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Local variability but landscape stability in coral reef communities following repeated hurricane impacts

Coral reef community structure has remained remarkably stable over a 10 yr period within a small protected marine area despite repeated hurricane impacts. Local community dynamics have been highly variable, however. Sites that were destroyed by disease in the 1970s are showing little or no recovery, while sites less than a kilometre away that were devastated by Hurricane Hugo in 1989 are recovering well. Strong coral recruitment has occurred in shallow, exposed areas that showed the greatest hurricane impacts, and these areas are now more species rich than in 1988, although coral cover has not reached pre-hurricane levels. Coral colony survivorship has been high throughout most of the study area. Partial mortality rates were elevated for several years following Hurricane Hugo, but significant whole coral-head mortality only occurred during periods with hurricane impacts and only at the most exposed sites. Overall, the coral community has proved resilient to closely repeated major hurricane impacts. From a single case study we cannot attribute this resilience to the relatively low level of human impacts, but grazing fish populations have apparently remained high enough to keep macroalgae in check despite the mass mortality of the herbivore Diadema antillarum in the 1980s.

St. Croix

Our evolving conceptual model of the coastal eutrophication problem

A primary focus of coastal science during the past 3 decades has been the question: How does anthropogenic nutrient enrichment cause change in the structure or function of nearshore coastal ecosystems? This theme of environmental science is recent, so our conceptual model of the coastal eutrophication problem continues to change rapidly. In this review, I suggest that the early (Phase I) conceptual model was strongly influenced by limnologists, who began intense study of lake eutrophication by the 1960s. The Phase I model emphasized changing nutrient input as a signal, and responses to that signal as increased phytoplankton biomass and primary production, decomposition of phytoplankton-derived organic matter, and enhanced depletion of oxygen from bottom waters. Coastal research in recent decades has identified key differences in the responses of lakes and coastal-estuarine ecosystems to nutrient enrichment. The contemporary (Phase II) conceptual model reflects those differences and includes explicit recognition of (1) system-specific attributes that act as a filter to modulate the responses to enrichment (leading to large differences among estuarine-coastal systems in their sensitivity to nutrient enrichment); and (2) a complex suite of direct and indirect responses including linked changes in: water transparency, distribution of vascular plants and biomass of macroalgae, sediment biogeochemistry and nutrient cycling, nutrient ratios and their regulation of phytoplankton community composition, frequency of toxic/harmful algal blooms, habitat quality for metazoans, reproduction/growth/survival of pelagic and benthic invertebrates, and subtle changes such as shifts in the seasonality of ecosystem functions. Each aspect of the Phase II model is illustrated here with examples from coastal ecosystems around the world. In the last section of this review I present one vision of the next (Phase III) stage in the evolution of our conceptual model, organized around 5 questions that will guide coastal science in the early 21st century: (1) How do system-specific attributes constrain or amplify the responses of coastal ecosystems to nutrient enrichment? (2) How does nutrient enrichment interact with other stressors (toxic contaminants, fishing harvest, aquaculture, nonindigenous species, habitat loss, climate change, hydrologic manipulations) to change coastal ecosystems? (3) How are responses to multiple stressors linked? (4) How does human-induced change in the coastal zone impact the Earth system as habitat for humanity and other species? (5) How can a deeper scientific understanding of the coastal eutrophication problem be applied to develop tools for building strategies at ecosystem restoration or rehabilitation?

