Abstract: The South American floating aquatic plant water hyacinth (Eichhornia crassipes) has a history of worldwide invasions, including a 1904 introduction into the Sacramento-San Joaquin Delta, California. The native pennywort (Hydrocotyle umbellata) occupies similar habitats in the Delta and is extensively used by resident invertebrates and fish. We sought to discover if an invader would be functionally equivalent to the native plant, by asking whether the encroadhing hyacinth modified the invertebrate assemblage structure and fish-invertebrate food web relative to pennywort. We sampled epiphytic, epibenthic, and benthic invertebrates, and plant canopy insects in patches of hyacinth and pennywort, and analyzed fish diets at three sites in the Delta during 1998. We also measured habitat structure (leaf density, root biomass, and surface area). In 1999, following control and absence of hyacinth, we again measured epiphytic invertebrates in pennywort. We found differences between hyacinth and pennywort in structure, associated invertebrates, and fish diets. Most measurements inferred functional non-equivalency between hyacinth and pennywort, although some functional equivalency and natural variation existed. Leaf and insect densities were significantly higher in pennywort and there were also significant differences in insect assemblage compositions. Hyacinth roots in the water column had significantly more surface area. Densities of epibenthic and benthic aquatic invertebrates were typically greater in pennywort and taxonomic compositions of aquatic invertebrate assemblages showed significant differences. Amphipods and isopods living epiphytically in the root masses were particularly abundant, including several newly discovered introduced species: the amphipodCrangonyx floridanus and the isopodsCaecidotea racovitai andAsellus hilgendorfii. The native amphipodHyalella azteca was more abundant in pennywort and heavily preyed upon by fish, while the non-indigenousC. floridanus was more abundant in hyacinth and not prevalent in fish diets. The introduction of hyacinth to the Delta has caused significant ecological alterations in the surrounding community, due to hyacinth being functionally different from native patches of pennywort.

Abstract: 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.

Abstract: We evaluate if the distribution and abundance ofThalassia testudinum, Syringodium filiforme, andHalodule wrightii within Biscayne Bay, Florida, are influenced by salinity regimes using, a combination of field surveys, salinity exposure experiments, and a seagrass simulation model. Surveys conducted in June 2001 revealed that whileT. testudinum is found throughout Biscayne Bay (84% of sites surveyed),S. filiforme andH wrightii have distributions limited mainly to the Key Biscayne area.H. wrightii can also be found in areas influenced by canal discharge. The exposure of seagrasses to short-term salinity pulses (14 d, 5–45‰) within microcosms showed species-specific susceptibility to the salinity treatments. Maximum growth rates forT testudinum were observed near oceanic salinity values (30–40‰) and lowest growth rates at extreme values (5‰ and 45‰).S. filiforme was the most susceptible seagrass species; maximum growth rates for this species were observed at 25‰ and dropped dramatically at higher and lower salinity.H. wrightii was the most tolerant, growing well at all salinity levels. Establishing the relationship between seagrass abundance and distribution and salinity is especially relevant in South Florida where freshwater deliveries into coastal bays are influenced by water management practices. The seagrass model developed by Fong and Harwell (1994) and modified here to include a shortterm salinity response function suggests that freshwater inputs and associated decreases in salinity in nearshore areas influence the distribution and growth of single species as well as modify competitive interactions so that species replacements may occur. Our simulations indicate that although growth rates ofT. testudinum decrease when salinity is lowered, this species can still be a dominant component of nearshore communities as confirmed by our surveys. Only when mean salinity values are drastically lowered in a hypothetical restoration scenario isH. wrightii able to outcompeteT. testudinum.

Abstract: In order to classify the trophic state of detritus sink systems, instead of the conventional indicators based on inorganic nutrient availability and algal biomass and productivity in the water column, we used new biochemical descriptors based on the amount of sedimentary organic carbon (C) and nitrogen (N) potentially available to heterotrophs. We investigated spatial and temporal changes in microphytobenthic biomass, organic matter biochemical composition, and enzymatically hydrolyzable protein and carbohydrate pools along a north-south transect in the Marsala lagoon (Mediterranean Sea, Italy) at three stations characterized by different hydrodynamic conditions and organic matter content in the sediment. In the Marsala lagoon water currents decreased from north to south and this pattern was reflected by organic matter distribution and composition. Sediment organic matter concentrations were among the highest reported in the literature and, in the central area where large meadows of the seagrassPosidonia oceanica were present, display a strong dominance of highly refractory carbohydrates. The protein to carbohydrate ratio was always < 1, indicating the dominance of aged organic detritus. Microphytobenthic biomass displayed an increasing pattern southward, and its contribution to the biopolymeric C pools ranged from negligible in the central sector of the lagoon to 50% in its northern part, indicating that sources of sediment organic C also changed along the hydrodynamic gradient. The percentage contribution of the enzymatically hydrolyzable fraction of proteins and carbohydrates was inversely related to total protein and total carbohydrate concentrations, respectively, suggesting that bioavailability of organic C and N increased with decreasing organic matter content in the sediment and with increasing hydrodynamic regime. Microphytobenthic contribution to biopolymeric C (as a proxy of autotrophic organic C) and the ratio of the enzymatically digestible fraction to biopolymeric C (as an indicator of organic matter liability) were significantly correlated, suggesting that chlorophylla sediment content might be used as an indicator of food promptly available to consumers. The present study also highlighted that the ratio of labile (i.e., enzymatically digestible) versus biopolymeric organic C in the sediments tends to decrease with increasing organic matter content, due to the increase of the refractory fraction of organic C.

