N. LeRoy Poff

The Natural Flow Regime

H umans have long been fascinated by the dynamism of free-flowing waters. Yet we have expended great effort to tame rivers for transportation, water supply, flood control, agriculture, and power generation. It is now recognized that harnessing of streams and rivers comes at great cost: Many rivers no longer support socially valued native species or sustain healthy ecosystems that provide important goods and services (Naiman et al. 1995, NRC 1992).

Functional organization of stream fish assemblages in relation to hydrological variability

Stream fish assmemblage data for 34 sites in Wisconsin and Minnesota were obtained from archived sources and were used in conjunction with long—term hydrological data to test the hypothesis that functional organization of fish communities is related to hydrological variability. For each of the 106 species present in the data set, six categories of species traits were derived to describe habitat, trophic, morphologica, and tolerance characteristics. A hierarchical clustering routine was used to identify two functionally similar groups of assemblages defined in terms of species presence/absence. Hydrological factors describing streamflow variability and predictability, as well as frequency and predictability of high flow and low flow extremes, were derived for each of the 34 sites and employed to explain differences among the functionally defined groups. Canonical discriminant analysis revealed that the hydrological data could clearly separate the two ecologically defined groups of assemblages, which were associated with either hydrologically variable streams (high coefficient of variation of daily flows, moderate frequency of spates) or hydrologically stable streams (high predictability of daily flows, stable baseflow conditions). Discriminant functions based on hydrological information classified the 34 fish assemblages into the correct ecological group with 85% accuracy. Assemblages from hydrologically variable sites had generalized feeding strategies, were associated with silt and general substrata, were characterized by slow—velocity species with headwater affinities, and were tolerant to silt. Proportions of species traits present at the 34 sites were regressed against an index of hydrological stability derived from a principal components analysis to test the hypothesis that functional organization of assemblages varied across a gradient of hydrological stability. Results were complementary with the discriminant analysis. Findings were in general agreement with theoretical predictions that variable should support resource generalists while stable habitats should be characterized by a higher proportion of specialist species. Several species of fish were identified as indicative of the variable—stable hydrological gradient among stream sites. A taxonomic analysis showed strong geographic patterns in species composition of the 34 assemblages. However, zoogeographic constraints did not explain the observed relationship between stream hydrology and functional organization of fish assemblages. The strong hydrological—assemblage relations found in the 34 midwestern sites suggest that hydrological factors are significant environmental variables influencing fish assemblage structure, and that hydrological alterations induced by climate change (or other anthropogenic disturbances) could modify stream fish assemblages structure in this region.

Ecological Thresholds: The Key to Successful Environmental Management or an Important Concept with No Practical Application?

An ecological threshold is the point at which there is an abrupt change in an ecosystem quality, property or phenomenon, or where small changes in an environmental driver produce large responses in the ecosystem. Analysis of thresholds is complicated by nonlinear dynamics and by multiple factor controls that operate at diverse spatial and temporal scales. These complexities have challenged the use and utility of threshold concepts in environmental management despite great concern about preventing dramatic state changes in valued ecosystems, the need for determining critical pollutant loads and the ubiquity of other threshold-based environmental problems. In this paper we define the scope of the thresholds concept in ecological science and discuss methods for identifying and investigating thresholds using a variety of examples from terrestrial and aquatic environments, at ecosystem, landscape and regional scales. We end with a discussion of key research needs in this area.

Homogenization of regional river dynamics by dams and global biodiversity implications.