Marine Ecology Progress Series

Influence of reactive sulfide (AVS) and supplementary food on Ag, Cd and Zn bioaccumulation in the marine polychaete Neanthes arenaceodentata

A laboratory bioassay determined the relative contribution of various pathways of Ag, Cd and Zn bioaccumulation in the marine polychaete Neanthes arenaceodentata exposed to moderately contaminated sediments. Juvenile worms were exposed for 25 d to experimental sediments containing 5 different reactive sulfide (acid volatile sulfides, AVS) concentrations (1 to 30 ??mol g-1), but with constant Ag, Cd, and Zn concentrations of 0.1, 0.1 and 7 ??mol g-1, respectively. The sediments were supplemented with contaminated food (TetraMin??) containing 3 levels of Ag-Cd-Zn (uncontaminated, 1?? or 5??1 metal concentrations in the contaminated sediment). The results suggest that bioaccumulation of Ag, Cd and Zn in the worms occurred predominantly from ingestion of contaminated sediments and contaminated supplementary food. AVS or dissolved metals (in porewater and overlying water) had a minor effect on bioaccumulation of the 3 metals in most of the treatments. The contribution to uptake from the dissolved source was most important in the most oxic sediments, with maximum contributions of 8% for Ag, 30% for Cd and 20% for Zn bioaccumulation. Sediment bioassays where uncontaminated supplemental food is added could seriously underestimate metal exposures in an equilibrated system; N. arenaceodentata feeding on uncontaminated food would be exposed to 40-60% less metal than if the food source was equilibrated (as occurs in nature). Overall, the results show that pathways of metal exposure are dynamically linked in contaminated sediments and shift as external geochemical characteristics and internal biological attributes vary.

Marine Ecology Progress Series

Food limitation and the recovery of sea otters following the 'Exxon Valdez' oil spill

We examined the potential role of food limitation in constraining the recovery of sea otters Enhydra lutris in Prince William Sound, Alaska, following the 'Exxon Valdez' oil spill. The spill resulted in the removal of a large number of sea otters in 1989, and as of 1998, the portion of the population in the heavily oiled northern Knight Island region had not fully recovered. Between 1996 and 1998, prey consumption rate was higher and the condition of sea otters was better at northern Knight Island than in an unoiled area of the sound (Montague Island). Estimates of prey energy available per unit mass of sea otter were about 4 times higher at Knight than Montague Island, albeit not significantly different between the 2 areas. Over this same period, the number of sea otters remained constant at northern Knight Island but increased at Montague Island. These data suggest that food was at least as abundant at Knight than at Montague Island, and that recovery of sea otters via intrinsic population growth was limited by factors other than food. However, the availability of food, the prey consumption rate, and the condition of sea otters were all much lower at both Knight and Montague Islands than in areas newly occupied by sea otters where the population growth rate was near the theoretical maximum. It is possible that the relatively short supply of food (compared to areas where sea otter population growth rate was high) may have inhibited immigration or interacted with other factors (e.g. oil-induced mortality or predation) to restrict sea otter population growth. Nonetheless, these data suggest that impacts of anthropogenic disturbances on large, often food-limited vertebrate predators can persist in spite of the availability of food resources that are sufficient for intrinsic population growth.

Alaska

Harlequin duck population recovery following the 'Exxon Valdez' oil spill: Progress, process and constraints

Following the 1989 'Exxon Valdez' oil spill in Prince William Sound, Alaska, we studied the status of recovery of harlequin duck Histrionicus histrionicus populations during 1995 to 1998. We evaluated potential constraints on full recovery, including (1) exposure to residual oil; (2) food limitation; and (3) intrinsic demographic limitations on population growth rates. In this paper, we synthesize the findings from our work and incorporate information from other harlequin duck research and monitoring programs to provide a comprehensive evaluation of the response of this species to the 'Exxon Valdez' spill. We conclude that harlequin duck populations had not fully recovered by 1998. Furthermore, adverse effects continued as many as 9 yr after the oil spill, in contrast to the conventional paradigm that oil spill effects on bird populations are short-lived. These conclusions are based on the findings that (1) elevated cytochrome P450 (CYP1A) induction on oiled areas indicated continued exposure to oil in 1998; (2) adult female winter survival was lower on oiled than unoiled areas during 1995 to 1998; (3) fall population surveys by the Alaska Department of Fish and Game indicated numerical declines in oiled areas during 1995 to 1997; and (4) densities on oiled areas in 1996 and 1997 were lower than expected using models that accounted for effects of habitat attributes. Based on hypothesized links between oil contamination and demography, we suggest that harlequin duck population recovery was constrained primarily by continued oil exposure. Full population recovery will also be delayed by the time necessary for intrinsic population growth to allow return to pre-spill numbers following cessation of residual oil spill effects. Although not all wildlife species were affected by the 'Exxon Valdez' oil spill, and some others may have recovered quickly from any effects, harlequin duck life history characteristics and benthic, nearshore feeding habits make them susceptible to both initial and long-term oil spill effects.

