Dietary overlap and selectivity among mountain steppe river fish in the United States and Mongolia.

Lotic systems in mountain regions have historically provided secure habitat for native fish populations because of their relative isolation from human settlement and lack of upstream disturbances. However, rivers of mountain ecoregions are currently experiencing heightened levels of disturbance due to the introduction of nonnative species impacting endemic fishes in these areas. We compared the fish assemblages and diets of mountain steppe fishes of the stocked rivers in Wyoming with rivers in northern Mongolia where stocking is absent. Using gut content analysis, we quantified the selectivity and diets of fishes collected in these systems. Nonnative species had more generalist diets with lower levels of selectivity than most native species and native species had high levels of dietary specificity and selectivity. High abundances of nonnative species and high levels of dietary overlaps in our Wyoming sites is a cause of concern for native Cutthroat Trout and overall system stability. In contrast, fish assemblages characterizing Mongolia mountain steppe rivers were composed of only native species with diverse diets and higher selectivity values, suggesting low probability for interspecific competition.

Intercontinental analysis of temperate steppe stream food webs reveals consistent autochthonous support of fishes.

Quantifying the trophic basis of production for freshwater metazoa at broad spatial scales is key to understanding ecosystem function and has been a research priority for decades. However, previous lotic food web studies have been limited by geographic coverage or methodological constraints. We used compound-specific stable carbon isotope analysis of amino acids (AAs) to estimate basal resource contributions to fish consumers in streams spanning grassland, montane and semi-arid ecoregions of the temperate steppe biome on two continents. Across a range of stream sizes and light regimes, we found consistent trophic importance of aquatic resources. Essential AAs of heterotrophic microbial origin generally provided secondary support for fishes, while terrestrial carbon did not seem to provide significant, direct support. These findings provide strong evidence for the dominant contribution of carbon to higher-order consumers by aquatic autochthonous resources (primarily) and heterotrophic microbial communities (secondarily) in temperate steppe streams.

Identifying monitoring information needs that support the management of fish in large rivers.

Management actions intended to benefit fish in large rivers can directly or indirectly affect multiple ecosystem components. Without consideration of the effects of management on non-target ecosystem components, unintended consequences may limit management efficacy. Monitoring can help clarify the effects of management actions, including on non-target ecosystem components, but only if data are collected to characterize key ecosystem processes that could affect the outcome. Scientists from across the U.S. convened to develop a conceptual model that would help identify monitoring information needed to better understand how natural and anthropogenic factors affect large river fishes. We applied the conceptual model to case studies in four large U.S. rivers. The application of the conceptual model indicates the model is flexible and relevant to large rivers in different geographic settings and with different management challenges. By visualizing how natural and anthropogenic drivers directly or indirectly affect cascading ecosystem tiers, our model identified critical information gaps and uncertainties that, if resolved, could inform how to best meet management objectives. Despite large differences in the physical and ecological contexts of the river systems, the case studies also demonstrated substantial commonalities in the data needed to better understand how stressors affect fish in these systems. For example, in most systems information on river discharge and water temperature were needed and available. Conversely, information regarding trophic relationships and the habitat requirements of larval fishes were generally lacking. This result suggests that there is a need to better understand a set of common factors across large-river systems. We provide a stepwise procedure to facilitate the application of our conceptual model to other river systems and management goals.

Geomorphology variables predict fish assemblages for forested and endorheic rivers of two continents.

Stream fishes are restricted to specific environments with appropriate habitats for feeding and reproduction. Interactions between streams and surrounding landscapes influence the availability and type of fish habitat, nutrient concentrations, suspended solids, and substrate composition. Valley width and gradient are geomorphological variables that influence the frequency and intensity that a stream interacts with the surrounding landscape. For example, in constrained valleys, canyon walls are steeply sloped and valleys are narrow, limiting the movement of water into riparian zones. Wide valleys have long, flat floodplains that are inundated with high discharge. We tested for differences in fish assemblages with geomorphology variation among stream sites. We selected rivers in similar forested and endorheic ecoregion types of the United States and Mongolia. Sites where we collected were defined as geomorphologically unique river segments (i.e., functional process zones; FPZs) using an automated ArcGIS-based tool. This tool extracts geomorphic variables at the valley and catchment scales and uses them to cluster stream segments based on their similarity. We collected a representative fish sample from replicates of FPZs. Then, we used constrained ordinations to determine whether river geomorphology could predict fish assemblage variation. Our constrained ordination approach using geomorphology to predict fish assemblages resulted in significance using fish taxonomy and traits in several watersheds. The watersheds where constrained ordinations were not successful were next analyzed with unconstrained ordinations to examine patterns among fish taxonomy and traits with geomorphology variables. Common geomorphology variables as predictors for taxonomic fish assemblages were river gradient, valley width, and valley slope. Significant geomorphology predictors of functional traits were valley width-to-floor width ratio, elevation, gradient, and channel sinuosity. These results provide evidence that fish assemblages respond similarly and strongly to geomorphic variables on two continents.

