Parallel shifts in trout feeding morphology suggest rapid adaptation to alpine lake environments.

Eco-evolutionary interactions following ecosystem change provide critical insight into the ability of organisms to adapt to shifting resource landscapes. Here we explore evidence for the rapid parallel evolution of trout feeding morphology following eco-evolutionary interactions with zooplankton in alpine lakes stocked at different points in time in the Wind River Range (Wyoming, USA). In this system, trout predation has altered the zooplankton species community and driven a decrease in average zooplankton size. In some lakes that were stocked decades ago, we find shifts in gill raker traits consistent with the hypothesis that trout have rapidly adapted to exploit available smaller-bodied zooplankton more effectively. We explore this morphological response in multiple lake populations across two species of trout (cutthroat trout, Oncorhynchus clarkii, and golden trout Oncorhynchus aguabonita) and examine the impact of resource availability on morphological variation in gill raker number among lakes. Furthermore, we present genetic data to provide evidence that historically stocked cutthroat trout populations likely derive from multiple population sources, and incorporate variation from genomic relatedness in our exploration of environmental predictors of feeding morphology. These findings describe rapid adaptation and eco-evolutionary interactions in trout and document an evolutionary response to novel, contemporary ecosystem change.

Simple statistical models can be sufficient for testing hypotheses with population time-series data.

Time-series data offer wide-ranging opportunities to test hypotheses about the physical and biological factors that influence species abundances. Although sophisticated models have been developed and applied to analyze abundance time series, they require information about species detectability that is often unavailable. We propose that in many cases, simpler models are adequate for testing hypotheses. We consider three relatively simple regression models for time series, using simulated and empirical (fish and mammal) datasets. Model A is a conventional generalized linear model of abundance, model B adds a temporal autoregressive term, and model C uses an estimate of population growth rate as a response variable, with the option of including a term for density dependence. All models can be fit using Bayesian and non-Bayesian methods. Simulation results demonstrated that model C tended to have greater support for long-lived, lower-fecundity organisms (K life-history strategists), while model A, the simplest, tended to be supported for shorter-lived, high-fecundity organisms (r life-history strategists). Analysis of real-world fish and mammal datasets found that models A, B, and C each enjoyed support for at least some species, but sometimes yielded different insights. In particular, model C indicated effects of predictor variables that were not evident in analyses with models A and B. Bayesian and frequentist models yielded similar parameter estimates and performance. We conclude that relatively simple models are useful for testing hypotheses about the factors that influence abundance in time-series data, and can be appropriate choices for datasets that lack the information needed to fit more complicated models. When feasible, we advise fitting datasets with multiple models because they can provide complementary information.

Hybridization decreases native cutthroat trout reproductive fitness.

Examining natural selection in wild populations is challenging, but crucial to understanding many ecological and evolutionary processes. Additionally, in hybridizing populations, natural selection may be an important determinant of the eventual outcome of hybridization. We characterized several components of relative fitness in hybridizing populations of Yellowstone cutthroat trout and rainbow trout in an effort to better understand the prolonged persistence of both parental species despite predictions of extirpation. Thousands of genomic loci enabled precise quantification of hybrid status in adult and subsequent juvenile generations; a subset of those data also identified parent-offspring relationships. We used linear models and simulations to assess the effects of ancestry on reproductive output and mate choice decisions. We found a relatively low number of late-stage (F3+) hybrids and an excess of F2 juveniles relative to the adult generation in one location, which suggests the presence of hybrid breakdown decreasing the fitness of F2+ hybrids later in life. Assessments of reproductive output showed that Yellowstone cutthroat trout are more likely to successfully reproduce and produce slightly more offspring than their rainbow trout and hybrid counterparts. Mate choice appeared to be largely random, though we did find statistical support for slight female preference for males of similar ancestry. Together, these results show that native Yellowstone cutthroat trout are able to outperform rainbow trout in terms of reproduction and suggest that management action to exclude rainbow trout from spawning locations may bolster the now-rare Yellowstone cutthroat trout.

Infection Status as the Basis for Habitat Choices in a Wild Amphibian.

