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.     

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.     

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 Chemical Connectivity of Streams and Riparian Wetlands to Downstream Waters: A Synthesis

Streams, riparian areas, floodplains, alluvial aquifers, and downstream waters (e.g., large rivers, lakes, and oceans) are interconnected by longitudinal, lateral, and vertical fluxes of water, other materials, and energy. Collectively, these interconnected waters are called fluvial hydrosystems. Physical and chemical connectivity within fluvial hydrosystems is created by the transport of nonliving materials (e.g., water, sediment, nutrients, and contaminants) which either do or do not chemically change (chemical and physical connections, respectively). A substantial body of evidence unequivocally demonstrates physical and chemical connectivity between streams and riparian wetlands and downstream waters. Streams and riparian wetlands are structurally connected to downstream waters through the network of continuous channels and floodplain form that make these systems physically contiguous, and the very existence of these structures provides strong geomorphologic evidence for connectivity. Functional connections between streams and riparian wetlands and their downstream waters vary geographically and over time, based on proximity, relative size, environmental setting, material disparity, and intervening units. Because of the complexity and dynamic nature of connections among fluvial hydrosystem units, a complete accounting of the physical and chemical connections and their consequences to downstream waters should aggregate over multiple years to decades.     

Biological Connectivity of Seasonally Ponded Wetlands across Spatial and Temporal Scales

Many species that inhabit seasonally ponded wetlands also rely on surrounding upland habitats and nearby aquatic ecosystems for resources to support life stages and to maintain viable populations. Understanding biological connectivity among these habitats is critical to ensure that landscapes are protected at appropriate scales to conserve species and ecosystem function. Biological connectivity occurs across a range of spatial and temporal scales. For example, at annual time scales many organisms move between seasonal wetlands and adjacent terrestrial habitats as they undergo life‐stage transitions; at generational time scales, individuals may disperse among nearby wetlands; and at multigenerational scales, there can be gene flow across large portions of a species’ range. The scale of biological connectivity may also vary among species. Larger bodied or more vagile species can connect a matrix of seasonally ponded wetlands, streams, lakes, and surrounding terrestrial habitats on a seasonal or annual basis. Measuring biological connectivity at different spatial and temporal scales remains a challenge. Here we review environmental and biological factors that drive biological connectivity, discuss implications of biological connectivity for animal populations and ecosystem processes, and provide examples illustrating the range of spatial and temporal scales across which biological connectivity occurs in seasonal wetlands.     

Southwestern Intermittent and Ephemeral Stream Connectivity

Ephemeral and intermittent streams are abundant in the arid and semiarid landscapes of the Western and Southwestern United States (U.S.). Connectivity of ephemeral and intermittent streams to the relatively few perennial reaches through runoff is a major driver of the ecohydrology of the region. These streams supply water, sediment, nutrients, and biota to downstream reaches and rivers. In addition, they provide runoff to recharge alluvial and regional groundwater aquifers that support baseflow in perennial mainstem stream reaches over extended periods when little or no precipitation occurs. Episodic runoff, as well as groundwater inflow to surface water in streams support limited naturally occurring riparian communities. This paper provides an overview and comprehensive examination of factors affecting the hydrologic, chemical, and ecological connectivity of ephemeral and intermittent streams on perennial or intermittent rivers in the arid and semiarid Southwestern U.S. Connectivity as influenced and moderated through the physical landscape, climate, and human impacts to downstream waters or rivers is presented first at the broader Southwestern scale, and secondly drawing on a specific and more detailed example of the San Pedro Basin due to its history of extensive observations and research in the basin. A wide array of evidence clearly illustrates hydrologic, chemical, and ecological connectivity of ephemeral and intermittent streams throughout stream networks.     

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.     

Lakes,Wetlands, and Streams as Predictors of Land Use/Cover Distribution

The importance of the surrounding landscape to aquatic ecosystems has been well established. Most research linking aquatic ecosystems to landscapes has focused on the one-way effect of land on water. However, to understand fully the complex interactions between aquatic and terrestrial ecosystems, aquatic ecosystems must be seen not only as receptors of human modification of the landscape, but also as potential drivers of these modifications. We hypothesized that the presence of aquatic ecosystems influences the spatial distribution of human land use/cover of the nearby landscape (≤1 km) and that this influence has changed through time from the 1930s to the 1990s. To test this hypothesis, we compared the distribution of residential, agricultural, and forested land use/cover around aquatic ecosystems (lakes, wetlands, and streams) to the overall regional land use/cover proportion in an area in southeast Michigan, USA; we also compared the distribution of land use/cover around county roads/highway and towns (known determinants of many land use/cover patterns) to the regional proportion. We found that lakes, wetlands, and streams were strongly associated with the distribution of land use/cover, that each ecosystem type showed different patterns, and that the magnitude of the association was at least as strong as the association with human features. We also found that the area closest to aquatic ecosystems (<500 m) was more strongly associated with land use/cover distribution than areas further away. Finally, we found that the strength of the association between aquatic ecosystems and land use/cover increased from 1938 to 1995, although the overall patterns were similar through time. Our results show that a more complete understanding is needed of the role of aquatic ecosystems on the distribution of land use/cover.     