Valley-scale hydrogeomorphology drives river fish assemblage variation in Mongolia.

River hydrogeomorphology is a major driver shaping biodiversity and community composition. Here, we examine how hydrogeomorphic heterogeneity expressed by Functional Process Zones (FPZs) in river networks is associated with fish assemblage variation. We examined this association in two distinct ecoregions in Mongolia expected to display different gradients of river network hydrogeomorphic heterogeneity. We delineated FPZs by extracting valley-scale hydrogeomorphic variables at 10 km sample intervals in forest steppe (FS) and in grassland (G) river networks. We sampled fish assemblages and examined variation associated with changes in gradients of hydrogeomorphology as expressed by the FPZs. Thus, we examined assemblage variation as patterns of occurrence- and abundance-based beta diversities for the taxonomic composition of assemblages and as functional beta diversity. Overall, we delineated 5 and 6 FPZs in river networks of the FS and G, respectively. Eight fish species were found in the FS river network and seventeen in the G, four of them common to both ecoregions. Functional richness was correspondingly higher in the G river network. Variation in the taxonomic composition of assemblages was driven by species turnover and was only significant in the G river network. Abundance-based taxonomic variation was significant in river networks of both ecoregions, while the functional beta diversity results were inconclusive. We show that valley-scale hydrogeomorphology is a significant driver of variation in fish assemblages at a macrosystem scale. Both changes in the composition of fish assemblages and the carrying capacity of the river network were driven by valley-scale hydrogeomorphic variables. River network hydrogeomorphology as accounted for in the study has, therefore, the potential to inform macrosystem scale community ecology research and conservation efforts.

Geomorphological characteristics of the Wabash River, USA: Influence on fish assemblages.

River hydrogeomorphology is a potential predictor of ecosystem and assemblage variation. We tested for fish assemblage variation as a function of hydrogeomorphology in a Midwestern US large river, the Wabash River. Fish data were classified by taxonomy and traits and we tested if assemblages varied with river hydrogeomorphology or river distance, defined into 10-km distinct reaches. Three unique geomorphological units, Functional Process Zones (FPZ), were identified using an ArcGIS hydrogeomorphic model, based primarily on channel width, floodplain width, and down valley slope. Five locations were identified as FPZ A with narrow stream channel, high down valley slope, and an expansive floodplain. Ten locations were identified as FPZ B with a wide river channel and wide floodplain. Thirty-five locations were identified as FPZ C with wide river channel and a constrained floodplain. The sites were categorized into three stream orders: 5, 6, and 7. We found hydrogeomorphology classified by unique FPZs or by river distance influenced taxonomic and functional fish assemblages for the Wabash River. There was high overlap among fish occurrences among FPZs, but nine species resulted as significant indicators of specific FPZs. Five traits were significant indicators of FPZs: an intermediate Swim Factor score, medium tolerance to silt, small-large stream size preference, and two Shape Factor categories. Our conclusions are that fish assemblages respond strongly to local geomorphology and river distance, fitting the riverine ecosystem synthesis and the river continuum concept.

Diet overlap among non-native trout species and native cutthroat Trout (Oncorhynchus clarkii) in two U.S. ecoregions.

The invasion of freshwater ecosystems by non-native species can constitute a significant threat to native species and ecosystem health. Non-native trouts have long been stocked in areas where native trouts occur and have negatively impacted native trouts through predation, competition, and hybridization. This study encompassed two seasons of sampling efforts across two ecoregions of the western United States: The Great Basin in summer 2016 and the Yellowstone River Basin in summer 2017. We found significant dietary overlaps among native and non-native trouts within the Great Basin and Yellowstone River Basin ecoregions. Three orders of invertebrates (Ephemeroptera, Trichoptera, and Diptera) composed the majority of stomach contents and were responsible for driving the observed patterns. Great Basin trout had higher body conditions (k), and non-native Great Basin trout had higher gut fullness values than Yellowstone River Basin trout, indicating a possible limitation of food in the Yellowstone River Basin. Native fishes were the least abundant and had the lowest body condition in each ecoregion. These findings may indicate a negative impact on native trouts by non-native trouts. We recommend additional monitoring of native and non-native trout diets, regular invertebrate surveys to identify the availability of diet items, and reconsidering stocking efforts that can result in overlap of non-native fishes with native cutthroat trout.

