John W. Fleeger

Indirect effects of contaminants in aquatic ecosystems.

Contaminants such as petroleum hydrocarbons, heavy metals and pesticides can cause direct toxic effects when released into aquatic environments. Sensitive species may be impaired by sublethal effects or decimated by lethality, and this ecological alteration may initiate a trophic cascade or a release from competition that secondarily leads to responses in tolerant species. Contaminants may exert direct effects on keystone facilitator and foundation species, and contaminant-induced changes in nutrient and oxygen dynamics may alter ecosystem function. Thus, populations and communities in nature may be directly and/or indirectly affected by exposure to pollutants. While the direct effects of toxicants usually reduce organism abundance, indirect effects may lead to increased or decreased abundance. Here we review 150 papers that reference indirect toxicant effects in aquatic environments. Studies of accidental contaminant release, chronic contamination and experimental manipulations have identified indirect contaminant effects in pelagic and benthic communities caused by many types of pollutants. Contaminant-induced changes in behavior, competition and predation/grazing rate can alter species abundances or community composition, and enhance, mask or spuriously indicate direct contaminant effects. Trophic cascades were found in 60% of the manipulative studies and, most commonly, primary producers increased in abundance when grazers were selectively eliminated by contaminants. Competitive release may also be common, but is difficult to distinguish from trophic cascades because few experiments are designed to isolate the mechanism(s) causing indirect effects. Indirect contaminant effects may have profound implications in environments with strong trophic cascades such as the freshwater pelagic. In spite of their undesirable environmental influence, contaminants can be useful manipulative tools for the study of trophic and competitive interactions in natural communities.

Coastal eutrophication as a driver of salt marsh loss

Salt marshes are highly productive coastal wetlands that provide important ecosystem services such as storm protection for coastal cities, nutrient removal and carbon sequestration. Despite protective measures, however, worldwide losses of these ecosystems have accelerated in recent decades. Here we present data from a nine-year whole-ecosystem nutrient-enrichment experiment. Our study demonstrates that nutrient enrichment, a global problem for coastal ecosystems, can be a driver of salt marsh loss. We show that nutrient levels commonly associated with coastal eutrophication increased above-ground leaf biomass, decreased the dense, below-ground biomass of bank-stabilizing roots, and increased microbial decomposition of organic matter. Alterations in these key ecosystem properties reduced geomorphic stability, resulting in creek-bank collapse with significant areas of creek-bank marsh converted to unvegetated mud. This pattern of marsh loss parallels observations for anthropogenically nutrient-enriched marshes worldwide, with creek-edge and bay-edge marsh evolving into mudflats and wider creeks. Our work suggests that current nutrient loading rates to many coastal ecosystems have overwhelmed the capacity of marshes to remove nitrogen without deleterious effects. Projected increases in nitrogen flux to the coast, related to increased fertilizer use required to feed an expanding human population, may rapidly result in a coastal landscape with less marsh, which would reduce the capacity of coastal regions to provide important ecological and economic services.

Microhabitat use by marsh-edge fishes in a Louisiana estuary

SynopsisWe used a drop sampler to characterize use of the marsh-edge ecotone by small fishes along two transects running inland from the Gulf of Mexico for ca. 25 km in Louisianas Barataria-Caminada Bay System. Monthly sampling was stratified among upper, middle, and lower reaches and within reaches to characterize fish responses to salinity, depth, distance from shore, substrate, dissolved oxygen concentration, temperature, turbidity, velocity, and emergent stem density. In 681 quantitative samples, covering 658 m2, collected between October 1987 and October 1989, we collected 57 fish species and 16 864 individuals, primarily larvae and juveniles. The 15 most abundant fishes, comprising 97.7% of all individuals, were concentrated near the marsh edge (i.e., 0 to 1.25 m distance). Some significant differences within species for seasonal variables (e.g., temperature and dissolved oxygen concentration) reflected the ephemeral duration of early life history stages. Other differences reflected ontogenetic microhabitat shifts (e.g., depth and distance from shore). Within ecological groups, characterized as demersal residents, nektonic transients, and demersal transients, spatial and temporal segregation reflected the particular habitat requirements of each species. In a principal component analysis of microhabitat use, the first three components were interpreted as seasonal, depth-and-distance, and salinity axes, respectively. The array of species and size classes in principal component space reflected the complex dimensionality of microhabitat use. The high density of fish larvae and juveniles near the marsh edge confirmed the importance of the marsh-edge ecotone as a nursery for many estuarine-dependent fishes.

