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.     

Physical and Temporal Isolation of Mountain Headwater Streams in the Western Mojave Desert,Southern California1

Abstract: Streams draining mountain headwater areas of the western Mojave Desert are commonly physically isolated from downstream hydrologic systems such as springs, playa lakes, wetlands, or larger streams and rivers by stream reaches that are dry much of the time. The physical isolation of surface flow in these streams may be broken for brief periods after rainfall or snowmelt when runoff is sufficient to allow flow along the entire stream reach. Despite the physical isolation of surface flow in these streams, they are an integral part of the hydrologic cycle. Water infiltrated from headwater streams moves through the unsaturated zone to recharge the underlying ground‐water system and eventually discharges to support springs, streamflow, isolated wetlands, or native vegetation. Water movement through thick unsaturated zones may require several hundred years and subsequent movement through the underlying ground‐water systems may require many thousands of years – contributing to the temporal isolation of mountain headwater streams.     

Recovery of lotic macroinvertebrate communities from disturbance

Ecosystem disturbances produce changes in macrobenthic community structure (abundances, biomass, and production) that persist for a few weeks to many decades. Examples of disturbances with extremely long-term effects on benthic communities include contamination by persistent toxic agents, physical changes in habitats, and altered energy inputs. Stream size, retention, and local geomorphology may ameliorate the influence of disturbances on invertebrates. Disturbances can alter food webs and may select for favorable genotypes (e.g., insecticidal resistance). Introductions of pesticides into lotic ecosystems, which do not result in major physical changes within habitats, illustrate several factors that influence invertebrate recovery time from disturbance. These include: (1) magnitude of original contamination, toxicity, and extent of continued use; (2) spatial scale of the disturbance; (3) persistence of the pesticide; (4) timing of the contamination in relation to the life history stages of the organisms; (5) vagility of populations influenced by pesticides; and (6) position within the drainage network. The ability of macroinvertebrates to recolonize denuded stream habitats may vary greatly depending on regional life histories, dispersal abilities, and position within the stream network (e.g., headwaters vs larger rivers). Although downstream drift is the most frequently cited mechanism of invertebrate recolonization following disturbance in middle- and larger-order streams, evidence is presented that shows aerial recolonization to be potentially important in headwater streams. There is an apparent stochastic element operating for aerial recolonization, depending on the timing of disturbance and flight periods of various taxa. Available evidence indicates that recolonization of invertebrate taxa without an aerial adult stage requires longer periods of time than for those that possess winged, terrestrial adult stages (i.e., most insects). Innovative, manipulative experiments are needed in order to address recolonization mechanisms of animals inhabiting streams that differ in size, latitude, disturbance frequency and magnitude, as well as the potential influence of early colonists on successional sequences of species following disturbance.     

ORGANIC MATTER DYNAMICS IN SMALL STREAMS OF THE PACIFIC NORTHWEST1

Small streams in forested landscapes are tightly coupled to the vegetation of the surrounding forest, and one of the key drivers of the stream ecosystem is the nature of organic matter supplied to it. This paper is focussed on three questions related to organic matter dynamics in small, forested streams of the conifer dominated Pacific Northwest: (1) How do small streams differ from large streams? (2) How do small streams of the Pacific Northwest differ from those of other regions? and (3) How do forest practices alter organic matter dynamics of small streams in the Pacific Northwest? The organic matter dynamics of small streams in this region differ from temperate deciduous forests in the nature of the organic matter deposited (protective chemicals, hard epidermis, slower loss rates), the timing of inputs (distributed throughout the year), and the transport rates (smaller, hard needles are more easily transported). The large amount and persistence of wood in these streams provides an additional source of organic matter that can be consumed by particular species and contributes to biofilm and fine particulate organic matter (FPOM) production. Logging is commonly practiced in many forests of the region. This practice has been shown to alter the type, amount, and timing of organic matter delivery to small streams and reduce the amount and size of large wood. Changes in channel complexity and water temperature after logging also can contribute to reduced organic matter storage. Many of the processes controlling organic matter dynamics in small streams are well described in other regions. However, the climate, vegetation, and topography of the Pacific Northwest suggest that the rates and nature of some processes affecting stream organic matter may differ considerably from other regions. Further research on small streams of this area will be required to better understand these differences.     