Global biodiversity in river and riparian ecosystems is generated and maintained by geographic variation in stream processes and fluvial disturbance regimes, which largely reflect regional differences in climate and geology. Extensive construction of dams by humans has greatly dampened the seasonal and interannual streamflow variability of rivers, thereby altering natural dynamics in ecologically important flows on continental to global scales. The cumulative effects of modification to regional-scale environmental templates caused by dams is largely unexplored but of critical conservation importance. Here, we use 186 long-term streamflow records on intermediate-sized rivers across the continental United States to show that dams have homogenized the flow regimes on third- through seventh-order rivers in 16 historically distinctive hydrologic regions over the course of the 20th century. This regional homogenization occurs chiefly through modification of the magnitude and timing of ecologically critical high and low flows. For 317 undammed reference rivers, no evidence for homogenization was found, despite documented changes in regional precipitation over this period. With an estimated average density of one dam every 48 km of third- through seventh-order river channel in the United States, dams arguably have a continental scale effect of homogenizing regionally distinct environmental templates, thereby creating conditions that favor the spread of cosmopolitan, nonindigenous species at the expense of locally adapted native biota. Quantitative analyses such as ours provide the basis for conservation and management actions aimed at restoring and maintaining native biodiversity and ecosystem function and resilience for regionally distinct ecosystems at continental to global scales.

The Network Dynamics Hypothesis: How Channel Networks Structure Riverine Habitats

Abstract Hierarchical and branching river networks interact with dynamic watershed disturbances, such as fires, storms, and floods, to impose a spatial and temporal organization on the nonuniform distribution of riverine habitats, with consequences for biological diversity and productivity. Abrupt changes in water and sediment flux occur at channel confluences in river networks and trigger changes in channel and floodplain morphology. This observation, when taken in the context of a river network as a population of channels and their confluences, allows the development of testable predictions about how basin size, basin shape, drainage density, and network geometry interact to regulate the spatial distribution of physical diversity in channel and riparian attributes throughout a river basin. The spatial structure of river networks also regulates how stochastic watershed disturbances influence the morphology and ages of fluvial features found at confluences.

How dams vary and why it matters for the emerging science of dam removal

D are structures designed by humans to capture water and modify the magnitude and timing of its movement downstream. The damming of streams and rivers has been integral to human population growth and technological innovation. Among other things, dams have reduced flood hazard and allowed humans to settle and farm productive alluvial soils on river floodplains; they have harnessed the power of moving water for commerce and industry; and they have created reservoirs to augment the supply of water during periods of drought. In the 5000 or so years that humans have been building dams, millions have been constructed globally, especially in the last 100 years (Smith 1971, WCD 2000). If dams have successfully met so many human needs, why is there a growing call for their removal? The answers to this question require an appreciation of society’s changing needs for, and concerns about, dams, including the emerging recognition that dams can impair river ecosystems (Babbit 2002). But decisions about dam removal are complex, in no small part because great scientific uncertainty exists over the potential environmental benefits of dam removal. Certainly, the scarcity of empirical knowledge on environmental responses to dam removal contributes to this uncertainty (Hart et al. 2002). More fundamentally, however, a scientific framework is lacking for considering how the tremendous variation in dam and river attributes determines the ecological impacts of dams and the restoration potential following removal. Such an ecological classification of dams is ultimately needed to support the emerging science of dam removal. In this article, we develop a conceptual foundation for the emerging science of dam removal by (a) reviewing the ways that dams impair river ecosystems, (b) examining criteria used to classify dams and describing how these criteria are of limited value in evaluating the environmental effects of dams, (c) quantifying patterns of variation in some environmentally relevant dam characteristics using governmental databases, (d) specifying a framework that can guide the development of an ecological classification of dams, and (e) evaluating the ways that dam characteristics affect removal decisions and the future of dam removals. We restrict our analysis to the United States, where dam removals are currently hotly debated; however, the ecological framework we advocate could also be generalized to other parts of the world.

Physical habitat template of lotic systems : Recovery in the context of historical pattern of spatiotemporal heterogeneity

