Spatial and Temporal Variability in Stream Thermal Regime Drivers for Three River Networks During the Summer Growing Season.

Many cold-water dependent aquatic organisms are experiencing habitat and population declines from increasing water temperatures. Identifying mechanisms which drive local and regional stream thermal regimes facilitates restoration at ecologically relevant scales. Stream temperatures vary spatially and temporally both within and among river basins. We developed a modeling process to identify statistical relationships between drivers of stream temperature and covariates representing landscape, climate, and management-related processes. The modeling process was tested in 3 study areas of the Pacific Northwest USA during the growing season (May [start], August [warmest], September [end]). Across all months and study systems, covariates with the highest relative importance represented the physical landscape (elevation [1st], catchment area [3rd], main channel slope [5th]) and climate covariates (mean monthly air temperature [2nd] and discharge [4th]). Two management covariates (ground water use [6th] and riparian shade [7th]) also had high relative importance. Across the growing season (for all basins) local reach slope had high relative importance in May, but transitioned to a regional main channel slope covariate in August and September. This modeling process identified regionally similar and locally unique relationships among drivers of stream temperature. High relative importance of management-related covariates suggested potential restoration actions for each system.

Indexing and partitioning the spatial linear model for large data sets.

We consider four main goals when fitting spatial linear models: 1) estimating covariance parameters, 2) estimating fixed effects, 3) kriging (making point predictions), and 4) block-kriging (predicting the average value over a region). Each of these goals can present different challenges when analyzing large spatial data sets. Current research uses a variety of methods, including spatial basis functions (reduced rank), covariance tapering, etc, to achieve these goals. However, spatial indexing, which is very similar to composite likelihood, offers some advantages. We develop a simple framework for all four goals listed above by using indexing to create a block covariance structure and nearest-neighbor predictions while maintaining a coherent linear model. We show exact inference for fixed effects under this block covariance construction. Spatial indexing is very fast, and simulations are used to validate methods and compare to another popular method. We study various sample designs for indexing and our simulations showed that indexing leading to spatially compact partitions are best over a range of sample sizes, autocorrelation values, and generating processes. Partitions can be kept small, on the order of 50 samples per partition. We use nearest-neighbors for kriging and block kriging, finding that 50 nearest-neighbors is sufficient. In all cases, confidence intervals for fixed effects, and prediction intervals for (block) kriging, have appropriate coverage. Some advantages of spatial indexing are that it is available for any valid covariance matrix, can take advantage of parallel computing, and easily extends to non-Euclidean topologies, such as stream networks. We use stream networks to show how spatial indexing can achieve all four goals, listed above, for very large data sets, in a matter of minutes, rather than days, for an example data set.

Riparian vegetation shade restoration and loss effects on recent and future stream temperatures.

River temperatures are expected to increase this century harming species requiring cold-water habitat unless restoration activities protect or improve habitat availability. Local shading by riparian vegetation can cool water temperatures, but uncertainty exists over the scaling of this local effect to larger spatial extents. We evaluate this issue using a regional spatial stream network temperature model with covariates representing shade effects to predict mean August stream temperatures across 78,195 km of tributaries flowing into the Columbia River in the northwestern US. We evaluate nine scenarios predicting stream temperatures for three riparian shade conditions (current, restored, and no riparian vegetation) within three different climate periods (2000s, 2040s, and 2080s). Results suggest riparian shade restoration (2000s climate) could decrease mean August stream temperatures by 0.62°C across the study network. Under the same restored shade conditions, temperature predictions for tributaries at their confluence with the Columbia River range from 0.02-2.08°C cooler than under current shade conditions. The climate warming effect predicted for the 2040s and 2080s, however, is greater than the cooling effect from restoring riparian shade. Streams less than 10m bankfull width cooled more frequently with riparian shade restoration. In Oregon, the proportion of fish habitat for salmon and trout rearing and migration that meet temperature numeric water quality criteria could be increased by 20% under restored shade conditions although net habitat declines may still occur in the future. We conclude riparian vegetation restoration could partially mitigate future warming and help maintain cold-water habitats that function as thermal refuges if implemented strategically.

Broad-scale eDNA sampling for describing aquatic species distributions in running waters: Pacific lamprey Entosphenus tridentatus in the upper Snake River, USA.