Abstract: The extensive spread ofPhragmites australis throughout brackish marshes on the East Coast of the United States is a major factor governing management and restoration decisions because it is assumed that biogeochemical functions are altered by the invasion. Microbial activity is important in providing wetland biogeochemical functions such as carbon and nitrogen cycling, but there is little known about sediment microbial communities inPhragmites marshes. Microbial populations associated with invasivePhragmites vegetation and with native salt marsh cordgrass,Spartina alterniflora, may differ in the relative abundance of microbial taxa (community structure) and in the ability of this biota to decompose organic substrates (community biogeochemical function). This study compares sediment microbial communities associated withPhragmites andSpartina vegetation in an undisturbed brackish marsh near Tuckerton, New Jersey (MUL), and in a brackish marsh in the anthropogenically affected Hackensack meadowlands (SMC). We use phospholipid fatty acid (PLFA) analysis and enzymataic activity to profile sediment microbial communities associated with both plants in each site. Sediment analyses include bulk density, total organic matter, and root biomass. PLFA profiles indicate that the microbial communities differ between sites with the undisturbed site exhibiting greater fatty acid richness (62 PLFA recovered from MUL versus 38 from SMC). Activity of the 5 enzymes analyzed (β-glucosidase, acid phosphatase, chitobiase, and 2 oxidases) was higher in the undisturbed site. Differences between vegetation species as measured by Principal Components Analysis were significantly greater at the undisturbed MUL site than at SMC, and patterns of enzyme activity and PLFAs did not correspond to patterns of root biomass. We suggest that in natural wetland sediments, macrophyte rhizosphere effects influence the community composition of sediment microbial populations. Physical and chemical site disturbances may impose limits on these rhizosphere effects, decreasing sediment microbial diversity and potentially, microbial biogeochemical functions.

Abstract: While it has long been known that Pacific salmon use estuarine habitat, it has proven much harder to establish that the loss of estuarine habitat results in reduced survival. We used coded-wire tagging of hatchery fish to estimate the survival from release until maturity and related this survival to several indicators of estuarine condition. We found a significant relationship between the survival of chinook salmon (Oncorhynchus tshawytscha) and the percentage of the estuary that is in pristine condition, but no significant relationship for coho salmon (Oncorhynchus kisutch). This supports field observations that chinook salmon use estuarine habitat much more than coho salmon and confirms that the loss of estuarine habitat results in lower survival of chinook salmon.

Abstract: In recent decades, marshes naturally dominated bySpartina spp. have been replaced byPhragmites australis throughout the northeastern United States. We suggest that early in this invasion there was little effect on the fish fauna. As the invasion proceeds, the marsh surface habitat became more altered (i.e., elevated, flattened, reduced water-filled depressions, and reduced standing water), which resulted in a reduction of feeding, reproduction, and nursery function for fishes, especiallyFundulus spp. These potential changes in marsh habitat and function have resulted in numerous attempts to removePhragmites and restoreSpartina spp. To evaluate the response of marsh surface fishes toPhragmites treatment, we examined fish use in the brackish water reaches of Alloway Creek in the Delaware Bay estuary. ReferencePhragmites habitats were compared with referenceSpartina alterniflora-dominated habitats and sites treated (1996–1998) to removePhragmites to restore former vegetation (i.e., restored, now comprised of 100%Spartina). Fish were sampled with an array (n=9 at each site) of shallow pit traps (rectangular glass dishes, 27.5×17.5×3.7 cm). Small individuals (mean=17.5, 5–45 mm TL) dominated all pit trap collections. Fish abundance was highest at the restored (catch per unit effort [CPUE]=2.16) andSpartina (CPUE=0.81) sites with significantly lower values atPhragmites (CPUE=0.05) habitats. Samples were dominated by young-of-the-year mummichog,Fundulus heteroclitus (98% of total fish, n=631). The only other fish species collected was spotfin killifish,Fundulus luciae (2% of total catch, n=14), which was only present in restored andSpartina habitats. These observations suggest that the restored marsh is providing habitat (water-filled depressions on the marsh surface) for young-of-the-yearFundulus spp. These marshes are responding favorably to the restoration based on the much greater abundance of fish in restored versusPhragmites habitats and the overall similarity between restored andSpartina habitats.

Abstract: Systematic morphological changes of the coastline of the outer Yangtze River mouth in response to storms versus calm weather were documented by daily surveys of tidal marshes and flats between April 1999 and May 2001 and by boat surveys offshore during this and earlier periods. The largest single event during 1999 to 2001 was Typhoon Paibaian, which eroded the unvegetated tidal flat and lower marsh and led to accretion on the middle-to-upper marsh and in the subtidal channel. The greatest erosion of 21 cm occurred at the border between the marsh and the unvegetated flat due to the landward retreat of the marsh edge during the storm. Strong waves on the flats increased suspended sediment concentration by 10-20 times. On the upper marsh, where the frequency of submergence by astronomical tides is only 3%, Typhoon Paibian led to 4 cm of accretion, accounting for 57% of the net accretion observed over the 2-yr study. Typhoon Paibian and other large storms in the 1990s caused over 50 cm of accretion along the deep axis of the river mouth outlet channel. During calm weather, when hydrodynamic energy was dominated by tides, deposition was centered on the unvegetated flats and lower marsh with little deposition on the high marsh and erosion in the subtidal channel. Depositional recovery of the tidal flat from typhoon-induced erosion took only several days, whereas recovery of the subtidal channel by erosion took several weeks. A conceptual model for the morphological responses of tidal marshes, flats, and subtidal channels to storms and calm weather is proposed such that sediment continually moves from regions of highest near-bed energy towards areas of lower energy.

