Spatiotemporal patterns and habitat associations of smallmouth bass (Micropterus dolomieu) invading salmon‐rearing habitat

1. Smallmouth bass (Micropterus dolomieu) have been widely introduced to fresh waters throughout the world to promote recreational fishing opportunities. In the Pacific Northwest (U.S.A.), upstream range expansions of predatory bass, especially into subyearling salmon‐rearing grounds, are of increasing conservation concern, yet have received little scientific inquiry. Understanding the habitat characteristics that influence bass distribution and the timing and extent of bass and salmon overlap will facilitate the development of management strategies that mitigate potential ecological impacts of bass. 2. We employed a spatially continuous sampling design to determine the extent of bass and subyearling Chinook salmon (Oncorhynchus tshawytscha) sympatry in the North Fork John Day River (NFJDR), a free‐flowing river system in the Columbia River Basin that contains an upstream expanding population of non‐native bass. Extensive (i.e. 53 km) surveys were conducted over 2 years and during an early and late summer period of each year, because these seasons provide a strong contrast in the river’s water temperature and flow condition. Classification and regression trees were applied to determine the primary habitat correlates of bass abundance at reach and channel‐unit scales. 3. Our study revealed that bass seasonally occupy up to 22% of the length of the mainstem NFJDR where subyearling Chinook salmon occur, and the primary period of sympatry between these species was in the early summer and not during peak water temperatures in late summer. Where these species co‐occurred, bass occupied 60–76% of channel units used by subyearling Chinook salmon in the early summer and 28–46% of the channel units they occupied in the late summer. Because these rearing salmon were well below the gape limitation of bass, this overlap could result in either direct predation or sublethal effects of bass on subyearling Chinook salmon. The upstream extent of bass increased 10–23 km (2009 and 2010, respectively) as stream temperatures seasonally warmed, but subyearling Chinook salmon were also found farther upstream during this time. 4. Our multiscale analysis suggests that bass were selecting habitat based on antecedent thermal history at a broad scale, and if satisfactory temperature conditions were met, mesoscale habitat features (i.e. channel‐unit type and depth) played an additional role in determining bass abundance. The upstream extent of bass in the late summer corresponded to a high‐gradient geomorphic discontinuity in the NFJDR, which probably hindered further upstream movements of bass. The habitat determinants and upstream extent of bass were largely consistent across years, despite marked differences in the magnitude and timing of spring peak flows prior to bass spawning. 5. The overriding influence of water temperature on smallmouth bass distribution suggests that managers may be able limit future upstream range expansions of bass into salmon‐rearing habitat by concentrating on restoration activities that mitigate climate‐ or land‐use‐related stream warming. These management activities could be prioritised to capitalise on survival bottlenecks in the life history of bass and spatially focused on landscape knick points such as high‐gradient discontinuities to discourage further upstream movements of bass.     

Thermal patterns in the surface waters of a glacial river corridor (Val Roseg, Switzerland)

SUMMARY 1. We examined the thermal patterns of the surface waters in the catchment of the Roseg River, which is fed by the meltwaters of two valley glaciers. One of the glaciers has a lake at its terminus. The river corridor comprised a proglacial stream reach below one glacier, the glacier lake outlet stream, a 2.5‐km long complex floodplain and a constrained reach extending to the end of the catchment. 2. Temperatures were continuously measured with temperature loggers at 27 sites between 1997 and 1998. Moreover, from 1997 to 1999, spot measurements were taken at 33–165 floodplain sites (depending on water level) at monthly intervals. 3. The temperature regime of glacial streams, including the glacier lake outlet, was characterised by rapidly increasing temperatures in April and May, a moderate decline from June to September (period of glacial melt) and a subsequent fast decline in autumn. During summer, the lake increased temperatures in the outlet stream by 2–4 °C, compared with the adjacent proglacial stream reach. 4. In the main channel (thalweg) of the Roseg River, annual degree‐days (DD) ranged from 176 DD in the upper proglacial reach to 1227 DD at the end of the catchment. 5. Thermal variation among different channels within the floodplain was higher than the variation along the entire main channel. Floodplain channels lacking surface connection to the main channel accumulated up to 1661 annual DDs. 6. Thermal heterogeneity within the floodplain was linked to the glacial flow pulse. With the onset of ice melt, temperatures in the main channel and in channels surface‐connected to the main channel began to decline, whereas in surface‐disconnected channels temperatures continued to increase; as a consequence, thermal heterogeneity at the floodplain scale rose slightly until September. 7. High thermal heterogeneity was not anticipated in the harsh environment of a largely glacierised alpine catchment. The relatively wide range of thermal environments may contribute to the highly diverse zoobenthic community.     