Alaska

Living on the edge: Distribution of Dungeness crab Cancer magister in a recently deglaciated fjord

Glacier Bay, Alaska, has supported a productive Dungeness crab fishery, although the area where the fishery occurred was small relative to the remainder of the Bay. We hypothesized that 1 or more abiotic limiting factors prevented crabs from surviving in the upper Bay. We tested this hypothesis by systematically sampling for relative abundance of Dungeness crabs from the mouth to the head of the Bay. We measured salinity, temperature and turbidity at each of the sampling sites and at permanent stations to characterize the habitat within and among years. We expected to find a completely truncated spatial distribution. Instead, we found that adult Dungeness crabs can survive in habitats heavily influenced by tidewater glaciers. There were significantly fewer crabs in the upper Bay and a sharp decrease in abundance around 40 km from the mouth of the Bay. The region of the Bay beyond 40 km from the mouth had a very low density of predominately adult male crabs. The narrow size frequency distribution and the lack of small crabs is consistent with poor survival in one of the pre-adult life stages in the upper Bay.

Alaska

Subcellular compartmentalization of Cd and Zn in two bivalves. I. Significance of metal-sensitive fractions (MSF) and biologically detoxified metal (BDM)

Many aspects of metal accumulation in aquatic invertebrates (i.e. toxicity, tolerance and trophic transfer) can be understood by examining the subcellular partitioning of accumulated metal. In this paper, we use a compartmentalization approach to interpret the significance of metal, species and size dependence in the subcellular partitioning of Cd and Zn in the bivalves Macoma balthica and Potamocorbula amurensis . Of special interest is the compartmentalization of metal as metal-sensitive fractions (MSF) (i.e. organelles and heat-sensitive proteins, termed Œenzymes¹ hereafter) and biologically detoxified metal (BDM) (i.e. metallothioneins [MT] and metal-rich granules [MRG]). Clams from San Francisco Bay, CA, were exposed for 14 d to seawater (20” salinity) containing 3.5 µg l -1 Cd and 20.5 µg l -1 Zn, including 109 Cd and 65 Zn as radiotracers. Uptake was followed by 21 d of depuration. The subcellular partitioning of metal within clams was examined following exposure and loss. P. amurensis accumulated ~22x more Cd and ~2x more Zn than M. balthica . MT played an important role in the storage of Cd in P. amurensis , while organelles were the major site of Zn accumulation. In M. balthica , Cd and Zn partitioned similarly, although the pathway of detoxification was metal-specific (MRG for Cd; MRG and MT for Zn). Upon loss, M. balthica depurated ~40% of Cd with Zn being retained; P. amurensis retained Cd and depurated Zn (~40%). During efflux, Cd and Zn concentrations in the MSF compartment of both clams declined with metal either being lost from the animal or being transferred to the BDM compartment. Subcellular compartmentalization was also size-dependent, with the importance of BDM increasing with clam size; MSF decreased accordingly. We hypothesized that progressive retention of metal as BDM (i.e. MRG) with age may lead to size dependency of metal concentrations often observed in some populations of M. balthica .