Abstract: Increased nutrient loadings have resulted in low dissolved oxygen (DO) concentrations in bottom waters of the Patuxent River, a tributary of Chesapeake Bay. We synthesize existing and newly collected data to examine spatial and temporal variation in bottom DO, the prevalence of hypoxia-induced mortality of fishes, the tolerance of Patuxent River biota to low DO, and the influence of bottom DO on the vertical distributions and spatial overlap of larval fish and fish eggs with their gelatinous predators and zooplankton prey. We use this information, as well as output from watershed-quality and water-quality models, to configure a spatially-explicit individual-based model to predict how chang- ing land use within the Patuxent watershed may affect survival of early life stages of summer breeding fishes through its effect on DO. Bottom waters in much of the mesohaline Patuxent River are below 50% DO saturation during summer. The system is characterized by high spatial and temporal variation in DO concentrations, and the current severity and extent of hypoxia are sufficient to alter distributions of organisms and trophic interactions in the river. Gelatinous zooplankton are among the most tolerant species of hypoxia, while several of the ecologically and economically important finfish are among the most sensitive. This variation in DO tolerances may make the Patuxent River, and similar estuaries, particularly susceptible to hypoxia-induced alterations in food web dynamics. Model simulations consistently predict high mortality of planktonic bay anchovy eggs (Anchoa mitchilli) under current DO, and increasing survival of fish eggs with increasing DO. Changes in land use that reduce nutrient loadings may either increase or decrease predation mortality of larval fish depending on the baseline DO conditions at any point in space and time. A precautionary approach towards fisheries and ecosystem management would recommend reducing nutrients to levels at which low oxygen effects on estuarine habitat are reduced and, where possible, eliminated.

Abstract: Over the past century, the distribution and abundance ofPhragmites australis (common reed) has dramatically increased in both freshwater and brackish wetlands throughout North America It has been hypothesized that the increased competitive ability ofPhragmites could be the result of an introduction of a more aggressive genotype Sequence data from 2 noncoding regions of the chloroplast genome show that, historically, 11 native haplotypes were found across North America and population-structuring distinguishing samples from the Atlantic Coast, Midwest, West, and Gulf Coast regions of the continent was evident Today a single genetically-distinct haplotype dominates the Atlantic Coast and is also found across the continent in lower frequencies; this type is common in Europe and Asia and has most likely been introduced to North America Comparisons of modern populations with historic samples show that along the Atlantic Coast, this cosmopolitan type has replaced native haplotypes and it is invading new sites throughout the rest of the country In the Midwest and West, native populations are still common but introduced populations are found along roadsides throughout the area Gulf Coast populations are dominated by another population type that is genetically distinct from all other North American population types

Abstract: The purpose of this study was to quantify the nitrogen (N) inputs to 34 estuaries on the Atlantic and Gulf Coasts of the United States. Total nitrogen (TN) inputs ranged from 1 kg N ha−1 yr−1 for Upper Laguna Madre, Texas, to 49 kg N ha−1 yr−1 for Massachusetts Bay, Massachusetts. TN inputs to 11 of the 34 estuaries were dominated by urban N sources (point sources and septic systems) and nonpoint source N runoff (5% of total); point sources accounted for 36–86% of the TN inputs to these 11 urban-dominated estuaries. TN inputs to 20 of the 34 estuaries were dominated by agricultural N sources; N fertilization was the dominant source (46% of the total), followed by manure (32% of the total) and N fixation by crops (16% of the total). Atmospheric deposition (runoff from watershed plus direct deposition to the surface of the estuary) was the dominant N source for three estuaries (Barnegat Bay, New Jersey: 64%; St. Catherines-Sapelo, Georgia: 72%; and Barataria Bay, Louisiana: 53%). Six estuaries had atmospheric contributions ≥30% of the TN inputs (Casco Bay, Maine: 43%; Buzzards Bay, Massachusetts: 30%; Great Bay, New Jersey: 40%; Chesapeake Bay: 30%; Terrebonne-Timbalier Bay, Louisiana: 59%; and Upper Laguna Madre: 41%). Results from our study suggest that reductions in N loadings to estuaries should be accomplished by implementing watershed specific programs that target the dominant N sources.

Abstract: The mummichog,Fundulus heteroclitus, is one of the most abundant macrofaunal components of salt marsh ecosystems along the east coast of the United States. During April–November 1998, we determined the habitat use and movement patterns of young-of-the-year (YOY) and adult mummichogs in a restored marsh, formerly a salt hay farm, and an adjacent creek in order to expand our understanding of the ecology of the species and evaluate the success of the restoration. Four major fish habitat types (large first-order natural creek, second-order created creek, linear drainage ditch, and marsh surface) were identified within the study site. Patterns of relative abundance and mark and recapture using coded wire tags were used to determine the habitat use, tidal movements, home range, and site fidelity of the species within these habitat types. A total of 14,784 fish, ranging from 20–100 mm SL, were captured with wire mesh traps and tagged, and 1,521 (10.3%) fish were recaptured. A variety of gears were used to attempt to recapture fish across all habitat types, including wire mesh traps, push nets, and otter trawls. Based on abundance and recaptures of tagged fish, the YOY and adults primarily used the shallow subtidal and intertidal areas of the created creek, the intertidal drainage ditches, and the marsh surface of the restored marsh but not the larger, first-order natural creek. At low tide, large numbers were found in the subtidal areas of the created creek; these then moved onto the marsh surface on the flooding tide. Elevation, and thus hydroperiod, appeared to influence the microscale use of the marsh surface. We estimated the home range of adults and large YOY (20–100 mm SL) to be 15 ha at high tide, which was much larger than previously quantified. There was strong site fidelity to the created creek at low tide. The habitat use and movement patterns of the mummichog appeared similar to that reported for natural marshes. Coupled with the results of other studies on the feeding, growth, and production of this species in this restoreh, the species appeared to have responded well to the restoration.