Oil Impacts on Coastal Wetlands: Implications for the Mississippi River Delta Ecosystem after the Deepwater Horizon Oil Spill

On 20 April 2010, the Deepwater Horizon explosion, which released a US government—estimated 4.9 million barrels of crude oil into the Gulf of Mexico, was responsible for the death of 11 oil workers and, possibly, for an environmental disaster unparalleled in US history. For 87 consecutive days, the Macondo well continuously released crude oil into the Gulf of Mexico. Many kilometers of shoreline in the northern Gulf of Mexico were affected, including the fragile and ecologically important wetlands of Louisianas Mississippi River Delta ecosystem. These wetlands are responsible for a third of the nations fish production and, ironically, help to protect an energy infrastructure that provides a third of the nations oil and gas supply. Here, we provide a basic overview of the chemistry and biology of oil spills in coastal wetlands and an assessment of the potential and realized effects on the ecological condition of the Mississippi River Delta and its associated flora and fauna.

Meiofaunal colonization of azoic estuarine sediment in Louisiana: Mechanisms of dispersal

Abstract Two mechanisms of muddy-bottom meiofaunal dispersal, waterborne suspended transport and holobenthic infaunal immigration, were compared as to their rate and effectiveness in mediating community reestablishment after small-scale defaunation. Colonizing meiofauna were quantitatively sampled in winter and summer from 16 replicates of two azoic sediment chamber designs on 2 and 29 days postplacement. The chambers were ≈ 3750 cm 3 ; one design allowed colonization via suspended movement through an open top, while the other design permitted entry only by infaunal crawling through subsurface open sides. After 48 h, mean harpacticoid copepod and naupliar densities in sediment chambers open to colonization exclusively by meiofauna in suspended transport were not significantly different from background sediment densities. Sediment chambers allowing colonization exclusively via infaunal immigration through the sediment, however, contained copepod and naupliar densities that were significantly less than densities in background sediments and suspension-colonized chambers. In contrast, nematode densities in both suspension- and infaunally colonized chambers were significantly less than in background sediments, but densities were not significantly different between the chamber treatments. Thus for a small-scale defaunation, copepods most rapidly and completely recolonize sediments via suspended transport. Nematode dispersal occurs equally well via suspended or infaunal movement; however nematodes never seemed to utilize the chambers fully because densities did not reach background levels even after 29 days.

Stable isotope indicators of movement and residency for brown shrimp (Farfantepenaeus aztecus) in coastal Louisiana marshscapes

Brown shrimp (Farfantepenaeus aztecus) are an important commercial aquatic species experiencing loss of inshore marsh nursery habitat in coastal Louisiana. To study inshore brown shrimp movements and identify aspects of essential habitat important for sustaining brown shrimp populations, we collected juvenile brown shrimp in April and May 2000, the time of annual maximum brown shrimp abundance, in a small 1-km2 marsh area on the central Louisiana coast. Drop sampling showed average shrimp densities of 1.6–2.4 m−2 in shallow marsh ponds and seining indicated lower densities of 0.5–0.9 m−2 in nearby shallow channel and open bay sites. Smaller shrimp (< 50 mm) fed disproportionately on benthic diatoms and small harpacticoid copepods, while large shrimp fed more frequently on larger-bodied amphipods and tanaids. We developed novel chemical approaches to estimate patterns of shrimp residency and movement using carbon and nitrogen stable isotopic determinations. Resident shrimp had isotopic values similar to average foods and showed consistent isotopic spacings between fast and slow turnover tissues. Residency was highest (47–55%) in ponds and shallow channel habitats and much less in open bays and deep channels (4–27%). There was sparse evidence for dietary specialization among individull shrimp. The results support the view that small 10–20 mm postlarval and juvenile brown shrimp arriving in estuaries from offshore waters continue movement through sub-optimal habitats (deep channels and open bays), but exhibit much less movement once an optimal habitat (marsh ponds or shallow channel margins) is reached. This study also indicated that combining estimates of shrimp densities, residency, growth rate, and mortality allows evaluation of the importance of different habitat types for shrimp production. Shallow ponds that in many ways resemble fertile aquaculture ponds appear to be hot spots for brown shrimp production, and coastal preservation and restoration efforts should focus on these areas as important for sustaining shrimp fisheries.

SUSCEPTIBILITY OF SALT MARSHES TO NUTRIENT ENRICHMENT AND PREDATOR REMOVAL

Salt marsh ecosystems have been considered not susceptible to nitrogen overloading because early studies suggested that salt marshes adsorbed excess nutrients in plant growth. However, the possible effect of nutrient loading on species composition, and the combined effects of nutrients and altered species composition on structure and function, was largely ignored. Failure to understand interactions between nutrient loading and species composition may lead to severe underestimates of the impacts of stresses. We altered whole salt marsh ecosystems (;60 000 m 2 /treatment) by addition of nutrients in flooding waters and by reduction of a key predatory fish, the mummichog. We added nutrients (N and P; 15-fold increase over ambient conditions) directly to the flooding tide to mimic the way anthropogenic nutrients are delivered to marsh ecosystems. Despite the high concentrations (70 mmol N/L) achieved in the water column, our annual N loadings (15-60 g Nm � 2 � yr � 1 ) were an order of magnitude less than most plot-level fertilization experiments, yet we detected responses at several trophic levels. Preliminary calculations suggest that 30-40% of the added N was removed by the marsh during each tidal cycle. Creek bank Spartina alterniflora and high marsh S. patens production increased, but not stunted high marsh S. alterniflora. Microbial production increased in the fertilized creek bank S. alterniflora habitat where benthic microalgae also increased. We found top-down control of benthic microalgae by killifish, but only under nutrient addition and in the opposite direction (increase) than that predicted by a fish-invertebrate-microalgae trophic cascade. Surprisingly, infauna declined in abundance during the first season of fertilization and with fish removal. Our results demonstrate ecological effects of both nutrient addition and mummichog reduction at the whole-system level, including evidence for synergistic interactions.