Marine Ecology Progress Series

Vertical structure of the phytoplankton community associated with a coastal plume in the Gulf of Mexico

Low salinity plumes of coastal origin are occasionally found far offshore, where they display a distinct color signature detectable by satellites. The impact of such plumes on carbon fixation and phytoplankton community structure in vertical profiles and on basin wide scales is poorly understood. On a research cruise in June 1999, ocean-color satellite-images (Sea-viewing Wide Field-of-view Sensor, SeaWiFS) were used in locating a Mississippi River plume in the eastern Gulf of Mexico . Profiles sampled within and outside of the plume were analyzed using flow cytometry, HPLC pigment analysis and primary production using 14 C incorporation. Additionally, RubisCO large subunit (rbcL) gene expression was measured by hybridization of extracted RNA using 3 full-length RNA gene probes specific for individual phytoplankton clades. We also used a combination of RT-PCR/PCR and TA cloning in order to generate cDNA and DNA rbcL clone libraries from samples taken in the plume . Primary productivity was greatest in the low salinity surface layer of the plume . The plume was also associated with high Synechococcus counts and a strong peak in Form IA rbcL expression. Form IB rbcL (green algal) mRNA was abundant at the subsurface chlorophyll maximum (SCM), whereas Form ID rbcL (chromophytic) expression showed little vertical structure . Phylogenetic analysis of cDNA libraries demonstrated the presence of Form IA rbcL Synechococcus phylotypes in the plume . Below the plume , 2 spatially separated and genetically distinct rbcL clades of Prochlorococcus were observed. This indicated the presence of the high- and low-light adapted clades of Prochlorococcus. A large and very diverse clade of Prymnesiophytes was distributed throughout the water column, whereas a clade of closely related prasinophytes may have dominated at the SCM. These data indicate that the Mississippi river plume may dramatically alter the surface picoplankton composition of the Gulf of Mexico , with Synechococcus displacing Prochlorococcus in the surface waters.

Gulf of Mexico

Sediment DIN fluxes and preferential recycling of benthic microalgal nitrogen in a shallow macrotidal estuary

Sediment -water fluxes of NH 4 + , NO 3 - , dissolved inorganic carbon, and O 2 were measured in cores collected from the upper Rowley River estuary , Massachusetts, and used to calculate rates of organic nitrogen (N) mineralization, nitrification, and coupled and direct denitrification (DNF). The cores contained 15 N label in benthic microalgae (BMA) and in NO 3 - in the overlying water as a result of an ongoing whole- estuary 15 NO 3 - enrichment study (NISOTREX II). The tracer allowed for estimation of gross NO 3 - regeneration in sediments and the contribution of BMA derived N to total mineralization. The mean mineralization rate between sites was 16.0 ± 2.0 mmol N m -2 d -1 . Approximately 13 to 56% of the mineralized N was nitrified at rates ranging from 1.8 to 10.1 mmol N m -2 d -1 . Total denitrification was dominated by direct DNF (3.6 mmol N m -2 d -1 ) furthest upstream, where NO 3 - concentrations were highest. Coupled DNF was most important (8.0 mmol N m -2 d -1 ) in the sediments with high nitrification and low water column NO 3 - . A gross NO 3 - flux from sediments to water of 0.9 to 2.1 mmol N m -2 d -1 was estimated from the isotope dilution of δ 15 NO 3 - in the overlying water of the cores. The isotope dilution seen in the cores was also detected as a deviation from conservative δ 15 NO 3 - mixing along estuarine transects. Incorporation of this NO 3 - regeneration into the DNF calculations effectively increased the estimate of direct DNF by up to 50% and decreased the coupled DNF estimate by up to 220%. Increasing δ 15 NH 4 + in the water of the cores indicated that the 15 N-labelled BMA were preferentially mineralized over bulk sediment organic N. Additional 15 N enrichments in the sediment bacterial biomarker diaminopimelic acid showed a link among 15 N-labeled BMA, active bacteria, and 15 NH 4 + released to the overlying water. Based on δ 15 NH 4 + enrichments in the cores, BMA accounted for approximately 50 to 100% of the N mineralized. An isotopic enrichment of δ 15 NH 4 + above background in the estuary was observed at a magnitude consistent with the core-based rates of BMA mineralization. These results provide further evidence that BMA are not unidirectional sinks for water column-dissolved organic nitrogen , but instead act to turn over N between sediments and estuarine water on the scale of days.