Thermal controls of Yellowstone cutthroat trout and invasive fishes under climate change

We combine large observed data sets and dynamically downscaled climate data to explore historic and future (2050–2069) stream temperature changes over the topographically diverse Greater Yellowstone Ecosystem (elevation range = 824–4017 m). We link future stream temperatures with fish growth models to investigate how changing thermal regimes could influence the future distribution and persistence of native Yellowstone cutthroat trout (YCT) and competing invasive species. We find that stream temperatures during the recent decade (2000–2009) surpass the anomalously warm period of the 1930s. Climate simulations indicate air temperatures will warm by 1 °C to >3 °C over the Greater Yellowstone by mid‐21st century, resulting in concomitant increases in 2050–2069 peak stream temperatures and protracted periods of warming from May to September (MJJAS). Projected changes in thermal regimes during the MJJAS growing season modify the trajectories of daily growth rates at all elevations with pronounced growth during early and late summer. For high‐elevation populations, we find considerable increases in fish body mass attributable both to warming of cold‐water temperatures and to extended growing seasons. During peak July to August warming, mid‐21st century temperatures will cause periods of increased thermal stress, rendering some low‐elevation streams less suitable for YCT. The majority (80%) of sites currently inhabited by YCT, however, display minimal loss (<10%) or positive changes in total body mass by midcentury; we attribute this response to the fact that many low‐elevation populations of YCT have already been extirpated by historical changes in land use and invasions of non‐native species. Our results further suggest that benefits to YCT populations due to warmer stream temperatures at currently cold sites could be offset by the interspecific effects of corresponding growth of sympatric, non‐native species, underscoring the importance of developing climate adaptation strategies that reduce limiting factors such as non‐native species and habitat degradation.     

Streams in an uninhabited watershed have predictably different thermal sensitivities to variable summer air temperatures

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Experimental whole‐stream warming alters community size structure

How ecological communities respond to predicted increases in temperature will determine the extent to which Earth's biodiversity and ecosystem functioning can be maintained into a warmer future. Warming is predicted to alter the structure of natural communities, but robust tests of such predictions require appropriate large‐scale manipulations of intact, natural habitat that is open to dispersal processes via exchange with regional species pools. Here, we report results of a two‐year whole‐stream warming experiment that shifted invertebrate assemblage structure via unanticipated mechanisms, while still conforming to community‐level metabolic theory. While warming by 3.8 °C decreased invertebrate abundance in the experimental stream by 60% relative to a reference stream, total invertebrate biomass was unchanged. Associated shifts in invertebrate assemblage structure were driven by the arrival of new taxa and a higher proportion of large, warm‐adapted species (i.e., snails and predatory dipterans) relative to small‐bodied, cold‐adapted taxa (e.g., chironomids and oligochaetes). Experimental warming consequently shifted assemblage size spectra in ways that were unexpected, but consistent with thermal optima of taxa in the regional species pool. Higher temperatures increased community‐level energy demand, which was presumably satisfied by higher primary production after warming. Our experiment demonstrates how warming reassembles communities within the constraints of energy supply via regional exchange of species that differ in thermal physiological traits. Similar responses will likely mediate impacts of anthropogenic warming on biodiversity and ecosystem function across all ecological communities.     