Connectivity of Streams and Wetlands to Downstream Waters: An Integrated Systems Framework

Interest in connectivity has increased in the aquatic sciences, partly because of its relevance to the Clean Water Act. This paper has two objectives: (1) provide a framework to understand hydrological, chemical, and biological connectivity, focusing on how headwater streams and wetlands connect to and contribute to rivers; and (2) briefly review methods to quantify hydrological and chemical connectivity. Streams and wetlands affect river structure and function by altering material and biological fluxes to the river; this depends on two factors: (1) functions within streams and wetlands that affect material fluxes; and (2) connectivity (or isolation) from streams and wetlands to rivers that allows (or prevents) material transport between systems. Connectivity can be described in terms of frequency, magnitude, duration, timing, and rate of change. It results from physical characteristics of a system, e.g., climate, soils, geology, topography, and the spatial distribution of aquatic components. Biological connectivity is also affected by traits and behavior of the biota. Connectivity can be altered by human impacts, often in complex ways. Because of variability in these factors, connectivity is not constant but varies over time and space. Connectivity can be quantified with field‐based methods, modeling, and remote sensing. Further studies using these methods are needed to classify and quantify connectivity of aquatic ecosystems and to understand how impacts affect connectivity.     

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.     

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.     

EFFECTS OF URBANIZATION ON CHANNEL INSTABILITY1

ABSTRACT: Channel instability and aquatic ecosystem degradation have been linked to watershed imperviousness in humid regions of the U.S. In an effort to provide a more process‐based linkage between observed thresholds of aquatic ecosystem degradation and urbanization, standard single event approaches (U.S. Geological Survey Flood Regression Equations and rational) and continuous hydrologic models (HSPF and CASC2D) were used to examine potential changes in flow regime associated with varying levels of watershed imperviousness. The predicted changes in flow parameters were then interpreted in concert with risk‐based models of channel form and instability. Although low levels of imperviousness (10 to 20 percent) clearly have the potential to destabilize streams, changes in discharge, and thus stream power, associated with increased impervious area are highly variable and dependent upon watershed‐specific conditions. In addition to the storage characteristics of the pre‐development watershed, the magnitude of change is sensitive to the connectivity and conveyance of impervious areas as well as the specific characteristics of the receiving channels. Different stream types are likely to exhibit varying degrees and types of instability, depending on entrenchment, relative erodibility of bed and banks, riparian condition, mode of sediment transport (bedload versus suspended load), and proximity to geomorphic thresholds. Nonetheless, simple risk‐based analyses of the potential impacts of land use change on aquatic ecosystems have the potential to redirect and improve the effectiveness of watershed management strategies by facilitating the identification of channels that may be most sensitive to changes in stream power.     

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.     

Identification of Putative Geographically Isolated Wetlands of the Conterminous United States

Geographically isolated wetlands (GIWs) are wetlands completely surrounded by uplands. While common throughout the United States (U.S.), there have heretofore been no nationally available, spatially explicit estimates of GIW extent, complicating efforts to understand the myriad biogeochemical, hydrological, and habitat functions of GIWs and hampering conservation and management efforts at local, state, and national scales. We used a 10‐m geospatial buffer as a proxy for hydrological or ecological connectivity of National Wetlands Inventory palustrine and lacustrine wetland systems to nationally mapped and available stream, river, and lake data. We identified over 8.3 million putative GIWs across the conterminous U.S., encompassing nearly 6.5 million hectares of wetland resources (average size 0.79 ± 4.81 ha, median size 0.19 ha). Putative GIWs thus represent approximately 16% of the freshwater wetlands of the conterminous U.S. The water regime for the majority of the putative GIWs was temporarily or seasonally flooded, suggesting a vulnerability to ditching or hydrologic abstraction, sedimentation, or alterations in precipitation patterns. Additional analytical applications of this readily available geospatially explicit mapping product (e.g., hydrological modeling, amphibian metapopulation, or landscape ecological analyses) will improve our understanding of the abundance and extent, effect, connectivity, and relative importance of GIWs to other aquatic systems of the conterminous U.S.     