Dietary specificity and overlap in endorheic river fishes: How do native and nonnative species compare?

The introduction of nonnative species is one of the most critical problems facing freshwater systems today. The rivers of the Great Basin (USA) have been particularly imperilled by nonnative species introductions and represent a valuable location to study the dietary trends of native and nonnative fishes in isolated, endorheic systems. We collected fish from 23 sites, spanning three Great Basin watersheds (Carson, Humboldt and Bear Rivers) and two elevation categories (upland and lowland). Only a single species (speckled dace Rhinichthys osculus) occurred in both elevation zones. Diet item analyses of over 500 fish stomachs indicated significant dietary overlaps between native and nonnative fishes and detailed dietary selectivity for all species. This finding, along with the low species diversity observed in the region, suggests low dietary niche diversity, which could have the potential to amplify the competitive impacts of nonnatives on native species. In upland sites, nonnative trouts were the dominant invaders, while in lowland sites warm-water nonnatives were prevalent. The management implications we recommend based on our results urge for continued monitoring of water temperature and species occurrences to predict if dietary overlaps observed in this study are likely to change in the future. SIGNIFICANCE STATEMENT: The Great Basin is an ideal endorheic region to study dietary trends in native and nonnative fishes. These trends are important in predicting competitive interactions among fishes. By looking at the diets of fishes within this region we were able to identify multiple significant overlaps among native and nonnative fishes. These results represent a baseline for future studies in the region as well as being comparable to other regions with similar invasive/native overlaps.

Correction: Long-term fish assemblages of the Ohio River: Altered trophic and life history strategies with hydrologic alterations and land use modifications.

[This corrects the article DOI: 10.1371/journal.pone.0211848.].

Long-term fish assemblages of the Ohio River: Altered trophic and life history strategies with hydrologic alterations and land use modifications.

Long-term monitoring of species assemblages provides a unique opportunity to test hypotheses regarding environmentally induced directional trajectories of freshwater species assemblages. We used 57 years of lockchamber fish rotenone and boat electrofishing survey data (1957-2014) collected by the Ohio River Valley Water Sanitation Commission (ORSANCO) to test for directional trajectories in taxonomy, trophic classifications, and life history strategies of freshwater fish assemblages in the Ohio River Basin. We found significant changes in taxonomic and trophic composition of freshwater fishes in the Ohio River Basin. Annual species richness varied from 31 to 90 species and generally increased with year. Temporal trajectories were present for taxonomic and trophic assemblages. Assemblage structure based on taxonomy was correlated with land use change (decrease in agriculture and increase in forest). Taxonomic assemblage structure was also correlated with altered hydrology variables of increased minimum discharge, decreased fall rate, and increased rise rate. Trophic composition of fish catch correlated with land use change (decrease in agriculture and increase in forest) and altered hydrology. Altered hydrology of increased minimum discharge, increased fall discharge, decreased base flows, and increased number of high pulse events was correlated with increased counts of herbivore-detritivores and decreased counts of piscivores and planktivores. We did not find directional changes in life history composition. We hypothesized a shift occurred from benthic to phytoplankton production throughout the basin that may have decreased secondary production of benthic invertebrates. This may also be responsible for lower trophic position of invertivore and piscivore fishes observed in other studies.

A century of morphological variation in Cyprinidae fishes.

BACKGROUND: Aquatic habitats have been altered over the past century due to a variety of anthropogenic influences. Ecomorphology is an area of aquatic ecology that can both directly and indirectly assess the effects of habitat alterations on organisms. However, few studies have explored long term trends in morphological variation. Long term changes in morphology can potentially impact niche and ultimately contribute to organismal success and the ecosystem. Therefore, in this study we assessed long term morphological variation with body size, sex, time, and hydrology using museum collections of five species of Cyprinidae (Minnows) from lentic and lotic systems over the past 100 years to gain insight into long term patterns in morphology. RESULTS: Variation in Cyprinidae morphology tended to relate to: body size-indicating strong allometric growth patterns with robustness of larger individuals; sex-indicating a level of fecundity selection for deeper bodies in females compared with males; and year-indirectly suggesting responses to habitat changes over the past century. In lotic ecosystems, Cyprinidae morphology tended to be more fusiform in conjunction with lower mean annual discharge or higher variation in discharge. In lentic ecosystems, change in morphology was observed but no historic habitat variables were available to discern potential mechanisms. Interestingly, not all species responded in the same magnitude or directionality. CONCLUSIONS: Long term changes in morphological variation provide a link to exploring functional relationships between taxa and their environment and have implications for understanding ecosystem attributes, community assembly patterns, and conservation.