An Ecologically Based Approach to Identifying Restoration Priorities in an Acid-Impacted Watershed

The extent of impairment to some Appalachian watersheds from acid precipitation is so extreme that watershed scale analytical tools are needed to help guide cost‐effective management decisions. The objective of this study was to develop a measure of the functional value of streams as potential areas for juvenile Brook trout recruitment. This measure, which we term “weighted potential recruitment area” (WPRA), is a function of the expected Brook trout spawning intensity and juvenile survivorship. Estimates of WPRA for each stream segment were then used to identify restoration priorities and optimal restoration programs in the upper Shavers Fork watershed in West Virginia, U.S.A. Using this approach, we determined that the watershed has lost nearly 80% of its historic juvenile recruitment potential as a result of acid precipitation. We also determined that of the 145 stream segments in the watershed, eight critical stream segments account for nearly 20% of the loss. The costs and ecological benefits of a series of five alternative restoration programs were then assessed using an ArcGIS model (Environmental Systems Research Institute, Redlands, CA, U.S.A.). This approach identified two “optimal” alternatives: (1) a low‐cost, moderate‐benefit approach that would use existing rail access to treat acidification in three critical headwater locations and (2) a high‐cost, high‐benefit approach that would use aerial limestone application to treat numerous acidic tributaries near their source. The measure of stream ecological value that we developed was effective in identifying critical restoration priorities and optimal restoration strategies in this watershed. A similar procedure could be used to guide watershed restoration decisions throughout the Appalachian region.     

Surface–subsurface water exchange rates along alluvial river reaches control the thermal patterns in an Alpine river network

1. Water temperature is a key characteristic of stream ecosystems that is gaining scientific and managerial relevance as maximum temperatures in aquatic ecosystems increase worldwide.
2. To assess the effect of surface–subsurface water exchange on stream water temperature patterns, four alluvial reaches in the Tagliamento River basin (NE Italy), constrained by geomorphic knickpoints at the upper and lower end, and two to four hyporheic flowpaths within each reach, were continuously studied during summer 2007 and winter 2007–08. Water temperature was continuously monitored at the upstream and downstream knickpoints of the floodplains, as well as at discrete upwelling areas within each reach. Discharge and vertical hydraulic gradient were measured along the alluvial reaches, and the residence time and chemistry of upwelling water were assessed four times during the study.
3. Discharge variation along the study reaches revealed that massive hyporheic exchange occurred in all sites, ranging from 21% in reach 2–52% in reach 1. End member mixing analysis showed little influence of ground water, as almost all upwelling water was freshly infiltrated hyporheic water. Importantly, hyporheic exchange flows shaped surface temperature at the upwelling locations in all study reaches, providing potential thermal refugia for aquatic biota. At sites with highest hyporheic flow rates, net temperature change was also reflected at the floodplain scale.
4. The magnitude of the thermal change along a hyporheic flowpath was not related to the flowpath length but to the estimated ²²²Rn water age. Reduction in the diel thermal amplitude by hyporheic flows rather than net temperature change, reduced temperature extremes. Therefore, restoration activities to create thermal refugia should consider the role of hyporheic flows and enhance the exchange between surface and hyporheic waters.     

