Trajectories and state changes of a grassland stream and riparian zone after a decade of woody vegetation removal.

Riparian zones and the streams they border provide vital habitat for organisms, water quality protection, and other important ecosystem services. These areas are under pressure from local (land use/land cover change) to global (climate change) processes. Woody vegetation is expanding in grassland riparian zones worldwide. Here we report on a decade-long watershed-scale mechanical removal of woody riparian vegetation along 4.5 km of stream channel in a before-after control impact experiment. Prior to this removal, woody plants had expanded into grassy riparian areas, associated with a decline in streamflow, loss of grassy plant species, and other ecosystem-scale impacts. We confirmed some expected responses, including rapid increases in stream nutrients and sediments, disappearance of stream mosses, and decreased organic inputs to streams via riparian leaves. We were surprised that nutrient and sediment increases were transient for 3 years, that there was no recovery of stream discharge, and that areas with woody removal did not shift back to a grassland state, even when reseeded with grassland species. Rapid expansion of shrubs (Cornus drummondii, Prunus americana) in the areas where trees were removed allowed woody vegetation to remain dominant despite repeating the cutting every 2 years. Our results suggest woody expansion can fundamentally alter terrestrial and aquatic habitat connections in grasslands, resulting in inexorable movement toward a new ecosystem state. Human pressures, such as climate change, atmospheric CO2 increases, and elevated atmospheric nitrogen deposition, could continue to push the ecosystem along a trajectory that is difficult to change. Our results suggest that predicting relationships between riparian zones and the streams they border could be difficult in the face of global change in all biomes, even in well-studied sites.

Assessing placement bias of the global river gauge network.

Knowing where and when rivers flow is paramount to managing freshwater ecosystems. Yet stream gauging stations are distributed sparsely across rivers globally and may not capture the diversity of fluvial network properties and anthropogenic influences. Here we evaluate the placement bias of a global stream gauge dataset on its representation of socioecological, hydrologic, climatic and physiographic diversity of rivers. We find that gauges are located disproportionally in large, perennial rivers draining more human-occupied watersheds. Gauges are sparsely distributed in protected areas and rivers characterized by non-perennial flow regimes, both of which are critical to freshwater conservation and water security concerns. Disparities between the geography of the global gauging network and the broad diversity of streams and rivers weakens our ability to understand critical hydrologic processes and make informed water-management and policy decisions. Our findings underscore the need to address current gauge placement biases by investing in and prioritizing the installation of new gauging stations, embracing alternative water-monitoring strategies, advancing innovation in hydrologic modelling, and increasing accessibility of local and regional gauging data to support human responses to water challenges, both today and in the future.

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.

Shifting stoichiometry: Long-term trends in stream-dissolved organic matter reveal altered C:N ratios due to history of atmospheric acid deposition.

Dissolved organic carbon (DOC) and nitrogen (DON) are important energy and nutrient sources for aquatic ecosystems. In many northern temperate, freshwater systems DOC has increased in the past 50 years. Less is known about how changes in DOC may vary across latitudes, and whether changes in DON track those of DOC. Here, we present long-term DOC and DON data from 74 streams distributed across seven sites in biomes ranging from the tropics to northern boreal forests with varying histories of atmospheric acid deposition. For each stream, we examined the temporal trends of DOC and DON concentrations and DOC:DON molar ratios. While some sites displayed consistent positive or negative trends in stream DOC and DON concentrations, changes in direction or magnitude were inconsistent at regional or local scales. DON trends did not always track those of DOC, though DOC:DON ratios increased over time for ~30% of streams. Our results indicate that the dissolved organic matter (DOM) pool is experiencing fundamental changes due to the recovery from atmospheric acid deposition. Changes in DOC:DON stoichiometry point to a shifting energy-nutrient balance in many aquatic ecosystems. Sustained changes in the character of DOM can have major implications for stream metabolism, biogeochemical processes, food webs, and drinking water quality (including disinfection by-products). Understanding regional and global variation in DOC and DON concentrations is important for developing realistic models and watershed management protocols to effectively target mitigation efforts aimed at bringing DOM flux and nutrient enrichment under control.

Characterizing Trophic State in Tropical/Subtropical Reservoirs: Deviations among Indexes in the Lower Latitudes.

