Differing Modes of Biotic Connectivity within Freshwater Ecosystem Mosaics

We describe a collection of aquatic and wetland habitats in an inland landscape, and their occurrence within a terrestrial matrix, as a “freshwater ecosystem mosaic” (FEM). Aquatic and wetland habitats in any FEM can vary widely, from permanently ponded lakes, to ephemerally ponded wetlands, to groundwater‐fed springs, to flowing rivers and streams. The terrestrial matrix can also vary, including in its influence on flows of energy, materials, and organisms among ecosystems. Biota occurring in a specific region are adapted to the unique opportunities and challenges presented by spatial and temporal patterns of habitat types inherent to each FEM. To persist in any given landscape, most species move to recolonize habitats and maintain mixtures of genetic materials. Species also connect habitats through time if they possess needed morphological, physiological, or behavioral traits to persist in a habitat through periods of unfavorable environmental conditions. By examining key spatial and temporal patterns underlying FEMs, and species‐specific adaptations to these patterns, a better understanding of the structural and functional connectivity of a landscape can be obtained. Fully including aquatic, wetland, and terrestrial habitats in FEMs facilitates adoption of the next generation of individual‐based models that integrate the principles of population, community, and ecosystem ecology.     

Biota Connect Aquatic Habitats throughout Freshwater Ecosystem Mosaics

Freshwater ecosystems are linked at various spatial and temporal scales by movements of biota adapted to life in water. We review the literature on movements of aquatic organisms that connect different types of freshwater habitats, focusing on linkages from streams and wetlands to downstream waters. Here, streams, wetlands, rivers, lakes, ponds, and other freshwater habitats are viewed as dynamic freshwater ecosystem mosaics (FEMs) that collectively provide the resources needed to sustain aquatic life. Based on existing evidence, it is clear that biotic linkages throughout FEMs have important consequences for biological integrity and biodiversity. All aquatic organisms move within and among FEM components, but differ in the mode, frequency, distance, and timing of their movements. These movements allow biota to recolonize habitats, avoid inbreeding, escape stressors, locate mates, and acquire resources. Cumulatively, these individual movements connect populations within and among FEMs and contribute to local and regional diversity, resilience to disturbance, and persistence of aquatic species in the face of environmental change. Thus, the biological connections established by movement of biota among streams, wetlands, and downstream waters are critical to the ecological integrity of these systems. Future research will help advance our understanding of the movements that link FEMs and their cumulative effects on downstream waters.     

Safeguarding Wetlands and Their Connections within Wetlandscapes to Improve Conservation Outcomes for Threatened Amphibian Species

Wetlands should not be considered as independent objects but as dynamically connected objects, collectively known as wetlandscapes. We developed a framework that analyzes the influences of wetland suitability and connectivity on amphibian distributions. We defined two indices: a Wetland Suitability Index describing wetland quality and a Movement Permeability Index characterizing wetland connectivity for amphibian population dynamics. These indices were calculated from raster datasets and time‐varying inundation estimates. The indices were used to define a wetlandscape and an amphibian model was used to simulate population dynamics within the wetlandscape. The framework was applied to the Nose Creek watershed, a highly modified wetlandscape in Alberta, Canada. Two amphibian species were selected with different habitat preferences: the Northern Leopard Frog that prefers wet habitats and has high mobility over land, and the Great Plains Toad that prefers terrestrial habitats and has low mobility over land. We found each amphibian species had a “preferred” wetlandscape, reflecting their life cycle traits and migration strategies which in turn were dependent on the hydrological and ecological connections within the wetlandscape. This study highlights the importance of investigating both individual wetlands and the wetlandscape and considering both wetland habitat quality and connectivity as non‐substitutable properties that act jointly, but differently, on population dynamics.     

