Nitrogen cycling and ecosystem exchanges in a Virginia tidal freshwater marsh

Tidal freshwater marshes are diverse habitats that differ both within and between marshes in terms of plant community composition, sediment type, marsh elevation, and nutrient status. Because our knowledge of the nitrogen (N) biogeochemistry of tidal freshwater systems is limited, it is difficult to assess how these marshes will respond to long-term progressive nutrient loading due to watershed development and urbanization. We present a process-based mass balance model of N cycling in Sweet Hall marsh, a pristine (i.e., low nutrient)Peltandra virginica-Pontederia cordata dominated tidal freshwater marsh in the York River estuary, Virginia. The model, which was based on a combination of field and literature data, revealed that N cycling in the system was largely conservative. The mineralization of organic N to NH₄ ⁺ provided almost twice as much inorganic N as was needed to support marsh macrophyte and benthic microalgal primary production. Efficient utilization of porewater NH₄ ⁺ by nitrifiers and other microbes resulted in low rates of tidal NH₄ ⁺ export from the marsh and little accumulation of NH₄ ⁺ in marsh porewaters. Inputs of N from the estuary and atmosphere were not critical in supporting marsh primary production, and served to balance N losses due to denitrification and burial. A comparison of these results with the literature suggests that the relative importance of tidal freshwater marsh N cycling processes, including plant productivity, organic matter mineralization, microbial immobilization, and coupled nitrification-denitrification, are largely independent of small changes in water column N loading. Although very high (millimolar) concentrations of dissolved inorganic N can affect processes including denitrification and plant productivity, the factors that cause the switch from efficient N recycling to a more open N cycle have not yet been identified.     

Interactive effects of animal disturbance and elevation on vegetation of a tidal freshwater marsh

We studied interactions between animal disturbance (geese, carp, and muskrat) and elevation in a field experiment in tidal freshwater marshes of the Patuxent River, Maryland, United States. Vegetation changes were recorded in fenced and unfenced plots in high and low marsh community types for 2 yr using measurements of areal cover and within-plot frequency (which were averaged to create a dominance index), Leaf Area Index (LAI), and aboveground biomass. We related light environment to differences in vegetation using below-canopy measurements of Photosynthetically Active Radiation (PAR). In the low marsh, total cover of all species, cover of annual species, biomass, and LAI were significantly higher in plots fenced to exclude animals (exclosures) than in unfenced plots (fenced/unfenced total cover=76/40%, annual cover=45/10%, biomass=936/352 g m^?2, LAI=3.3/1.4). PAR was significantly lower in fenced than unfenced plots (fenced/unfenced=115/442 μmol s-1 m^?2). Despite the strong effect of fencing on biomass, species richness per plot (i.e., the number of species per plot, or species density) was not affected significantly by fencing in the low marsh. Most of the observed differences in cover, biomass, LAI, and PAR were due to variation in the abundance of the herbaceous annual speciesBidens laevis (dominance index fenced/unfenced=45/10%) andZizania aquatica (30/12%). In the high marsh community, fencing had only minor effects on plant community composition and did not significantly affect species richness, cover, biomass, PAR, or LAI. Our results show that animals can dramatically affect low marsh vegetation, primarily via physical disturbance or herbivory of shallowly rooted seedlings of annual species.     

The impact of vertebrate herbivores on wetland vegetation in Atchafalaya Bay, Louisiana

Delta islands in the Atchafalaya and Wax Lake deltas in Atchafalaya Bay, Louisiana, are in an extremely dynamic successional phase. These islands initially supported large marshes dominated by the pioneering plant species Sagittaria latifolia and Sagittaria platyphylla. A general decrease in vegetated areas has occurred in the delta island marshes in the Atchafalaya Delta since about 1980, while in the Wax Lake Delta portion of the complex the vegetation still flourished. The Atchafalaya Delta provides an interesting setting for the study of herbivory because of the complex interaction of biotic and physical factors operating in this delta. We hypothesized that grazing by herbivores has a marked effect on vegetation in these developing marshes. To test this hypothesis, exclosures were erected on islands in both deltas in September 1985 and January 1986. Each set of exclosure treatments included an openly-grazed control area, an ungrazed area, an area allowing nutria grazing, and one allowing waterfowl grazing in each site. Results of the experiment, based on field sampling of vegetation, indicated decreases in plant biomass and changes in plant species composition in grazed treatments. Waterfowl and nutria reduced biomass aboul equally, but there was a more marked effect in the openly grazed areas. These findings may be extrapolated to sediment diversion areas along the Mississippi River.     

