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.

Critical Role for hierarchical geospatial analyses in the design of fluvial research, assessment, and management.

River science and management can be conducted at a range of spatiotemporal scales from reach to basin levels as long as the project goals and questions are matched correctly with the study design's spatiotemporal scales and dependent variables. These project goals should also incorporate information on the hydrogeomorphically patchy nature of riverine macrosystems which is only partially predictable in type and location from a river's headwaters to its terminus. This patchiness significantly affects a river's habitat template, and thus community structure, ecosystem function, and responses to perturbations. Our manuscript is designed for use by senior administrators at government agencies through entry-level river scientists. It analyzes common challenges in project design and recommends solutions based partially on hierarchical analyses that combine geographic information systems and multivariate statistical analysis to enable self-emergence of a stream's patchy structure. These approaches are useful at all spatial levels and can vary from primary reliance on geospatial techniques at the valley level to a greater dependence on field-based measurements and expert opinion at the reach level. Comparative uses of functional process zones (FPZs = valley-scale hydrogeomorphic patches), ecoregions, hydrologic unit codes, and reaches in project designs are discussed along with other comparative approaches for stream classification and analysis of species distributions (e.g., GAP analysis). Use of hierarchical classification of patch structure for sample stratification, reference site selection, ecosystem services, rehabilitation, and mitigation are briefly explored.

Automated riverine landscape characterization: GIS-based tools for watershed-scale research, assessment, and management.

River systems consist of hydrogeomorphic patches (HPs) that emerge at multiple spatiotemporal scales. Functional process zones (FPZs) are HPs that exist at the river valley scale and are important strata for framing whole-watershed research questions and management plans. Hierarchical classification procedures aid in HP identification by grouping sections of river based on their hydrogeomorphic character; however, collecting data required for such procedures with field-based methods is often impractical. We developed a set of GIS-based tools that facilitate rapid, low cost riverine landscape characterization and FPZ classification. Our tools, termed RESonate, consist of a custom toolbox designed for ESRI ArcGIS®. RESonate automatically extracts 13 hydrogeomorphic variables from readily available geospatial datasets and datasets derived from modeling procedures. An advanced 2D flood model, FLDPLN, designed for MATLAB® is used to determine valley morphology by systematically flooding river networks. When used in conjunction with other modeling procedures, RESonate and FLDPLN can assess the character of large river networks quickly and at very low costs. Here we describe tool and model functions in addition to their benefits, limitations, and applications.

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.

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.

Communities associated with the Functional Process Zone scale: A case study of stream macroinvertebrates in endorheic drainages.

Rivers are being increasingly analyzed from a holistic scale focus, imposing the challenge to establish a clear sampling framework that integrates complex valley-to-reach hydrogeomorphic features. Here, we address this challenge by examining macroinvertebrate communities of different hydrogeomorphic patches, or Functional Process Zones (FPZs), established by the GIS-based model RESonate. We delineated FPZs across three endorheic drainages in the Great Basin, USA, using a self-emerging clustering method that classifies segments of rivers with similar hydrogeomorphic characteristics. We sampled macroinvertebrate communities across different FPZs. We examined the taxonomic and functional organizations of these communities, and we assessed the relative contributions of in-stream and watershed-scale environmental filters in structuring these communities. We found discreet macroinvertebrate communities associated with FPZs across drainages, where elevation prevailed on valley confinement in structuring these communities. Communities of upland FPZs exhibited a higher heterogeneity suggested by higher ß-diversity and nested structure of communities, while lowland FPZs showed a higher pairwise abundance agreement across communities. Eltonian trait composition, primarily describing bionomic traits, showed a higher degree of niche differentiation in upland FPZs, thereby increasing the overall ecosystem function. Differences in variance partitioning among environmental filters acting at different spatial scales show a strong spatial structure in the response of communities in different FPZs. Overall, environmental filters had a stronger control of the communities' functional organization than the taxonomic composition. Our results support the paradigm of different FPZs having distinct communities that express different ecosystem properties. Findings of this study constitute a fruitful avenue for expanding community-based research using the FPZ template as a tool for riverine ecology. However, the unique nature of rivers in endorheic basins needs to be considered when applying our conclusions to other systems, as some findings (e.g., the higher community homogenization in lowland FPZs) might be specific to this rarely examined type of river systems.

Multidimensional metrics of niche space for use with diverse analytical techniques.