Hydrologic Connectivity and the Contribution of Stream Headwaters to Ecological Integrity at Regional Scales1

Abstract: Cumulatively, headwater streams contribute to maintaining hydrologic connectivity and ecosystem integrity at regional scales. Hydrologic connectivity is the water‐mediated transport of matter, energy and organisms within or between elements of the hydrologic cycle. Headwater streams compose over two‐thirds of total stream length in a typical river drainage and directly connect the upland and riparian landscape to the rest of the stream ecosystem. Altering headwater streams, e.g., by channelization, diversion through pipes, impoundment and burial, modifies fluxes between uplands and downstream river segments and eliminates distinctive habitats. The large‐scale ecological effects of altering headwaters are amplified by land uses that alter runoff and nutrient loads to streams, and by widespread dam construction on larger rivers (which frequently leaves free‐flowing upstream portions of river systems essential to sustaining aquatic biodiversity). We discuss three examples of large‐scale consequences of cumulative headwater alteration. Downstream eutrophication and coastal hypoxia result, in part, from agricultural practices that alter headwaters and wetlands while increasing nutrient runoff. Extensive headwater alteration is also expected to lower secondary productivity of river systems by reducing stream‐system length and trophic subsidies to downstream river segments, affecting aquatic communities and terrestrial wildlife that utilize aquatic resources. Reduced viability of freshwater biota may occur with cumulative headwater alteration, including for species that occupy a range of stream sizes but for which headwater streams diversify the network of interconnected populations or enhance survival for particular life stages. Developing a more predictive understanding of ecological patterns that may emerge on regional scales as a result of headwater alterations will require studies focused on components and pathways that connect headwaters to river, coastal and terrestrial ecosystems. Linkages between headwaters and downstream ecosystems cannot be discounted when addressing large‐scale issues such as hypoxia in the Gulf of Mexico and global losses of biodiversity.     

Biological Responses to Contrasting Hydrology in Backwaters of Upper Mississippi River Navigation Pool 25

Water level management in Mississippi River Pool 25 differentially influences off-channel habitats in the mid-pool and lower pool. Hydrologic models indicate lower pool off-channel habitats dry with greater frequency and duration compared to similar habitats at mid-pool. We examined the influence of this contrasting hydrology on substrate characteristics, organic matter, macroinvertebrate, and fish communities in off-channel habitats during 2001–2003. Benthic organic matter standing stocks were stable in mid-pool habitats but lower pool values were variable because of annual differences in moist-soil vegetation production. Generally, small-bodied and multivoltine invertebrate taxa had high community biomass and dominated lower pool habitats, whereas longer-lived and large-bodied taxa were more abundant and had higher community biomass in mid-pool habitats having longer hydroperiods. Fish communities were dominated by cyprinids in both habitats, and mid-pool habitats tended to be higher in overall species richness. Unique fish taxa were collected in each pool, with primarily rheophilic forms in mid-pool habitats and limnophilic forms in lower pool habitats. Results indicate that contrasting hydrology associated with a mid-pool control point directly and indirectly influences biological communities in off-channel habitats. Further, management regimes that promote hydrologic diversity in off-channel habitats may enhance biological diversity at larger spatial and temporal scales.     

Land use in Korean tidal wetlands: impacts and management strategies

The coastal landscapes in southwestern Korea include a diverse array of tidal wetlands and salt marshes. These coastal zones link the ecological functions of marine tidal wetlands and freshwater ecosystems with terrestrial ecosystems. They are rich in biological diversity and play important roles in sustaining ecological health and processing environmental pollutants. Korean tidal wetlands are particularly important as nurseries for economically important fishes and habitats for migratory birds. Diking, draining, tourism, and conversion to agricultural and urban uses have adversely affected Korean tidal wetlands. Recent large development projects have contributed to further losses. Environmental impact assessments conducted for projects affecting tidal wetlands and their surrounding landscapes should be customized for application to these special settings. Adequate environmental impact assessments will include classification of hydrogeomorphic units and consideration of their responses to biological and environmental stressors. As is true worldwide, Korean laws and regulations are changing to be more favorable to the conservation and protection of tidal wetlands. More public education needs to be done at the local level to build support for tidal wetland conservation. Some key public education points include the role of tidal wetlands in maintaining healthy fish populations and reducing impacts of nonpoint source pollution. There is also a need to develop procedures for integrating economic and environmental objectives within the overall context of sustainable management and land uses.     