Nutrient transport in a riverine-influenced,tidal freshwater bayou in Louisiana

Transport of ammonium (NH₄ ⁺), nitrate + nitrite (NO₃ ^?), total Kjeldahl nitrogen (TKN), soluble reactive phosphate (SRP), and total suspended solids (TSS) was measured in a freshwater tidal bayou located in a marsh system near the mouth of the Atchafalaya River in Louisiana. Sampling was conducted six times over one year and was timed to assess effects of seasonal variation in river flow and mean sea level of the Gulf of Mexico on material fluxes. Net fluxes of all materials were large and ebb directed in all seasons except fall, when net transport was 2 to 3 orders-of-magnitude smaller than in any other season. These results demonstrate that riverine forcing was the primary influence on materials transport in all seasons except fall when tidal forcing was most important. The range of net fluxes (g s^?1) for each nutrient was as follows (a negative sign indicates a net export toward the Gulf): NO₃ ^?, ?0.006 to ?6.69; TKN, 0.09 to ?10.41; NH₄ ⁺, ?0.02 to ?1.36; SRP, ?0.001 to ?0.53; TSS, ?2 to ?81. Analysis of nutrient concentrations indicated the marsh/aquatic system removed NO₃ ^?, SRP, and TSS from the water column from late spring through early fall and released NH₄ ⁺ and TKN in summer. The results of this study show that net materials export per unit cross section channel area increased as riverine influence increased.     

Nutrient and metal accumulation in a freshwater tidal marsh

Stratigraphic records from sediment cores collected in a freshwater tidal marsh and in the estuary upstream and downstream from the marsh were used to determine the accumulation of nutrients and trace metals over long time periods. Analysis of pollen and seeds show that the high marsh has formed only within the past 100 yr, following increased sedimentation rates in the area. Variations in nutrient and trace metal accumulations over several decades show that pollutants from agricultural runoff and wastewater discharge are stored to a greater extent in high-marsh than in low-marsh sediments. Greater accumulation rates in the high marsh are probably related to its greater sedimentary organic carbon concentration.     

A comparison of water-column macroinvertebrate communities in beds of differing submersed aquatic vegetation in the tidal freshwater Potomac River

Macroinvertebrates are a major food source for fish species and macrophyte beds are hypothesized to harbor a rich community of these organisms. Macroinvertebrates inhabiting the water column in two macrophyte beds and an adjacent open area were sampled in a small embayment of the tidal freshwater Potomac River. One macrophyte bed consisted of an almost complete monoculture ofHydrilla verticillata, while the second community was a more diverse mixture of plant species. In samples with substantial amounts of submersed aquatic vegetation (SAV), macroinvertebrate density was two orders of magnitude higher than and substantially more taxa were found than at the open water site. Total macroinvertebrate abundance was significantly greater at theH. verticillata site than at the mixed site in July, but no significant difference was observed in August. Taxa richness did not vary between the two vegetated sites in July but was higher in the mixed bed in August. While the two vegetated sites shared similar taxa, they differed in their abundance. TheH. verticillata site harbored more hydrobiid snails, and the mixed site was characterized by more chironomids and hydroptilid caddisflies. Differences between July and August collections were even greater than between sites. Numbers of hydroptilid caddisflies, baetid mayflies, and coenagrionid damselflies were substantially higher in August, while oligochaetes, hydrobiids, and chironomids were reduced. Results support the hypothesis that water-column macroinvertebrates are greatly enhanced in the presence of macrophytes. The ecological significance of the less substantial differences in macroinvertebrates between macrophyte beds requires further study.     