Abstract: Several recent studies indicate that the replacement of extant species withPhragmites australis can alter the size of nitrogen (N) pools and fluxes within tidal marshes. Some common effects ofP. australis expansion are increased standing stocks of N, greater differentiation of N concentrations between plant tissues (high N leaves and low N stems), and slower whole-plant decay rates than competing species (e.g.,Spartina, Typha spp.). Some of the greater differences between marsh types involveP. australis effects on extractable and porewater pools of dissolved inorganic nitrogen (DIN) and N mineralization rates. Brackish and salt marshes show higher concentrations of DIN in porewater beneathSpartina spp. relative toP. australis, but this is not observed in freshwater tidal marshes whenP. australis is compared withTypha spp. or mixed plant assemblages. With few studies of concurrent N fluxes, the net effect ofP. australis on marsh N budgets is difficult to quantify for single sites and even more so between sites. The magnitude and direction of impacts ofP. australis on N cycles appears to be system-specific, driven more by the system and species being invaded than byP. australis itself. WhereP. australis is found to affect N pools and fluxes, we suggest these alterations result from increased biomass (both aboveground and belowground) and increased allocation of that biomass to recalcitrant stems. Because N pools are commonly greater inP. australis than in most other communities (due to plant and litter uptake), one of the most critical questions remaining is “From where is the extra N inP. australis communities coming?” It is important to determine if the source of the new N is imported (e.g., anthropogenic) or internallyproduced (e.g., fixed, remineralized organic matter). In order to estimate net impacts ofP. australis on marsh N budgets, we suggest that further research be focused on the N source that supports high standing stocks of N inP. australis biomass (external input versus internal cycling) and the relative rates of N loss from different marshes (burial versus subsurface flow versus denitrification).

Abstract: The invasion ofSpartina marshes by the common reed,Phragmites australis, along the east coast of the United States over the last several decades has been well documented, although we know little about the impact of this invasion on the fish fauna and the few published papers seem contradictory. During 1999–2000 (May–September) we evaluated the fish response to vegetation type (Phragmites australis veersusSpartina alterniflora) by monitoring several aspects of fish early life history (egg deposition, embryonic development, hatching success, and larval and juvenile abundance) in low salinity marshes in the Mullica River in southern New Jersey. The dominant fish species using the marsh surface,Fundulus heteroclitus (93% of total catch, n=996 individuals), reproduced in both vegetation types with eggs deposited in leaf axils near the base of the plant inSpartina and in broken stems ofPhragmites during both years. These eggs also undergo successful embryonic development to hatching in both vegetation types. Larval and juvenile (5–75 mm total length, but 95% < 34 mm TL) abundance of this species is much reduced onPhragmites-dominated (mean CUPE=0.02, n=7 ind) marsh surface relative toSpartina (mean CPUE=2.31). These findings, and similar results for fish abundance in 1997 and 1998, indicate that theSpartima marsh surface is likely essential fish habitat for this species because it provides habitat for larvae and small juveniles, whilePhragmites does not. ThePhragmites invasion in brackish marshes may be having deleterious effects on fish populations and possibly on predators that prey uponF. heteroclitus, and as a result, marsh secondary production.

Abstract: Ecological and paleoecological studies from the Patuxent River mouth reveal dynamic variations in benthic ostracode assemblages over the past 600 years due to climatic and anthropogenic factors. Prior to the late 20th century, centennial-scale changes in species dominance were influenced by climatic and hydrological factors that primarily affected salinity and at times led to oxygen depletion. Decadal-scale droughts also occurred resulting in higher salinities and migration of ostracode species from the deep chanel (Loxoconcha sp.,Cytheromorpha newportensis) into shallower water along the flanks of the bay. During the 19th century the abundance ofLeptocythere nikraveshae andPerissocytheridea brachyforma suggest increased turbidity and decreased salinity. Unprecedented changes in benthic ostracodes at the Patuxent mouth and in the deep channel of the bay occurred after the 1960s whenCythermorpha curta became the dominant species, reflecting seasonal anoxia. The change in benthic assemblages coicided with the appearance of deformities in foraminifers. A combination of increased nitrate loading due to greater fertilize use and increased fresh-water flow explains this shift. A review of the geochemical and paleoecological evidence for dissolved oxygen indicates that seasonal oxygen depletion in the main channel of Chesapeake Bay varies over centennial and decadal timescales. Prior to 1700 AD, a relatively wet climate and high freshwater runoff led to oxygen depletion but rarely anoxia. Between 1700 and 1700, progressive eutrophication occurred related to land clearance and increased sedimentation, but this was superimposed on the oscillatory pattern of oxygen depleton most likely driven by climatological and hydrological factors. It also seems probable that the four-to five-fold increase in sedimentation due to agricultural and timber activity could have contributed to an increased natural nutrient load, likely fueling the early periods (1700–1900) of hypoxia prior to widespread fertilizer use. Twentieth-century anoxia worsened in the late 1930s–1940s and again around 1970, reaching unprecedented levels in the past few decades. Decadal and interannual variability in oxygen depletion even in the 20th century is still strongly influenced by climatic processes influencing precipitation and freshwater runoff.