One of the most fundamental yet challenging tasks for aquatic ecologists is to precisely delineate the range of species, particularly those that are broadly distributed, require specialized sampling methods, and may be simultaneously declining and increasing in different portions of their range. An exemplar is the Pacific lamprey Entosphenus tridentatus, a jawless anadromous fish of conservation concern that is actively managed in many coastal basins in western North America. To efficiently determine its distribution across the accessible 56,168 km of the upper Snake River basin in the north-western United States, we first delimited potential habitat by using predictions from a species distribution model based on conventionally collected historical data and from the distribution of a potential surrogate, Chinook salmon Oncorhynchus tshawytscha, which yielded a potential habitat network of 10,615 km. Within this area, we conducted a two-stage environmental DNA survey involving 394 new samples and 187 archived samples collected by professional biologists and citizen scientists using a single, standardized method from 2015 to 2021. We estimated that Pacific lamprey occupied 1875 km of lotic habitat in this basin, of which 1444 km may have been influenced by recent translocation efforts. Pacific lamprey DNA was consistently present throughout most river main stems, although detections became weaker or less frequent in the largest and warmest downstream channels and near their headwater extent. Pacific lamprey were detected in nearly all stocked tributaries, but there was no evidence of indigenous populations in such habitats. There was evidence of post-stocking movement because detections were 1.8-36.0 km upstream from release sites. By crafting a model-driven spatial sampling template and executing an eDNA-based sampling campaign led by professionals and volunteers, supplemented by previously collected samples, we established a benchmark for understanding the current range of Pacific lamprey across a large portion of its range in the interior Columbia River basin. This approach could be tailored to refine range estimates for other wide-ranging aquatic species of conservation concern.

Climate Change Shrinks and Fragments Salmon Habitats in a Snow-Dependent Region.

Climate change threatens biodiversity through global alteration of habitats, but efficient conservation responses are often hindered by imprecise downscaling of impacts. Besides thermal effects, warming also drives important ancillary environmental changes, such as when river hydrology evolves in response to climate forcing. Earlier snowmelt runoff and summer flow declines are broadly manifested in snow-dependent regions and relevant to socioeconomically important cold-water fishes. Here, we mechanistically quantify how climate-induced summer flow declines during historical and future periods cause complex local changes in Chinook salmon (Oncorhynchus tshawytscha) habitats for juveniles and spawning adults. Changes consisted of large reductions in useable habitat area and connectivity between the main channel and adjacent off-channel habitats. These reductions decrease the capacity of freshwater habitats to support historical salmon abundances and could pose risks to population persistence in some areas.

Do metapopulations and management matter for relict headwater bull trout populations in a warming climate?

Mountain headwater streams have emerged as important climate refuges for native cold-water species due to their slow climate velocities and extreme physical conditions that inhibit non-native invasions. Species persisting in refuges often do so as fragmented, relict populations from broader historical distributions that are subject to ongoing habitat reductions and increasing isolation as climate change progresses. Key for conservation planning is determining where remaining populations will persist and how habitat restoration strategies can improve biological resilience to enhance the long-term prospects for species of concern. Studying bull trout, a headwater species in the northwestern USA, we developed habitat occupancy models using a data set of population occurrence in 991 natal habitat patches with a suite of novel geospatial covariates derived from high-resolution hydroclimatic scenarios and other sources representing watershed and instream habitat conditions, patch geometry, disturbance, and biological interactions. The best model correctly predicted bull trout occupancy status in 82.6% of the patches and included effects for: patch size estimated as habitat volume, extent of within-patch reaches <9°C mean August temperature, distance to nearest occupied patch, road density, invasive brook trout prevalence, patch slope, and frequency of high winter flows. The model was used to assess 16 scenarios of bull trout occurrence within the study streams that represented a range of restoration strategies under three climatic conditions (baseline, moderate change, and extreme change). Results suggested that regional improvements in bull trout status were difficult to achieve in realistic restoration strategies due to the pervasive nature of climate change and the limited extent of restoration actions given their high costs. However, occurrence probabilities in a subset of patches were highly responsive to restoration actions, suggesting that targeted investments to improve the resilience of some populations may be contextually beneficial. A possible strategy, therefore, is focusing effort on responsive populations near more robust population strongholds, thereby contributing to local enclaves where dispersal among populations further enhances resilience. Equally important, strongholds constituted a small numerical percentage of patches (5%-21%), yet encompassed the large majority of occupied habitat by volume (72%-89%) and their protection could have significant conservation benefits for bull trout.

