Alan D. Steinman

MEETING ECOLOGICAL AND SOCIETAL NEEDS FOR FRESHWATER

Human society has used freshwater from rivers, lakes, groundwater, and wetlands for many different urban, agricultural, and industrial activities, but in doing so has overlooked its value in supporting ecosystems. Freshwater is vital to human life and societal well-being, and thus its utilization for consumption, irrigation, and transport has long taken precedence over other commodities and services provided by freshwater ecosystems. However, there is growing recognition that functionally intact and biologically complex aquatic ecosystems provide many economically valuable services and long-term benefits to society. The short-term benefits include ecosystem goods and services, such as food supply, flood control, purification of human and industrial wastes, and habitat for plant and animal life—and these are costly, if not impossible, to replace. Long-term benefits include the sustained provision of those goods and services, as well as the adaptive capacity of aquatic ecosystems to respond to future environmental alterations, such as climate change. Thus, maintenance of the processes and properties that support freshwater ecosystem integrity should be included in debates over sustainable water resource allocation. The purpose of this report is to explain how the integrity of freshwater ecosystems depends upon adequate quantity, quality, timing, and temporal variability of water flow. Defining these requirements in a comprehensive but general manner provides a better foundation for their inclusion in current and future debates about allocation of water resources. In this way the needs of freshwater ecosystems can be legitimately recognized and addressed. We also recommend ways in which freshwater ecosystems can be protected, maintained, and restored. Freshwater ecosystem structure and function are tightly linked to the watershed or catchment of which they are a part. Because riverine networks, lakes, wetlands, and their connecting groundwaters, are literally the “sinks” into which landscapes drain, they are greatly influenced by terrestrial processes, including many human uses or modifications of land and water. Freshwater ecosystems, whether lakes, wetlands, or rivers, have specific requirements in terms of quantity, quality, and seasonality of their water supplies. Sustainability normally requires these systems to fluctuate within a natural range of variation. Flow regime, sediment and organic matter inputs, thermal and light characteristics, chemical and nutrient characteristics, and biotic assemblages are fundamental defining attributes of freshwater ecosystems. These attributes impart relatively unique characteristics of productivity and biodiversity to each ecosystem. The natural range of variation in each of these attributes is critical to maintaining the integrity and dynamic potential of aquatic ecosystems; therefore, management should allow for dynamic change. Piecemeal approaches cannot solve the problems confronting freshwater ecosystems. Scientific definitions of the requirements to protect and maintain aquatic ecosystems are necessary but insufficient for establishing the appropriate distribution between societal and ecosystem water needs. For scientific knowledge to be implemented science must be connected to a political agenda for sustainable development. We offer these recommendations as a beginning to redress how water is viewed and managed in the United States: (1) Frame national and regional water management policies to explicitly incorporate freshwater ecosystem needs, particularly those related to naturally variable flow regimes and to the linking of water quality with water quantity; (2) Define water resources to include watersheds, so that freshwaters are viewed within a landscape, or systems context; (3) Increase communication and education across disciplines, especially among engineers, hydrologists, economists, and ecologists to facilitate an integrated view of freshwater resources; (4) Increase restoration efforts, using well-grounded ecological principles as guidelines; (5) Maintain and protect the remaining freshwater ecosystems that have high integrity; and (6) Recognize the dependence of human society on naturally functioning ecosystems.

Biomass and Pigments of Benthic Algae

Abstract Biomass is one of the most fundamental measurements made in ecology. In stream ecology, biomass is frequently used to estimate the abundance of benthic primary producers, both autotrophic and heterotrophic. In this chapter, we (1) provide a context for the study of benthic algal biomass; (2) discuss in detail some of the more commonly used approaches to measure benthic algal biomass; and (3) describe a field exercise to examine the influence of irradiance on algal biomass, whereby these approaches can be employed and compared with each other to assess their individual performance.

Effects of herbivore type and density on taxonomic structure and physiognomy of algal assemblages in laboratory streams

Four densities of a snail (Juga silicula) and a caddisfly (Dicosmoecus gilvipes) were introduced into separate laboratory streams, and their effects on algal biomass and community structure were monitored for 32 d. Tiles in an ungrazed control stream were covered by thick algal mats by day 32, and were composed primarily of Scenedesmus spp., Characium, and a variety of diatoms. Biomass and community structure of algal assemblages in the stream with the lowest density of snails were very similar to those in the control stream. In the other streams with snails, an inverse relationship developed between algal biomass and snail density after day 16. By day 32, the algal assemblages in the streams with high snail densities were dominated by adnate diatoms (e.g., Achnanthes lanceolata), and basal cells and short filaments of Stigeoclorium tenue. In contrast to the streams with snails, algal biomass was relatively low in all streams with caddisflies. The differences in algal biomass and structure between the streams with the lowest and highest densities of caddisflies were much smaller than those between streams with the lowest and highest densities of snails. On day 32, the taxonomic and physiognomic structure of the algal assemblages in all the streams with caddisflies resembled that in the streams with higher densities of snails. Scanning electron micrographs showed that even at the highest densities, neither snails nor caddisflies could completely remove the algal assemblage. It is concluded that grazing can substantially influence algal growth form and assemblage physiognomy in lotic ecosystems.