Abstract: The invasion ofPhragmites australis into tidal marshes formerly dominated bySpartina alterniflora has resulted in considerable interest in the consequences of this invasion for the ecological functions of marsh habitat. We examined the provision of trophic support for a resident marsh fish,Fundulus heteroclitus, in marshes dominated byP. australis, byS. alterniflora, and in restored marshes, using multiple stable isotope analysis. We first evaluated our ability to distinguish among potential primary producers using the multiple stable isotope approach. Within a tidal creek system we found significant marsh and elevation effects on microalgal isotope values, and sufficient variability and overlap in primary producer isotope values to create some difficulty in identifying unique end members. The food webs supportingF. heteroclitus production were examined using dual isotope plots. At both sites, the δ13C values ofF. heteroclitus were clustered over values for benthic microalgae (BMI) and approximately midway between δ13C values ofSpartina andPhragmites. Based on comparisons of fish and primary producer δ13C, δ15N, and δ34S values, and consideration ofF. heteroclitus feeding habits, we conclude that BMI were a significant component of the food web supportingF. heteroclitus in these brackish marshes, especially recently-hatched fish occupying pools on the marsh surface. A 2‰ difference in δ13C betweenFundulus occupying nearly adjacentSpartina andPhragmites marshes may be indicative of relatively less reliance on BMI and greater reliance onPhragmites production inPhragmites-dominated marshes, a conclusion consistent with the reduced BMI biomass found inPhragmites marshes. The mean δ13C value ofF. heteroclitus from restored marshes was intermediate between values of fish from naturally occurringSpartina marshes and areas invaded byPhragmites. We also examined the isotopic evidence for ontogenetic changes in the trophic position of larval and juvenileF. heteroclitus. We found significant positive relationships betweenF. heteroclitus δ15N values and total length, reflective of an increase in trophic position as fish grow.F. heteroclitus δ15N values indicate that these fish are feeding approximately two trophic levels above primary producers.

Abstract: We quantified annual nutrient inputs to the Patuxent River estuary from point and nonpoint sources and from direct atmospheric deposition. We also compared nonpoint source (NPS) discharges from Piedmont and Coastal Plain regions and from agricultural and developed lands. Using continuous automated-sampling, we measured discharges of water, nitrogen, phosphorus, organic carbon (C), and suspended solids from a total of 23 watersheds selected to represent various proportions of developed land and cropland in the Patuxent River basin and the neighboring Rhode River basin. The sampling period spanned two years that differed in annual precipitation by a factor of 1.7. Water discharge from the watershed to the Patuxent River estuary was 3.4 times higher in the wet year than in the dry year. Annual water discharges from the study watersheds increased as the proportion of developed land increased. As the proportion of cropland increased, there were increases in the annual flow-weighted mean concentrations of nitrate (NO3 −), total nitrogen (TN), dissolved silicate (Si), total phosphate (TPO4 3−), total organic phosphorus (TOP), total P (TP), and total suspended solids (TSS) in NPS discharges. The effect of cropland on the concentrations of NO3 − and TN was stronger for Piedmont watersheds than for Coastal Plain watersheds. As the proportion of developed land increased, there were increases in annual mean concentrations of NO3 −, total ammonium (TNH4 +), total organic N (TON), TN, total organic C (TOC), TPO4 3−, TOP, TP, and TSS and decreases in concentrations of Si. Annual mean concentrations of TON, TOC, forms of P, and TSS were highest in the wet year. Annual mean concentrations of NO3 −, TNH4 +, TN, and Si did not differ significantly between years. We directly measured NPS discharges from about half of the Patuxent River basin and estimated discharges from the other half of the basin using statistical models that related annual water flow and material concentrations to land cover and physiographic province. We compared NPS discharges to public data on point source (PS) discharges. We estimated direct atmospheric deposition of forms of N, P, and organic C to the Patuxent River estuary based on analysis of bulk deposition near the Rhode River. During the wet year, most of the total terrestrial and atmospheric inputs of forms of N and P came from NPS discharges. During the dry year, 53% of the TNH4 + input was from atmospheric deposition and 58% of the NO3 − input was from PS discharges; NPS and PS discharges were about equally important in the total inputs of TN and TPO4 3−. During the entire 2-yr period, the Coastal Plain portion of the Patuxent basin delivered about 80% of the NPS water discharges to the estuary and delivered similar proportions of the NPS TNH4 +, TN, TOP, and TSS. The Coastal Plain delivered greater proportions of the NPS TON, TOC, Si, and TP (89%, 90%, 93%, and 95%, respectively) than of water, and supplied nearly all of the NPS TPO4 3− (99%). The Piedmont delivered 33% of the NPS NO3 − while delivering only 20% of the NPS water to the stuary. We used statistical models to infer the percentages of NPS discharges supplied by croplands, developed lands, and other lands. Although cropland covers only 10% of the Patuxent River basin, it was the most important source of most materials in NPS discharge, supplying about 84% of the total NPS discharge of NO3 −; about three quarters of the TPO4 3−, TOP, TP, and TSS; and about half of the TNH4 + and TN. Compared to developed land, cropland supplied a significantly higher percentage of the NPS discharges of NO3 −, TN, TPO4 3−, TOP, TP, and TSS, despite the fact development land covered 12% of the basin.