Trophic state indexes (TSI) guide management strategies regarding eutrophication control worldwide. Such indexes usually consider chlorophyll-a (Chl-a), total phosphorus (TP), and Secchi disk depth (SDD) as independent variables for estimating aquatic productivity and the degree of impairment. TSIs for each of these components are frequently averaged to produce a single TSI value associated with a trophic state classification (e.g., oligotrophic, mesotrophic, or eutrophic). The potential divergence among equations and classification systems originally developed for temperate lakes or tropical/subtropical reservoirs might be particularly relevant in the tropics, where there is a lack of data and the use of equations originally developed for temperate systems may be inappropriate. We calculated two widely used TSIs for temperate lakes (TSItemp) or tropical reservoirs (TSItrop) and explored the deviations among TSI components in Brazilian reservoirs. When applied to our tropical/subtropical reservoirs, the TSItemp provided a conservative approach, with lower limits anticipating increasing trophic state classification. TSI components for Chl-a and SDD significantly deviated for both sets of equations, and these discrepancies were related to turbidity, water temperature, and cyanobacterial biomass. For TSItemp, but not for TSItrop, TSI values in relation to Chl-a and TP were also significantly different. All such deviations have important management implications especially when Chl-a, TP, and SDD are averaged in a single TSI, representing loss of information and less useful trophic state classifications. Our results demonstrate that tropical water bodies may respond to drivers of eutrophication differently than temperate systems, highlighting the need for more data to better inform management of these understudied ecosystems. As managers collect data from more tropical water bodies, regional models may offer even better understanding of factors influencing trophic state.

Fate and toxicity of engineered nanomaterials in the environment: A meta-analysis.

The global environment annually receives thousands of tons of engineered nanomaterials (ENMs, particles less than 100 nm diameter). These particles have high active surface area, unique chemical properties, and can enter cells. Humanity uses many ENMs for their biological reactivity (e.g. microbicides), but their environmental effects are complex. We cataloged 2102 experimental results on whole organisms for 22 particle classes (mainly on Ag, Zn, Ti, and Cu) to assess biological responses, effective and lethal concentrations, and bioaccumulation of ENMs. Most responses were negative and varied significantly by particle type, functional group of organism, and type of response. Smaller particles tended to be more toxic. Aquatic organisms responded more negatively than did terrestrial organisms. Animals generally were most sensitive and plants least. Silver ENMs generally had the strongest negative effects. Effective and lethal concentrations generally exceeded modeled environmentally relevant concentrations and organisms usually did not accumulate or biomagnify to concentrations above those in their environment. However, most experiments lasted less than a week and were not field studies. Research to date is probably insufficient to understand chronic effects and long-term biomagnification. Numerous unique and untested ENMs continue to enter environments at accelerating rates, and our analysis indicates potential for negative effects. Our data suggest substantial research is still required to understand the ultimate influence of ENMs as they continue to accumulate in the environment. Around 40% of the papers with experimental data for ENMs failed with respect to reporting means, sample sizes, or experimental error, or they did not have proper experimental design (e.g. lack of true controls). We need more high-quality experiments that are more realistic (field or mesocosm), longer duration, contain a wider range of organisms, and account for complex food web structure.

River ecosystem conceptual models and non-perennial rivers: A critical review.

Conceptual models underpin river ecosystem research. However, current models focus on continuously flowing rivers and few explicitly address characteristics such as flow cessation and drying. The applicability of existing conceptual models to nonperennial rivers that cease to flow (intermittent rivers and ephemeral streams, IRES) has not been evaluated. We reviewed 18 models, finding that they collectively describe main drivers of biogeochemical and ecological patterns and processes longitudinally (upstream-downstream), laterally (channel-riparian-floodplain), vertically (surface water-groundwater), and temporally across local and landscape scales. However, perennial rivers are longitudinally continuous while IRES are longitudinally discontinuous. Whereas perennial rivers have bidirectional lateral connections between aquatic and terrestrial ecosystems, in IRES, this connection is unidirectional for much of the time, from terrestrial-to-aquatic only. Vertical connectivity between surface and subsurface water occurs bidirectionally and is temporally consistent in perennial rivers. However, in IRES, this exchange is temporally variable, and can become unidirectional during drying or rewetting phases. Finally, drying adds another dimension of flow variation to be considered across temporal and spatial scales in IRES, much as flooding is considered as a temporally and spatially dynamic process in perennial rivers. Here, we focus on ways in which existing models could be modified to accommodate drying as a fundamental process that can alter these patterns and processes across spatial and temporal dimensions in streams. This perspective is needed to support river science and management in our era of rapid global change, including increasing duration, frequency, and occurrence of drying.

Experimental nitrogen and phosphorus enrichment stimulates multiple trophic levels of algal and detrital-based food webs: a global meta-analysis from streams and rivers.