The paleoecological development of a late holocene,tidal freshwater marsh of the Upper Delaware River estuary

Changes in groundwater tables brought about by sea level increases in the Delaware River Basin (near Philadelphia) about 2,500 years B.P., initiated wetland development at the Princeton-Jefferson Branch of the Woodbury Creek marshes. Continual increases in sea level pushed groundwater tables further upward, and by approximately 800 years B.P., groundwater tables had risen to the upper limits for woody vegetation at the site. By the time European settlers arrived in the late 1600s nontidal sedge marshes dominated the site. Upon arriving colonists began manipulating the hydrology of the Delaware River Basin by constructing dams and dikes for flood control. Soon many areas were cut off from direct contact with the river. During the next one and one-half centuries sea level continued to rise, and because of channelization of the Delaware River the tidal range doubled. During the early 1900s flood control structures began to fail allowing tidal waters to periodically inundate these protected sites. At that time the site was dominated by a Quercus-Castanea swamp forest with hummocks of Cyperaceae interspersed throughout. In 1940 the dike surrounding the Princeton-Jefferson marsh collapsed and the site was immediately inundated with tidal waters on a regular basis. Within a short period of time tidal freshwater marsh developed and has continued to the present day. It is clear from this investigation that changes in hydrology brought about by cultural modifications have been directly responsible for the ontogeny of this tidal marsh. The influence cultural impacts have had on wetland development at the Princeton-Jefferson marsh suggest that it may be necessary to reevaluate the extent humans have modified the development and structure of the present day upper Delaware River estuary. Although the ability to discern historic vegetation zonation patterns is limited, these marshes can record individual events that have shaped these wetlands through time. Due to differences in the structure of the plant community, rates of decomposition, and processes of accretion, Redfield’s model (1972) of tidal salt marsh development does not apply to the Princeton-Jefferson marsh. Along a submerging coast, the development of tidal freshwater marsh in many estuaries may be necessary for the establishment of brackish and salt marshes by creating and maintaining a suitable habitat for the eventual colonization of more salt-tolerant plant species. The roles these wetlands have played in the development of the estuaries has been underestimated in the past.     

Nitrogen and phosphorus in the sediments of a tidal,freshwater marsh in Massachusetts

Total organic nitrogen (TON) and phosphorus (TOP) were measured as a function of depth in 14 cores taken from a New England, tidal, freshwater marsh. TON and TOP ranged from 1.56 to 1.97% and 0.11 to 0.30% of dry weight sediments, respectively. The variation in both pool sizes over time was small and TON varied inconsistently with depth; however, TOP decreased regularly down to 20 cm. Consequently, the TON: TOP ratio increased linearly from 14∶1 at the surface to 32∶1 at 20 cm, then was nearly constant to 70 cm. This pattern may be a general feature of marsh sediments and may indicate 1) that phosphorus is recycled less efficiently than nitrogen, 2) that over time proportionately more introgen than phosphorus is incorporated into recalcitrant compounds, or 3) that phosphorus is more mobile than nitrogen in these marsh sediments. The total inorganic nitrogen pool was measured in this marsh also and was dominated by ammonium (97% of total). The annual average free ammonium concentration was 3.70±0.64 mg N per 1 at the surface and decreased to 0.92±0.18 mg N per 1 at 20 to 22 cm in the sediments. Sorptiondesorption studies showed that, on a fresh sediment volume basis, sediment sorbed ammonium was roughly equivalent to free porewater ammonium (K=0.8). The relationship between free and sorbed ammonium was linear between 0.4 and 24.0 mg NH₄·N per 1 of pore water. The depth distribution of ammonium in these sediments is probably maintained by a dynamic balance between net microbial mineralization of litter, plant uptake, transpiration, diffusion, and porewater advection.     