ASSESSING RELATIVE RISKS TO AQUATIC ECOSYSTEMS: A MID-APPALACHIAN CASE STUDY1

ABSTRACT: Aquatic monitoring aims to assess the condition of aquatic habitats and biota. To make statements about condition, the range of human activities and the risks they pose to aquatic ecosystems must be identified. Assessing relative risk and placing sample sites on a human disturbance gradient is necessary for interpreting biological response and distinguishing human disturbance from natural controls in aquatic systems. We describe a process that uses readily available sources, such as topographic maps, aerial photographs, and field information, to identify and prioritize stream reach and watershed stressors for 102 streams in the mid-Appalachian region of the United States. All perceptible human alterations to riparian and upland areas along with their number, type, intensity, and extent of impact were recorded and ranked; a relative risk index was developed to assign scores to the watersheds. The resulting risk index scores were consistent with measures of stream condition based on water chemistry and benthic macroinvertebrates. The risk index gives a cost-effective, regional picture of the relative risk of impairment to aquatic ecosystems in the mid-Appalachian region of the USA and could be modified for other regions or ecosystem types.     

An Assessment of U.S. Stream Compensatory Mitigation Policy: Necessary Changes to Protect Ecosystem Functions and Services

Compensatory mitigation of impacted streams and wetlands has increased over the past two decades, with the associated industry spending over US$2.9 billion in aquatic restoration annually. Despite these expenditures, evaluations by the National Research Council and U.S. Government Accountability Office have provided evidence that compensatory mitigation practices are failing to protect aquatic resource functions and services, and vague federal policy and inadequate evaluation of compensatory mitigation projects are to blame. To address these weaknesses, an update to federal regulations on compensatory mitigation was released in 2008. Additionally, the 2012 Reissuance of Nationwide Permits, some of which affects compensatory stream mitigation, was recently published. Current policy, as reflected in these documents, still uses nonspecific language to direct compensatory stream mitigation leaving most implementation decisions to the local U.S. Army Corps of Engineers district. The majority of federal mitigation policy has focused on wetland compensation, with other aquatic resources receiving less attention (e.g., streams). In this article, weaknesses of current policy are discussed, as are suggested policy changes to minimize the loss of stream ecosystem functions and services. Compensatory mitigation policy should clearly define key terms, incorporate adaptive management procedures, and provide guidelines for determining mitigation costs and compensation ratio requirements.     

URBAN WATER MANAGEMENT AND COASTAL WETLAND PROTECTION IN COLLIER COUNTY,FLORIDA1

ABSTRACT: Traditional development in South Florida has in many cases resulted in undesirable degradation of terrestrial and aquatic ecosystems, due to overdrainage and overenrichment of surface waters. The study described in this paper was undertaken in order to establish guidelines under which urban development may take place in coastal areas, while minimizing unwanted environmental changes. The study area consists of approximately 70 square miles of relatively flat land in Collier County, Florida. The coastal wetlands of the region are a highly valued natural resource containing the Rookery Bay Wildlife Sanctuary. The upland properties are mainly pine woodlands and have great potential for development. A master plan was developed which will (1) provide adequate drainage for existing and projected development within the study area and (2) maintain the integrity of the estuarine zone. The major recommendations of the plan relate to land use, physical control of surface waters, including construction and maintenance of the water management system, and implementation of the plan.     

An Ecological Integrity Index for Littoral Wetlands in Agricultural Catchments of Semiarid Mediterranean Regions

The main goal of the present study was to develop an ecological integrity index for littoral wetland management and conservation in semiarid Mediterranean areas that have been highly impacted by agriculture, including the selection of pressure and state indicators at landscape and wetlands scales that reflect the status, condition, and trends of wetlands ecosystems. We used a causality framework based on the relationship between pressure of anthropogenic activities and the ecological state of wetlands and their catchments, integrating environmental, biologic, economic, and social issues. From the application of 51 indicators in 7 littoral wetlands in the southeastern Iberian Peninsula, we selected 12 indicators (5 at catchment scale and 7 at wetland scale) to constitute the ecological integrity index proposed. The potential nitrogen export per area at catchment scale and the potential relative nitrogen export from the area surrounding the wetlands were the best pressure single predictors of state indicators with a causal relationship with environmental meaning. Wetlands in catchments with more agriculture had less ecological integrity than those in less impacted areas. A wide riparian zone in some wetlands acts as a buffer area, diminishing the effects of intensive agriculture. The index of ecological integrity developed here has a number of essential characteristics that make it a useful tool for ecosystem managers and decision-makers. The index can be used to (1) assess and control ecological integrity, (2) diagnose probable causes of ecological impairment, (3) establish criteria for protecting and restoring wetland ecosystems, and (4) integrate catchment management. Published online     

TIMBER MANAGEMENT INFLUENCES ON AQUATIC ECOSYSTEMS AND RECOMMENDATIONS FOR FUTURE RESEARCH1

ABSTRACT: Recent watershed research indicates that many timber management practices have profound effects on the water quality of small headwater streams. These streams often support fisheries of high value. Current knowledge seems to indicate that a definite potential exists for a symbiotic relationship between timber and fisheries management. Maximum development of both resources is attainable only if further research efforts recognize the mutually benefiting aspects of these heretofore separate disciplines. Future research should carefully examine the complex interrelationships between small headwater aquatic ecosystems and the riparian forest environment.