Non-native species limit stream restoration benefits for brook trout

Success of stream restoration can be difficult to define because many interacting abiotic and biotic factors across spatio-temporal scales can have measurable effects. Consequently, failure in habitat restoration to achieve targeted biological goals may reflect interactions of habitat restoration with unaccounted risks that have yet to be addressed on the landscape. This is particularly true within invaded landscapes, where habitat restoration can benefit non-native competitors as much as the native fishes for which restoration is designed. We tested for interacting effects of a reach scale habitat restoration effort and non-native trout competition on habitat use by a brook trout (Salvelinus fontinalis) metapopulation within a productive main stem corridor of the Shavers Fork watershed, West Virginia. We used a joint species occupancy model within a BACI sampling design to show that brook trout occupancy of main stem habitat was highest post-restoration within restored sampling reaches, but this benefit to native brook trout was conditional on brown trout (Salmo trutta) not being present within the main stem habitat. Collectively these results indicate that habitat restoration was only beneficial for native brook trout when non-native trout were absent from the restored sampling area. Proactive approaches to restoration will be integral for supporting resilient ecosystems in response to future anthropogenic threats (e.g. climate change), and we have shown that such actions will only be successful if non-native competitors do not also benefit from the restoration actions.     

Behavioral Thermoregulation by Juvenile Spring and Fall Chinook Salmon, Oncorhynchus Tshawytscha, during Smoltification

Fall chinook salmon evolved to emigrate during the summer months. The shift in the temperature preference we observed in smolting fall chinook but not spring chinook salmon may reflect a phylogenetic adaptation to summer emigration by (1) providing directional orientation as fall chinook salmon move into the marine environment, (2) maintaining optimal gill function during emigration and seawater entry, and/or (3) resetting thermoregulatory set-points to support physiological homeostasis once smolted fish enter the marine environment. Phylogenetically determined temperature adaptations and responses to thermal stress may not protect fall chinook salmon from the recent higher summer water temperatures, altered annual thermal regimes, and degraded cold water refugia that result from hydropower regulation of the Columbia and Snake rivers. The long-term survival of fall chinook salmon will likely require restoration of normal annual thermographs and rigorous changes in land use practices to protect critical thermal refugia and control maximum summer water temperatures in reservoirs.     

Thermal adaptation of soil microbial respiration to elevated temperature

In the short‐term heterotrophic soil respiration is strongly and positively related to temperature. In the long‐term, its response to temperature is uncertain. One reason for this is because in field experiments increases in respiration due to warming are relatively short‐lived. The explanations proposed for this ephemeral response include depletion of fast‐cycling, soil carbon pools and thermal adaptation of microbial respiration. Using a > 15 year soil warming experiment in a mid‐latitude forest, we show that the apparent ‘acclimation’ of soil respiration at the ecosystem scale results from combined effects of reductions in soil carbon pools and microbial biomass, and thermal adaptation of microbial respiration. Mass‐specific respiration rates were lower when seasonal temperatures were higher, suggesting that rate reductions under experimental warming likely occurred through temperature‐induced changes in the microbial community. Our results imply that stimulatory effects of global temperature rise on soil respiration rates may be lower than currently predicted.     

In‐stream habitat and macroinvertebrate responses to riparian corridor length in rangeland streams

Conservation and restoration of riparian vegetation in agricultural landscapes has had mixed success at protecting in‐stream habitat, potentially due to the mismatch between watershed‐scale impacts and reach‐scale restoration. Prioritizing contiguous placement of small‐scale restoration interventions may effectively create larger‐scale restoration projects and improve ecological outcomes. We performed a multi‐site field study to evaluate whether greater linear length of narrow riparian tree corridors resulted in measurable benefits to in‐stream condition. We collected data at 41 sites with varying upstream tree cover nested within 13 groups in rangeland streams in coastal northern California, United States. We evaluated the effect of riparian tree corridor length on benthic macroinvertebrate communities, as well as food resources, water temperature, and substrate size. Sites with longer riparian corridors had higher percentages of invertebrates sensitive to disturbance (including clingers and EPT taxa) as well as lower water temperatures and less fine sediment, two of the most important aquatic stressors. Despite marked improvement, we found no evidence that macroinvertebrate communities fully recovered, suggesting that land use continued to constrain conditions. The restoration of long riparian corridors may be an economically viable and rapidly implementable technique to improve habitat, control sediment, and counter increasing water temperatures expected with climate change within the context of ongoing land use.     