Using submerged aquatic vegetation to establish minimum and maximum freshwater inflows to the Caloosahatchee estuary, Florida

Species of submerged aquatic vegetation (SAV) are frequently used in the management of estuarine systems to set restoration goals, nutrient load reduction goals, and water quality targets. As human need for water increases, the amount of freshwater required by estuaries has become an increasingly important issue. While the, science of establishing the freshwater needs of estuaries is not well developed, recent attempts have emphasized the freshwater requirements of fisheries. We evaluate the hypothesis that SAV can be used to establish freshwater inflow needs. Salinity tolerance data from laboratory and field studies of SAV in the Caloosahatchee estuary, Florida, are used to estimate a minimum flow required to maintain the salt-tolerant freshwater species,Vallisneria americana, at the head of the estuary and a maximum flow required to prevent mortality, of the marine speciesHalodule wrightii at its mouth. ForV. americana, laboratory experiments showed that little or no growth occurred between 10‰ and 15‰ In the field, lower shoot densities (<400 shoots m^?2) were associated with salinities greater than 10‰. Results forH. wrightii were more variable than forV. americana. Laboratory experiments indicated that mortality could occur at salinities <6‰, with little growth occurring between 6‰ and 12‰. Field data indicated that higher blade densities (>600 blades m^?2) tend to occur at salinities greater than 12‰ Relationships between salinity in the estuary and discharge from the Caloosahatchee River indicated that flows>8.5 m³ s^?1 would produce tolerable salinity (<10‰) forV. americana and flows<89 m³ s^?1 would avoid lethal salinities (<6‰) forH. wrightii.     

Nekton use of salt marsh creeks in the upper Tejo estuary

The use of the Tejo estuary, Portugal, salt marsh creeks by nekton was examined based on sampling surveys with a fyke net from September 1998 until August 2001. From the 20 taxa (14 fish species, 5 decapod crustacean species, and 1 cephalopod species) identified in the studied creeks, 16 were regularly caught throughout the sampling period. The shrimpPalaemonetes varians was the most numerically abundant species in the creeks, while the biomass was dominated by the mulletLiza ramada. The nekton assemblage was mainly represented by marine-estuarine opportunist species, comprising 85% of the total. A high seasonality was detected on the species abundance patterns: the most abundant species (P. varians, Crangon crangon, L. ramada, Pomatoschistus microps, Syngnathus sp., andAnguilla anguilla) occurred throughout the sampling period,Sardina pilchardus, Dicentrarchus, labrax, andAtherina boyeri were particularly abundant in spring and summer, andEngraulis encrasicholus, Liza aurata, Gambusia holbrooki, Palaemon longirostris, andPalaemon serratus were most abundant in autumn and winter.L. ramada occurred in the tidal creeks in high numbers during neap tides, while the majority of the remaining taxa were most abundant during spring tides, suggesting a differential pattern of habitat use occording to species.     

Seasonal and spatial water quality changes in the outflow plume of the Atchafalaya River,Louisiana, USA

The objective of this study was to examine the interaction between the Atchafalaya River and the Atchafalaya Delta estuarine complex. Measurements of suspended sediments, inorganic nutrients (NO₃ ^?, NH₄ ⁺, PO₄ ^3?), chlorophylla (chla), and-salinity were taken monthly from December 1996 to January 1998. These data were compiled by season, and the Atchafalaya River plume data were also analyzed using the Generalized Additive Model technique. There were significant decreases in NO₃ ^? concentrations during summer, fall, and winter as river water passed through the estuary, that were attributable to chemical and biological processes rather than dilution with ambient water. In some regions there were higher chla concentrations during summer and fall compared to winter and spring, when river discharge and the introduction of inorganic nutrients were highest, suggesting biological processes were active during this study. The presence of NH₄ ⁺, as a percentage of available dissolved inorganic nitrogen, increased with distance from the Atchafalaya River, indicative of remineralization processes and NO₃ ^? reduction. Mean PO₄ ^3? concentrations were often higher in the estuarine regions compared to the Atchafalaya River. During summer total suspended solid (TSS) concentrations increased with distance from the river mouth, suggesting a turbidity maximum. Highest chla concentrations were found in the bayous and shallow water bodies of the Terrebonne marshes, as were the lowest TSS concentrations. The low chla concentrations found in other areas of this study, despite high inorganic nutrient concentrations, suggest light limitation as the major control of phytoplankton growth. Salinity reached near seawater concentrations at the outer edge of the Atchafalaya River plume, but much lower salinities (<10 psu) were observed at all other regions. The Atchafalaya Delta estuarine complex buffers the impact of the Atchafalaya River on the Louisiana coastal shelf zone, with a 41% of 47% decrease in Atchafalaya River NO₃ ^? concentrations before reaching Gulf waters.     