Transport and metabolic fate of sewage particles in a recipient stream

Although the implementation of wastewater treatment plants (WWTP) has dramatically increased the quality of surface waters in urbanized areas, WWTPs can still discharge noticeable amounts of solutes and particles to recipient streams. Although the fate of WWTP nutrients has received considerable attention, transport and in-stream transformation of sewage-derived particulate organic matter (SDPOM) have not. To investigate the transport and transformation of SDPOM in recipient streams, we experimentally injected fluorescently labeled SDPOM into a headwater stream and tracked its downstream fate at baseflow. Most SDPOM disappeared from the streamwater within a 160-m long reach with an average deposition velocity of 0.14 mm s(-1). We further coupled hydrometric measurements of specific water fluxes through the streambed interface with a mixing model to estimate streambed oxygen removal, and found significantly higher oxygen removal in the deposition (0.75 g O2 m(-2) d(-1)) than in the downstream post-deposition (0.36 g O2 m(-2) d(-1)) subreach. Contrary to our expectations, we did not detect any apparent effect of SDPOM deposition on streambed clogging. Our results show the capacity of a recipient stream to retain SDPOM and to reduce its downstream export, and thus contribute to a better understanding of ecosystem services of human-altered streams.     

The Role of Headwater Streams in Downstream Water Quality1

Abstract: Knowledge of headwater influences on the water‐quality and flow conditions of downstream waters is essential to water‐resource management at all governmental levels; this includes recent court decisions on the jurisdiction of the Federal Clean Water Act (CWA) over upland areas that contribute to larger downstream water bodies. We review current watershed research and use a water‐quality model to investigate headwater influences on downstream receiving waters. Our evaluations demonstrate the intrinsic connections of headwaters to landscape processes and downstream waters through their influence on the supply, transport, and fate of water and solutes in watersheds. Hydrological processes in headwater catchments control the recharge of subsurface water stores, flow paths, and residence times of water throughout landscapes. The dynamic coupling of hydrological and biogeochemical processes in upland streams further controls the chemical form, timing, and longitudinal distances of solute transport to downstream waters. We apply the spatially explicit, mass‐balance watershed model SPARROW to consider transport and transformations of water and nutrients throughout stream networks in the northeastern United States. We simulate fluxes of nitrogen, a primary nutrient that is a water‐quality concern for acidification of streams and lakes and eutrophication of coastal waters, and refine the model structure to include literature observations of nitrogen removal in streams and lakes. We quantify nitrogen transport from headwaters to downstream navigable waters, where headwaters are defined within the model as first‐order, perennial streams that include flow and nitrogen contributions from smaller, intermittent and ephemeral streams. We find that first‐order headwaters contribute approximately 70% of the mean‐annual water volume and 65% of the nitrogen flux in second‐order streams. Their contributions to mean water volume and nitrogen flux decline only marginally to about 55% and 40% in fourth‐ and higher‐order rivers that include navigable waters and their tributaries. These results underscore the profound influence that headwater areas have on shaping downstream water quantity and water quality. The results have relevance to water‐resource management and regulatory decisions and potentially broaden understanding of the spatial extent of Federal CWA jurisdiction in U.S. waters.     

Decreased carbon and nutrient input to boreal lakes from particulate organic matter following riparian clear-cutting

The plankton communities of oligotrophic Canadian Shield lakes are strongly regulated by the allochthonous supply of total phosphorus (TP) and dissolved organic carbon (DOC), a proportion of both of which originate from particulate organic matter. Although decreased inputs of allochthonous leaf litter have been documented for small streams whose riparian forests have been removed, no such data exist for boreal lakes. Through estimates of airborne litter input from forested and clear-cut shorelines and laboratory measurements of concentrations released from leaf leachate, we determined that riparian deforestation resulted in reductions of DOC from 17.8 to 0.4 g/m shoreline/yr and of TP from 2.9 to 0.3 g/m shoreline/yr. Previous predictive models indicate that such reductions may be substantial enough to decrease basic metabolic processes of lake plankton communities by as much as 9% in primary production and 17% in respiration.     