Some (Fish Might) Like It Hot: Habitat Quality and Fish Growth from Past to Future Climates.

Current expectation is that projected climate change may have adverse effects on fish habitats and survival. The analysis leading to these concerns is typically done at large scale with limited possibility to quantify the local biological response and compare with previous conditions. Our research investigated the effects of recorded climate conditions on Chinook salmon (Oncorhynchus tshawytscha) spawning and rearing habitats and growth responses to the local climate and compared those conditions to predicted responses to a climate change. The study site was a 7 km long reach of Bear Valley Creek, an important spawning stream for this US Endangered Species Act listed species, in the Pacific Northwest of United States. We used 2D numerical modeling supported by accurate, high-resolution survey data to calculate flow hydraulics at various discharges from base to bankfull flows. For past and future conditions, computed flow hydraulics were combined with habitat suitability indices (SI) to compute spawning and rearing habitat suitability. Information on habitat suitability along with fish density and stream water temperature informed a growth model to quantify the potential fish size, an index of survival rates and fitness. Our results indicate that yearly-averaged rearing habitat quality remains similar to historic, but the timing of high- and low-quality habitat periods shift within the calendar year. Future spawning habitat quality may be significantly reduced during the seasonal period to which Chinook have currently adapted their spawning behavior. The growth model indicates an increase in anticipated size of Chinook salmon for predicted future climate conditions due to water temperature increase. Consequently, future climate conditions may have a substantial negative impact on spawning and limited impact on rearing conditions due to flow reduction and thus quality and extent of available habitat. However, the expected warmer stream water temperatures may benefit rearing, because of increased fish size in these high elevation streams.

Application of multiple-population viability analysis to evaluate species recovery alternatives.

Population viability analysis (PVA) is a powerful conservation tool, but it remains impractical for many species, particularly species with multiple, broadly distributed populations for which collecting suitable data can be challenging. A recently developed method of multiple-population viability analysis (MPVA), however, addresses many limitations of traditional PVA. We built on previous development of MPVA for Lahontan cutthroat trout (LCT) (Oncorhynchus clarkii henshawi), a species listed under the U.S. Endangered Species Act, that is distributed broadly across habitat fragments in the Great Basin (U.S.A.). We simulated potential management scenarios and assessed their effects on population sizes and extinction risks in 211 streams, where LCT exist or may be reintroduced. Conservation populations (those managed for recovery) tended to have lower extinction risks than nonconservation populations (mean = 19.8% vs. 52.7%), but not always. Active management or reprioritization may be warranted in some cases. Eliminating non-native trout had a strong positive effect on overall carrying capacities for LCT populations but often did not translate into lower extinction risks unless simulations also reduced associated stochasticity (to the mean for populations without non-native trout). Sixty fish or 5-10 fish/km was the minimum reintroduction number and density, respectively, that provided near-maximum reintroduction success. This modeling framework provided crucial insights and empirical justification for conservation planning and specific adaptive management actions for this threatened species. More broadly, MPVA is applicable to a wide range of species exhibiting geographic rarity and limited availability of abundance data and greatly extends the potential use of empirical PVA for conservation assessment and planning.