Featured Collection Introduction: Connectivity of Streams and Wetlands to Downstream Waters

Connectivity is a fundamental but highly dynamic property of watersheds. Variability in the types and degrees of aquatic ecosystem connectivity presents challenges for researchers and managers seeking to accurately quantify its effects on critical hydrologic, biogeochemical, and biological processes. However, protecting natural gradients of connectivity is key to protecting the range of ecosystem services that aquatic ecosystems provide. In this featured collection, we review the available evidence on connections and functions by which streams and wetlands affect the integrity of downstream waters such as large rivers, lakes, reservoirs, and estuaries. The reviews in this collection focus on the types of waters whose protections under the U.S. Clean Water Act have been called into question by U.S. Supreme Court cases. We synthesize 40+ years of research on longitudinal, lateral, and vertical fluxes of energy, material, and biota between aquatic ecosystems included within the Act's frame of reference. Many questions about the roles of streams and wetlands in sustaining downstream water integrity can be answered from currently available literature, and emerging research is rapidly closing data gaps with exciting new insights into aquatic connectivity and function at local, watershed, and regional scales. Synthesis of foundational and emerging research is needed to support science‐based efforts to provide safe, reliable sources of fresh water for present and future generations.     

Application of ecological theory to determining recovery potential of disturbed lotic ecosystems: Research needs and priorities

This article summarizes the views of aquatic scientists who gathered to assess the ability of stream ecosystem theory to predict recovery from disturbance. Two views of disturbance were evident: a discrete removal of organisms vs an unusual deviation from normal. These were perceived as applying to different scales and/or objectives. Long-term information is required from both points of view to define recovery. Recovery also may be defined in different ways, but it is clear that recovery has both spatial and temporal components, and includes both physical and biological processes. Consensus was very strong that a major role (and challenge) for theory lies in the understanding of spatial aspects, temporal scales, coupling of physics and biology, and the interaction of these features in recovery processes. Some progress is evident in the articles of this volume, but among the topics identified as critical for further theoretical contributions were: homogeneous vs heterogeneous distribution of disturbance, local extent of disturbance relative to a regional context, critical vs noncritical patches (size and location) of disturbance at different spatial scales and temporal frequencies, delineation of reversible and nonreversible processes, and physical and biological constraints on the time frame for recovery. Such concepts need attention across different types of lotic ecosystems. Thus, there was strong consensus that a national monitoring system of representative lotic ecosystems within ecological regions be established. The purpose of this monitoring system would be to acquire long-term data on natural variability, to establish viable indicators of spatial and temporal aspects of recovery, and to develop and test emerging theoretical developments.     

Using Fly Ash as a Marker to Quantify Culturally‐Accelerated Sediment Accumulation in Playa Wetlands

Wetlands in the Rainwater Basin in Nebraska are vulnerable to sediment accumulation from the surrounding watershed. Sediment accumulation has a negative impact on wetland quality by decreasing the depth and volume of water stored, and the plant community species composition and density growing in the wetland. The objective of this study was to determine the amount of sediment that has accumulated in five selected wetlands in the Rainwater Basin in Nebraska. Soil cores were taken at five or six locations along transects across each wetland. This study used the fly ash, which is generated by coal‐burning locomotives that were present generally in the late 1800s and early 1900s, as a marker to quantify the sediment deposition rates. The cores were divided into 5 cm sections and the soils were analyzed using a fly ash extraction and identification technique. Results indicate that the average depth of sediment ranged from 23.00 to 38.00 cm. The annual average depth of sediment accumulation ranged from 0.18 cm/yr to 0.29 cm/yr. The annual sediment accumulation rate from both wind erosion and water erosion in these five sampling wetlands was between 1.946 and 3.225 kg/m²/yr. The results of this research can be used to develop restoration plans for wetlands. The fly ash testing technology can also be applied to other areas with the railroads across the United States.     

A landscape ecological approach to address scaling problems in conservation management and monitoring

Management of many African game reserves is today often still an art based on experience and intuition, rather than a science. Decision-making is based on an informal integration of accumulated individual knowledge and keen field observations. Data are generally poorly captured and curated. Until fairly recently, denominators of biological parameters (such as the unit of land or unit of plant production used as measurement) have generally been treated as being homogenous. The patchiness of landscapes and the issue of ecological scaling were ignored, often because of a lack of appropriate technical tools. The ecological data available on the 49,000-ha Songimvelo Game Reserve (SGR) result from a number of discrete survey and monitoring projects undertaken by different researchers, with different objectives, at different spatial and temporal scales. A landscape ecological approach towards research and monitoring is appropriate for an area of the size and diversity of the SGR. A combination of a database approach and spatial representation was used to consolidate and integrate data across temporal and spatial scales. Herbivore spatial and temporal distribution patterns were explored across three spatial scales. An understanding was achieved of the importance of landscape patchiness in controlling resource availability for herbivores. This insight is important in guiding management and monitoring of the SGR by placing perceived patch overutilization in its proper landscape context. The landscape ecological approach bridges the traditional scale-independent view to a more contemporary scale-related understanding of ecosystem diversity and functioning.     

