David T. Fischer

ZEBRA MUSSEL INVASION IN A LARGE, TURBID RIVER: PHYTOPLANKTON RESPONSE TO INCREASED GRAZING

Changes in the biomass of benthic bivalves can cause dramatic changes in total grazing pressure in aquatic systems, but few studies document ecosystem-level impacts of these changes. This study documents a massive decline in phytoplankton biomass con- current with the invasion of an exotic benthic bivalve, the zebra mussel ( Dreissena poly- morpha), and demonstrates that the zebra mussel actually caused this decline. In the fall of 1992 the zebra mussel became established at high biomass in the Hudson River Estuary, and biomass of mussels remained high during 1993 and 1994. During these 2 yr, grazing pressure on phytoplankton was over 10-fold greater than it had been prior to the zebra mussel invasion. This increased grazing was associated with an 85% decline in phyto- plankton biomass. Between 1987 and 1991 (pre-invasion), summertime chlorophyll aver- aged 30 mg/m 3 ; during 1993 and 1994 summertime concentrations were ,5 mg/m 3 . Over this same period, light availability increased, phosphate concentrations doubled, some planktonic grazers declined, and average flow was not different from the pre-invasion period. Thus, changes in these other factors were not responsible for phytoplankton declines. We developed a mechanistic model that reproduces the spatial and temporal dynamics of phytoplankton prior to the invasion of the zebra mussel (1987-1991). The model ac- curately predicts extreme declines in phytoplankton biomass after the invasion (1993-1994). The model demonstrates that zebra mussel grazing was sufficient to cause the observed phytoplankton decline. The model also allows us to test which features make the Hudson River sensitive to the impact of benthic grazers. The model suggests that the fate of light- scattering inorganic particles (turbidity) is a key feature determining the impact of benthic grazers in aquatic systems.

Original Articles: Sources of Dissolved Organic Carbon Supporting Planktonic Bacterial Production in the Tidal Freshwater Hudson River

ABSTRACT Planktonic bacterial production in the tidal freshwater Hudson River is a major component of secondary productivity and is uncoupled from planktonic primary productivity. There are several major sources of allochthonous dissolved organic carbon (DOC) whose potential contribution to heterotrophic bacterial growth was examined with bioassays. Supply of DOC from the upper Hudson drainage basin and a large tributary in the mid-Hudson together comprise 70 kT DOC/year, which is the bulk of the DOC load to the tidal freshwater Hudson River. Two contrasting tidal wetlands contribute DOC to the main-stem river but were only a few percent of the tributary load even during summer low-flow conditions. The quantity of DOC released from fine sediments was intermediate to the other two loadings considered. Bacterial growth in bioassays receiving water from the sources varied, but differences in thymidine incorporation between reference and DOC sources were small, usually less than 2 nmol/L/h. Similarity in thymidine incorporation suggests that all sources of DOC were capable of supporting bacterial growth at approximately equal rates. Seasonal shifts in carbon availability were clear in several cases, for example, greater growth on wetland-derived DOC at times of peak plant productivity. Seasonal differences in tributary DOC bioavailability were not large despite the well-known seasonality of tributary inputs. Activities of a suite of extracellular enzymes were used as a biologically based characterization of DOC from the various sources. Shifts in allocation among enzymes were apparent, indicating that there are biologically relevant differences in composition among the sources. Fluorescence characteristics and absorbance per unit carbon also varied among sources, providing an independent confirmation of compositional differences among sources. The absence of large differences in bacterial productivity among sources suggests that growth is supported by a wide range of DOC, and the relative importance of the sources is probably related to the quantitative differences in inputs. Efforts to classify carbon supplies to ecosystems must recognize that organism plasticity in carbon use and physical mixing processes will both act to homogenize what might initially appear to be quite distinctive carbon inputs.

Consumption, Selectivity, and Use of Zooplankton by Larval Striped Bass and White Perch in a Seasonally Pulsed Estuary

Abstract Many estuaries exhibit seasonal pulses of phytoplankton and zooplankton production. Larval fishes that co-occur with these “blooms” should be at a growth and survival advantage compared with larvae that occur before or after the bloom, although this has been difficult to observe in many systems. We tested this potential advantage for larval (< 10 mm notochord length) striped bass Morone saxatilis and white perch M. americana in the Hudson River by examining consumption and feeding selectivities with respect to zooplankton blooms. The cladoceran Bosmina longirostris and large copepodite and adult copepods together composed 97.4%, and 90.9% of larval striped bass and white perch diets, respectively. Peak consumption rates of Bosmina coincided with the bloom. whereas copepod consumption rates continued to increase throughout the sampling period. Selectivity for copepods was inversely related to selectivity for Bosmina and was high, except at those sites and times when Bosmina densities exceeded 14 a...