Spatial and temporal environmental heterogeneity in lotic ecosystems can be quantitatively described and identified with characteristic levels of ecological organization. The long-term pattern of physicochemical variability in conjunction with the complexity and stability of the substratum establishes a physical habitat template that theoretically influences which combinations of behavioral, physiological and life history characteristics constitute appropriate “ecological strategies” for persistence in the habitat. The combination of strategies employed will constrain ecological response to and recovery from disturbance. Physical habitat templates and associated ecological attributes differ geographically because of biogeoclimatic processes that constrain lotic habitat structure and stability and that influence physicochemical variability and disturbance patterns (frequency, magnitude, and predictability). Theoretical considerations and empirical studies suggest that recovery from natural and anthropogenic disturbance also will vary among lotic systems, depending on historical temporal variability regime, degree of habitat heterogeneity, and spatial scale of the perturbation. Characterization of physical habitat templates and associated ecological dynamics along gradients of natural disturbance would provide a geographic framework for predicting recovery from anthropogenic disturbance for individual streams. Description of lotic environmental templates at the appropriate spatial and temporal scale is therefore desirable to test theoretical expectations of biotic recovery rate from disturbance and to guide selection of appropriate reference study sites for monitoring impacts of anthropogenic disturbance. Historical streamflow data, coupled with stream-specific thermal and substratum-geomorphologic characteristics, are suggested as minimum elements needed to characterize physical templates of lotic systems.

Functional trait niches of North American lotic insects: traits-based ecological applications in light of phylogenetic relationships

Abstract The use of species traits to characterize the functional composition of benthic invertebrate communities has become well established in the ecological literature. This approach holds much potential for predicting changes of both species and species assemblages along environmental gradients in terms of traits that are sensitive to local environmental conditions. Further, in the burgeoning field of biomonitoring, a functional approach provides a predictive basis for understanding community-level responses along gradients of environmental alteration caused by humans. Despite much progress in recent years, the full potential of the functional traits-based approach is currently limited by several factors, both conceptual and methodological. Most notably, we lack adequate understanding of how individual traits are intercorrelated and how this lack of independence among traits reflects phylogenetic (evolutionary) constraint. A better understanding is needed if we are to make the transition from a largely univariate approach that considers single-trait responses along single environmental gradients to a multivariate one that more realistically accounts for the responses of many traits across multiple environmental gradients characteristic of most human-dominated landscapes. Our primary objective in this paper is to explore the issue of inter-trait correlations for lotic insects and to identify opportunities and challenges for advancing the theory and application of traits-based approaches in stream community ecology. We created a new database on species-trait composition of North American lotic insects. Using published accounts and expert opinion, we collected information on 20 species traits (in 59 trait states) that fell into 4 broad categories: life-history, morphological, mobility, and ecological. First, we demonstrate the importance of considering how the linkage of specific trait states within a taxon is critical to developing a more-robust traits-based community ecology. Second, we examine the statistical correlations among traits and trait states for the 311 taxa to identify trait syndromes and specify which traits provide unique (uncorrelated) information that can be used to guide trait selection in ecological studies. Third, we examine the evolutionary associations among traits by mapping trait states onto a phylogentic tree derived from morphological and molecular analyses and classifications from the literature. We examine the evolutionary lability of individual traits by assessing the extent to which they are unconstrained by phylogenic relationships across the taxa. By focusing on the lability of traits within lotic genera of Ephemeroptera, Plecoptera, and Trichoptera, taxa often used as water-quality indicators, we show how a traits-based approach can allow a priori expectations of the differential response of these taxa to specific environmental gradients. We conclude with some ideas about how specific trait linkages, statistical correlations among traits, and evolutionary lability of traits can be used in combination with a mechanistic understanding of trait response along environmental gradients to select robust traits useful for a more predictive community ecology. We indicate how these new insights can direct the research in statistical modeling that is necessary to achieve the full potential of models that can predict how multiple traits will respond along multiple environmental gradients.

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.

River flows and water wars: emerging science for environmental decision making

Real and apparent conflicts between ecosystem and human needs for fresh water are contributing to the emergence of an alternative model for conducting river science around the world. The core of this new paradigm emphasizes the need to forge new partnerships between scientists and other stakeholders where shared ecological goals and river visions are developed, and the need for new experimental approaches to advance scientific understanding at the scales relevant to whole-river management. We identify four key elements required to make this model succeed: existing and planned water projects represent opportunities to conduct ecosystem-scale experiments through controlled river flow manipulations; more cooperative interactions among scientists, managers, and other stakeholders are critical; experimental results must be synthesized across studies to allow broader generalization; and new, innovative funding partnerships are needed to engage scientists and to broadly involve the government, the private sector, and NGOs.