Abstract: We developed an empirical model integrating nonpoint source (NPS) runoff, point sources (PS), and reservoir management to predict watershed discharges of water, sediment, organic carbon, silicate, nitrogen, and phosphorus to the Patuxent River in Maryland. We estimated NPS discharges with linear models fit to measurements of weekly flow and 10 material concentrations from 22 study watersheds. The independent variables were the proportions of cropland and developed land, physiographic province (Coastal Plain or Piedmont), and time (week). All but one of the NPS models explained between 62% and 83% of the variability among concentration or flow measurements. Geographic factors (land cover and physiographic province) accounted for the explained variability in largely dissolved material concentrations (nitrate [NO3], silicate [Si], and total nitrogen [TN]), but the explained variability in flow and particulates (sediment and forms of phosphorus) was more strongly related to temporal variability or its interactions with land cover and province. Average concentrations of all materials increased with cropland proportion and also with developed land (except Si), but changes in cropland produced larger concentration shifts than equivalent changes in developed land proportion. Among land cover transitions, conversions between cropland and forest-grassland cause the greatest changes in material discharges, cropland and developed land conversions are intermediate, and developed land and forest-grassland conversions have the weakest effects. Changing land cover has stronger effects on NO3 and TN in the Piedmont than in the coastal Plain, but for all other materials, the effects of land-use change are greater in the Coastal Plain. We predicted the changes in nutrient load to the estuary under several alternate land cover configurations, including a state planning scenario that extrapolates current patterns of population growth and land development to the year 2020. In that scenario, declines in NPS discharges from reducing cropland are balanced by NPS discharge increases from developing an area almost six times larger than the lost cropland. When PS discharges are included, there are net increases in total water, total phosphorus, and TN discharges.

Abstract: The flooding-drying process over the intertidal zone of the Satilla River estuary of Georgia was examined using a three-dimensional (3-D) primitive equations numerical model with Mellor and Yamada's (1982) level 2.5 turbulent closure scheme. The model was forced by the semi-diurnal M2, S2, and N2 tides and freshwater discharge at the upstream end of the estuary. The intertidal salt marsh was treated using a 3-D wet-dry point treatment technique that was developed for the σ-coordinate transformation estuary model. Good agreement was found between model-data comparison at anchor monitoring sites and also along the estuary that suggested that the model provided a reasonable simulation of the temporal and spatial distribution of the 3-D tidal current and salinity in the Satilla River estuary. Numerical experiments have shown that the flooding-drying process plays a key role in the simulation of tidal currents in the main river channel and in water transport over the estuarine-salt marsh complex. Ignoring this process could lead to a 50% under-estimation of the amplitude of tidal currents. The model results also revealed a complex spatial structure of the residual flow in the main channel of the river, with characteristics of multiple eddy-like cell circulations. These complicated residual currents are formed due to tidal rectification over variable topography with superimposition of inertial effects, asymmetry of tidal currents, and baroclinic pressure gradients. Water exchanges over the estuary-intertidal salt marsh complex are asymmetric across the estuary, and tend to vary periodically on the northern side while quickly washing out of the marsh zone on the southern side. Strong Stokes’ drifting velocity was predicted in the estuary, so that the Lagrangian trajectories of particles were characterized by strong nonlinear processes that differ significantly from those estimated by the Eulerian residual currents.

Abstract: Environmental factors that influence annual variability and spatial differences (within and between estuaries) in eelgrass meadows (Zostera marine L.) were examined within Willapa Bay, Washington, and Coos Bay, Oregon, over a period of 4 years (1998–2001). A suite of eelgrass metrics were recorded annually at field sites that spanned the estuarine gradient from the marine-dominated to mesohaline region of each estuary. Plant density (shoots m−2) of eelgrass was positively correlated with summer estuarine salinity and inversely correlated with water temperature gradients in the estuaries. Eelgrass density, biomass, and the incidence of flowering plants all increased substantially in Willapa Bay, and less so in Coos Bay, over the duration of the study. Warmer winters and cooler summers associated with the transition from El Nino to La Nina ocean conditions during the study period corresponded with this increase in eelgrass abundance and flowering. Large-scale changes in climate and nearshore ocean conditions may exert a strong regional influence on eelgrass abundance that can vary annually by as much as 700% in Willapa Bay. Lower levels of annual variability observed in Coos Bay may be due to the stronger and more direct influence of the nearshore Pacific Ocean on the Coos Bay study sites. The results suggest profound effects of climate variation on the abundance and flowering of eelgrass in Pacific Northwest coastal estuaries.

Abstract: We examined the vascular plant species richness and the extent, density, and height ofSpartina species of ten Narragansett Bay, Rhode Island (United States) fringe salt marshes which had a wide range of residential land development and N-loadings associated with their watersheds. Significant inverse relationships of tallS. alterniflora with species richness and with the extent and density ofS. patens and shortS. alterniflora were observed. Extent and density ofS. patens and extent of shortS. alterniflora were positively and significantly related with plant species richness. Marsh elevation and area did not significantly correlate with plant structure. Flood tide height significantly and inversely correlated withS. patens, but did not significantly relate toS. alterniflora or plant species richness. Marsh width significantly and positively correlated with plant species richness andS. patens and inversely correlated with tallS. alterniflora. Significant inverse relationships were observed for N-load, % residential development, and slope withS. patens, shortS. alterniflora, and species richness, and significant positive relationships with tallS. alterniflora. The marsh slope and width were significantly correlated with N-load and residential development that made it difficult to determine to what extent anthropogenic stressors were contributing to the variation in the plant structure among the marshes. At five marhes with similar slopes, there were significant inverse relationships of N-load withS. patens (density and extent) and a positive relationship with tallS. alterniflora (extent). Although there were no significant relationships of slope with the plant metrics among the five sites, other physical factors, such as the flood tide height and marsh width, significantly correlated with the extent and density ofSpartina species. Significant relationships of N-load with plant structure (albeit confounded by the effect of the physical characteristics) support the hypothesis of competitive displacement of dominant marsh plants under elevated nitrogen. It is likely that the varying plant structure in New England marshes is a response to a combination of natural factors and multiple anthropogenic stressors (e.g., eutrophication and sea level rise).