Spatial and temporal variability in the lower food web of the tidal freshwater Hudson River

The mid Hudson River is a heterotrophic system where allochthonous inputs apparently fuel the largest proportion of secondary production and ecosystem metabolism. We have analyzed a 6-yr dataset collected quarterly at six stations spanning a 150-km reach to assess variability at inter- and intra-annual time scales and regional spatial scales. The major components of the lower food web: bacterial biomass, detrital particulate organic carbon (POC), and dissolved organic carbon (DOC), show surprisingly discordant patterns in temporal and spatial variability. Bacterial abundance shows significant variability at all three scales, but the interannual variability is by far the greatest. DOC concentrations showed greatest variability among years, with intra-annual and spatial variability roughly equal. Freshwater flow is commonly considered a major driving force in river-estuarine variability but simple discharge was not a strong predictor of any component of suspended matter or DOC. For organisms in the Hudson River food web, these multiple scales of variability indicate highly unpredictable food resources in time and space, and these fluctuations may contribute to the variability in higher trophic levels.

Biodiversity in Hudson River shore zones: influence of shoreline type and physical structure

The shore zones of the Hudson River, like those of many developed waterways, are highly varied, containing a mix of seminatural and highly engineered shores. Our goal was to document the biodiversity supported by different kinds of shore zones in the Hudson. We chose six common types of shore zones, three of them “natural” (sand, unconsolidated rock, and bedrock), and three of them engineered (riprap, cribbing, and bulkheads). We measured selected physical characteristics (shore zone width, exposure, substrate roughness and grain size, shoreline complexity) of three examples of each of these shore types, and also sampled communities of terrestrial plants, fishes, and aquatic and terrestrial invertebrates. Community composition of most taxa differed across shore types, and frequently differed between wide, sheltered shores and narrow, exposed shores. Alien plant species were especially well represented along engineered shores. Nevertheless, a great deal of variation in biological communities was not explained by our six-class categorization of shore zones or the physical variables that we measured. No single shore type supported the highest values of all kinds of biodiversity, but engineered shore zones (especially cribbing and bulkheads) tended to have less desirable biodiversity characteristics than “natural” shore zones.

Predicting Carbon and Nutrient Transformations in Tidal Freshwater Wetlands of the Hudson River

The exchange of water between the main channel of the tidal freshwater Hudson River and its tidal wetlands is a large proportion of the whole-river water volume and causes large changes in concentrations of some dissolved and suspended constituents. Ten representative wetlands were assessed for their ability to alter quantities of inorganic nutrients, suspended particles, dissolved organic carbon (DOC), and dissolved oxygen during tidal exchange. The majority of sites acted as sinks for oxygen and nitrate and as sources of DOC. For other variables such as phosphate and pigments, individual wetlands varied broadly in both the direction and magnitude of change. For some variables (oxygen, DOC) we found mechanistically plausible predictors for the magnitude of alteration. The proportional coverage of submerged vegetation or intertidal marsh graminoid vegetation was related to the degree of change in oxygen and DOC. For most cases, however, we did not find strong predictors and we attribute this to the spatial positioning of “hot spots” or redundancy in the processes actually responsible for the transformation. Our ability to predict ecosystem performance from whole-ecosystem attributes may be impeded by lack of consideration of within-system spatial contingencies or lack of knowledge of which process is actually responsible for the observed alteration in material flux.

Multi-Scale Controls on Water Quality Effects of Submerged Aquatic Vegetation in the Tidal Freshwater Hudson River

Patches of submerged vegetation can be important sites of primary production and habitat for organisms in many aquatic ecosystems. In the tidal freshwater Hudson River they make up about 6% of the river bottom area. Direct sampling of water masses passing through patches of vegetation and week-long continuous monitoring of water characteristics were used to determine plant effects on dissolved oxygen and suspended sediments. Vegetated areas could have dissolved oxygen concentrations substantially higher than in the main channel and suspended sediments and turbidity were frequently higher in vegetated areas. Patches of Vallisneria americana had variable capacity to maintain super-saturated oxygen concentrations; patch size accounted for some of the variability whereas larger-scale differences in main-channel influent water also contributed. Differences in turbidity among sites were harder to account for; width of plant beds and abundance of neighboring vegetated areas contributed weakly to predictions of local turbidity. Functional heterogeneity within ecosystems is common and attempts to explain variability may require understanding different controlling factors for different functions and appreciating that factors operate at multiple scales.

Submersed Macrophyte Effects on Nutrient Exchanges in Riverine Sediments

Submersed macrophytes are important in nutrient cycling in marine and lacustrine systems, although their in nutrient exchange in tidally-influenced riverine systems is not well studied. In the laboratory, plants significantly lowered porewater nutrient pools of riverine sediments compared with bare controls. Deep-rootedVallisneria americana lowered the porewater nutrients to a greater extent than the shallow-rootedPotamogeton pectinatus. V. americana showed significantly higher tissue nutrient content (N in roots, P in leaves) thanP. pectinatus. porewater nutrients in the river increased from spring to summer (1995) when vegetation was at its peak (for porewater PO4-P, p<0.05). In 1996, porewater nutrients were higher during peak plant biomass in the summer than in the fall (for porewater PO4-P, p<0.05). In the summer (1995) vegetated patches had significantly greater porewater PO4-P than bare patches. We hypothesize that the concentrating of particulates in riverine grassbeds and subsequent microbial processing may provide an indirect source of nutrients for submersed macrophytes. In tidally-influenced riverine systems, biological mechanisms such as root uptake of nutrients and lateral oxygen release may be masked by the interaction of physical forces (i.e., tides, currents) with the structure of the grassbeds.