Investigations of the availability and survival of submersed aquatic vegetation propagules in the tidal Potomac River

The establishment of submersed aquatic vegetation (SAV) at unvegetated sites in the freshwater tidal Potomac River was limited primarily by factors other than propagule availability. For two years, traps were used to quantify the amount of plant material reaching three unvegetated sites over the growing season. The calculated flux values provided a gross estimate of the flux of propagules that could potentially survive if other site factors were suitable. The mean flux ofHydrilla verticillata and all other species (≥0.01 gdw m^?2 d^?1) appeared sufficient to favor the establishment of vegetation, particularly considering the high viability (70–100%) of whole plants and fragments under controlled conditions. However, median water clarity values (i.e., for light attenuation, Secchi depth, total suspended solids, and chlorophylla) were below SAV restoration goals at all unvegetated sites. Additionally, sediments from unvegetated sites showed a potential for nitrogen limitation of the growth ofH. verticillata. Our findings support the hypothesis that in the tidal Potomac River, water clarity and nutrient (especially nitrogen) levels in sediment are key to plant community establishment.     

Primary production and seasonal aspects of emergent plants in a tidal freshwater marsh

Seasonal changes in aboveground plant biomass, cover, and frequency were monitored in Sweet Hall Marsh, a tidal freshwater marsh located on the Pamunkey River, Virginia, during the 1974 growing season.Peltandra virginica accumulated the most biomass, 423.40 g per m², followed byLeersia oryzoides at 67.75 g per m². Annual net community production was estimated to be 775.74 g per m² by using a multiple-harvest technique. Comparisons with other studies revealed that production was somewhat low for tidal freshwater marshes but mostly higher than production in Virginia brackish and saline wetlands. Measurements revealed an annual succession of plant species from spring to fall. The pattern observed was early dominance byPeltandra followed by a rise in importance ofPolygonum spp.,Impatients capensis andLeersia.     

The value of salt marsh edge vs interior as a habitat for fish and decapod crustaceans in a Louisiana tidal marsh

Flume nets of various lengths and a 3-m seine were used to sample the fishes and macrocrustaceans using a flooded Louisiana salt marsh and the adjacent tidal creek. The experiment allowed for species-specific comparisons of the flooded marsh at the creek edge versus the interior. Of the 37,667 organisms collected in flume nets from January through November 1989, 89% were decapods (nine species) and 11% were fish (29 species). An additional 18,539 organisms (75% decapods and 25% fish) were collected from concurrent seine samples taken from July through November. Comparison of catches among different flume lengths and low tide versus high tide seine collections revealed distinct patterns of marsh habitat utilization. Densities of most organisms were highest within 3 m of the water’s edge, but significant numbers of marsh-resident fish species used the interior marshes. The edge marshes appeared to be used by both transient and resident species; however, the interior marshes were used primarily by marsh-resident species (Cyprinodontiformes andPalaemonetes sp.) that are excellent food sources for adult transient-species. Four zonations of marsh use are described for transients, residents, and rare species.     

Changing elevation,accretion, and tidal marsh plant assemblages in a South San Francisco Bay tidal marsh

Analyses of organic content, pollen, and the carbon-isotopic composition of a 3.5-m sediment core collected from a subsided tidal marsh located in South San Francisco Bay, California, have provided a 500-yr record of sediment accretion and vegetation change before, during, and after a rapid 1 m increase in sea level. Core chronology was established using¹⁴C dating of fossil plant material, the first appearance of pollen types produced by plants not native to California, and changes in lead concentrations coincident with anthropogenic contamination. Prior to the mid 19th century, rates of sediment accretion were between 1 and 4 mm yr⁻¹; sediment accretion accelerated to an average of 22 mm yr⁻¹ following the initiation of subsidence. Changes in tidal marsh vegetation also accompanied this depositional change. Vegetation shifted from a high to low marsh assemblage, as indicated by a larger percentage of grass pollen, rhizomes ofSpartina foliosa, and a strong C₄ signal. Between 1980 and 2001, Triangle marsh again developed high marsh vegetation, as indicated by higher percentages of the Amaranthaceane pollen type, seed deposition, includingSalicornia spp., and more negative carbon isotopic ratios.     

