Closing the gap between science and management of cold-water refuges in rivers and streams.

Human activities and climate change threaten coldwater organisms in freshwater ecosystems by causing rivers and streams to warm, increasing the intensity and frequency of warm temperature events, and reducing thermal heterogeneity. Cold-water refuges are discrete patches of relatively cool water that are used by coldwater organisms for thermal relief and short-term survival. Globally, cohesive management approaches are needed that consider interlinked physical, biological, and social factors of cold-water refuges. We review current understanding of cold-water refuges, identify gaps between science and management, and evaluate policies aimed at protecting thermally sensitive species. Existing policies include designating cold-water habitats, restricting fishing during warm periods, and implementing threshold temperature standards or guidelines. However, these policies are rare and uncoordinated across spatial scales and often do not consider input from Indigenous peoples. We propose that cold-water refuges be managed as distinct operational landscape units, which provide a social and ecological context that is relevant at the watershed scale. These operational landscape units provide the foundation for an integrated framework that links science and management by (1) mapping and characterizing cold-water refuges to prioritize management and conservation actions, (2) leveraging existing and new policies, (3) improving coordination across jurisdictions, and (4) implementing adaptive management practices across scales. Our findings show that while there are many opportunities for scientific advancement, the current state of the sciences is sufficient to inform policy and management. Our proposed framework provides a path forward for managing and protecting cold-water refuges using existing and new policies to protect coldwater organisms in the face of global change.

Riverscape approaches in practice: perspectives and applications.

Landscape perspectives in riverine ecology have been undertaken increasingly in the last 30 years, leading aquatic ecologists to develop a diverse set of approaches for conceptualizing, mapping and understanding 'riverscapes'. Spatiotemporally explicit perspectives of rivers and their biota nested within the socio-ecological landscape now provide guiding principles and approaches in inland fisheries and watershed management. During the last two decades, scientific literature on riverscapes has increased rapidly, indicating that the term and associated approaches are serving an important purpose in freshwater science and management. We trace the origins and theoretical foundations of riverscape perspectives and approaches and examine trends in the published literature to assess the state of the science and demonstrate how they are being applied to address recent challenges in the management of riverine ecosystems. We focus on approaches for studying and visualizing rivers and streams with remote sensing, modelling and sampling designs that enable pattern detection as seen from above (e.g. river channel, floodplain, and riparian areas) but also into the water itself (e.g. aquatic organisms and the aqueous environment). Key concepts from landscape ecology that are central to riverscape approaches are heterogeneity, scale (resolution, extent and scope) and connectivity (structural and functional), which underpin spatial and temporal aspects of study design, data collection and analysis. Mapping of physical and biological characteristics of rivers and floodplains with high-resolution, spatially intensive techniques improves understanding of the causes and ecological consequences of spatial patterns at multiple scales. This information is crucial for managing river ecosystems, especially for the successful implementation of conservation, restoration and monitoring programs. Recent advances in remote sensing, field-sampling approaches and geospatial technology are making it increasingly feasible to collect high-resolution data over larger scales in space and time. We highlight challenges and opportunities and discuss future avenues of research with emerging tools that can potentially help to overcome obstacles to collecting, analysing and displaying these data. This synthesis is intended to help researchers and resource managers understand and apply these concepts and approaches to address real-world problems in freshwater management.

Integrating regional and local monitoring data and assessment tools to evaluate habitat conditions and inform river restoration.

Restoring degraded rivers requires initial assessment of the fluvial landscape to identify stressors and riverine features that can be enhanced. We associated local-scale river habitat data collected using standardized national monitoring tools with modeled regional water temperature and flow data on mid-sized northwest U.S. rivers (30-60 m wide). We grouped these rivers according to quartiles of their modeled mean August water temperature and examined their physical habitat structure and flow. We then used principal components analysis to summarize the variation in several dimensions of physical habitat. We also compared local conditions in the Priest River, a river targeted for restoration of native salmonid habitat in northern Idaho, with those in other rivers of the region to infer potential drivers controlling water temperature. The warmest rivers had physical structure and fluvial characteristics typical of thermally degraded rivers, whereas the coldest rivers had higher mean summer flows and greater channel planform complexity. The Priest River sites had approximately twice as many deep residual pools (>50, >75, and >100 cm) and incision that averaged approximately twice that in the coldest rivers. Percentage fines and natural cover in the Priest were also more typical of the higher-temperature river groups. We found generally low instream cover and low levels of large wood both across the region and within the Priest River. Our approach enabled us to consider the local habitat conditions of a river in the context of other similarly sized rivers in the surrounding region. Understanding this context is important for identifying potential influences on river water temperature within the focal basin and for defining attainable goals for management and restoration of thermal and habitat conditions.