Anthropogenic effects on the biodiversity of riparian wetlands of a northern temperate landscape

Land uses such as forestry and agriculture are presumed to degrade the biodiversity of riparian wetlands in the northern temperate regions of the United States. In order to improve land use decision making in this landscape, floral and faunal communities of 15 riparian wetlands associated with low-order streams were related to their surrounding land cover to establish which organismal groups are affected by anthropogenic disturbance and whether these impacts are scale-specific. Study sites were chosen to represent a gradient of disturbance. Vascular plants of wet meadow and shrub carr communities, aquatic macro-invertebrates, amphibians, fish and birds were surveyed, and total abundance, species richness and Shannon diversity were calculated. For each site, anthropogenic disturbances were evaluated at local and landscape scales (500, 1000, 2500 and 5000 m from the site and the site catchment) from field surveys and a geographic information system (GIS). Land use data were grouped into six general land use types: urban, cultivated, rangeland, forest, wetland and water. Shrub carr vegetation, bird and fish diversity and richness generally decrease with increasing cultivation in the landscape. Amphibian abundance decreases and fish abundance increases as the proportions of open water and rangeland increases; bird diversity and richness increase with forest and wetland extent in the landscape. Wet meadow vegetation, aquatic macro-invertebrates, amphibians and fish respond to local disturbances or environmental conditions. Shrub carr vegetation, amphibians and birds are influenced by land use at relatively small landscape scales (500 and 1000 m), and fish respond to land use at larger landscape scales (2500, 5000 m and the catchment). Effective conservation planning for these riparian wetlands requires assessment of multiple organismal groups, different types of disturbance and several spatial scales.1998 Academic Press     

Spatial and Temporal Patterns in Fish Assemblages Following an Artificially Extended Floodplain Inundation Event, Northern Murray-Darling Basin, Australia

Water extraction from dryland rivers is often associated with declines in the health of river and floodplain ecosystems due to reduced flooding frequency and extent of floodplain inundation. Following moderate flooding in early 2008 in the Narran River, Murray-Darling Basin, Australia, 10,423 ML of water was purchased from agricultural water users and delivered to the river to prolong inundation of its terminal lake system to improve the recruitment success of colonial waterbirds that had started breeding in response to the initial flooding. This study examined the spatial and temporal patterns of fish assemblages in river and floodplain habitats over eight months following flooding to assess the possible ecological benefits of flood extension. Although the abundances of most fish species were greater in river channel habitats, the fish assemblage used floodplain habitats when inundated. Young-of-the-year (4–12 months age) golden perch (Macquaria ambigua) and bony bream (Nematalosa erebi) were consistently sampled in floodplain sites when inundated, suggesting that the floodplain provides rearing habitat for these species. Significant differences in the abundances of fish populations between reaches upstream and downstream of a weir in the main river channel indicates that the effectiveness of the environmental water release was limited by restricted connectivity within the broader catchment. Although the seasonal timing of flood extension may have coincided with sub-optimal primary production, the use of the environmental water purchase is likely to have promoted recruitment of fish populations by providing greater access to floodplain nursery habitats, thereby improving the ability to persist during years of little or no flow.     

Effects of Habitat Disturbance from Residential Development on Breeding Bird Communities in Riparian Corridors

This study assessed the relationship among land use, riparian vegetation, and avian populations at two spatial scales. Our objective was to compare the vegetated habitat in riparian corridors with breeding bird guilds in eight Rhode Island subwatersheds along a range of increasing residential land use. Riparian habitats were characterized with fine-scale techniques (used field transects to measure riparian vegetation structure and plant species richness) at the reach spatial scale, and with coarse-scale landscape techniques (a Geographic Information System to document land-cover attributes) at the subwatershed scale. Bird surveys were conducted in the riparian zone, and the observed bird species were separated into guilds based on tolerance to human disturbance, habitat preference, foraging type, and diet preference. Bird guilds were correlated with riparian vegetation metrics, percent impervious surface, and percent residential land use, revealing patterns of breeding bird distribution. The number of intolerant species predominated below 12% residential development and 3% impervious surface, whereas tolerant species predominated above these levels. Habitat guilds of edge, forest, and wetland bird species correlated with riparian vegetation. This study showed that the application of avian guilds at both stream reach and subwatershed scales offers a comprehensive assessment of effects from disturbed habitat, but that the subwatershed scale is a more efficient method of evaluation for environmental management.     