AbstractAnimals challenged with disease may select specific habitat conditions that help prevent or reduce infection. Whereas preinfection avoidance of habitats with a high risk of disease exposure has been documented in both captive and free-ranging animals, evidence of switching habitats after infection to support the clearing of the infection is limited to laboratory experiments. The extent to which wild animals proximately modify habitat choices in response to infection status thus remains unclear. We investigated preinfection behavioral avoidance and postinfection habitat switching using wild, radio-tracked boreal toads (Anaxyrus boreas boreas) in a population challenged with Batrachochytrium dendrobatidis (Bd), a pathogenic fungus responsible for a catastrophic panzootic affecting hundreds of amphibian species worldwide. Boreal toads did not preemptively avoid microhabitats with conditions conducive to Bd growth. Infected individuals, however, selected warmer, more open habitats, which were associated with elevated body temperature and the subsequent clearing of infection. Our results suggest that disease can comprise an important selective pressure on animal habitat and space use. Habitat selection models, therefore, may be greatly improved by including variables that quantify infection risk and/or the infection status of individuals through time.

Warmer temperatures interact with salinity to weaken physiological facilitation to stress in freshwater fishes.

Management of stressors requires an understanding of how multiple stressors interact, how different species respond to those interactions and the underlying mechanisms driving observed patterns in species' responses. Salinization and rising temperatures are two pertinent stressors predicted to intensify in freshwater ecosystems, posing concern for how susceptible organisms achieve and maintain homeostasis (i.e. allostasis). Here, glucocorticoid hormones (e.g. cortisol), responsible for mobilizing energy (e.g. glucose) to relevant physiological processes for the duration of stressors, are liable to vary in response to the duration and severity of salinization and temperature rises. With field and laboratory studies, we evaluated how both salinity and temperature influence basal and stress-reactive cortisol and glucose levels in age 1+ mottled sculpin (Cottus bairdii), mountain sucker (Catostomus platyrhynchus) and Colorado River cutthroat trout (Oncorhynchus clarki pleuriticus). We found that temperature generally had the greatest effect on cortisol and glucose concentrations and the effect of salinity was often temperature dependent. We also found that when individuals were chronically exposed to higher salinities, baseline concentrations of cortisol and glucose usually declined as salinity increased. Reductions in baseline concentrations facilitated stronger stress reactivity for cortisol and glucose when exposed to additional stressors, which weakened as temperatures increased. Controlled temperatures near the species' thermal maxima became the overriding factor regulating fish physiology, resulting in inhibitory responses. With projected increases in freshwater salinization and temperatures, efforts to reduce the negative effects of increasing temperatures (i.e. increased refuge habitats and riparian cover) could moderate the inhibitory effects of temperature-dependent effects of salinization for freshwater fishes.

Variable hybridization outcomes in trout are predicted by historical fish stocking and environmental context.

Hybridization can profoundly affect the genomic composition and phenotypes of closely related species, and provides an opportunity to identify mechanisms that maintain reproductive isolation between species. Recent evidence suggests that hybridization outcomes within a species pair can vary across locations. However, we still do not know how variable outcomes of hybridization are across geographic replicates, and what mechanisms drive that variation. In this study, we described hybridization outcomes across 27 locations in the North Fork Shoshone River basin (Wyoming, USA) where native Yellowstone cutthroat trout and introduced rainbow trout co-occur. We used genomic data and hierarchical Bayesian models to precisely identify ancestry of hybrid individuals. Hybridization outcomes varied across locations. In some locations, only rainbow trout and advanced backcrossed hybrids towards rainbow trout were present, while trout in other locations had a broader range of ancestry, including both parental species and first-generation hybrids. Later-generation intermediate hybrids were rare relative to backcrossed hybrids and rainbow trout individuals. Using an individual-based simulation, we found that outcomes of hybridization in the North Fork Shoshone River basin deviate substantially from what we would expect under null expectations of random mating and no selection against hybrids. Since this deviation implies that some mechanisms of reproductive isolation function to maintain parental taxa and a diversity of hybrid types, we then modelled hybridization outcomes as a function of environmental variables and stocking history that are likely to affect prezygotic barriers to hybridization. Variables associated with history of fish stocking were the strongest predictors of hybridization outcomes, followed by environmental variables that might affect overlap in spawning time and location.

Multiple approaches to surface water quality assessment provide insight for small streams experiencing oil and natural gas development.