Evidence of counter‐gradient growth in western pond turtles (Actinemys marmorata) across thermal gradients

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Fish recruitment in rivers with modified discharge depends on the interacting effects of flow and thermal regimes

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Restoring Stream Ecosystems: Lessons from a Midwestern State

Reach‐scale stream restorations are becoming a common approach to repair degraded streams, but the effectiveness of these projects is rarely evaluated or reported. We surveyed governmental, private, and nonprofit organizations in the state of Indiana to determine the frequency and nature of reach‐scale stream restorations in this midwestern U.S. state. For 10 attempted restorations in Indiana, questionnaires and on‐site assessments were used to better evaluate current designs for restoring stream ecosystems. At each restoration site, habitat and water quality were evaluated in restored and unrestored reaches. Our surveys identified commonalities across all restorations, including the type of restoration, project goals, structures installed, and level of monitoring conducted. In general, most restorations were described as stream‐relocation projects that combined riparian and in‐stream enhancements. Fewer than half of the restorations conducted pre‐ or post‐restoration monitoring, and most monitoring involved evaluations of riparian vegetation rather than aquatic variables. On‐site assessments revealed that restored reaches had significantly lower stream widths and greater depths than did upstream unrestored reaches, but riparian canopy cover often was lower in restored than in unrestored reaches. This study provides basic information on midwestern restoration strategies, which is needed to identify strengths and weaknesses in current practices and to better inform future stream restorations.     

Natural‐Channel‐Design Restorations That Changed Geomorphology Have Little Effect on Macroinvertebrate Communities in Headwater Streams

Stream restorations that increase geomorphic stability can improve habitat quality, which should benefit selected species and local aquatic ecosystems. This assumption is often used to define primary restoration goals; yet, biological responses to restoration are rarely monitored or evaluated methodically. Macroinvertebrate communities were inventoried at 6 study reaches within 5 Catskill Mountain streams between 2002 and 2006 to characterize their responses to natural‐channel‐design (NCD) restoration. Although bank stability increased significantly at most restored reaches, analyses of variation showed that NCD restorations had no significant effect on 15 of 16 macroinvertebrate community metrics. Multidimensional scaling ordination indicated that communities from all reach types within a stream were much more similar to each other within any given year than they were in the same reaches across years or within any type of reach across streams. These findings indicate that source populations and watershed‐scale factors were more important to macroinvertebrate community characteristics than were changes in channel geomorphology associated with NCD restoration. Furthermore, the response of macroinvertebrates to restoration cannot always be used to infer the response of other stream biota to restoration. Thus, a broad perspective is needed to characterize and evaluate the full range of effects that restoration can have on stream ecosystems.     

Host thermoregulatory constraints predict growth of an amphibian chytrid pathogen (Batrachochytrium dendrobatidis)

1. The course and outcome of many wildlife diseases are context-dependent, and therefore change depending on the behaviour of hosts and environmental response of the pathogen.2. Contemporary declines in amphibian populations are widely attributed to chytridiomycosis, caused by the pathogenic fungus Batrachochytrium dendrobatidis. Despite the thermal sensitivity of the pathogen and its amphibian hosts, we do not understand how host thermal regimes experienced by frogs in the wild directly influence pathogen growth.3. We tested how thermal regimes experienced by the rainforest frog Litoria rheocola in the wild influence pathogen growth in the laboratory, and whether these responses differ from pathogen growth under available environmental thermal regimes.4. Frog thermal regimes mimicked in the laboratory accelerated pathogen growth during conditions representative of winter at high elevations more so than if temperatures matched air or stream water temperatures. By contrast, winter frog thermal regimes at low elevations slowed pathogen growth relative to air temperatures, but not water temperatures.5. The growth pattern of the fungus under frog thermal regimes matches field prevalence and intensity of infections for this species (high elevation winter > high elevation summer > low elevation winter > low elevation summer), whereas pathogen growth trajectories under environmental temperatures did not match these patterns.6. If these laboratory results translate into field responses, tropical frogs may be able to reduce disease impacts by regulating their body temperatures to limit pathogen growth (e.g., by using microhabitats that facilitate basking to reach high temperatures); in other cases, the environment may limit the ability of frogs to thermoregulate such that individuals are more vulnerable to this pathogen (e.g., in dense forests at high elevations).7. Species-specific thermoregulatory behaviour, and interactions with and constraints imposed by the environment, are therefore essential to understanding and predicting the spatial and temporal impacts of this global disease.     