Subtidal sea level variability in a shallow Mississippi River Deltaic estuary, Louisiana

The relative roles of river, atmospheric, and tidal forcings on estuarine sea level variability are examined in Breton Sound, a shallow (0.7 m) deltaic estuary situated in an interdistributary basin on the Mississippi River deltaic plain. The deltaic landscape contains vegetated marshes, tidal flats, circuitous channels, and other features that frictionally dissipate waves propagating through the system. Direct forcing by local wind stress over the surface of the estuary is minimal, owing to the lack of significant fetch due to landscape features of the estuary. Atmospheric forcing occurs almost entirely through remote forcing, where alongshore winds facilitate estuary-shelf exchange through coastal Ekman convergence. The highly frictional nature of the deltaic landscape causes the estuary to act as a low-pass filter to remote atmospheric forcing, where high-frequency, coastally-induced fluctuations are significantly damped, and the damping increases with distance from the estuary mouth. During spring, when substantial quantities of controlled Mississippi River inputs (?q = 62 m³ s^-1) are discharged into the estuary, upper estuary subtidal sea levels are forced by a combination of river and remote atmospheric forcings, while river effects are less clear downestuary. During autumn (?q = 7 m³ s^-1) sea level variability throughout the estuary is governed entirely by coastal variations at the marine boundary. A frequency-dependent analytical model, previously used to describe sea level dynamics forced by local wind stress and coastal forcing in deeper, less frictional systems, is applied in the shallow Breton Sound estuary. In contrast to deeper systems where coastally-induced fluctuations exhibit little or no frictional attenuation inside the estuary, these fluctuations in the shallow Breton Sound estuary show strong frequency dependent amplitude reductions that extend well into the subtidal frequency spectrum.     

Macroinvertebrates of a rocky shore in the freshwater tidal Hudson River

We studied the macroinvertebrate fauna of a rocky shore in the freshwater tidal Hudson River during 1992–1994, the early years of the zebra mussel (Dreissena polymorpha) invasion. The macroinvertebrate community was numerically dominated by chironomids, nematodes, oligochaetes, gastropods, zebra mussels, and planarian flatworms. The community was a mixture of species typical of stony warm water rivers and lake shores, freshwater generalists, and semiterrestrial species. Overall macroinvertebrate densities were moderate to low (2,800–14,600 m⁻²). Density was a strong function of season and elevation, with consistently low densities in the early spring and in the intertidal zone. This pattern suggests that physical harshness (alternating submergence and desiccation;ice and low temperatures) limits the distribution of invertebrates at this site. Zebra mussels occurred at our study site chiefly below the low tide mark, but only at moderate abundance (usually <1,000 m⁻²). A weak correlation between the densities of zebra mussels and those of other macroinvertebrates nonetheless suggests that the zebra mussel invasion may have affected community structure.     

Sources, input pathways, and distributions of Fe, Cu, and Zn in a Chesapeake Bay tidal freshwater marsh

Tidal freshwater marshes exist at the interface between watersheds and estuaries, and thus may serve as critical buffers protecting estuaries from anthropogenic metal pollution. Bi-weekly samples of newly deposited marsh sediments were collected and analyzed for Cu, Zn, and Fe concentrations over 21 months from July 1995 to March 1997 in five distinct habitats at the head of Bush River, Maryland. Bi-weekly anthropogenic metal enrichments ranged from 0.9–4.7. Anthropogenic excess metal loadings averaged over 1996 ranged from 6–306 and 25–1302 μg cm⁻² year⁻¹ between sites for Cu and Zn, respectively. Based on Fe-normalized trace metal signatures, Susquehanna River sediment does not significantly contribute to upper Bush River. Organic matter was found to dilute total metal concentrations, whereas past studies suggested organics enhance labile metal content. Analysis of metal input pathways shows that marsh metals are primarily imported from nearby subtidal accumulations of historic watershed material by tidal flushing. Received: 29 April 1999 / Accepted: 7 December 1999     