Multidimensional data are integral to many community-ecological studies and come in various forms, such as stable isotopes, compound specific analyses (e.g., amino acids and fatty acids), and both biodiversity and life history traits. Scientists employing such data often lack standardized metrics to evaluate communities in niche space where more than 2 dimensions are involved. To alleviate this problem, we developed a graphing and analytical approach for use with more than two variables, based on previously established stable isotope bi-plot metrics. We introduce here our community metrics as R scripts. By extending the original metrics to multiple dimensions, we created n-dimensional plots and metrics to characterize any set of quantitative measurements of a community. We demonstrate the utility of these metrics using stable isotope data; however, the approaches are applicable to many types of data. The resulting metrics provide more and better information compared to traditional analytic frameworks. The approach can be applied in many branches of community ecology, and it offers accessible metrics to quantitatively analyze the structure of communities across ecosystems and through time.

Patches and the responses of lake benthos to sunfish nest-building.

Males of three sunfish species (Centrarchidae) construct nests for spawning and often share them sequentially in the littoral zone of a 4-hectare lake in New York State. To determine spatial and temporal effects of this reproductive behavior on zoobenthos in "patches," I sampled bottom assemblages from inside nests and from adjacent (<1 m from nest perimeter), relatively undisturbed sites during the reproductive season of 1986 and immediately prior to nest-building in 1987. The reproductive behaviors of largemouth bass (Micropterus salmoides), redbreast sunfish (Lepomis auritus), and pumpkinseeds (L. gibbosus) altered relative abundances and significantly decreased benthic invertebrate diversity and density. These effects were extremely pronounced during the reproductive season and were partially detectable the following year. Community changes were probably the result of both bioturbation (modification of sediment size and organic content) and predation. The ecosystem-wide effects of nest-building are evaluated in terms of the number, distribution, and longevity of patches.

Agonistic behavior in crayfish in relation to temperature and reproductive period.

I examined the relationship between temperature and agonistic behavior in the crayfish Cambarus latimanus LeConte a species which is reproductively active in winter. Crayfish were acclimated for 2 wks at 9.5, 14, 22 and 30°C in summer and at 9.5 and 22°C in winter. Agonistic behavior of pairs of the same sex was recorded for 1 h following acclimation. The entire acclimation, testing, and temperature-readjustment procedure was repeated until all crayfish pairs had been tested at each acclimation temperature.Published information on lobsters and catfish indicate that agonistic behavior is directly related to temperature. In contrast, for C. latimanus in the present study, duration of total agonistic behavior, maximum duration of a single agonistic interaction, and average length of a agonistic encounter were all inversely related to acclimation temperature. I hypothesized that for ectothermic species whose agonistic behavior is closely associated with reproductive processes (including competition for mates) the level of agonism will vary either directly or inversely depending upon whether the reproductive period is cued by increasing or decreasing seasonal temperatures, respectively.

Field experiments on interactions between vertebrate predators and larval midges (Diptera: Chironomidae) in the littoral zone of a reservoir.

A field experiment was designed to test a frequent assumption in the literature that vertebrate predators (in this case, fish and turtles) are capable of regulating the seasonal abundance and diversity of benthic communities in the littoral zone of lentic environments. Effects of thermal effluents from a nuclear reactor on predator-prey relationships were also examined. Benthic samples were removed after each of three, 3-month test periods from 36 predator exclusion cages (4m2) and 36 control plots located along a thermal gradient in Par Pond, an 1,100 ha freshwater reservoir in the southeastern United States.Results of our field experiments provide little evidence to suggest that either a single "keystone" species or vertebrate predators as a group were capable of regulating the abundance, diversity or productivity of chironomids in Par Pond. The relationship between predator treatment and community response (changes in density and species richness) was generally unaffected by either plot location or temperature fluctuations. When data from caged and control plots were pooled, however, both location and water temperature individually had significant impacts on the chironomid community. Alternative hypotheses are proposed to explain the lack of regulatory control of the benthic community by individual species or guilds of predators.

The influence of temperature on the functional response of the dragonfly Celithemis fasciata (Odonata: Libellulidae).

1) Functional response curves were constructed for Celithemis fasciata larvae feeding on 6 different densities of midge larvae at 10, 15, 20, and 25°C. Values for attack rate and handling time were estimated with Rogers's random predator equation. 2) Polynomial regression revealed that the functional response curves were linear although a tendency toward decreasing consumption rates at higher densities was shown. 3) The mean number of prey eaten increased with temperature; however, temperature did not alter the fundamental shape of the functional response curve. 4) The attack rate and handling time were linearly related to temperature in general, but changed relatively little between 15 and 20° C.

Isotopic analysis of three food web theories in constricted and floodplain regions of a large river.