Integrating thermal infrared stream temperature imagery and spatial stream network models to understand natural spatial thermal variability in streams.

Under a warmer future climate, thermal refuges could facilitate the persistence of species relying on cold-water habitat. Often these refuges are small and easily missed or smoothed out by averaging in models. Thermal infrared (TIR) imagery can provide empirical water surface temperatures that capture these features at a high spatial resolution (<1 m) and over tens of kilometers. Our study examined how TIR data could be used along with spatial stream network (SSN) models to characterize thermal regimes spatially in the Middle Fork John Day (MFJD) River mainstem (Oregon, USA). We characterized thermal variation in seven TIR longitudinal temperature profiles along the MFJD mainstem and compared them with SSN model predictions of stream temperature (for the same time periods as the TIR profiles). TIR profiles identified reaches of the MFJD mainstem with consistently cooler temperatures across years that were not consistently captured by the SSN prediction models. SSN predictions along the mainstem identified ~80% of the 1-km reach scale temperature warming or cooling trends observed in the TIR profiles. We assessed whether landscape features (e.g., tributary junctions, valley confinement, geomorphic reach classifications) could explain the fine-scale thermal heterogeneity in the TIR profiles (after accounting for the reach-scale temperature variability predicted by the SSN model) by fitting SSN models using the TIR profile observation points. Only the distance to the nearest upstream tributary was identified as a statistically significant landscape feature for explaining some of the thermal variability in the TIR profile data. When combined, TIR data and SSN models provide a data-rich evaluation of stream temperature captured in TIR imagery and a spatially extensive prediction of the network thermal diversity from the outlet to the headwaters.

Conceptualizing Ecological Responses to Dam Removal: If You Remove It, What's to Come?

One of the desired outcomes of dam decommissioning and removal is the recovery of aquatic and riparian ecosystems. To investigate this common objective, we synthesized information from empirical studies and ecological theory into conceptual models that depict key physical and biological links driving ecological responses to removing dams. We define models for three distinct spatial domains: upstream of the former reservoir, within the reservoir, and downstream of the removed dam. Emerging from these models are response trajectories that clarify potential pathways of ecological transitions in each domain. We illustrate that the responses are controlled by multiple causal pathways and feedback loops among physical and biological components of the ecosystem, creating recovery trajectories that are dynamic and nonlinear. In most cases, short-term effects are typically followed by longer-term responses that bring ecosystems to new and frequently predictable ecological condition, which may or may not be similar to what existed prior to impoundment.

Modeling intrinsic potential for beaver (Castor canadensis) habitat to inform restoration and climate change adaptation.

Through their dam-building activities and subsequent water storage, beaver have the potential to restore riparian ecosystems and offset some of the predicted effects of climate change by modulating streamflow. Thus, it is not surprising that reintroducing beaver to watersheds from which they have been extirpated is an often-used restoration and climate-adaptation strategy. Identifying sites for reintroduction, however, requires detailed information about habitat factors-information that is not often available at broad spatial scales. Here we explore the potential for beaver relocation throughout the Snohomish River Basin in Washington, USA with a model that identifies some of the basic building blocks of beaver habitat suitability and does so by relying solely on remotely sensed data. More specifically, we developed a generalized intrinsic potential model that draws on remotely sensed measures of stream gradient, stream width, and valley width to identify where beaver could become established if suitable vegetation were to be present. Thus, the model serves as a preliminary screening tool that can be applied over relatively large extents. We applied the model to 5,019 stream km and assessed the ability of the model to correctly predict beaver habitat by surveying for beavers in 352 stream reaches. To further assess the potential for relocation, we assessed land ownership, use, and land cover in the landscape surrounding stream reaches with varying levels of intrinsic potential. Model results showed that 33% of streams had moderate or high intrinsic potential for beaver habitat. We found that no site that was classified as having low intrinsic potential had any sign of beavers and that beaver were absent from nearly three quarters of potentially suitable sites, indicating that there are factors preventing the local population from occupying these areas. Of the riparian areas around streams with high intrinsic potential for beaver, 38% are on public lands and 17% are on large tracts of privately-owned timber land. Thus, although there are a large number of areas that could be suitable for relocation and restoration using beavers, current land use patterns may substantially limit feasibility in these areas.