Aplicación de un Análisis de Viabilidad Multi-Poblacional para Evaluar Alternativas de Recuperación de Especies Resumen El análisis de viabilidad poblacional (AVP) es una herramienta poderosa de conservación, que desafortunadamente sigue siendo impráctica para muchas especies, en particular para aquellas con poblaciones múltiples distribuidas ampliamente, para las cuales puede ser un reto la recolección de datos apropiados. Sin embargo, un método recientemente desarrollado de análisis de viabilidad multi-poblacional (AVMP) aborda muchas de las limitaciones de los AVP tradicionales. Partimos del desarrollo previo de un AVMP para la trucha degollada lahontana (LCT, en inglés) (Oncorhynchus clarkii henshawi), una especie enlistada bajo el Acta de Especies en Peligro de los Estados Unidos, la cual está distribuida ampliamente a lo largo de los fragmentos de hábitat que se encuentran en la Gran Cuenca (E.U.A.). Simulamos los escenarios potenciales de manejo y evaluamos sus efectos sobre el tamaño de las poblaciones y los riesgos de extinción en 211 arroyos en donde existe la LCT o en donde podría ser reintroducida. Las poblaciones de conservación (aquellas manejadas para su recuperación) tuvieron una tendencia hacia un riesgo de extinción más bajo que las poblaciones sin conservación (media = 19.8% vs. 52.7%), pero no en todos los casos. El manejo activo o la repriorización podrían ser justificadas en algunos casos. La eliminación de las truchas no nativas tuvo un fuerte efecto positivo generalizado sobre las capacidades de carga de las poblaciones de LCT, aunque frecuentemente esto no se transformó en un riesgo de extinción más bajo a menos que las simulaciones también redujeran la estocasticidad asociada (para la media de las poblaciones sin truchas no nativas). Para proporcionar un éxito de reintroducción cercano al máximo, el número mínimo de reintroducción debió ser de 60 peces o una densidad de 5-10 peces/km. Este marco de trabajo para el modelo proporcionó una percepción muy importante y una justificación empírica para la planeación de la conservación y para las acciones de manejo adaptativo para esta especie amenazada. En términos más generales, el AVMP puede aplicarse a una gama amplia de especies que exhiban una rareza geográfica y una disponibilidad limitada de datos de abundancia, además de que expande enormemente el uso potencial de AVP empíricos para la evaluación y planeación de la conservación.

Hierarchical multi-population viability analysis.

Population viability analysis (PVA) uses concepts from theoretical ecology to provide a powerful tool for quantitative estimates of population dynamics and extinction risks. However, conventional statistical PVA requires long-term data from every population of interest, whereas many species of concern exist in multiple isolated populations that are only monitored occasionally. We present a hierarchical multi-population viability analysis model that increases inference power from sparse data by sharing information among populations to assess extinction risks while accounting for incomplete detection and sampling biases with explicit observation and sampling sub-models. We present a case study in which we customized this model for historical population monitoring data (1985-2015) from federally threatened Lahontan cutthroat trout populations in the Great Basin, USA. Data were counts of fish captured during backpack electrofishing surveys from locations associated with 155 isolated populations. Some surveys (25%) included multi-pass removal sampling, which provided valuable information about capture efficiency. GIS and remote sensing were used to estimate August stream temperatures, peak flows, and riparian vegetation condition in each population each year. Field data were used to derive an annual index of nonnative trout densities. Results indicated that population growth rates were higher in colder streams and that nonnative trout reduced carrying capacities of native trout. Extinction risks increased with more environmental stochasticity and were also related to population extent, water temperatures, and nonnative densities. We developed a graphical user interface to interact with the fitted model results and to simulate future habitat scenarios and management actions to assess their influence on extinction risks in each population. Hierarchical multi-population viability analysis bridges the gap between site-level field observations and population-level processes, making effective use of existing datasets to support management decisions with robust estimates of population dynamics, extinction risks, and uncertainties.

Ecological segregation moderates a climactic conclusion to trout hybridization.

For decades, it has been assumed that introgressive hybridization between introduced rainbow trout and native cutthroat trout in western North America will lead to genomic extinction of the latter. A broad-scale re-examination of their interaction indicates that ecological differences between these species and demographic processes are dictating the location and extent of their hybrid zones, and that runaway introgression between these taxa is unlikely.

Big biology meets microclimatology: defining thermal niches of ectotherms at landscape scales for conservation planning.

Temperature profoundly affects ecology, a fact ever more evident as the ability to measure thermal environments increases and global changes alter these environments. The spatial structure of thermalscapes is especially relevant to the distribution and abundance of ectothermic organisms, but the ability to describe biothermal relationships at extents and grains relevant to conservation planning has been limited by small or sparse data sets. Here, we combine a large occurrence database of >23 000 aquatic species surveys with stream microclimate scenarios supported by an equally large temperature database for a 149 000-km mountain stream network to describe thermal relationships for 14 fish and amphibian species. Species occurrence probabilities peaked across a wide range of temperatures (7.0-18.8°C) but distinct warm- or cold-edge distribution boundaries were apparent for all species and represented environments where populations may be most sensitive to thermal changes. Warm-edge boundary temperatures for a native species of conservation concern were used with geospatial data sets and a habitat occupancy model to highlight subsets of the network where conservation measures could benefit local populations by maintaining cool temperatures. Linking that strategic approach to local estimates of habitat impairment remains a key challenge but is also an opportunity to build relationships and develop synergies between the research, management, and regulatory communities. As with any data mining or species distribution modeling exercise, care is required in analysis and interpretation of results, but the use of large biological data sets with accurate microclimate scenarios can provide valuable information about the thermal ecology of many ectotherms and a spatially explicit way of guiding conservation investments.