Historic, current, and future oil and natural gas development can affect water quality in streams flowing through developed areas. We compared small stream drainages in a semiarid landscape with varying amounts of disturbance from oil and natural gas development to examine potential effects of this development on surface water quality. We used physical, chemical, and biological approaches to assess water quality and found several potential avenues of degradation. Surface disturbance likely contributed to elevated suspended sediment concentrations and spill history likely led to elevated stream polycyclic aromatic hydrocarbon concentrations. In combination, these environmental stressors could explain the loss of aquatic macroinvertebrate taxon at sites highly affected by oil and natural gas development. Our results provide insight into advantages and disadvantages of approaches for assessing surface water quality in areas affected by oil and natural gas development. Integr Environ Assess Manag 2019;00:000-000. © 2019 SETAC.

The interaction of exposure and warming tolerance determines fish species vulnerability to warming stream temperatures.

Species vulnerability to climate change involves an interaction between the magnitude of change (exposure) and a species's tolerance to change. We evaluated fish species vulnerability to predicted stream temperature increases by examining warming tolerances across the Wyoming fish assemblage. Warming tolerance combines stream temperature with a thermal tolerance metric to estimate how much warming beyond current conditions a species can withstand. Brown trout, rainbow trout and burbot had the lowest warming tolerances and the highest proportion of currently occupied sites that will become unsuitable under predicted temperature increases. These most vulnerable species were coldwater species, but had neither the lowest thermal tolerances nor would they experience the greatest temperature increases. Our results highlight the importance of considering the interaction of exposure and warming tolerance when predicting climate change vulnerability and demonstrate an approach that can be applied broadly.

The effectiveness of surrogate taxa to conserve freshwater biodiversity.

Establishing protected areas has long been an effective conservation strategy and is often based on readily surveyed species. The potential of any freshwater taxa to be a surrogate for other aquatic groups has not been explored fully. We compiled occurrence data on 72 species of freshwater fishes, amphibians, mussels, and aquatic reptiles for the Great Plains, Wyoming (U.S.A.). We used hierarchical Bayesian multispecies mixture models and MaxEnt models to describe species' distributions and the program Zonation to identify areas of conservation priority for each aquatic group. The landscape-scale factors that best characterized aquatic species' distributions differed among groups. There was low agreement and congruence among taxa-specific conservation priorities (<20%), meaning no surrogate priority areas would include or protect the best habitats of other aquatic taxa. Common, wideranging aquatic species were included in taxa-specific priority areas, but rare freshwater species were not included. Thus, the development of conservation priorities based on a single freshwater aquatic group would not protect all species in the other aquatic groups.

Stream Vulnerability to Widespread and Emergent Stressors: A Focus on Unconventional Oil and Gas.

Multiple stressors threaten stream physical and biological quality, including elevated nutrients and other contaminants, riparian and in-stream habitat degradation and altered natural flow regime. Unconventional oil and gas (UOG) development is one emerging stressor that spans the U.S. UOG development could alter stream sedimentation, riparian extent and composition, in-stream flow, and water quality. We developed indices to describe the watershed sensitivity and exposure to natural and anthropogenic disturbances and computed a vulnerability index from these two scores across stream catchments in six productive shale plays. We predicted that catchment vulnerability scores would vary across plays due to climatic, geologic and anthropogenic differences. Across-shale averages supported this prediction revealing differences in catchment sensitivity, exposure, and vulnerability scores that resulted from different natural and anthropogenic environmental conditions. For example, semi-arid Western shale play catchments (Mowry, Hilliard, and Bakken) tended to be more sensitive to stressors due to low annual average precipitation and extensive grassland. Catchments in the Barnett and Marcellus-Utica were naturally sensitive from more erosive soils and steeper catchment slopes, but these catchments also experienced areas with greater UOG densities and urbanization. Our analysis suggested Fayetteville and Barnett catchments were vulnerable due to existing anthropogenic exposure. However, all shale plays had catchments that spanned a wide vulnerability gradient. Our results identify vulnerable catchments that can help prioritize stream protection and monitoring efforts. Resource managers can also use these findings to guide local development activities to help reduce possible environmental effects.

Species- and community-level responses combine to drive phenology of lake phytoplankton.