Nutrient retention efficiency in streams receiving inputs from wastewater treatment plants

We tested the effect of nutrient inputs from wastewater treatment plants (WWTPs) on stream nutrient retention efficiency by examining the longitudinal patterns of ammonium, nitrate, and phosphate concentrations downstream of WWTP effluents in 15 streams throughout Catalonia (Spain). We hypothesized that large nutrient loadings would saturate stream communities, lowering nutrient retention efficiency (i.e., nutrient retention relative to nutrient flux) relative to less polluted streams. Longitudinal variation in ambient nutrient concentration reflected the net result of physical, chemical, or biological uptake and release processes. Therefore, gradual increases in nutrient concentration indicate that the stream acts as a net source of nutrients to downstream environments, whereas gradual declines indicate that the stream acts as a net sink. In those streams where gradual declines in nutrient concentration were observed, we calculated the nutrient uptake length as an indicator of the stream nutrient retention efficiency. No significant decline was found in dilution-corrected concentrations of dissolved inorganic nitrogen (DIN) and phosphate in 40 and 45% of streams, respectively. In the remaining streams, uptake length (estimated based on the decline of nutrient concentrations at ambient levels) ranged from 0.14 to 29 km (DIN), and from 0.14 to 14 km (phosphate). Overall, these values are longer (lower retention efficiency) than those from nonpolluted streams of similar size, supporting our hypothesis, and suggest that high nutrient loads affect fluvial ecosystem function. This study demonstrates that the efficiency of stream ecosystems to remove nutrients has limitations because it can be significantly altered by the quantity and quality of the receiving water.     

Hydrological Connectivity Between Headwater Streams and Downstream Waters: How Science Can Inform Policy1

Abstract: In January 2001, the U.S. Supreme Court ruled that the U.S. Army Corps of Engineers exceeded its statutory authority by asserting Clean Water Act (CWA) jurisdiction over non‐navigable, isolated, intrastate waters based solely on their use by migratory birds. The Supreme Court’s majority opinion addressed broader issues of CWA jurisdiction by implying that the CWA intended some “connection” to navigability and that isolated waters need a “significant nexus” to navigable waters to be jurisdictional. Subsequent to this decision (SWANCC), there have been many lawsuits challenging CWA jurisdiction, many of which are focused on headwater, intermittent, and ephemeral streams. To inform the legal and policy debate surrounding this issue, we present information on the geographic distribution of headwater streams and intermittent and ephemeral streams throughout the U.S., summarize major findings from the scientific literature in considering hydrological connectivity between headwater streams and downstream waters, and relate the scientific information presented to policy issues surrounding the scope of waters protected under the CWA. Headwater streams comprise approximately 53% (2,900,000 km) of the total stream length in the U.S., excluding Alaska, and intermittent and ephemeral streams comprise approximately 59% (3,200,000 km) of the total stream length and approximately 50% (1,460,000 km) of the headwater stream length in the U.S., excluding Alaska. Hillslopes, headwater streams, and downstream waters are best described as individual elements of integrated hydrological systems. Hydrological connectivity allows for the exchange of mass, momentum, energy, and organisms longitudinally, laterally, vertically, and temporally between headwater streams and downstream waters. Via hydrological connectivity, headwater, intermittent and ephemeral streams cumulatively contribute to the functional integrity of downstream waters; hydrologically and ecologically, they are a part of the tributary system. As this debate continues, scientific input from multiple fields will be important for policymaking at the federal, state, and local levels and to inform water resource management regardless of the level at which those decisions are being made. Strengthening the interface between science, policy, and public participation is critical if we are going to achieve effective water resource management.     

Influence of Restoration Age and Riparian Vegetation on Reach‐Scale Nutrient Retention in Restored Urban Streams

In urban watersheds, stormwater inputs largely bypass the buffering capacity of riparian zones through direct inputs of drainage pipes and lowered groundwater tables. However, vegetation near the stream can still influence instream nutrient transformations via maintenance of streambank stability, input of woody debris, modulation of organic matter sources, and temperature regulation. Stream restoration seeks to mimic many of these functions by engineering channel complexity, grading stream banks to reconnect incised channels, and replanting lost riparian vegetation. The goal of this study was to quantify these effects by measuring nitrate and phosphate uptake in five restored streams in Charlotte and Raleigh, North Carolina, with a range of restoration ages. Using nutrient spiraling methods, uptake velocity of nitrate (0.02‐3.56 mm/min) and phosphate (0.14‐19.1 mm/min) was similar to other urban restored streams and higher than unimpacted forested streams with variability influenced by restoration age and geomorphology. Using a multiple linear regression approach, reach‐scale phosphate uptake was greater in newly restored sites, which was attributed to assimilation by algal biofilms, whereas nitrate uptake was highest in older sites potentially due to greater channel stability and establishment of microbial communities. The patterns we observed highlight the influence of riparian vegetation on energy inputs (e.g., heat, organic matter) and thereby on nutrient retention.     