Climate, Demography, and Zoogeography Predict Introgression Thresholds in Salmonid Hybrid Zones in Rocky Mountain Streams.

Among the many threats posed by invasions of nonnative species is introgressive hybridization, which can lead to the genomic extinction of native taxa. This phenomenon is regarded as common and perhaps inevitable among native cutthroat trout and introduced rainbow trout in western North America, despite that these taxa naturally co-occur in some locations. We conducted a synthetic analysis of 13,315 genotyped fish from 558 sites by building logistic regression models using data from geospatial stream databases and from 12 published studies of hybridization to assess whether environmental covariates could explain levels of introgression between westslope cutthroat trout and rainbow trout in the U.S. northern Rocky Mountains. A consensus model performed well (AUC, 0.78-0.86; classification success, 72-82%; 10-fold cross validation, 70-82%) and predicted that rainbow trout introgression was significantly associated with warmer water temperatures, larger streams, proximity to warmer habitats and to recent sources of rainbow trout propagules, presence within the historical range of rainbow trout, and locations further east. Assuming that water temperatures will continue to rise in response to climate change and that levels of introgression outside the historical range of rainbow trout will equilibrate with those inside that range, we applied six scenarios across a 55,234-km stream network that forecast 9.5-74.7% declines in the amount of habitat occupied by westslope cutthroat trout populations of conservation value, but not the wholesale loss of such populations. We conclude that introgression between these taxa is predictably related to environmental conditions, many of which can be manipulated to foster largely genetically intact populations of westslope cutthroat trout and help managers prioritize conservation activities.

Slow climate velocities of mountain streams portend their role as refugia for cold-water biodiversity.

The imminent demise of montane species is a recurrent theme in the climate change literature, particularly for aquatic species that are constrained to networks and elevational rather than latitudinal retreat as temperatures increase. Predictions of widespread species losses, however, have yet to be fulfilled despite decades of climate change, suggesting that trends are much weaker than anticipated and may be too subtle for detection given the widespread use of sparse water temperature datasets or imprecise surrogates like elevation and air temperature. Through application of large water-temperature databases evaluated for sensitivity to historical air-temperature variability and computationally interpolated to provide high-resolution thermal habitat information for a 222,000-km network, we estimate a less dire thermal plight for cold-water species within mountains of the northwestern United States. Stream warming rates and climate velocities were both relatively low for 1968-2011 (average warming rate = 0.101 °C/decade; median velocity = 1.07 km/decade) when air temperatures warmed at 0.21 °C/decade. Many cold-water vertebrate species occurred in a subset of the network characterized by low climate velocities, and three native species of conservation concern occurred in extremely cold, slow velocity environments (0.33-0.48 km/decade). Examination of aggressive warming scenarios indicated that although network climate velocities could increase, they remain low in headwaters because of strong local temperature gradients associated with topographic controls. Better information about changing hydrology and disturbance regimes is needed to complement these results, but rather than being climatic cul-de-sacs, many mountain streams appear poised to be redoubts for cold-water biodiversity this century.

The cold-water climate shield: delineating refugia for preserving salmonid fishes through the 21st century.

The distribution and future fate of ectothermic organisms in a warming world will be dictated by thermalscapes across landscapes. That is particularly true for stream fishes and cold-water species like trout, salmon, and char that are already constrained to high elevations and latitudes. The extreme climates in those environments also preclude invasions by most non-native species, so identifying especially cold habitats capable of absorbing future climate change while still supporting native populations would highlight important refugia. By coupling crowd-sourced biological datasets with high-resolution stream temperature scenarios, we delineate network refugia across >250 000 stream km in the Northern Rocky Mountains for two native salmonids-bull trout (BT) and cutthroat trout (CT). Under both moderate and extreme climate change scenarios, refugia with high probabilities of trout population occupancy (>0.9) were predicted to exist (33-68 BT refugia; 917-1425 CT refugia). Most refugia are on public lands (>90%) where few currently have protected status in National Parks or Wilderness Areas (<15%). Forecasts of refuge locations could enable protection of key watersheds and provide a foundation for climate smart planning of conservation networks. Using cold water as a 'climate shield' is generalizable to other species and geographic areas because it has a strong physiological basis, relies on nationally available geospatial data, and mines existing biological datasets. Importantly, the approach creates a framework to integrate data contributed by many individuals and resource agencies, and a process that strengthens the collaborative and social networks needed to preserve many cold-water fish populations through the 21st century.