Global change is leading to shifts in the seasonal timing of growth and maturation for primary producers. Remote sensing is increasingly used to measure the timing of primary production in both aquatic and terrestrial ecosystems, but there is often a poor correlation between these results and direct observations of life-history responses of individual species. One explanation may be that, in addition to phenological shifts, global change is also causing shifts in community composition among species with different seasonal timing of growth and maturation. We quantified how shifts in species phenology and in community composition translated into phenological change in a diverse phytoplankton community from 1962 to 2000. During this time, the aggregate community spring-summer phytoplankton peak has shifted 63 days earlier. The mean taxon shift was only 3 days earlier, and shifts in taxa phenology explained only 40% of the observed community phenological shift. The remaining community shift was attributed to dominant early-season taxa increasing in abundance while a dominant late-season taxon decreased in abundance. In diverse producer communities experiencing multiple stressors, changes in species composition must be considered to fully understand and predict shifts in the seasonal timing of primary production.

Interactive effects of water diversion and climate change for juvenile chinook salmon in the lemhi river basin (USA.).

The combined effects of water diversion and climate change are a major conservation challenge for freshwater ecosystems. In the Lemhi Basin, Idaho (U.S.A.), water diversion causes changes in streamflow, and climate change will further affect streamflow and temperature. Shifts in streamflow and temperature regimes can affect juvenile salmon growth, movement, and survival. We examined the potential effects of water diversion and climate change on juvenile Chinook salmon (Oncorhynchus tshawytscha), a species listed as threatened under the U.S. Endangered Species Act (ESA). To examine the effects for juvenile survival, we created a model relating 19 years of juvenile survival data to streamflow and temperature and found spring streamflow and summer temperature were good predictors of juvenile survival. We used these models to project juvenile survival for 15 diversion and climate-change scenarios. Projected survival was 42-58% lower when streamflows were diverted than when streamflows were undiverted. For diverted streamflows, 2040 climate-change scenarios (ECHO-G and CGCM3.1 T47) resulted in an additional 11-39% decrease in survival. We also created models relating habitat carrying capacity to streamflow and made projections for diversion and climate-change scenarios. Habitat carrying capacity estimated for diverted streamflows was 17-58% lower than for undiverted streamflows. Climate-change scenarios resulted in additional decreases in carrying capacity for the dry (ECHO-G) climate model. Our results indicate climate change will likely pose an additional stressor that should be considered when evaluating the effects of anthropogenic actions on salmon population status. Thus, this type of analysis will be especially important for evaluating effects of specific actions on a particular species. Efectos Interactivos de la Desviación del Agua y el Cambio Climático en Individuos Juveniles de Salmón Chinook en la Cuenca del Río Lemhi (E.U.A.).

How low can you go? Impacts of a low-flow disturbance on aquatic insect communities.

The natural hydrology of streams and rivers is being extensively modified by human activities. Water diversion, dam construction, and climate change have the potential to increase the frequency and intensity of low-flow events. Flow is a dominant force structuring stream aquatic insect communities, but the impacts of water diversion are poorly understood. Here we report results of an experimental stream flow diversion designed to test how aquatic insect communities respond to a low-flow disturbance. We diverted 40% to 80% of the water in three replicate streams for three summers, leading to summer flow exceedance probabilities of up to 99.9%. Shifts in habitat availability appeared to be a major driver of aquatic insect community responses. Responses also varied by habitat type: total insect density decreased in riffle habitats, but there was no change in pool habitats. Overall, the total biomass of aquatic insects decreased sharply with lowered flow. Collector-filterers, collector-gatherers, and scrapers were especially susceptible, while predatory insects were more resistant. Despite extremely low flow levels, there was no shift in aquatic insect family richness. The experimental water withdrawal did not increase water temperature or decrease water quality, and some wetted habitat was always maintained, which likely prevented more severe impacts on aquatic insect communities.

An experimental disturbance alters fish size structure but not food chain length in streams.

Streams experience frequent natural disturbance and are undergoing considerable anthropogenic disturbance due to dam construction and water diversion. Disturbance is known to impact community structure, but its effect on food chain length is still a matter of considerable debate. Theoretical models show that longer food chains are less resilient to disturbance, so food chain length is predicted to be shorter following a disturbance event. Here we experimentally test the effect of disturbance on food chain length in streams by diverting stream flow. We found that our experimental low-flow disturbance did not alter food chain length. We did see an effect on body-size structure in our food webs suggesting that food chain length may be an insensitive indicator of disturbance. We suggest that habitat heterogeneity and food web complexity buffer the effect of disturbance on food chain length. The theoretical predictions of disturbance on food chain length are only likely to be seen in homogeneous systems that closely approximate the linear food chains the models are based upon.