Probing whole‐stream metabolism: influence of spatial heterogeneity on rate estimates

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Climate Change Simulations Predict Altered Biotic Response in a Thermally Heterogeneous Stream System

Climate change is predicted to increase water temperatures in many lotic systems, but little is known about how changes in air temperature affect lotic systems heavily influenced by groundwater. Our objectives were to document spatial variation in temperature for spring-fed Ozark streams in Southern Missouri USA, create a spatially explicit model of mean daily water temperature, and use downscaled climate models to predict the number of days meeting suitable stream temperature for three aquatic species of concern to conservation and management. Longitudinal temperature transects and stationary temperature loggers were used in the Current and Jacks Fork Rivers during 2012 to determine spatial and temporal variability of water temperature. Groundwater spring influence affected river water temperatures in both winter and summer, but springs that contributed less than 5% of the main stem discharge did not affect river temperatures beyond a few hundred meters downstream. A multiple regression model using variables related to season, mean daily air temperature, and a spatial influence factor (metric to account for groundwater influence) was a strong predictor of mean daily water temperature (r² = 0.98; RMSE = 0.82). Data from two downscaled climate simulations under the A2 emissions scenario were used to predict daily water temperatures for time steps of 1995, 2040, 2060, and 2080. By 2080, peak numbers of optimal growth temperature days for smallmouth bass are expected to shift to areas with more spring influence, largemouth bass are expected to experience more optimal growth days (21 – 317% increase) regardless of spring influence, and Ozark hellbenders may experience a reduction in the number of optimal growth days in areas with the highest spring influence. Our results provide a framework for assessing fine-scale (10 s m) thermal heterogeneity and predict shifts in thermal conditions at the watershed and reach scale.     

Thermal adaptation and phenotypic plasticity in a warming world: Insights from common garden experiments on Alaskan sockeye salmon

An important unresolved question is how populations of coldwater‐dependent fishes will respond to rapidly warming water temperatures. For example, the culturally and economically important group, Pacific salmon (Oncorhynchus spp.), experience site‐specific thermal regimes during early development that could be disrupted by warming. To test for thermal local adaptation and heritable phenotypic plasticity in Pacific salmon embryos, we measured the developmental rate, survival, and body size at hatching in two populations of sockeye salmon (Oncorhynchus nerka) that overlap in timing of spawning but incubate in contrasting natural thermal regimes. Using a split half‐sibling design, we exposed embryos of 10 families from each of two populations to variable and constant thermal regimes. These represented both experienced temperatures by each population, and predicted temperatures under plausible future conditions based on a warming scenario from the downscaled global climate model (MIROC A1B scenario). We did not find evidence of thermal local adaptation during the embryonic stage for developmental rate or survival. Within treatments, populations hatched within 1 day of each other, on average, and among treatments, did not differ in survival in response to temperature. We did detect plasticity to temperature; embryos developed 2.5 times longer (189 days) in the coolest regime compared to the warmest regime (74 days). We also detected variation in developmental rates among families within and among temperature regimes, indicating heritable plasticity. Families exhibited a strong positive relationship between thermal variability and phenotypic variability in developmental rate but body length and mass at hatching were largely insensitive to temperature. Overall, our results indicated a lack of thermal local adaptation, but a presence of plasticity in populations experiencing contrasting conditions, as well as family‐specific heritable plasticity that could facilitate adaptive change.