Use of tidal freshwater marshes by fishes and macrofaunal crustaceans along a marsh stream-order gradient

Fishes and invertebrate macrofauna (nekton) were sampled biweekly (July through October 1985) from the surface of tidal freshwater marshes. Samples were collected with flume nets at three different stream orders (orders 2, 3 and 4+) along a marsh stream order gradient. Twenty-five species of fishes (5,610 individuals, 17.072 kg preserved wet weight) representing 13 families, and three species of invertebrates (19,570 individuals, 13.026 kg preserved wet weight) were collected. The most abundant species were grass shrimp (Palaemonetes pugio), mummichogs (Fundulus heteroclitus), banded killifish (F. diaphanus), inland silversides (Menidia beryllina), and blue crabs (Callinectes sapidus). Invertebrate catches (mostly grass shrimp and blue crabs) were not significantly different among stations. Total numbers of fishes were significantly greater at both headwater (order 2) and main creek (order 3) stations than river (order 4+) stations, but catches of headwater and main creek stations were not significantly different. The relationship between marsh stream order and fish abundance may partly be related to the distribution of submerged aquatic vegetation (SAV) within marsh tidal creeks. Submerged aquatic vegetation decreases in abundance with increasing stream order. Some species may use SAV as a refuge from predators or as a foraging area during low tide when the marsh surface is inaccessible. The presence of SAV in tidal creeks may enhance the habitat value of adjacent marshes.     

Metabolism and organic carbon fluxes in the tidal freshwater Hudson River

We summarize rates of metabolism and major sources and sinks of organic carbon in the 148-k long, tidally influenced, freshwater Hudson River. The river is strongly heterotrophic, with respiration exceeding gross primary production (GPP). The P:R ration averages 0.57 (defined as the ratio of GPP to total ecosystem respiration) if only the aquatic portion of the ecosystem is considered and 0.70 if the emergent marshes are also included. Gross primary production (GPP) by photoplankton averages approximately 300 g C m^?2 yr^?1 and is an order of magnitude greater than that by submersed macrophytes. However, the river is deep, well mixed, and turbid, and phytoplankton spend a majority of their time in the dark. As a result, respiration by living phytoplankton is extremely high and net primary production (NPP) by phytoplankton is estimated to be only some 6% of GPP. NPP by phytoplankton and submersed macrophytes are roughly equal (approximately 20 g C m^?2 yr^?1 each) when averaged over the river. Emergent marshes are quite productive, but probably less than 16 g C m^?2 yr^?1 enters the aquatic portion of the ecosystem from these marshes. Heterotrophic respiration and secondary production in the river are driven primarily by allochthonous inputs of organic matter from terrestrial sources. Rates of metabolism vary along the river, with depth being a critical controlling factor. The P:R ratio for the aquatic portion of the ecosystem varies from 1 in the mid-river to 0.2 in the deeper waters. NPP is actually negative in the downstream waters where average depths are greater since phytoplankton respiration exceeds GPP there; the positive rates of NPP occurring upriver support a downstream advection of phytoplankton to the deeper waters where this C is largely respired away by the algae themselves. This autotrophic respiration contributes significantly to oxygen depletion in the deeper waters of the Hudson. The tidally influenced freshwater Hudson largely fits the patterns predicted by the river continuum model for larger rivers. However, we suggest that the continuum model needs to more clearly distinguish between GPP and NPP and should include the importance of autotrophic respiration by phytoplankton that are advected along a river. The organic carbon budget for the tidally influenced freshwater Hudson is balanced to within a few percent. Respiration (54%) and downstream advection into the saline estuary (41%) are the major losses of organic carbon from the ecosystem. Allochthonous inputs from nonpoint sources on land (61%) and GPP by phytoplankton (28%) are the major sources to the system. Agricultural erosion is the major source of allochthonous inputs. Since agricultural land use increased dramatically in the last century, and has fallen in this century, the carbon cycle of the tidally influenced freshwater Hudson River has probably changed markedly over time. Before human disturbance, the Hudson was probably a less heterotrophic system and may even have been autotrophic, with gross primary production exceeding ecosystem respiration.