Recent Land Cover History and Nutrient Retention in Riparian Wetlands

Wetland ecosystems are profoundly affected by altered nutrient and sediment loads received from anthropogenic activity in their surrounding watersheds. Our objective was to compare a gradient of agricultural and urban land cover history during the period from 1949 to 1997, with plant and soil nutrient concentrations in, and sediment deposition to, riparian wetlands in a rapidly urbanizing landscape. We observed that recent agricultural land cover was associated with increases in Nitrogen (N) and Phosphorus (P) concentrations in a native wetland plant species. Conversely, recent urban land cover appeared to alter receiving wetland environmental conditions by increasing the relative availability of P versus N, as reflected in an invasive, but not a native, plant species. In addition, increases in surface soil Fe content suggests recent inputs of terrestrial sediments associated specifically with increasing urban land cover. The observed correlation between urban land cover and riparian wetland plant tissue and surface soil nutrient concentrations and sediment deposition, suggest that urbanization specifically enhances the suitability of riparian wetland habitats for the invasive species Japanese stiltgrass [Microstegium vimenium (Trinius) A. Camus].     

Predicting Fish Growth Potential and Identifying Water Quality Constraints: A Spatially-Explicit Bioenergetics Approach

Anthropogenic impairment of water bodies represents a global environmental concern, yet few attempts have successfully linked fish performance to thermal habitat suitability and fewer have distinguished co-varying water quality constraints. We interfaced fish bioenergetics, field measurements, and Thermal Remote Imaging to generate a spatially-explicit, high-resolution surface of fish growth potential, and next employed a structured hypothesis to detect relationships among measures of fish performance and co-varying water quality constraints. Our thermal surface of fish performance captured the amount and spatial-temporal arrangement of thermally-suitable habitat for three focal species in an extremely heterogeneous reservoir, but interpretation of this pattern was initially confounded by seasonal covariation of water residence time and water quality. Subsequent path analysis revealed that in terms of seasonal patterns in growth potential, catfish and walleye responded to temperature, positively and negatively, respectively; crappie and walleye responded to eutrophy (negatively). At the high eutrophy levels observed in this system, some desired fishes appear to suffer from excessive cultural eutrophication within the context of elevated temperatures whereas others appear to be largely unaffected or even enhanced. Our overall findings do not lead to the conclusion that this system is degraded by pollution; however, they do highlight the need to use a sensitive focal species in the process of determining allowable nutrient loading and as integrators of habitat suitability across multiple spatial and temporal scales. We provide an integrated approach useful for quantifying fish growth potential and identifying water quality constraints on fish performance at spatial scales appropriate for whole-system management.     

Building spectral libraries for wetlands land cover classification and hyperspectral remote sensing

Recent advances in remote sensing provide opportunities to map plant species and vegetation within wetlands at management relevant scales and resolutions. Hyperspectral imagers, currently available on airborne platforms, provide increased spectral resolution over existing space-based sensors that can document detailed information on the distribution of vegetation community types, and sometimes species. Development of spectral libraries of wetland species is a key component needed to facilitate advanced analytical techniques to monitor wetlands. Canopy and leaf spectra at five sites in California, Texas, and Mississippi were sampled to create a common spectral library for mapping wetlands from remotely sensed data. An extensive library of spectra (n=1336) for coastal wetland communities, across a range of bioclimatic, edaphic, and disturbance conditions were measured. The wetland spectral libraries were used to classify and delineate vegetation at a separate location, the Pacheco Creek wetland in the Sacramento Delta, California, using a PROBE-1 airborne hyperspectral data set (5m pixel resolution, 128 bands). This study discusses sampling and collection methodologies for building libraries, and illustrates the potential of advanced sensors to map wetland composition. The importance of developing comprehensive wetland spectral libraries, across diverse ecosystems is highlighted. In tandem with improved analytical tools these libraries provide a physical basis for interpretation that is less subject to conditions of specific data sets. To facilitate a global approach to the application of hyperspectral imagers to mapping wetlands, we suggest that criteria for and compilation of wetland spectral libraries should proceed today in anticipation of the wider availability and eventual space-based deployment of advanced hyperspectral high spatial resolution sensors.     

