to maintain high effective quantum yields at high light intensities.     

Simulated green turtle grazing affects nutrient composition of the seagrass <Emphasis Type="Italic">Thalassia testudinum</Emphasis>

Before populations of green turtles (Chelonia mydas) were severely reduced by human overexploitation, the seagrass Thalassia testudinum was intensively grazed by green turtles in the Caribbean. To explore how nutrient composition of T. testudinum pastures responds to intense grazing pressure, we simulated green turtle grazing in 15 plots (each 3 m × 3 m) for 16 months in the central Bahamas. Comparisons of clipped plots with 15 adjacent control (unclipped) plots revealed that simulated grazing resulted in significantly higher energy, nitrogen, phosphorus, lignin, cutin, and condensed tannin content in blades in clipped plots, but sediment organic content was not affected. By continually re-cropping blades in grazing plots, turtles ingest young, actively growing blade tissue with higher energy, nitrogen, and phosphorus concentrations. Our 16-month clipping trial did not generate the expected decline in nutrient content in T. testudinum blades under intensive grazing. However, significant decreases in nitrogen and organic matter reserves in rhizomes, with declines apparent after 16 and 11 months, respectively, indicate that nutrient content of blades and/or blade productivity may decline under continued clipping.     

Consumers affect prey biomass and diversity through resource partitioning

Consumer presence and nutrient availability can have contrasting and interactive effects on plant diversity. In a factorial experiment, we manipulated two levels of nutrient supply and the presence of two moderately specialized grazers in different combinations (no grazers, two species in monoculture, and both in combination). We tested how nutrients and grazers regulated the biomass of marine coastal epiphytes and the diversity of algal assemblages, based on the prediction that the effect of consumers on prey diversity depends on productivity and consumer specialization. Nutrient enrichment increased the epiphytic load, while monocultures of single grazer species partly prevented epiphyte growth. However, only the presence of two species with complementary feeding preferences effectively prevented epiphyte overgrowth. The epiphytes comprised micro- and macroalgal species, and the diversity of these algal assemblages differed, depending on grazer identity. For the microalgae, diversity was reduced by nutrient addition when grazer control was inefficient, but not when specialist microalgal grazers were present. Macroalgal diversity was reduced in ambient water with specialist macroalgal grazers compared to the treatment with inefficient ones. These results indicate that grazer composition and productivity are crucial in determining whether consumer pressure will have a positive or negative effect on algal diversity.     

Variable effects of symbiotic snapping shrimps on their sponge hosts

Mutualistic relationships are ubiquitous in tropical coral reefs, but the costs and benefits to partner species are often poorly known. In Caribbean coral reefs, several species of snapping shrimp (Synalpheus spp.) dwell exclusively in marine sponges, which serve as both habitat and food source. A paired experimental design was used to examine the effects of Synalpheus occupancy on predation, morphology, and growth of their sponge host Lissodendoryx colombiensis in Bocas del Toro, Panama (9.351°N, 82.258°W) in June 2009. Shrimp occupancy significantly decreased consumption of sponges by a predatory sea star (Oreaster reticulatus) and also affected sponge morphology; sponges grown without shrimps decreased in canal size, in both the laboratory and the field. Shrimp occupancy had more ambiguous effects on sponge growth. In laboratory experiments, shrimp occupancy benefited sponge growth, although all sponges experienced overall decreases in mass. In field experiments, there were no significant differences in growth between occupied and empty sponges. However, the benefits of shrimp occupancy on sponge growth were negatively correlated with overall increases in sponge size; sponges that decreased in mass during the experiment benefited more from shrimp occupancy than sponges that increased in mass. These costs and benefits suggest that Synalpheus has variable effects on sponges: positive effects on sponges in the presence of predators, and/or when sponges are decreasing in mass (e.g., during periods of physical stress), but a negative effect on sponges during periods of active sponge growth.     

A stochastic simulation model of brown shrimp (Penaeus aztecus ives) growth,movement, and survival in Galveston Bay,Texas

A stochastic simulation model of brown shrimp (Penaeus aztecus Ives) population dynamics in Galveston Bay, Texas, is described, validated, and used to evaluate the effects of management alternatives and changing environmental conditions on shrimp dynamics. The model is composed of submodels representing: (1) recruitment, (2) growth, (3) natural mortality, (4) fishing mortality, and (5) emigration of brown shrimp. The model predicts significant changes in total annual harvest from the food shrimp, bait, and recreational fisheries resulting from (1) closure of the bay system to all fishing except during the spring and fall open seasons, (2) two-week postponement of the opening and closing of the open seasons for the food shrimp fishery, (3) a 2.5°C increase and (4) a 2.5°C decrease in mean water temperature, (5) an 80% increase and (6) an 80% decrease in fishing effort. No significant change in the total annual harvest is predicted when the food shrimp fishing season is extended from May 15 through December 15. Sensitivity analysis suggests that field experimentation designed specifically to test the hypothesis of a 60-day time lag between brown shrimp recruitment into the bays and exposure to the fishery should receive high priority. Simulation results are discussed within a management framework.     

Dietary-induced responses in the phagotrophic flagellate Oxyrrhis marina

Primary producers may be limited by different nutrients as well as by light availability, which in turn affects their quality as food for higher trophic levels. Typically, algae with high C:N and/or C:P ratios are low-quality food for consumers. Heterotrophic protists are important grazers on these autotrophes, but despite their importance as grazers, knowledge of food quality effects on heterotrophic protists is sparse. In the present study, we examined how differently grown Rhodomonas salina (nutrient replete, N-limited and P-limited) affected the phagotrophic flagellate Oxyrrhis marina. The functional response of O. marina (based on ingested biovolume) did not show significant differences between food sources, thus food uptake was independent of food quality. O. marina was weakly homoeostatic which means that its C:N:P ratio still reflected the elemental composition of its food to some extent. Food quality had a significantly negative effect on the numerical response of O. marina. Whereas N-limited R. salina and nutrient replete R. salina resulted in similar growth rates, P-limited algae had a significantly negative effect on the specific growth rate of O. marina. Hence, the lack of elemental phosphorus of O. marina feeding on P-limited algae caused a reduction in growth. Thus, despite their weaker homoeostasis, heterotrophic protists are also affected by high C:P food in a similar way to crustacean zooplankton.