Landscape context and the biophysical response of rivers to dam removal in the United States.

Dams have been a fundamental part of the U.S. national agenda over the past two hundred years. Recently, however, dam removal has emerged as a strategy for addressing aging, obsolete infrastructure and more than 1,100 dams have been removed since the 1970s. However, only 130 of these removals had any ecological or geomorphic assessments, and fewer than half of those included before- and after-removal (BAR) studies. In addition, this growing, but limited collection of dam-removal studies is limited to distinct landscape settings. We conducted a meta-analysis to compare the landscape context of existing and removed dams and assessed the biophysical responses to dam removal for 63 BAR studies. The highest concentration of removed dams was in the Northeast and Upper Midwest, and most have been removed from 3rd and 4th order streams, in low-elevation (< 500 m) and low-slope (< 5%) watersheds that have small to moderate upstream watershed areas (10-1000 km2) with a low risk of habitat degradation. Many of the BAR-studied removals also have these characteristics, suggesting that our understanding of responses to dam removals is based on a limited range of landscape settings, which limits predictive capacity in other environmental settings. Biophysical responses to dam removal varied by landscape cluster, indicating that landscape features are likely to affect biophysical responses to dam removal. However, biophysical data were not equally distributed across variables or clusters, making it difficult to determine which landscape features have the strongest effect on dam-removal response. To address the inconsistencies across dam-removal studies, we provide suggestions for prioritizing and standardizing data collection associated with dam removal activities.

Simulated juvenile salmon growth and phenology respond to altered thermal regimes and stream network shape.

It is generally accepted that climate change will stress coldwater species like Pacific salmon. However, it is unclear what aspect of altered thermal regimes (e.g., warmer winters, springs, summers, or increased variability) will have the greatest effect, and what role the spatial properties of river networks play. Thermally diverse habitats may afford protection from climate change by providing opportunities for aquatic organisms to find and use habitats with optimal conditions for growth. We hypothesized that climate-altered thermal regimes will change growth and timing of life history events such as emergence or migration but that changes will be moderated in topologically complex stream networks where opportunities to thermoregulate are more readily available to mobile animals. Because climate change effects on populations are spatially variable and contingent upon physiological optima, assessments of risk must take a spatially explicit approach. We developed a spatially-structured individual based model for Chinook Salmon (Oncorhynchus tshawytscha) in which movement decisions and growth were governed by water temperature and conspecific density. We evaluated growth and phenology (timing of egg emergence and smolting) under a variety of thermal regimes (each having a different minimum, rate of warming, maximum, and variability) and in three network shapes of increasing spatial complexity. Across networks, fish generally grew faster and were capable of smolting earlier in warmer scenarios where water temperatures experienced by fish were closer to optimal; however, growth decreased for some fish. We found that salmon size and smolt date responded more strongly to warmer springs and summers than to warmer winters or increased variability. Fish in the least complex network grew faster and were ready to smolt earlier than fish in the more spatially complex network shapes in the contemporary thermal regime; patterns were similar but less clear in warmer thermal regimes. Our results demonstrate that network topology may influence how fish respond to thermal landscapes, and this information will be useful for incorporating a spatiotemporal context into conservation decisions that promote long-term viability of salmon in a changing climate.

The interactive effects of climate change, riparian management, and a nonnative predator on stream-rearing salmon.