Probabilistic accounting of uncertainty in forecasts of species distributions under climate change.

Forecasts of species distributions under future climates are inherently uncertain, but there have been few attempts to describe this uncertainty comprehensively in a probabilistic manner. We developed a Monte Carlo approach that accounts for uncertainty within generalized linear regression models (parameter uncertainty and residual error), uncertainty among competing models (model uncertainty), and uncertainty in future climate conditions (climate uncertainty) to produce site-specific frequency distributions of occurrence probabilities across a species' range. We illustrated the method by forecasting suitable habitat for bull trout (Salvelinus confluentus) in the Interior Columbia River Basin, USA, under recent and projected 2040s and 2080s climate conditions. The 95% interval of total suitable habitat under recent conditions was estimated at 30.1-42.5 thousand km; this was predicted to decline to 0.5-7.9 thousand km by the 2080s. Projections for the 2080s showed that the great majority of stream segments would be unsuitable with high certainty, regardless of the climate data set or bull trout model employed. The largest contributor to uncertainty in total suitable habitat was climate uncertainty, followed by parameter uncertainty and model uncertainty. Our approach makes it possible to calculate a full distribution of possible outcomes for a species, and permits ready graphical display of uncertainty for individual locations and of total habitat.

Stream isotherm shifts from climate change and implications for distributions of ectothermic organisms.

Stream ecosystems are especially vulnerable to climate warming because most aquatic organisms are ectothermic and live in dendritic networks that are easily fragmented. Many bioclimatic models predict significant range contractions in stream biotas, but subsequent biological assessments have rarely been done to determine the accuracy of these predictions. Assessments are difficult because model predictions are either untestable or so imprecise that definitive answers may not be obtained within timespans relevant for effective conservation. Here, we develop the equations for calculating isotherm shift rates (ISRs) in streams that can be used to represent historic or future warming scenarios and be calibrated to individual streams using local measurements of stream temperature and slope. A set of reference equations and formulas are provided for application to most streams. Example calculations for streams with lapse rates of 0.8 °C/100 m and long-term warming rates of 0.1-0.2 °C decade(-1) indicate that isotherms shift upstream at 0.13-1.3 km decade(-1) in steep streams (2-10% slope) and 1.3-25 km decade(-1) in flat streams (0.1-1% slope). Used more generally with global scenarios, the equations predict isotherms shifted 1.5-43 km in many streams during the 20th Century as air temperatures increased by 0.6 °C and would shift another 5-143 km in the first half of the 21st Century if midrange projections of a 2 °C air temperature increase occur. Variability analysis suggests that short-term variation associated with interannual stream temperature changes will mask long-term isotherm shifts for several decades in most locations, so extended biological monitoring efforts are required to document anticipated distribution shifts. Resampling of historical sites could yield estimates of biological responses in the short term and should be prioritized to validate bioclimatic models and develop a better understanding about the effects of temperature increases on stream biotas.

Flow regime, temperature, and biotic interactions drive differential declines of trout species under climate change.

Broad-scale studies of climate change effects on freshwater species have focused mainly on temperature, ignoring critical drivers such as flow regime and biotic interactions. We use downscaled outputs from general circulation models coupled with a hydrologic model to forecast the effects of altered flows and increased temperatures on four interacting species of trout across the interior western United States (1.01 million km(2)), based on empirical statistical models built from fish surveys at 9,890 sites. Projections under the 2080s A1B emissions scenario forecast a mean 47% decline in total suitable habitat for all trout, a group of fishes of major socioeconomic and ecological significance. We project that native cutthroat trout Oncorhynchus clarkii, already excluded from much of its potential range by nonnative species, will lose a further 58% of habitat due to an increase in temperatures beyond the species' physiological optima and continued negative biotic interactions. Habitat for nonnative brook trout Salvelinus fontinalis and brown trout Salmo trutta is predicted to decline by 77% and 48%, respectively, driven by increases in temperature and winter flood frequency caused by warmer, rainier winters. Habitat for rainbow trout, Oncorhynchus mykiss, is projected to decline the least (35%) because negative temperature effects are partly offset by flow regime shifts that benefit the species. These results illustrate how drivers other than temperature influence species response to climate change. Despite some uncertainty, large declines in trout habitat are likely, but our findings point to opportunities for strategic targeting of mitigation efforts to appropriate stressors and locations.