Anthropogenic increases in nitrogen (N) and phosphorus (P) concentrations can strongly influence the structure and function of ecosystems. Even though lotic ecosystems receive cumulative inputs of nutrients applied to and deposited on land, no comprehensive assessment has quantified nutrient-enrichment effects within streams and rivers. We conducted a meta-analysis of published studies that experimentally increased concentrations of N and/or P in streams and rivers to examine how enrichment alters ecosystem structure (state: primary producer and consumer biomass and abundance) and function (rate: primary production, leaf breakdown rates, metabolism) at multiple trophic levels (primary producer, microbial heterotroph, primary and secondary consumers, and integrated ecosystem). Our synthesis included 184 studies, 885 experiments, and 3497 biotic responses to nutrient enrichment. We documented widespread increases in organismal biomass and abundance (mean response = +48%) and rates of ecosystem processes (+54%) to enrichment across multiple trophic levels, with no large differences in responses among trophic levels or between autotrophic or heterotrophic food-web pathways. Responses to nutrient enrichment varied with the nutrient added (N, P, or both) depending on rate versus state variable and experiment type, and were greater in flume and whole-stream experiments than in experiments using nutrient-diffusing substrata. Generally, nutrient-enrichment effects also increased with water temperature and light, and decreased under elevated ambient concentrations of inorganic N and/or P. Overall, increased concentrations of N and/or P altered multiple food-web pathways and trophic levels in lotic ecosystems. Our results indicate that preservation or restoration of biodiversity and ecosystem functions of streams and rivers requires management of nutrient inputs and consideration of multiple trophic pathways.

What's in a Name? Patterns, Trends, and Suggestions for Defining Non-Perennial Rivers and Streams.

Rivers that cease to flow are globally prevalent. Although many epithets have been used for these rivers, a consensus on terminology has not yet been reached. Doing so would facilitate a marked increase in interdisciplinary interest as well as critical need for clear regulations. Here we reviewed literature from Web of Science database searches of 12 epithets to learn (Objective 1-O1) if epithet topics are consistent across Web of Science categories using latent Dirichlet allocation topic modeling. We also analyzed publication rates and topics over time to (O2) assess changes in epithet use. We compiled literature definitions to (O3) identify how epithets have been delineated and, lastly, suggest universal terms and definitions. We found a lack of consensus in epithet use between and among various fields. We also found that epithet usage has changed over time, as research focus has shifted from description to modeling. We conclude that multiple epithets are redundant. We offer specific definitions for three epithets (non-perennial, intermittent, and ephemeral) to guide consensus on epithet use. Limiting the number of epithets used in non-perennial river research can facilitate more effective communication among research fields and provide clear guidelines for writing regulatory documents.

Drivers of nitrogen transfer in stream food webs across continents.

Studies of trophic-level material and energy transfers are central to ecology. The use of isotopic tracers has now made it possible to measure trophic transfer efficiencies of important nutrients and to better understand how these materials move through food webs. We analyzed data from thirteen 15 N-ammonium tracer addition experiments to quantify N transfer from basal resources to animals in headwater streams with varying physical, chemical, and biological features. N transfer efficiencies from primary uptake compartments (PUCs; heterotrophic microorganisms and primary producers) to primary consumers was lower (mean 11.5%, range <1% to 43%) than N transfer efficiencies from primary consumers to predators (mean 80%, range 5% to >100%). Total N transferred (as a rate) was greater in streams with open compared to closed canopies and overall N transfer efficiency generally followed a similar pattern, although was not statistically significant. We used principal component analysis to condense a suite of site characteristics into two environmental components. Total N uptake rates among trophic levels were best predicted by the component that was correlated with latitude, DIN:SRP, GPP:ER, and percent canopy cover. N transfer efficiency did not respond consistently to environmental variables. Our results suggest that canopy cover influences N movement through stream food webs because light availability and primary production facilitate N transfer to higher trophic levels.

Global synthesis of the temperature sensitivity of leaf litter breakdown in streams and rivers.