Abstract: Fish-habitat relationships on the shallow inner continental shelf were quantified with video sled and metered beam trawl on Fenwick and Weaver shoals offshore of Maryland and Delaware, U.S. These areas provide megascale physical relief and habitat complexity, but for juvenile fishes, mesoscale and microscale habitat is very important particularly as refuge from predation. At these smaller scales, much of the relief on the inner continental shelf is contributed by bedforms or sand waves and biogenic structures such as tubes, shell beds, or pits. A quantitative association for juvenile fishes between and within benthic habitats was found and related primarily to bedform size and amount of biogenic structure. The incidence of fishes was about four-times higher for large bedforms (> 30 cm wavelength and about 10 cm crest height) relative to smaller bedforms (<30 cm wavelength and about 5 cm crest height). For biogenic structure, going from high patch-mat tube densities to lower densities or no biogenic structure increased fish incidence by 5.4 and 3.3 times, respectively. The significant relationships of fishes with bedform size and density of biogenic structure indicated that seemingly small differences in physical structure of a habitat can make the difference between unacceptable and essential habitat for juvenile fishes. Proximity of complex and simple habitats was important in the diel use of habitat and in balancing pressure of refuge from predation provided by complex habitats with foraging for increased resources available in simpler habitats. During the day, spatially complex habitats comprised ofDiopatra andAsabellides tube mats had about twice as many fishes relative to bare sandy habitats (8.3–9.9 versus 4.0–4.1 fishes 100 m−2, respectively). At night, the pattern was reversed with more fishes present in the bare sandy habitats (12.4–13.5 versus 5.6–8.7 fishes 100 m−2). Some fish, such asAmmodytes spp., were very habitat specific and occurred only on dynamic coarser sands near the top of the shoals. Others, such asUrophycis regia, showed less habitat preference and occurred in all habitats during both day and night. Combining the effects of physical relief and biogenics, the habitat with the highest incidence of fishes had large bedforms with some biogenic structure. More emphasis needs to be placed on quantifying the relationship between fishes and their habitats for the fisheries management concept of essential fish habitat to develop into an effective tool on the inner continental shelf. The juvenile life history stages need to be emphasized because fish-habitat interactions are the strongest for these stages and may be the most ecologically important.

Abstract: Oxygen air-water gas exchange was measured using floating chambers in two shallow tidal estuaries of differing bathymetry and local terrain, near Waquoit Bay, Massachusetts (United States). The specific chamber design permitted measurements of gas flux in 15 min, allowing analysis of the relationship with wind speed and tidal stage. Exchange coefficients ranged from 0.5 to 2.5 g O2·m−2 h−1 atm−1 (equivalent to piston velocities of 1.5 to 7 cm h−1) for wind speeds of 0.3 to 9 m s−1 at 10 m elevation. While the relationships for each estuary appear linear (significant linear regressions with wind speed were shown for each estuary, and the slopes were different at the 99.5% confidence level), the range of speeds differed at the two sites and an exponential function of wind speed was consistent with the combined data from both estuaries. A power function of wind speed was not an acceptable model. The exchange coefficients for our estuaries are from 57% to as low as 9% of that predicted by previously published generic equations. Because the atmospheric correction can be significant in shallow, metabolically active coastal waters, we suggest that empirically determined relationships for gas exchange versus wind for a specific estuary are preferable to the predictions of the general equations. While the floating chamber method should be used cautiously, at low winds speeds (below 8 m s−1) and in slowly flowing waters, it provides a convenient approach for quantifying these site-specific differences. The differences, especially those between shallow sheltered systems and the open waters best fit by some published relationships, are ecologically important and do not appear yet to be measurable by other methods.

Abstract: Decapod crustaceans occupying seagrass, salt marsh edge, and oyster habitats within the St. Martins Aquatic Preserve along the central Gulf coast of Florida were quantitatively sampled using a 1-m2 throw trap during July–August 1999 and March–April 2000. Relative abundance and biomass were used as the primary measures to compare patterns of occupancy among the three habitat types. Representative assemblages of abundant and common species from each habitat were compared using Schoener's Percent Similarity Index (PSI). In all, 17,985 decapods were sampled, representing 14 families and 28 species. In the summer sampling period, mean decapod density did not differ between oyster and seagrass habitats, which both held greater densities of decapods than marsh-edge. In the spring sampling period oyster reef habitat supported greater mean decapod density than both seagrass and marsh-edge, which had similar densities of decapods. Habitat-specific comparisons of decapod density between the two sampling periods indicated no clear seasonal effect. In summer 1999, when seagrasses were well established, decapod biomass among the three habitats was not significantly different. During spring 2000, decapod biomass in oyster (41.40 gm−2) was greater than in marshedge (4.20 gm−2), but did not differ from that of seagrass (9.73 g m−2). There was no significant difference in decapod biomas between seagrass and marsh-edge habitats during the spring 2000 sampling period. The assemblage analysis using Schoener's PSI indicated that decapod assemblages associated with oyster were distinct from seagrass and marshedge habitats (which were similar). The results of this study suggest that in comparison to seagrass and marsh-edge habitats, oyster reef habitats and the distinct assemblage of decapod crustaceans that they support represent an ecologically important component of this estuarine system.