Effects of climate change and wildfire on stream temperatures and salmonid thermal habitat in a mountain river network.

Mountain streams provide important habitats for many species, but their faunas are especially vulnerable to climate change because of ectothermic physiologies and movements that are constrained to linear networks that are easily fragmented. Effectively conserving biodiversity in these systems requires accurate downscaling of climatic trends to local habitat conditions, but downscaling is difficult in complex terrains given diverse microclimates and mediation of stream heat budgets by local conditions. We compiled a stream temperature database (n = 780) for a 2500-km river network in central Idaho to assess possible trends in summer temperatures and thermal habitat for two native salmonid species from 1993 to 2006. New spatial statistical models that account for network topology were parameterized with these data and explained 93% and 86% of the variation in mean stream temperatures and maximas, respectively. During our study period, basin average mean stream temperatures increased by 0.38 degrees C (0.27 degrees C/decade), and maximas increased by 0.48 degrees C (0.34 degrees C/decade), primarily due to long-term (30-50 year) trends in air temperatures and stream flows. Radiation increases from wildfires accounted for 9% of basin-scale temperature increases, despite burning 14% of the basin. Within wildfire perimeters, however, stream temperature increases were 2-3 times greater than basin averages, and radiation gains accounted for 50% of warming. Thermal habitat for rainbow trout (Oncorhynchus mykiss) was minimally affected by temperature increases, except for small shifts towards higher elevations. Bull trout (Salvelinus confluentus), in contrast, were estimated to have lost 11-20% (8-16%/decade) of the headwater stream lengths that were cold enough for spawning and early juvenile rearing, with the largest losses occurring in the coldest habitats. Our results suggest that a warming climate has begun to affect thermal conditions in streams and that impacts to biota will be specific to both species and context. Where species are at risk, conservation actions should be guided based on considerations of restoration opportunity and future climatic effects. To refine predictions based on thermal effects, more work is needed to understand mechanisms associated with biological responses, climate effects on other habitat features, and habitat configurations that confer population resilience.

Chinook salmon use of spawning patches: relative roles of habitat quality, size, and connectivity.

Declines in many native fish populations have led to reassessments of management goals and shifted priorities from consumptive uses to species preservation. As management has shifted, relevant environmental characteristics have evolved from traditional metrics that described local habitat quality to characterizations of habitat size and connectivity. Despite the implications this shift has for how habitats may be prioritized for conservation, it has been rare to assess the relative importance of these habitat components. We used an information-theoretic approach to select the best models from sets of logistic regressions that linked habitat quality, size, and connectivity to the occurrence of chinook salmon (Oncorhynchus tshawytscha) nests. Spawning distributions were censused annually from 1995 to 2004, and data were complemented with field measurements that described habitat quality in 43 suitable spawning patches across a stream network that drained 1150 km2 in central Idaho. Results indicated that the most plausible models were dominated by measures of habitat size and connectivity, whereas habitat quality was of minor importance. Connectivity was the strongest predictor of nest occurrence, but connectivity interacted with habitat size, which became relatively more important when populations were reduced. Comparison of observed nest distributions to null model predictions confirmed that the habitat size association was driven by a biological mechanism when populations were small, but this association may have been an area-related sampling artifact at higher abundances. The implications for habitat management are that the size and connectivity of existing habitat networks should be maintained whenever possible. In situations where habitat restoration is occurring, expansion of existing areas or creation of new habitats in key areas that increase connectivity may be beneficial. Information about habitat size and connectivity also could be used to strategically prioritize areas for improvement of local habitat quality, with areas not meeting minimum thresholds being deemed inappropriate for pursuit of restoration activities.