Streams and rivers are important conduits of terrestrially derived carbon (C) to atmospheric and marine reservoirs. Leaf litter breakdown rates are expected to increase as water temperatures rise in response to climate change. The magnitude of increase in breakdown rates is uncertain, given differences in litter quality and microbial and detritivore community responses to temperature, factors that can influence the apparent temperature sensitivity of breakdown and the relative proportion of C lost to the atmosphere vs. stored or transported downstream. Here, we synthesized 1025 records of litter breakdown in streams and rivers to quantify its temperature sensitivity, as measured by the activation energy (Ea , in eV). Temperature sensitivity of litter breakdown varied among twelve plant genera for which Ea could be calculated. Higher values of Ea were correlated with lower-quality litter, but these correlations were influenced by a single, N-fixing genus (Alnus). Ea values converged when genera were classified into three breakdown rate categories, potentially due to continual water availability in streams and rivers modulating the influence of leaf chemistry on breakdown. Across all data representing 85 plant genera, the Ea was 0.34 ± 0.04 eV, or approximately half the value (0.65 eV) predicted by metabolic theory. Our results indicate that average breakdown rates may increase by 5-21% with a 1-4 °C rise in water temperature, rather than a 10-45% increase expected, according to metabolic theory. Differential warming of tropical and temperate biomes could result in a similar proportional increase in breakdown rates, despite variation in Ea values for these regions (0.75 ± 0.13 eV and 0.27 ± 0.05 eV, respectively). The relative proportions of gaseous C loss and organic matter transport downstream should not change with rising temperature given that Ea values for breakdown mediated by microbes alone and microbes plus detritivores were similar at the global scale.

Spatial and successional dynamics of microbial biofilm communities in a grassland stream ecosystem.

Biofilms represent a metabolically active and structurally complex component of freshwater ecosystems. Ephemeral prairie streams are hydrologically harsh and prone to frequent perturbation. Elucidating both functional and structural community changes over time within prairie streams provides a general understanding of microbial responses to environmental disturbance. We examined microbial succession of biofilm communities at three sites in a third-order stream at Konza Prairie over a 2- to 64-day period. Microbial abundance (bacterial abundance, chlorophyll a concentrations) increased and never plateaued during the experiment. Net primary productivity (net balance of oxygen consumption and production) of the developing biofilms did not differ statistically from zero until 64 days suggesting a balance of the use of autochthonous and allochthonous energy sources until late succession. Bacterial communities (MiSeq analyses of the V4 region of 16S rRNA) established quickly. Bacterial richness, diversity and evenness were high after 2 days and increased over time. Several dominant bacterial phyla (Beta-, Alphaproteobacteria, Bacteroidetes, Gemmatimonadetes, Acidobacteria, Chloroflexi) and genera (Luteolibacter, Flavobacterium, Gemmatimonas, Hydrogenophaga) differed in relative abundance over space and time. Bacterial community composition differed across both space and successional time. Pairwise comparisons of phylogenetic turnover in bacterial community composition indicated that early-stage succession (≤16 days) was driven by stochastic processes, whereas later stages were driven by deterministic selection regardless of site. Our data suggest that microbial biofilms predictably develop both functionally and structurally indicating distinct successional trajectories of bacterial communities in this ecosystem.

Patch-Burn Grazing Effects on the Ecological Integrity of Tallgrass Prairie Streams.

Conversion to agriculture, habitat fragmentation, and the loss of native grazers have made tallgrass prairie one of the most endangered ecosystems. One management option for the remaining prairie parcels, patch-burn grazing (PBG), applies a controlled burn to a portion of the prairie to attract cattle, creating a mosaic of more- and less-grazed patches. Although beneficial to cattle and grassland birds, the potential impacts of PBG on streams have not been studied, and a holistic approach is needed to ensure against adverse effects. We used a Before-After-Control-Impact design to assess potential impacts of PBG with and without riparian protection on tallgrass prairie headwater streams. We sampled stream macroinvertebrates and benthic organic matter 2 yr before and 2 yr during PBG treatments on two grazed watersheds with riparian fencing (fenced), two unfenced grazed watersheds (unfenced), and two ungrazed (control) watersheds. Very fine benthic organic matter increased significantly (51%) in unfenced streams compared with controls ( < 0.007), and fine particulate organic matter (<1 mm and >250 µm) increased 3-fold in the unfenced streams compared with controls ( = 0.008). The contribution of fine inorganic sediments to total substrata increased 28% in unfenced streams during PBG, which was significantly different from controls ( = 0.03). Additionally, the abundance of Ephemeroptera, Plecoptera, and Trichoptera taxa decreased from 7635 to 687 individuals m in unfenced streams, which was significantly lower than in control streams ( = 0.008). Our results indicate that PBG adversely influences prairie streams through sediment inputs and reductions in sensitive invertebrate taxa, but riparian fencing can alleviate these impacts.

Woody plant encroachment, and its removal, impact bacterial and fungal communities across stream and terrestrial habitats in a tallgrass prairie ecosystem.