Analyses of stable isotope (δ13C and δ15N) and C:N ratios of food webs within a floodplain and a constricted-channel region of the Ohio River during October 1993 and July 1994 indicate that the increasingly influential flood pulse concept (FPC) does not, for either location, adequately address food web structure for this very large river. Furthermore, results of this study suggest that the riverine productivity model (RPM) is more appropriate than the widely known river continuum concept (RCC) for the constricted region of this river. These␣conclusions are based on stable isotope analyses of potential sources of organic matter (riparian C3 trees, riparian C4 grasses and agricultural crops, submerged macrophytes, benthic filamentous algae, benthic particulate organic matter, and transported organic matter containing detritus and phytoplankton) and various functional feeding groups of invertebrate and fish consumers. The FPC, which stresses the key contribution of organic matter, particularly terrestrial organic matter, originating from the floodplain to riverine food webs, was judged inappropriate for the floodplain region of the Ohio River for hydrodynamic and biotic reasons. The rising limb and peak period of discharge typically occur in November through March when temperatures are low (generally much less than 10°C) and greater than bank-full conditions are relatively unpredictable and short-lived. The major food potentially available to riverine organisms migrating into the floodplain would be decaying vegetation because autotrophic production is temperature and light limited and terrestrial insect production is minimal at that time. It is clear from our data that terrestrial C4 plants contribute little, if anything, to the consumer food web (based on δ13C values), and δ15N values for C3 plants, coarse benthic organic matter, and fine benthic organic matter were too depleted (∼7-12‰ lower than most invertebrate consumer values) for this organic matter to be supporting the food web. The RPM, which emphasizes the primary role of autotrophic production in large rivers, is the most viable of the remaining two ecosystem models for the constricted-channel region of the Ohio based on stable isotope linkage between sources and consumers of organic matter in the food web. The most important form of food web organic matter is apparently transported (suspended) fine (FTOM) and ultra-fine particulate organic matter. We propose that phytoplankton and detritus of an autochthonous origin in the seston would represent a more usable energy source for benthic (bivalve molluscs, hydropsychid caddisflies) and planktonic (microcrustaceans) suspension feeders than the more refractory allochthonous materials derived from upstream processing of terrestrial organic matter. Benthic grazers depend heavily on nonfilamentous benthic algae (based on gut analysis from a separate study), but filamentous benthic algae have no apparent connection to invertebrate consumers (based on δ13C values). Amphipod and crayfish show a strong relationship to aquatic macrophytes (possibly through detrital organic matter rather than living plant tissue). These observations contrast with the prediction of the RCC that food webs in large rivers are based principally on refractory FTOM and dissolved organic matter from upstream inefficiencies in organic-matter processing and the bacteria growing upon these suspended or dissolved detrital compounds. The conclusions drawn here for the Ohio River cannot yet be extended to other floodplain and constricted-channel rivers in temperate and tropical latitudes until more comparable data are available on relatively pristine and moderately regulated rivers.

Significance of instream autotrophs in trophic dynamics of the Upper Mississippi River.

Trophic dynamics of large river-floodplain ecosystems are still not well understood despite development of several conceptual models over the last 25 years. To help resolve questions about the relative contribution of algal and detrital organic matter to food webs in the Upper Mississippi River, we (1) separated living and detrital components of ultrafine and fine transported organic matter (UTOM and FTOM, respectively) by colloidal silica centrifugation; (2) identified stable isotope signatures (delta(13)C and delta(15)N) for these two portions of transported organic matter and other potential organic matter sources; and (3) employed a multiple source, dual-isotope mixing model to determine the relative contribution of major energy sources to primary consumers and the potential contribution of basal sources to the biomass of secondary consumers. The delta(13)C and delta(15)N of living and detrital fractions of UTOM and FTOM were distinct, indicating clear differences in isotopic composition of the algal and detrital fractions of transported organic matter. Living and detrital transported organic matter also differed from other potential organic matter sources by either delta(13)C or delta(15)N. A six-source mixing model using both delta(13)C and delta(15)N indicated that algal transported organic matter was the major resource assimilated by primary consumers. The contribution of detrital transported organic matter was small in most cases, but there were a small number of taxa for which it could potentially contribute to more than half the assimilated diet. Colloidal dissolved organic matter, which includes heterotrophic bacteria, accounted for only a small fraction of the organic matter assimilated by most primary consumers, indicating that coupling between microbial processes and metazoan production is minimal. Terrestrial C(3) litter from the floodplain forest floor and aquatic macrophytes were also relatively unimportant to the assimilated diet of primary consumers. Application of the mixing model to compare basal source isotopic ratios to secondary consumers revealed that most organic matter moving from primary to secondary consumers originated from algal TOM. Our findings indicate that autochthonous organic matter is the major energy source supporting metazoan production in the main channel of this large river, at least during the summer. This study joins a number of other investigations performed globally that indicate organic matter originating from instream production of sestonic and benthic microalgae is a major driver in the trophic dynamics of large river ecosystems.