Effects of spatial and temporal scales on cultural services valuation

The monetary valuation of ecosystem services has proved to be a powerful tool for influencing management decisions, as it can be used to quantify the relative importance of various ecosystem functions. However, these valuation methods are rarely implemented with adequate care; in particular, they do not consider spatial and temporal variations in the services offered and demanded. This paper examines the effect of heterogeneity on the valuation of cultural services offered by the Doñana natural protected area in Spain. We apply the zonal travel cost method at various spatial scales, and the individual travel cost method at various temporal scales. We conclude that economic valuation techniques must account for spatial and temporal heterogeneity if they are to provide accurate and realistic information.     

Modeling the stream water nitrate dynamics in a 60,000-km2 European catchment, the Garonne, southwest France

The spatial and temporal dynamics in the stream water NO(3)-N concentrations in a major European river-system, the Garonne (62,700 km(2)), are described and related to variations in climate, land management, and effluent point-sources using multivariate statistics. Building on this, the Hydrologiska Byr?ns Vattenbalansavdelning (HBV) rainfall-runoff model and the Integrated Catchment Model of Nitrogen (INCA-N) are applied to simulate the observed flow and N dynamics. This is done to help us to understand which factors and processes control the flow and N dynamics in different climate zones and to assess the relative inputs from diffuse and point sources across the catchment. This is the first application of the linked HBV and INCA-N models to a major European river system commensurate with the largest basins to be managed under the Water Framework Directive. The simulations suggest that in the lowlands, seasonal patterns in the stream water NO(3)-N concentrations emerge and are dominated by diffuse agricultural inputs, with an estimated 75% of the river load in the lowlands derived from arable farming. The results confirm earlier European catchment studies. Namely, current semi-distributed catchment-scale dynamic models, which integrate variations in land cover, climate, and a simple representation of the terrestrial and in-stream N cycle, are able to simulate seasonal NO(3)-N patterns at large spatial (>300 km(2)) and temporal (> or = monthly) scales using available national datasets.     

Marine Habitat Classification for Ecosystem-Based Management: A Proposed Hierarchical Framework

Creating a habitat classification and mapping system for marine and coastal ecosystems is a daunting challenge due to the complex array of habitats that shift on various spatial and temporal scales. To meet this challenge, several countries have, or are developing, national classification systems and mapping protocols for marine habitats. To be effectively applied by scientists and managers it is essential that classification systems be comprehensive and incorporate pertinent physical, geological, biological, and anthropogenic habitat characteristics. Current systems tend to provide over-simplified conceptual structures that do not capture biological habitat complexity, marginalize anthropogenic features, and remain largely untested at finer scales. We propose a multi-scale hierarchical framework with a particular focus on finer scale habitat classification levels and conceptual schematics to guide habitat studies and management decisions. A case study using published data is included to compare the proposed framework with existing schemes. The example demonstrates how the proposed framework’s inclusion of user-defined variables, a combined top-down and bottom-up approach, and multi-scale hierarchical organization can facilitate examination of marine habitats and inform management decisions.     

The Contribution of Headwater Streams to Biodiversity in River Networks1

Abstract: The diversity of life in headwater streams (intermittent, first and second order) contributes to the biodiversity of a river system and its riparian network. Small streams differ widely in physical, chemical, and biotic attributes, thus providing habitats for a range of unique species. Headwater species include permanent residents as well as migrants that travel to headwaters at particular seasons or life stages. Movement by migrants links headwaters with downstream and terrestrial ecosystems, as do exports such as emerging and drifting insects. We review the diversity of taxa dependent on headwaters. Exemplifying this diversity are three unmapped headwaters that support over 290 taxa. Even intermittent streams may support rich and distinctive biological communities, in part because of the predictability of dry periods. The influence of headwaters on downstream systems emerges from their attributes that meet unique habitat requirements of residents and migrants by: offering a refuge from temperature and flow extremes, competitors, predators, and introduced species; serving as a source of colonists; providing spawning sites and rearing areas; being a rich source of food; and creating migration corridors throughout the landscape. Degradation and loss of headwaters and their connectivity to ecosystems downstream threaten the biological integrity of entire river networks.