Woody plant encroachment has become a global threat to grasslands and has caused declines in aboveground richness and changes in ecosystem function; yet we have a limited understanding on the effects of these phenomena on belowground microbial communities. We completed riparian woody plant removals at Konza Prairie Biological Station, Kansas and collected soils spanning land-water interfaces in removal and woody vegetation impacted areas. We measured stream sediments and soils for edaphic variables (C and N pools, soil water content, pH) and bacterial (16S rRNA genes) and fungal (ITS2 rRNA gene repeat) communities using Illumina MiSeq metabarcoding. Bacterial richness and diversity decreased with distance from streams. Fungal richness decreased with distance from the stream in wooded areas, but was similar across landscape position while Planctomycetes and Basidiomycota relative abundance was lower in removal areas. Cyanobacteria, Ascomycota, Chytridiomycota and Glomeromycota relative abundance was greater in removal areas. Ordination analyses indicated that bacterial community composition shifted more across land-water interfaces than fungi yet both were marginally influenced by treatment. This study highlights the impacts of woody encroachment restoration on grassland bacterial and fungal communities which likely subsequently affects belowground processes and plant health in this ecosystem.

Fire and grazing influences on rates of riparian woody plant expansion along grassland streams.

Grasslands are threatened globally due to the expansion of woody plants. The few remaining headwater streams within tallgrass prairies are becoming more like typical forested streams due to rapid conversion of riparian zones from grassy to wooded. Forestation can alter stream hydrology and biogeochemistry. We estimated the rate of riparian woody plant expansion within a 30 m buffer zone surrounding the stream bed across whole watersheds at Konza Prairie Biological Station over 25 years from aerial photographs. Watersheds varied with respect to experimentally-controlled fire and bison grazing. Fire frequency, presence or absence of grazing bison, and the historical presence of woody vegetation prior to the study time period (a proxy for proximity of propagule sources) were used as independent variables to predict the rate of riparian woody plant expansion between 1985 and 2010. Water yield was estimated across these years for a subset of watersheds. Riparian woody encroachment rates increased as burning became less frequent than every two years. However, a higher fire frequency (1-2 years) did not reverse riparian woody encroachment regardless of whether woody vegetation was present or not before burning regimes were initiated. Although riparian woody vegetation cover increased over time, annual total precipitation and average annual temperature were variable. So, water yield over 4 watersheds under differing burn frequencies was quite variable and with no statistically significant detected temporal trends. Overall, burning regimes with a frequency of every 1-2 years will slow the conversion of tallgrass prairie stream ecosystems to forested ones, yet over long time periods, riparian woody plant encroachment may not be prevented by fire alone, regardless of fire frequency.

Human impact on freshwater ecosystem services: a global perspective.

Human environmental change influences freshwaters as well as the regulating, provisioning, and cultural services that ecosystems provide worldwide. Here, we assess the global human impact on the potential value of six freshwater ecosystem services (ES) and estimate the proportion of each used globally (the mean value across all countries is in parentheses): biodiversity (0.37), disturbance regulation (0.24), commodities (0.39), greenhouse gases (0.09), water availability (0.10), and water quality (0.33). We also created a composite index of the impact. Using different valuation schemes, we found that humans have used potential global freshwater ES scaled by a relative value of roughly 4-20%, with a median of 16%. All countries use a considerable amount of the potential ES value, invalidating the idea that wealthier countries have less impact on their ES once they have developed. The data suggest that humans have diminished the potential ES provided by freshwaters across the globe and that factors associated with high population growth rates are related to the overall degradation.

Ecosystem characteristics of remnant, headwater tallgrass prairie streams.

North America has lost >95% of its native tallgrass prairie due to land conversion, making prairie streams one of the most endangered ecosystems. Research on the basic ecosystem characteristics of the remaining natural prairie streams will inform conservation and management. We examined the structure and function of headwater streams draining tallgrass prairie tracts at Osage Prairie in Missouri and the Konza Prairie Biological Station in Kansas and compared those values with literature values for streams draining agricultural watersheds in the region. We quantified physicochemical and biological characteristics for 2 yr. Streams at Osage and Konza were characterized by low nutrients and low suspended sediments (substantially lower than impacted sites in the region), slight heterotrophic status, and high temporal variability. Suspended sediments and nutrient concentrations were generally low in all prairie streams, but storms increased concentrations of both by 3- to 12-fold. Spring prescribed burns were followed by a slight increase in chlorophyll and decreased nutrients, potentially due to greater light availability. Benthic macroinvertebrate communities at Osage showed seasonal patterns that were probably linked to variable hydrology. We found nine amphibian species using the Osage streams as habitat or breeding sites, but little usage at Konza was probably due to dry conditions and low discharge. Our study indicates that two remnant tallgrass prairie streams along a longitudinal gradient are fairly similar in terms of physicochemical features and have good water quality relative to agricultural watersheds but can differ considerably in macroinvertebrate and amphibian abundance.