Joint analysis of stressors and ecosystem services to enhance restoration effectiveness

With increasing pressure placed on natural systems by growing human populations, both scientists and resource managers need a better understanding of the relationships between cumulative stress from human activities and valued ecosystem services. Societies often seek to mitigate threats to these services through large-scale, costly restoration projects, such as the over one billion dollar Great Lakes Restoration Initiative currently underway. To help inform these efforts, we merged high-resolution spatial analyses of environmental stressors with mapping of ecosystem services for all five Great Lakes. Cumulative ecosystem stress is highest in near-shore habitats, but also extends offshore in Lakes Erie, Ontario, and Michigan. Variation in cumulative stress is driven largely by spatial concordance among multiple stressors, indicating the importance of considering all stressors when planning restoration activities. In addition, highly stressed areas reflect numerous different combinations of stressors rather than a single suite of problems, suggesting that a detailed understanding of the stressors needing alleviation could improve restoration planning. We also find that many important areas for fisheries and recreation are subject to high stress, indicating that ecosystem degradation could be threatening key services. Current restoration efforts have targeted high-stress sites almost exclusively, but generally without knowledge of the full range of stressors affecting these locations or differences among sites in service provisioning. Our results demonstrate that joint spatial analysis of stressors and ecosystem services can provide a critical foundation for maximizing social and ecological benefits from restoration investments.

Role of Nutrient Cycling and Herbivory in Regulating Periphyton Communities in Laboratory Streams

In this study we examined the role of nutrient cycling and herbivory in regulating stream periphyton communities. Population, community, and ecosystem—level properties were studied in laboratory stream channels that had nutrient inputs reduced compared to channels where ambient nutrient levels were maintained. We reduced nutrient inputs in four of eight channels by recirculating 90% of the flow, whereas the other four channels received once—through flow of spring water. We examined the interaction between herbivory and nutrients by varying the number of snails (Elimia clavaeformis) among streams with different nutrient input (circulation) regimes. Reduction in nutrient input viar recirculation resulted in lower concentrations of nutrients in the water but did not result in significant differences in biomass, carbon fixation, or algal taxonomic composition. However, herbivory had large effects on these characteristics by reducing biomass and areal rates of carbon fixation and simplifying periphyton taxonom...

Effects of three herbivores on periphyton communities in laboratory streams

The effects of grazing on algal assemblages by three different stream herbivores, the mayfly Centroptilum elsa, the snail Juga silicula, and the caddisfly Dicosmoecus gilvipes, were studied during a 48-d experiment in six laboratory streams. Compared with ungrazed control streams, grazing by Centroptilum (500/m2) modified algal community structure slightly but had little effect on periphyton biomass and chlorophyll a. Grazing by Juga (350/m2) reduced periphyton biomass and chlorophyll a by nearly 50%, but increased the rate of primary production by up to 25%. Juga also prevented significant accumulation of cyanophytes and some diatom species. Grazing by Dicosmoecus (200/m2) reduced periphyton biomass and chlorophyll a to less than 5% of the ungrazed levels, but primary production declined by only 50%. Only adnate algal cells and short filaments persisted on substrates grazed by Dicosmoecus. Algal export rates were increased by all three herbivores. Modification of algal growth patterns by both consumption and dislodgement, and dampening of temporal fluctuations were key mechanisms by which these herbivores altered periphyton communities. Primary production was stimulated by low rates of grazing by Juga in the laboratory streams, possibly as a result of increased light intensity in lower strata of the periphyton or removal of senescent algal cells. Algal assemblages displayed both community-level responses (e.g., biomass, production) and species-level responses (e.g., taxonomic composition) that should be considered in other studies of stream herbivory.

Effect of periphyton biomass on hydraulic characteristics and nutrient cycling in streams

The effect of periphyton biomass on hydraulic characteristics and nutrient cycling was studied in laboratory streams with and without snail herbivores. Hydraulic characteristics, such as average water velocity, dispersion coefficients, and relative volume of transient storage zones (zones of stationary water), were quantified by performing short-term injections of a conservative tracer and fitting an advection-dispersion model to the conservative tracer concentration profile downstream from the injection site. Nutrient cycling was quantified by measuring two indices: (1) uptake rate of phosphorus from stream water normalized to gross primary production (GPP), a surrogate measure of total P demand, and (2) turnover rate of phosphorus in the periphyton matrix. These measures indicate the importance of internal cycling (within the periphyton matrix) in meeting the P demands of periphyton. Dense growths of filamentous diatoms and blue-green algae accumulated in the streams with no snails (high-biomass streams), whereas the periphyton communities in streams with snails consisted almost entirely of a thin layer of basal cells of Stigeoclonium sp. (low-biomass streams). Dispersion coefficients were significantly greater and transient storage zones were significantly larger in the high-biomass streams compared to the low-biomass streams. Rates of GPP-normalized P uptake from water and rates of P turnover in periphyton were significantly lower in high biomass than in low biomass periphyton communities, suggesting that a greater fraction of the P demand was met by recycling in the high biomass communities. Increases in streamwater P concentration significantly increased GPP-normalized P uptake in high biomass communities, suggesting diffusion limitation of nutrient transfer from stream water to algal cells in these communities. Our results demonstrate that accumulations of periphyton biomass can alter the hydraulic characteristics of streams, particularly by increasing transient storage zones, and can increase internal nutrient cycling. They suggest a close coupling of hydraulic characteristics and nutrient cycling processes in stream ecosystems.