Marine Ecology Progress Series

Subcellular compartmentalization of Cd and Zn in two bivalves. II. Significance of trophically available metal (TAM)

This paper examines how the subcellular partitioning of Cd and Zn in the bivalves Macoma balthica and Potamocorbula amurensis may affect the trophic transfer of metal to predators. Results show that the partitioning of metals to organelles, Œenzymes¹ and metallothioneins (MT) comprise a subcellular compartment containing trophically available metal (TAM; i.e. metal trophically available to predators), and that because this partitioning varies with species, animal size and metal, TAM is similarly influenced. Clams from San Francisco Bay, California, were exposed for 14 d to 3.5 µg l -1 Cd and 20.5 µg l -1 Zn, including 109 Cd and 65 Zn as radiotracers, and were used in feeding experiments with grass shrimp Palaemon macrodatylus , or used to investigate the subcellular partitioning of metal. Grass shrimp fed Cd-contaminated P. amurensis absorbed ~60% of ingested Cd, which was in accordance with the partitioning of Cd to the bivalve¹s TAM compartment (i.e. Cd associated with organelles, Œenzymes¹ and MT); a similar relationship was found in previous studies with grass shrimp fed Cd-contaminated oligochaetes. Thus, TAM may be used as a tool to predict the trophic transfer of at least Cd. Subcellular fractionation revealed that ~34% of both the Cd and Zn accumulated by M. balthica was associated with TAM, while partitioning to TAM in P. amurensis was metal-dependent (~60% for TAM-Cd%, ~73% for TAM-Zn%). The greater TAM-Cd% of P. amurensis than M. balthica is due to preferential binding of Cd to MT and Œenzymes¹, while enhanced TAM-Zn% of P. amurensis results from a greater binding of Zn to organelles. TAM for most species-metal combinations was size-dependent, decreasing with increased clam size. Based on field data, it is estimated that of the 2 bivalves, P. amurensis poses the greater threat of Cd exposure to predators because of higher tissue concentrations and greater partitioning as TAM; exposure of Zn to predators would be similar between these species.

Marine Ecology Progress Series

Benthic sulfate reduction along the Chesapeake Bay central channel. II. Temporal controls

Seasonal and interannual controls of benthic sulfate reduction (SR) were examined at 3 sites (upper [UB], mid- [MB] and lower [LB] bay) along the Chesapeake Bay central channel, from early spring through fall, for 6 yr (1989 to 1994). The combined influences of temperature, sulfate, organic loading and bioturbation affected seasonal SR rates differently in the 3 regions. Consistently low SR rates at UB resulted from low overlying-water sulfate concentrations and the dominance of refractory organic terrestrial material. Combined seasonal variation in temperature and sulfate accounted for 50% of the annual variability in 0 to 2 cm depth interval SR rates, while sediment organic content had no significant seasonal influence. In contrast, MB and LB sites had high rates of SR fostered by high levels of overlying water SO42- and organic input dominated by labile phytoplankton detritus. New organic loading (measured as chl a) stimulated 0 to 2 cm SR during spring at both sites. Combined organic quantity (as particulate C and/or N) and temperature accounted for > 75% of the variability in 0 to 2 cm SR at MB during spring and fall. Molecular diffusion supplied 25 to 45% of the SO 42- needed to fuel 0 to 12 cm depth interval SR at MB, with the balance presumably supplied by S-recycling. Interannual differences in summertime SR rates were linked to the extent of freshwater flow during spring, with high-flow years associated with high SR rates at UB and MB, and low rates at LB. The negative trend between benthic SR and river flow at LB may result from the up-estuary transport of senescing organic matter in bottom water, which increases in the lower reach of the estuary with increasing freshwater inflow.