Predicting how climate change is likely to interact with myriad other stressors that threaten species of conservation concern is an essential challenge in aquatic ecosystems. This study provides a framework to accomplish this task in salmon-bearing streams of the northwestern United States, where land-use-related reductions in riparian shading have caused changes in stream thermal regimes, and additional warming from projected climate change may result in significant losses of coldwater fish habitat over the next century. Predatory, nonnative smallmouth bass have also been introduced into many northwestern streams, and their range is likely to expand as streams warm, presenting an additional challenge to the persistence of threatened Pacific salmon. The goal of this work was to forecast the interactive effects of climate change, riparian management, and nonnative species on stream-rearing salmon and to evaluate the capacity of restoration to mitigate these effects. We intersected downscaled global climate forecasts with a local-scale water temperature model to predict mid- and end-of-century temperatures in streams in the Columbia River basin. We compared one stream that is thermally impaired due to the loss of riparian vegetation and another that is cooler and has a largely intact riparian corridor. Using the forecasted stream temperatures in conjunction with fish-habitat models, we predicted how stream-rearing chinook salmon and bass distributions would change as each stream warmed. In the highly modified stream, end-of-century warming may cause near total loss of chinook salmon-rearing habitat and a complete invasion of the upper watershed by bass. In the less modified stream, bass were thermally restricted from the upstream-most areas. In both systems, temperature increases resulted in higher predicted spatial overlap between stream-rearing chinook salmon and potentially predatory bass in the early summer (two- to fourfold increase) and greater abundance of bass. We found that riparian restoration could prevent the extirpation of chinook salmon from the more altered stream and could also restrict bass from occupying the upper 31 km of salmon-rearing habitat. The proposed methodology and model predictions are critical for prioritizing climate-change adaptation strategies before salmonids are exposed to both warmer water and greater predation risk by nonnative species.

Network analysis reveals multiscale controls on streamwater chemistry.

By coupling synoptic data from a basin-wide assessment of streamwater chemistry with network-based geostatistical analysis, we show that spatial processes differentially affect biogeochemical condition and pattern across a headwater stream network. We analyzed a high-resolution dataset consisting of 664 water samples collected every 100 m throughout 32 tributaries in an entire fifth-order stream network. These samples were analyzed for an exhaustive suite of chemical constituents. The fine grain and broad extent of this study design allowed us to quantify spatial patterns over a range of scales by using empirical semivariograms that explicitly incorporated network topology. Here, we show that spatial structure, as determined by the characteristic shape of the semivariograms, differed both among chemical constituents and by spatial relationship (flow-connected, flow-unconnected, or Euclidean). Spatial structure was apparent at either a single scale or at multiple nested scales, suggesting separate processes operating simultaneously within the stream network and surrounding terrestrial landscape. Expected patterns of spatial dependence for flow-connected relationships (e.g., increasing homogeneity with downstream distance) occurred for some chemical constituents (e.g., dissolved organic carbon, sulfate, and aluminum) but not for others (e.g., nitrate, sodium). By comparing semivariograms for the different chemical constituents and spatial relationships, we were able to separate effects on streamwater chemistry of (i) fine-scale versus broad-scale processes and (ii) in-stream processes versus landscape controls. These findings provide insight on the hierarchical scaling of local, longitudinal, and landscape processes that drive biogeochemical patterns in stream networks.

Diel horizontal migration in streams: juvenile fish exploit spatial heterogeneity in thermal and trophic resources.

Vertical heterogeneity in the physical characteristics of lakes and oceans is ecologically salient and exploited by a wide range of taxa through diel vertical migration to enhance their growth and survival. Whether analogous behaviors exploit horizontal habitat heterogeneity in streams is largely unknown. We investigated fish movement behavior at daily timescales to explore how individuals integrated across spatial variation in food abundance and water temperature. Juvenile coho salmon made feeding forays into cold habitats with abundant food, and then moved long distances (350-1300 m) to warmer habitats that accelerated their metabolism and increased their assimilative capacity. This behavioral thermoregulation enabled fish to mitigate trade-offs between trophic and thermal resources by exploiting thermal heterogeneity. Fish that exploited thermal heterogeneity grew at substantially faster rates than did individuals that assumed other behaviors. Our results provide empirical support for the importance of thermal diversity in lotic systems, and emphasize the importance of considering interactions between animal behavior and habitat heterogeneity when managing and restoring ecosystems.

Modelling dendritic ecological networks in space: an integrated network perspective.

Dendritic ecological networks (DENs) are a unique form of ecological networks that exhibit a dendritic network topology (e.g. stream and cave networks or plant architecture). DENs have a dual spatial representation; as points within the network and as points in geographical space. Consequently, some analytical methods used to quantify relationships in other types of ecological networks, or in 2-D space, may be inadequate for studying the influence of structure and connectivity on ecological processes within DENs. We propose a conceptual taxonomy of network analysis methods that account for DEN characteristics to varying degrees and provide a synthesis of the different approaches within the context of stream ecology. Within this context, we summarise the key innovations of a new family of spatial statistical models that describe spatial relationships in DENs. Finally, we discuss how different network analyses may be combined to address more complex and novel research questions. While our main focus is streams, the taxonomy of network analyses is also relevant anywhere spatial patterns in both network and 2-D space can be used to explore the influence of multi-scale processes on biota and their habitat (e.g. plant morphology and pest infestation, or preferential migration along stream or road corridors).