Complex interactions between autotrophs in shallow marine and freshwater ecosystems: implications for community responses to nutrient stress.

The relative biomass of autotrophs (vascular plants, macroalgae, microphytobenthos, phytoplankton) in shallow aquatic ecosystems is thought to be controlled by nutrient inputs and underwater irradiance. Widely accepted conceptual models indicate that this is the case both in marine and freshwater systems. In this paper we examine four case studies and test whether these models generally apply. We also identify other complex interactions among the autotrophs that may influence ecosystem response to cultural eutrophication. The marine case studies focus on macroalgae and its interactions with sediments and vascular plants. The freshwater case studies focus on interactions between phytoplankton, epiphyton, and benthic microalgae. In Waquoit Bay, MA (estuary), controlled experiments documented that blooms of macroalgae were responsible for the loss of eelgrass beds at nutrient-enriched locations. Macroalgae covered eelgrass and reduced irradiance to the extent that the plants could not maintain net growth. In Hog Island Bay, VA (estuary), a dense lawn of macroalgae covered the bottom sediments. There was reduced sediment-water nitrogen exchange when the algae were actively growing and high nitrogen release during algal senescence. In Lakes Brobo (West Africa) and Okeechobee (FL), there were dramatic seasonal changes in the biomass and phosphorus content of planktonic versus attached algae, and these changes were coupled with changes in water level and abiotic turbidity. Deeper water and/or greater turbidity favored dominance by phytoplankton. In Lake Brobo there also was evidence that phytoplankton growth was stimulated following a die-off of vascular plants. The case studies from Waquoit Bay and Lake Okeechobee support conceptual models of succession from vascular plants to benthic algae to phytoplankton along gradients of increasing nutrients and decreasing under-water irradiance. The case studies from Hog Island Bay and Lake Brobo illustrate additional effects (modified sediment-water nutrient fluxes, allelopathy or nutrient release during plant senescence) that could play a role in ecosystem response to nutrient stress.

Periphyton Function in Lake Ecosystems

Periphyton communities have received relatively little attention in lake ecosystems. However, evidence is increasing that they play a key role in primary productivity, nutrient cycling, and food web interactions. This review summarizes those findings and places them in a conceptual framework to evaluate the functional importance of periphyton in lakes. The role of periphyton is conceptualized based on a spatial hierarchy. At the coarsest scale, landscape properties such as lake morphometry, influence the amount of available habitat for periphyton growth. Watershed-related properties, such as loading of dissolved organic matter, nutrients, and sediments influence light availability and hence periphyton productivity. At the finer scale of within the lake, both habitat availability and habitat type affect periphyton growth and abundance. In addition, periphyton and phytoplankton compete for available resources at the within-lake scale. Our review indicates that periphyton plays an important functional role in lake nutrient cycles and food webs, especially under such conditions as relatively shallow depths, nutrient-poor conditions, or high water-column transparency. We recommend more studies assessing periphyton function across a spectrum of lake morphometry and trophic conditions.

Research in Artificial Streams: Applications, Uses, and Abuses

Increased use of artificial streams in aquatic research over the last 20 years has not been accompanied by a systematic, critical analysis of their advantages and disadvantages. A symposium held at the 1992 annual meeting of the North American Benthological Society in Louisville, Kentucky, attempted specifically to provide this information. We define an artificial stream as any constructed channel that has a controlled flow of water and that is used to study a physical, chemical, or biological property of natural streams. The following aspects of artificial streams were covered in the symposium: historical perspectives, hydrodynamics, algal-nutrient dynamics, macroinvertebrate growth, grazer-algal interactions, fish ecology, disturbance, ecotoxicology, small- and large-scale artificial streams, and longitudinal linkages. Although the symposium addressed a wide variety of subjects, each contribution was linked by a common desire to ascertain the strengths and weaknesses of artificial streams relative to that subject. Major conclusions that emerged from the symposium include: (1) there is no single best design for artificial streams; appropriate stream design is contingent on the question being asked; (2) research geared to mechanistic understanding of lotic processes is particularly well-suited for artificial streams; and (3) generation of testable hypotheses, which can then be validated in natural stream ecosystems, is a useful application of research in artificial streams.