Experimental investigation of the effects of polynuclear aromatic hydrocarbons on an estuarine sediment food web

Abstract The influence of polynuclear aromatic hydrocarbons (PAH) on a benthic estuarine sedimentary salt-marsh food web was examined using a microcosm system to simulate natural conditions. Microcosms were dosed with sublethal concentrations of PAH-contaminated sediment collected from a produced-water site at Pass Fourchon, Louisiana, USA (final PAH concentrations ranged from 0.3 to 3 mg PAH/kg dry sediment). Bacterial activity, physiological condition and abundance were not influenced by PAH, but microalgal activity and physiological condition were. Grazing by meiobenthic copepods on benthic microalgae was not significantly influenced by PAH concentration, nor was the physiological condition of copepods, as determined by their lipid-storage material. Meiofaunal community composition was influenced by PAH, as nematodes became disproportionately abundant, and the nauplius copepod ratio increased in High-PAH treatments. Overall, however, sublethal effects of PAH were not pronounced at the concentrations examined. Considering that coastal Louisiana has been exposed to chronic contamination by petroleum hydrocarbons for decades, it is suggested that the sedimentary microbial/meiofaunal community may have adapted to elevated PAH concentrations.

Food, density, and microhabitat: factors affecting growth and recruitment potential of juvenile saltmarsh fishes

The relationships among microhabitat use, food habits, conspecific density and recent growth rate for estuarine-dependent juvenile spotted seatrout, Cynoscion nebulosus, and red drum, Sciaenops ocellatus, were studied to determine how nursery habitat influences early growth. Juvenile spotted seatrout and red drum were quantified along the marsh-edge ecotone from multiple drop samples, and their immediate environments characterized by a suite of physical and chemical variables along with substrate type and Spartina stem density. Recent daily growth of individual fish was modeled in a series of multiple regression analyses that considered the relative contributions of food, microhabitat, and conspecific density. The spotted seatrout model (p < 0.0001) included four independent variables, otolith radius, prey diversity, salinity, and a salinity-DO interaction term, and explained 67.9% of the variation in daily growth. All variables were significant (p < 0.05), and regression slopes were positive for all variables except salinity. The red drum model (p > 0.0001) included five independent variables, otolith radius, temperature, salinity, depth and substrate, and explained 63.3% of the variation in daily growth. All variables were significant (p > 0.05), and all regression slopes were positive. Fish size (as estimated by otolith radius) accounted for most of the variance in the spotted seatrout (60.2%) and red drum (44.8%) models, while the remaining environmental variables were significant and responsible for 7.7 and 18.5%, respectively. Density variables were not selected for either model, suggesting that density-dependence was not an important influence on recent daily growth. Generally, physico-chemical variables such as temperature, salinity, and dissolved oxygen contributed more to growth than diet or extrinsic factors such as grass stem density.

Sustained mass culture of Amphiascoides atopus a marine harpacticoid copepod in a recirculating system

Abstract A sustainable mass-culture system (4 m2 basal surface area) of the meiobenthic harpacticoid copepod Amphiascoides atopus, capable of producing in excess of one million individuals and over 5 g dry weight biomass per day, was successfully designed and implemented. The system consists of culture tanks containing a shallow layer of limestone cobble facilitating naupliar and juvenile copepodite development. Adults and older copepodites swim into the overlying water which is recirculated and filtered, expediting their clean capture. A. atopus ranges in length from 0.19 mm (for Nauplius I) to 0.92 mm in the adult, and up to 5 μg dry weight. Copepods were fed either cultured algae (Chaetocerous muelleri, strain Chaet 10) or commercial fish-flake food, and grew well on each diet or a mixture. Amphiascoides atopus life history patterns are favorable for mass culture. Clutch size averages 24 eggs per ovigerous female, and new egg clutches are produced every 3–4 days. When cultured at 23 °C, life expectancy is about 3 mo, and generation time is less than one month. Preliminary laboratory feeding experiments with various shrimp and fishes and suggest that A. atopus may prove to be an excellent food source for larvae, post-larvae and juveniles and may serve as an alternative live prey to brine shrimp in mariculture applications.Key words: harpacticoid copepod culture, meiobenthos, fish and shellfish mariculture.