Shift in a large river fish assemblage: body-size and trophic structure dynamics.

As the intensity and speed of environmental change increase at both local and global scales it is imperative that we gain a better understanding of the ecological implications of community shifts. While there has been substantial progress toward understanding the drivers and subsequent responses of community change (e.g. lake trophic state), the ecological impacts of food web changes are far less understood. We analyzed Wabash River fish assemblage data collected from 1974-2008, to evaluate temporal variation in body-size structure and functional group composition. Two parameters derived from annual community size-spectra were our major response variables: (1) the regression slope is an index of ecological efficiency and predator-prey biomass ratios, and (2) spectral elevation (regression midpoint height) is a proxy for food web capacity. We detected a large assemblage shift, over at least a seven year period, defined by dramatic changes in abundance (measured as catch-per-unit-effort) of the dominant functional feeding groups among two time periods; from an assemblage dominated by planktivore-omnivores to benthic invertivores. There was a concurrent increase in ecological efficiency (slopes increased over time) following the shift associated with an increase in large-bodied low trophic level fish. Food web capacity remained relatively stable with no clear temporal trends. Thus, increased ecological efficiency occurred simultaneous to a compensatory response that shifted biomass among functional feeding groups.

Assessing changes in the Presque Isle Bay watershed fish community using a modified index of biotic integrity: before and after the elimination of combined sewer overflows.

An index of biotic integrity and species richness were used to assess changes in the Presque Isle Bay watershed fish community before and after the elimination of combined sewer overflows (CSOs). The fish community was sampled with a backpack electrofisher in 2011 at 12 stream locations on 4 tributaries of Presque Isle Bay, Erie County, Pennsylvania. All sites were previously sampled in 2001. Significant increases in species richness and index of biotic integrity (IBI) scores were observed in 2011 compared to 2001. The significant increases in species richness and IBI scores occurred following the elimination of 10 CSOs to Garrison Run, 7 CSOs to Cascade Creek, and 37 CSOs to Mill Creek. Despite the increased richness and IBI scores, the fish community remains in poor condition, which may be related to the highly urbanized land use of the watershed. Urban land uses comprise 77% of the Presque Isle Bay watershed, and in both 2011 and 2001, the watershed as a whole did not meet warm-water habitat criteria. It is unlikely that the fish community will continue to recover without addressing urbanization throughout the watershed.

Role of the fish Astyanax aeneus (Characidae) as a keystone nutrient recycler in low-nutrient neotropical streams.

Nutrient recycling by animals is a potentially important biogeochemical process in both terrestrial and aquatic ecosystems. Stoichiometric traits of individual species may result in some taxa playing disproportionately important roles in the recycling of nutrients relative to their biomass, acting as keystone nutrient recyclers. We examined factors controlling the relative contribution of 12 Neotropical fish species to nutrient recycling in four streams spanning a range of phosphorus (P) levels. In high-P conditions (135 microg/L soluble reactive phosphorus, SRP), most species fed on P-enriched diets and P excretion rates were high across species. In low-P conditions (3 microg/L SRP), aquatic food resources were depleted in P, and species with higher body P content showed low rates of P recycling. However, fishes that were subsidized by terrestrial inputs were decoupled from aquatic P availability and therefore excreted P at disproportionately high rates. One of these species, Astyanax aeneus (Characidae), represented 12% of the total population and 18% of the total biomass of the fish assemblage in our focal low-P study stream but had P excretion rates > 10-fold higher than other abundant fishes. As a result, we estimated that P excretion by A. aeneus accounted for 90% of the P recycled by this fish assemblage and also supplied approximately 90% of the stream P demand in this P-limited ecosystem. Nitrogen excretion rates showed little variation among species, and the contribution of a given species to ecosystem N recycling was largely dependent upon the total biomass of that species. Because of the high variability in P excretion rates among fish species, ecosystem-level P recycling could be particularly sensitive to changes in fish community structure in P-limited systems.