A process-based hierarchical framework for monitoring glaciated alpine headwaters.

Recent studies have demonstrated the geomorphic complexity and wide range of hydrologic regimes found in alpine headwater channels that provide complex habitats for aquatic taxa. These geohydrologic elements are fundamental to better understand patterns in species assemblages and indicator taxa and are necessary to aquatic monitoring protocols that aim to track changes in physical conditions. Complex physical variables shape many biological and ecological traits, including life history strategies, but these mechanisms can only be understood if critical physical variables are adequately represented within the sampling framework. To better align sampling design protocols with current geohydrologic knowledge, we present a conceptual framework that incorporates regional-scale conditions, basin-scale longitudinal profiles, valley-scale glacial macroform structure, valley segment-scale (i.e., colluvial, alluvial, and bedrock), and reach-scale channel types. At the valley segment- and reach-scales, these hierarchical levels are associated with differences in streamflow and sediment regime, water source contribution and water temperature. Examples of linked physical-ecological hypotheses placed in a landscape context and a case study using the proposed framework are presented to demonstrate the usefulness of this approach for monitoring complex temporal and spatial patterns and processes in glaciated basins. This approach is meant to aid in comparisons between mountain regions on a global scale and to improve management of potentially endangered alpine species affected by climate change and other stressors.

Projected climate-induced habitat loss for salmonids in the John Day River network, Oregon, U.S.A.

Climate change will likely have profound effects on cold-water species of freshwater fishes. As temperatures rise, cold-water fish distributions may shift and contract in response. Predicting the effects of projected stream warming in stream networks is complicated by the generally poor correlation between water temperature and air temperature. Spatial dependencies in stream networks are complex because the geography of stream processes is governed by dimensions of flow direction and network structure. Therefore, forecasting climate-driven range shifts of stream biota has lagged behind similar terrestrial modeling efforts. We predicted climate-induced changes in summer thermal habitat for 3 cold-water fish species-juvenile Chinook salmon, rainbow trout, and bull trout (Oncorhynchus tshawytscha, O. mykiss, and Salvelinus confluentus, respectively)-in the John Day River basin, northwestern United States. We used a spatially explicit statistical model designed to predict water temperature in stream networks on the basis of flow and spatial connectivity. The spatial distribution of stream temperature extremes during summers from 1993 through 2009 was largely governed by solar radiation and interannual extremes of air temperature. For a moderate climate change scenario, estimated declines by 2100 in the volume of habitat for Chinook salmon, rainbow trout, and bull trout were 69-95%, 51-87%, and 86-100%, respectively. Although some restoration strategies may be able to offset these projected effects, such forecasts point to how and where restoration and management efforts might focus.

Temperature-associated population diversity in salmon confers benefits to mobile consumers.

Habitat heterogeneity can generate intraspecific diversity through local adaptation of populations. While it is becoming increasingly clear that population diversity can increase stability in species abundance, less is known about how population diversity can benefit consumers that can integrate across population diversity in their prey. Here we demonstrate cascading effects of thermal heterogeneity on trout-salmon interactions in streams where rainbow trout rely heavily on the seasonal availability of anadromous salmon eggs. Water temperature in an Alaskan stream varied spatially from 5 degrees C to 17.5 degrees C, and spawning sockeye salmon showed population differentiation associated with this thermal heterogeneity. Individuals that spawned early in cool regions of the 5 km long stream were genetically differentiated from those spawning in warmer regions later in the season. Sockeye salmon spawning generates a pulsed resource subsidy that supports the majority of seasonal growth in stream-dwelling rainbow trout. The spatial and temporal structuring of sockeye salmon spawn timing in our focal stream extended the duration of the pulsed subsidy compared to a thermally homogeneous stream with a single population of salmon. Further, rainbow trout adopted movement strategies that exploited the multiple pulses of egg subsidies in the thermally heterogeneous stream. Fish that moved to track the resource pulse grew at rates about 2.5 times higher than those that remained stationary or trout in the reference stream with a single seasonal pulse of eggs. Our results demonstrate that habitat heterogeneity can have important effects on the population diversity of dominant species, and in turn, influence their value to species that prey upon them. Therefore, habitat homogenization may have farther-reaching ecological effects than previously considered.