Chesapeake Bay

Mobile fishing gear reduces benthic megafaunal production on Georges Bank

This study addresses the effect of mobile fishing gear disturbance on benthic megafaunal production on the gravel pavement of northern Georges Bank. From 1994 to 2000, we sampled benthic megafauna with a 1 m Naturalists' dredge at shallow (47 to 62 m) and deep (80 to 90 m) sites. The cessation of fishing in large areas of Georges Bank in January 1995 allowed us to monitor changes in production at a previously disturbed site. Production at a shallow disturbed site varied little over the sampling period (32 to 57 kcal m-2 yr-1) and was markedly lower than production at the nearby recovering site, where production increased from 17 kcal m-2yr -1 in 1994 before the closure to 215 kcal m-2 yr -1 in 2000. Atlantic sea scallops Placopecten magellanicus and green sea urchins Strongylocentrotus droebachiensis dominated production at the recovering site. The community production:biomass ratio decreased over time at the recovering site as the sea scallop population matured. At the deep sites, production remained significantly higher at undisturbed sites (174 to 256 kcal m-2 yr-1) than at disturbed sites (30 to 52 kcal m -2 yr-1). The soft-bodied tube-building polychaete Thelepus cincinnatus dominated production at the undisturbed site, while hard-shelled bivalve molluscs Astarte spp. and P. magellanicus were prevalent at the disturbed site. Mobile fishing gear disturbance has a conspicuous effect on benthic megafaunal production in this hard-bottom habitat. Cessation of mobile fishing has resulted in a marked increase in benthic megafaunal production. These findings should help fishery managers to gauge the costs and benefits of management tools such as area closures and low-impact fishing gears.

Georges Banks

Hydrologic aspects of marsh ponds during winter on the Gulf Coast Chenier Plain, USA: Effects of structural marsh management

The hydrology of marsh ponds influences aquatic invertebrate and waterbird communities. Hydrologic variables in marsh ponds of the Gulf Coast Chenier Plain are potentially affected by structural marsh management (SMM: levees, water control structures and impoundments) that has been implemented since the 1950s. Assuming that SMM restricts tidal flows and drainage of rainwater, we predicted that SMM would increase water depth, and concomitantly decrease salinity and transparency in impounded marsh ponds. We also predicted that SMM would increase seasonal variability in water depth in impounded marsh ponds because of the potential incapacity of water control structures to cope with large flooding events. In addition, we predicted that SMM would decrease spatial variability in water depth. Finally, we predicted that ponds of impounded freshwater (IF), oligohaline (IO), and mesohaline (IM) marshes would be similar in water depth, temperature, dissolved oxygen ( O 2 ), and transparency. Using a priori multivariate analysis of variance (MANOVA) contrast, we tested these predictions by comparing hydrologic variables within ponds of impounded and unimpounded marshes during winters 1997-1998 to 1999-2000 on Rockefeller State Wildlife Refuge, near Grand Chenier, Louisiana. Specifically, we compared hydrologic variables (1) between IM and unimpounded mesohaline marsh ponds (UM); and (2) among IF, IO, and IM marshes ponds. As predicted, water depth was higher and salinity and O 2 were lower in IM than in UM marsh ponds. However, temperature and transparency did not differ between IM and UM marsh ponds. Water depth varied more among months in IM marsh ponds than within those of UM marshes, and variances among and within ponds were lower in IM than UM marshes. Finally, all hydrologic variables, except salinity, were similar among IF, IO, and IM marsh ponds. Hydrologic changes within marsh ponds due to SMM should (1) promote benthic invertebrate taxa that tolerate low levels of O 2 and salinity; (2) deter waterbird species that cannot cope with increased water levels; and (3) reduce waterbird species diversity by decreasing spatial variability in water depth among and within marsh ponds.