Abstract: We compared nekton densities over a range of measured flooding conditions and locations withinPhragmites australis andSpartina alterniflora (salt marsh cordgrass) at the Charles Wheeler Salt Marsh, located on the lower Housatonic River estuary in southwestern Connecticut. Nekton were sampled on nine spring high tide events from May to October 2000 using bottomless lift nets positioned between 0–5 and 10–20 m from the creek edge. Flooding depth, duration, and frequency were measured from each vegetation type during each sampling month. Benthic macroinvertebrate density was also measured within each vegetation type in May, July, and September. Frequency of flooding was 52% lower and flooding depth and duration were also significantly reduced inP. australis relative toS. alterniflora. A total of 4,197 individuals representing 7 species, mostlyPalaemonetes pugio (dagger-blade grass shrimp) andFundulus heteroclitus (common mummichog), were captured.P. pugio densities were significantly greater inS. alterniflora as were benthic macroinvertebrate density and taxa richness during May, but not during June or October. Total fish density was not significantly different betweenP. australis andS. alterniflora and was independent of location on the marsh. Significantly more juvenileF. heteroclitus were collected withinS. alterniflora relative toP. australis in June and July, suggesting that recruitment of this species may be lower inP. australis habitat. Fish density generally did not vary predictably across the range of flooding depth and duration; there was a positive relationship between flooding depth and fish density inS. alterniflora. The measured reduction in flooding frequency (52%) withinP. australis at the Housatonic site would result in an average total monthly fish use, expressed as density, of 447 ind m−2 forP. australis and 947 ind m−2 forS. alterniflora. WhenP. australis expansion results in reduction of flooding frequency and duration, nekton community composition can change, access to the marsh surface is reduced twofold, and nursery habitat function may be impaired.

Abstract: Microbial mercury (Hg) methylation and methylmercury (MeHg) degradation processes were examined using radiolabled model Hg compounds in San Francisco Bay-Delta surface sediments during three seasonal periods: late winter, spring, and fall. Strong seasonal and spatial differences were evident for both processes. MeHg production rates were positively correlated with microbial sulfate reduction rates during late winter only. MeHg production potential was also greatest during this period and decreased during spring and fall. This temporal trend was related both to an increase in gross MeHg degradation, driven by increasing temperature, and to a build-up in pore water sulfide and solid phase reduced sulfur driven by increased sulfate reduction during the warmer seasons. MeHg production decreased sharply with depth at two of three sites, both of which exhibited a corresponding increase in reduced sulfur compounds with depth. One site that was comparatively oxidized and alkaline exhibited little propensity for net MeHg production. These results support the hypothesis that net MeHg production is greatest when and where gross MeHg degradation rates are low and dissolved and solid phase reduced sulfur concentrations are low.

Abstract: A large data set, collected under the national Danish monitoring program, was used to evaluate the importance of photon flux density (PFD), relative wave exposure (REI), littoral slope, and salinity in regulating eelgrass cover at different depth intervals in Danish coastal waters. Average eelgrass cover exhibited a bell-shaped pattern with depth, reflecting that different factors regulate eelgrass cover at shallow- and deep-water sites. The multiple logistic regression analysis was used to identify regulating factors and determine their role in relation to eelgrass cover at different depth intervals. PFD, REI, and salinity were main factors affecting eelgrass cover while littoral slope had no significant effect. Eelgrass cover increased with increasing PFD at water depths of more than 2 m, while cover was in versely related to REI in shallow water. This pattern favored eelgrass cover at intermediate depths where levels of PFD and REI were moderate. Salinity had a minor, but significant, effect on eelgrass cover that is most likely related to the varying costs of osmoregulation with changing salinity. The analysis provided a useful conceptual framework for understanding the factors that regulate eelgrass abundance with depth. Although the regression model was statistically significant and included the factors generally considered most important in regulating eelgrass cover, its explanatory power was low, especially in shallow water. The largest discrepancies between predicted and observed values of cover appeared in cases where no eelgrass occurred despite sufficient light and moderate levels of exposure (almost 50% of all observations). These discrepancies suggest that population losses due to stochastic phenomena, such as extreme wind events, played an important regulating role that is not adequately described by average exposure levels. A more thorough knowledge of the importance of such loss processes and the time scales involved in recovery of seagrass populations after a severe disturbance are necessary if we are to understand the regulation of seagrass distribution in shallow coastal areas more fully.

Abstract: We investigated the independent and interactive effects of nutrient loading and summer water temperature on phytoplankton, drift macroalgae, and eelgrass (Zostera marina) in a coastal lagoon mesocosm experiment conducted from May through August 1999. Temperature treatments consisted of controls that approximated the 9-yr mean daily temperatures for Ninigret and Point Judith Lagoons in Rhode Island (United States) and treatments approximately 4°C above and 4°C below the controls. Nutrient treatments consisted of the addition of 6 mmol N m−2d−1 and 0.5 mmol P m−2 d−1 to mesocosms 4°C above and 4°C below the 9-yr daily mean. Nutrient enrichment produced marked phytoplankton blooms in both cool and warm treatments during early summer. These were replaced after midsummer by dramatic growths of macroalgal mats ofEnteromorpha flexuosa and, to a lesser degree,Cladophora sericea. No phytoplankton blooms were observed in the cool unenriched treatments, but blooms did develop in the mean temperature and warm mesocosms during the second half of the summer that were similar in intensity, though of shorter duration, than those observed earlier in the enriched systems. Macroalgal blooms did not occur in the unenriched mesocosms. Sustained warm water temperatures markedly decreased eelgrass density and belowground production and increased the time interval between the initiation of new leaves, particuarly when the biomass of macroalgae was high. The negative effect of elevated water temperature on eelgrass was significantly increased under conditions of elevated inorganic nutrient input. By the end of summer, virtually all of the measures of eelgrass health declined in rank order from cool, to mean, to cool enriched, to warm, to warm enriched treatments. It is likely that the marked declines in eelgrass abundance observed during recent decades in the Northeast have resulted from an interaction of increasing nutrient enrichment combined with increasing summer water temperatures.