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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) 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.

Context Quantifying variability in landscape-scale surface water connectivity can help improve our understanding of the multiple effects of wetlands on downstream waterways. Objectives We examined how wetland merging and the coalescence of wetlands with streams varied both spatially (among ecoregions) and interannually (from drought to deluge) across parts of the Prairie Pothole Region. Methods Wetland extent was derived over a time series (1990-2011) using Landsat imagery. Changes in landscape-scale connectivity, generated by the physical coalescence of wetlands with other surface water features, were quantified by fusing static wetland and stream datasets with Landsat-derived wetland extent maps, and related to multiple wetness indices. The usage of Landsat allows for decadal-scale analysis, but limits the types of surface water connections that can be detected.Results Wetland extent correlated positively with the merging of wetlands and wetlands with streams. Wetness conditions, as defined by drought indices and runoff, were positively correlated with wetland extent, but less consistently correlated with measures of surface water connectivity. The degree of wetlandwetland merging was found to depend less on total wetland area or density, and more on climate conditions, as well as the threshold for how wetland/upland was defined. In contrast, the merging of wetlands with streams was positively correlated with stream density, and inversely related to wetland density. Conclusions Characterizing the degree of surface water connectivity within the Prairie Pothole Region in North America requires consideration of (1) climate-driven variation in wetness conditions and (2) within-region variation in wetland and stream spatial arrangements.

[1] The forest area in the western United States that burns annually is increasing with warmer temperatures, more frequent droughts, and higher fuel densities. Studies that examine fire effects for regional carbon balances have tended to either focus on individual fires as examples or adopt generalizations without considering how forest type, fire severity, and regional climate influence carbon legacies. This study provides a more detailed characterization of fire effects and quantifies the full carbon impacts in relation to direct emissions, slow release of fire-killed biomass, and net carbon uptake from forest regrowth. We find important variations in fire-induced mortality and combustion across carbon pools (leaf, live wood, dead wood, litter, and duff) and across low-to high-severity classes. This corresponds to fire-induced direct emissions from 1984 to 2008 averaging 4 TgC yr À1 and biomass killed averaging 10.5 TgC yr À1, with average burn area of 2723 km 2 yr À1 across the western United States. These direct emission and biomass killed rates were 1.4 and 3.7 times higher, respectively, for high-severity fires than those for low-severity fires. The results show that forest regrowth varies greatly by forest type and with severity and that these factors impose a sustained carbon uptake legacy. The western U.S. fires between 1984 and 2008 imposed a net source of 12.3 TgC yr

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Understanding hydrologic connectivity between wetlands and perennial streams is critical to understanding the reliance of stream flow on inputs from wetlands. We used the isotopic evaporation signal in water and remote sensing to examine wetland‐stream hydrologic connectivity within the Pipestem Creek watershed, North Dakota, a watershed dominated by prairie‐pothole wetlands. Pipestem Creek exhibited an evaporated‐water signal that had approximately half the isotopic‐enrichment signal found in most evaporatively enriched prairie‐pothole wetlands. Groundwater adjacent to Pipestem Creek had isotopic values that indicated recharge from winter precipitation and had no significant evaporative enrichment, indicating that enriched surface water did not contribute significantly to groundwater discharging into Pipestem Creek. The estimated surface water area necessary to generate the evaporation signal within Pipestem Creek was highly dynamic, varied primarily with the amount of discharge, and was typically greater than the immediate Pipestem Creek surface water area, indicating that surficial flow from wetlands contributed to stream flow throughout the summer. We propose a dynamic range of spilling thresholds for prairie‐pothole wetlands across the watershed allowing for wetland inputs even during low‐flow periods. Combining Landsat estimates with the isotopic approach allowed determination of potential (Landsat) and actual (isotope) contributing areas in wetland‐dominated systems. This combined approach can give insights into the changes in location and magnitude of surface water and groundwater pathways over time. This approach can be used in other areas where evaporation from wetlands results in a sufficient evaporative isotopic signal.

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.

Globally, hydrologic modifications such as ditching and subsurface drainage have significantly reduced wetland water storage capacity (i.e., volume of surface water a wetland can retain) and consequent wetland functions. While wetland area has been well documented across many landscapes and used to guide restoration efforts, few studies have directly quantified the associated wetland storage capacity. Here, we present a novel raster-based approach to quantify both contemporary and potential (i.e., restorable) storage capacities of individual depressional basins across landscapes. We demonstrate the utility of this method by applying it to the Delmarva Peninsula, a region punctuated by both depressional wetlands and drainage ditches. Across the entire peninsula, we estimated that restoration (i.e., plugging ditches) could increase storage capacity by 80%. Focusing on an individual watershed, we found that over 59% of restorable storage capacity occurs within 20 m of the drainage network, and that 93% occurs within 1 m elevation of the drainage network. Our demonstration highlights widespread ditching in this landscape, spatial patterns of both contemporary and potential storage capacities, and clear opportunities for hydrologic restoration. In Delmarva and more broadly, our novel approach can inform targeted landscape-scale conservation and restoration efforts to optimize hydrologically mediated wetland functions.