Louisiana

Role of a large marine protected area for conserving landscape attributes of sand habitats on Georges Bank (NW Atlantic)

Mobile fishing gear reduces seafloor habitat complexity through the removal of structure-building fauna, e.g. emergent organisms that create pits and burrows, as well as by smoothing of sedimentary bedforms (e.g. sand ripples). In this study, we compared the relative abundance of microhabitat features (the scale at which individual fish associate with seafloor habitat) inside and outside of a large fishery closed area (6917 km 2 ) on Georges Bank. Starting in late 1994, the closed area excluded all bottom tending fishing gear capable of capturing demersal fishes. A total of 32 stations were selected inside and outside of the closed area in sand habitats. Video and still photographic transects were conducted at each station using the Seabed Observation and Sampling System (SEABOSS). Seven common (i.e. featureless sand, rippled sand, sand with emergent fauna, bare gravelly sand, gravelly sand with attached-erect fauna, whole shell, shell fragment) and 2 rare (sponges, biogenic depressions) microhabitat types were compared separately. Results showed significant differences in the relative abundance of the shell fragment and sponge microhabitat types between fished and unfished areas. The lack of differences for the other microhabitats may indicate that the level of fishing activity in the area is matched by the system's ability to recover.

Georges Bank

Uptake pathway for Ag bioaccumulation in three benthic invertebrates exposed to contaminated sediments

We exposed 3 benthic invertebrates, the clam Macoma balthica , the polychaete Neanthes arenaceodentata and the amphipod Leptocheirus plumulosus , to Ag-contaminated sediments to evaluate the relative importance of various uptake routes (sediments, porewater or overlying water, and supplementary food) for Ag bioaccumulation. Silver bioaccumulation was evaluated at 4 levels of sediment Ag (0.1, 0,3, 1,2 and 3.3 µmol Ag g -1 ) and 2 levels of acid-volatile sulfide (AVS), <0.5 or ~40 µmol g -1 , and compared among food treatments with or without Ag contamination, or with different food rations. L. plumulosus were incubated for 35 d in the Ag-contaminated sediments after 3 mo of Ag-sediment equilibration, and M. balthica and N. arenaceodentata for 19 d after 5 mo equilibration. Ag bioaccumulation in the 3 organisms was significantly correlated with 1N HCl-extractable Ag concentrations (Ag-SEM: simultaneously extracted Ag with AVS) in sediments. The Ag concentrations in porewater and overlying water were greatest in the sediments with least AVS, consistent with previous studies. Nevertheless, the amphipod and clam exposed to oxic sediments (<0.5 µmol AVS g -1 ) accumulated amounts of Ag similar to those accumulated by organisms exposed to anoxic sediments (~40 µmol AVS g -1 ), when Ag-SEM levels were comparable. The dissolved Ag source was important for bioaccumulation in the polychaete N. arenaceodentata . Amphipods fed Ag-contaminated food contained ~1.8-fold more tissue Ag concentrations than those fed uncontaminated food. As suggested in kinetic (DYMBAM) modeling studies, ingestion of contaminated sediments and food were the principle routes of Ag bioaccumulation by the benthic invertebrates during chronic exposure, but the relative importance of each uptake route differed among species.

Marine Ecology Progress Series

Effects of marine reserves and urchin disease on southern Californian rocky reef communities