A Spatially Explicit Model for Mapping Headwater Streams

Headwater streams are the primary sources of water in a drainage network and serve as a critical hydrologic link between the surrounding landscape and larger, downstream surface waters. Many states, including North Carolina, regulate activity in and near headwater streams for the protection of water quality and aquatic resources. A fundamental tool for regulatory management is an accurate representation of streams on a map. Limited resources preclude field mapping every headwater stream and its origin across a large region. It is more practical to develop a model for headwater streams based on a sample of field data that can then be extrapolated to a larger area of interest. The North Carolina Division of Water Quality has developed a cost‐effective method for modeling and mapping the location, length, and flow classification (intermittent and perennial) of headwater streams. We used a multiple logistic regression approach that combined field data and terrain derivatives for watersheds located in the Triassic Basins ecoregion. Field data were collected using a standard methodology for identifying headwater streams and origins. Terrain derivatives were generated from digital elevation models interpolated from bare‐earth Light Detection and Range data. Model accuracies greater than 80% were achieved in classifying stream presence and absence, stream length and perennial stream length, but were not as consistent in predicting intermittent stream length.     

Using GIS to Delineate Headwater Stream Origins in the Appalachian Coalfields of Kentucky

Headwater streams have a significant nexus or physical, chemical, and/or biological connection to downstream reaches. Generally, defined as 1st‐3rd order with ephemeral, intermittent, or perennial flow regimes, these streams account for a substantial portion of the total stream network particularly in mountainous terrain. Due to their often remote locations, small size, and large numbers, conducting field inventories of headwater streams is challenging. A means of estimating headwater stream location and extent according to flow regime type using publicly available spatial data is needed to simplify this complex process. Using field‐collected headwater point of origin data from three control watersheds, streams were characterized according to a set of spatial parameters related to topography, geology, and soils. These parameters were (1) compared to field‐collected point of origin data listed in three nearby Jurisdictional Determinations, (2) used to develop a geographic information system (GIS)‐based stream network for identifying ephemeral, intermittent, and perennial streams, and (3) applied to a larger watershed and compared to values obtained using the high‐resolution National Hydrography Dataset (NHD). The parameters drainage area and local valley slope were the most reliable predictors of flow regime type. Results showed the high‐resolution NHD identified no ephemeral streams and 9 and 65% fewer intermittent and perennial streams, respectively, than the GIS model.     

Hydrological,Physical, and Chemical Functions and Connectivity of Non‐Floodplain Wetlands to Downstream Waters: A Review

We reviewed the scientific literature on non‐floodplain wetlands (NFWs), freshwater wetlands typically located distal to riparian and floodplain systems, to determine hydrological, physical, and chemical functioning and stream and river network connectivity. We assayed the literature for source, sink, lag, and transformation functions, as well as factors affecting connectivity. We determined NFWs are important landscape components, hydrologically, physically, and chemically affecting downstream aquatic systems. NFWs are hydrologic and chemical sources for other waters, hydrologically connecting across long distances and contributing compounds such as methylated mercury and dissolved organic matter. NFWs reduced flood peaks and maintained baseflows in stream and river networks through hydrologic lag and sink functions, and sequestered or assimilated substantial nutrient inputs through chemical sink and transformative functions. Landscape‐scale connectivity of NFWs affects water and material fluxes to downstream river networks, substantially modifying the characteristics and function of downstream waters. Many factors determine the effects of NFW hydrological, physical, and chemical functions on downstream systems, and additional research quantifying these factors and impacts is warranted. We conclude NFWs are hydrologically, chemically, and physically interconnected with stream and river networks though this connectivity varies in frequency, duration, magnitude, and timing.     