While the species level effects of marine reserves are widely recognized, community level shifts due to marine reserves have only recently been documented. Protection from fishing of top predators may lead to trophic cascades, which have community-wide implications. Disease may act in a similar manner, regulating population levels of dominant species within a community. Two decades of data from the Channel Islands National Park Service's Kelp Forest Monitoring database allowed us to compare the effects of fishing and urchin disease on rocky reef community patterns and dynamics. Different size-frequency distributions of urchins inside and outside of reserves indicated reduced predation on urchins at sites where fishing removes urchin predators. Rocky reefs inside reserves were more likely to support kelp forests than were fished areas. We suggest that this results from cascading effects of the fishery on urchin predators outside the reserves, which releases herbivores (urchins) from predation. After periods of prevalent urchin disease, the reef community shifted more towards kelp forest assemblages. Specific groups of algae and invertebrates were associated with kelp forest and barrens communities. The community dynamics leading to transitions between kelp forests and barrens are driven by both fishing and disease; however the fishery effect was of greater magnitude. This study further confirms the importance of marine reserves not only for fisheries conservation, but also for the conservation of historically dominant community types.

Marine Ecology Progress Series

Phytoplankton community ecology: Principles applied in San Francisco Bay

In his seminal 1961 paper 'The paradox of the plankton' Am Nat 95:137-147, G. E. Hutchinson asked why many species of phytoplankton can coexist while competing for a small number of limiting resources in an unstructured habitat. Hutchinson anticipated the resolution of his paradox, recognizing that communities are organized by processes beyond resource competition including species interactions, habitat variability and dispersal. Since 1961 we have made fundamental discoveries that have revolutionized our conceptual understanding of pelagic ecology, including (1) habitat heterogeneity at all scales relevant to plankton population dynamics, (2) community shifts in response to global climate cycles, (3) fast and selective predation as a powerful top-down force to shape phytoplankton communities, (4) turbulent mixing as a physical process that selects species on the basis of their size and form, (5) mixotrophy that allows some algal species to tap organic nutrient pools and function at multiple trophic levels, (6) taxon-specific life cycles including alternating vegetative and resting stages, and (7) the pelagic as an open system where communities are continually reshaped by species immigration. Here we synthesize these discoveries to show how they validate and amplify Hutchinson's hypothesis that phytoplankton communities are assembled by many processes. Our synthesis is built around observations of phytoplankton species composition from a decade of study in San Francisco Bay, used as a case study to illustrate the contemporary principles of phytoplankton community ecology. We apply these principles to address 2 central questions: (1) What processes assemble phytoplankton communities? (2) How does phytoplankton community composition influence ecosystem functions such as production in pelagic and benthic food webs?

California

Grazing by the intertidal gastropod Melampus coffeus greatly increases mangrove leaf litter degradation rates

Melampus coffeus , a pulmonate gastropod, forages for mangrove leaf litter at low tide and climbs tree trunks to avoid inundation during high tide. Unlike many grazers, these snails can assimilate mangrove leaf material. At Boca Ceiga Bay, Florida, densities of adult snails were high (>100 snails m –2 ) throughout a 130 m wide intertidal zone. A mark–recapture study indicated that over the course of 1 mo 48.6 ± 6.1% of snails returned to their initial tree during high tides, while those that did relocate moved 6.66 ± 0.60 m and were recorded from 33 new trees at high tide. A field experiment comparing tethered leaves with small and large litter bags showed that snail grazing greatly increased the rate of red ( Rhizophora mangle ) and black ( Avicennia germinans ) leaf litter breakdown. Grazing by M. coffeus resulted in 90% weight loss in <4 wk ( A. germinans ) and 7 wk ( R. mangle ), compared to slower breakdown in litter bags (12 to 26 wk, A. germinans ; >26 wk, R. mangle ). Another experiment showed greater leaf litter accumulation on the forest floor in plots where M. coffeus was excluded. The decomposition coefficient ( k ) was 10× greater when M. coffeus grazing was allowed than k for any studies that enclosed leaves in litterbags. During our experiment, M. coffeus consumed an estimated 40.5% of mangrove leaf fall, and 19.8% of leaf litter was exported as particulate or dissolved material. This M. coffeus population can produce an estimated 3 × 10 6 larvae m –2 yr –1 , which suggests that the larval pathway is an important conduit of mangrove leaf resources to the estuarine food web.

Florida