The Role of Headwater Wetlands in Altering Streamflow and Chemistry in a Maine,USA Catchment1

Morley, Terry R., Andrew S. Reeve, and Aram J.K. Calhoun, 2011. The Role of Headwater Wetlands in Altering Streamflow and Chemistry in a Maine, USA Catchment. Journal of the American Water Resources Association (JAWRA) 1‐13. DOI: 10.1111/j.1752‐1688.2011.00519.x Abstract: Headwater wetlands, including hillside seeps, may contribute to downstream systems disproportionately to their relatively small size. We quantified the hydrology and chemistry of headwater wetlands in a central Maine, USA, catchment from 2003 to 2005 to determine their role in maintaining headwater streamflow and in affecting stream chemistry. A few of these headwater wetlands, commonly referred to as “seeps,” were characterized by relatively high groundwater discharge. During summer base flow, seeps were the primary source of surface water to the stream, contributing between 40 and 80% of stream water. Comparisons of groundwater and surface water dominant ion chemistry revealed only slight differences at the bedrock interface; however, significant changes occurred at the shallow groundwater‐surface water interface where we found decreases in total and individual cation concentrations with decreasing depth. Seep outflows significantly increased total cation and calcium concentrations in streams. Outflows at two seeps produced relatively high nitrate concentrations (88 ± 15 and 93 ± 15 μg/l respectively), yet did not correspond to higher nitrate in stream water below seep outflows (2 ± 1 μg/l). We demonstrate that small wetlands (< 1,335 m²) can contribute to headwater stream processes by linking groundwater and surface‐water systems, increasing the duration and magnitude of stream discharge, and by affecting stream chemistry, particularly during periods of base flow.     

Summer Temperature Patterns in Headwater Streams of the Oregon Coast Range1

Abstract: Cool summertime stream temperature is an important component of high quality aquatic habitat in Oregon coastal streams. Within the Oregon Coast Range, small headwater streams make up a majority of the stream network; yet, little information is available on temperature patterns and the longitudinal variability for these streams. In this paper we describe preharvest spatial and temporal patterns in summer stream temperature for small streams of the Oregon Coast Range in forests managed for timber production. We also explore relationships between stream and riparian attributes and observed stream temperature conditions and patterns. Summer stream temperature, channel, and riparian data were collected on 36 headwater streams in 2002, 2003, and 2004. Mean stream temperatures were consistent among summers and generally warmed in a downstream direction. However, longitudinal trends in maximum temperatures were more variable. At the reach scale of 0.5‐1.7 km, maximum temperatures increased in 17 streams, decreased in seven streams and did not change in three reaches. At the subreach scale (0.1‐1.5 km), maximum temperatures increased in 28 subreaches, decreased in 14, and did not change in 12 subreaches. Models of increasing temperature in a downstream direction may oversimplify fine‐scale patterns in small streams. Stream and riparian attributes that correlated with observed temperature patterns included cover, channel substrate, channel gradient, instream wood jam volume, riparian stand density, and geology type. Longitudinal patterns of stream temperature are an important consideration for background characterization of water quality. Studies attempting to evaluate stream temperature response to timber harvest or other modifications should quantify variability in longitudinal patterns of stream temperature prior to logging.     

Transport and fate of nitrate in headwater agricultural streams in Illinois

Nitrogen inputs to the Gulf of Mexico have increased during recent decades and agricultural regions in the upper Midwest, such as those in Illinois, are a major source of N to the Mississippi River. How strongly denitrification affects the transport of nitrate (NO(3)-N) in Illinois streams has not been directly assessed. We used the nutrient spiraling model to assess the role of in-stream denitrification in affecting the concentration and downstream transport of NO(3)-N in five headwater streams in agricultural areas of east-central Illinois. Denitrification in stream sediments was measured approximately monthly from April 2001 through January 2002. Denitrification rates tended to be high (up to 15 mg N m(-2) h(-1)), but the concentration of NO(3)-N in the streams was also high (>7 mg N L(-1)). Uptake velocities for NO(3)-N (uptake rate/concentration) were lower than reported for undisturbed streams, indicating that denitrification was not an efficient N sink relative to the concentration of NO(3)-N in the water column. Denitrification uptake lengths (the average distance NO(3)-N travels before being denitrified) were long and indicated that denitrification in the streambed did not affect the transport of NO(3)-N. Loss rates for NO(3)-N in the streams were <5% d(-1) except during periods of low discharge and low NO(3)-N concentration, which occurred only in late summer and early autumn. Annually, most NO(3)-N in these headwater sites appeared to be exported to downstream water bodies rather than denitrified, suggesting previous estimates of N losses through in-stream denitrification may have been overestimated.     

CLIMATIC AND HYDROLOGIC VARIABILITY IN A COASTAL WATERSHED OF SOUTHWESTERN BRITISH COLUMBIA1

ABSTRACT: Climate data from the Malcolm Knapp Research Forest (MKRF) in the Coast Range mountains of southwestern British Columbia were used to examine relationships between climate and hydrology and variations in the El Niño Southern Oscillation (ENSO) and Pacific Decadal Oscillation (PDO). Air and water temperatures were higher and precipitation was lower during in‐phase or warm PDO/E1 Niño events than in other years. In contrast, in‐phase or cool PDO/La Niña years were generally cooler and wetter than other years. Precipitation and East Creek discharge were positively related to the Southern Oscillation Index (SOI) and negatively related to the PDO index. Conversely, air and water temperatures were negatively related to the SOI and positively related to the PDO index. Differences in precipitation and air temperature were also evident at longer time scales when separated by PDO phase. Because of drier conditions during in‐phase El Niño events, the flow of organic matter from East Creek to downstream portions of the channel network was lower compared to other years. This reduction has implications for downstream communities, as these subsidies provide a major source of energy for stream food webs. Therefore, short term and long term shifts in climate, discharge, and water temperature may have profound impacts on the ecology of Pacific Northwest (PNW) watersheds due to changes in a number of ecosystem processes such as altered flux of organic matter from headwater streams to larger rivers.     

GRASS VERSUS TREES: MANAGING RIPARIAN AREAS TO BENEFIT STREAMS OF CENTRAL NORTH AMERICA1

ABSTRACT: Forestation of riparian areas has long been promoted to restore stream ecosystems degraded by agriculture in central North America. Although trees and shrubs in the riparian zone can provide many benefits to streams, grassy or herbaceous riparian vegetation can also provide benefits and may be more appropriate in some situations. Here we review some of the positive and negative implications of grassy versus wooded riparian zones and discuss potential management outcomes. Compared to wooded areas, grassy riparian areas result in stream reaches with different patterns of bank stability, erosion, channel morphology, cover for fish, terrestrial runoff, hydrology, water temperature, organic matter inputs, primary production, aquatic macroinvertebrates, and fish. Of particular relevance in agricultural regions, grassy riparian areas may be more effective in reducing bank erosion and trapping suspended sediments than wooded areas. Maintenance of grassy riparian vegetation usually requires active management (e.g., mowing, burning, herbicide treatments, and grazing), as successional processes will tend ultimately to favor woody vegetation. Riparian agricultural practices that promote a dense, healthy, grassy turf, such as certain types of intensively managed livestock grazing, have potential to restore degraded stream ecosystems.     

RIPARIAN COMMUNITIES ASSOCIATED WITH PACIFIC NORTHWEST HEADWATER STREAMS: ASSEMBLAGES,PROCESSES, AND UNIQUENESS1

Riparian areas of large streams provide important habitat to many species and control many instream processes — but is the same true for the margins of small streams? This review considers riparian areas alongside small streams in forested, mountainous areas of the Pacific Northwest and asks if there are fundamental ecological differences from larger streams and from other regions and if there are consequences for management from any differences. In the moist forests along many small streams of the Pacific Northwest, the contrast between the streamside and upslope forest is not as strong as that found in drier regions. Small streams typically lack floodplains, and the riparian area is often constrained by the hillslope. Nevertheless, riparian‐associated organisms, some unique to headwater areas, are found along small streams. Disturbance of hillslopes and stream channels and microclimatic effects of streams on the riparian area provide great heterogeneity in processes and diversity of habitats. The tight coupling of the terrestrial riparian area with the aquatic system results from the closed canopy and high edge‐to‐area ratio for small streams. Riparian areas of the temperate, conifer dominated forests of the Pacific Northwest provide a unique environment. Forest management guidelines for small streams vary widely, and there has been little evaluation of the local or downstream consequences of forest practices along small streams.