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The Clean Water Act (CWA) of 1972 regulates water quality in U.S. inland waters under a system of cooperative federalism in which states are delegated implementation and enforcement authority of CWA provisions by the U.S. Environmental Protection Agency. We leveraged heterogeneity in state implementation of the CWA to evaluate the efficacy of its nonpoint source provisions in reducing nutrient pollution, the leading cause of water quality impairment in U.S. inland waters. We used national survey data to estimate changes in nutrient concentrations over a decade and evaluated the effect of state-level policy implementation. We found no evidence to support an effect of (i) grant spending on nonpoint source pollution remediation, (ii) nutrient criteria development, or (iii) water quality monitoring intensity on 10-year trends in nutrient concentrations. These results suggest that the current federal policy paradigm for improving water quality is not creating desired outcomes. [ABSTRACT FROM AUTHOR]

Nutrient uptake, storage, and release are critical ecosystem functions that affect carbon processing and food web dynamics. Yet, mechanisms controlling when ecosystems are net sinks or sources of nutrients are uncertain. Specifically, how nutrient supply ratios alter rates and ratios of net nutrient uptake and release is unclear. To assess whether net nitrogen (N) and phosphorus (P) uptake and release are linked to supply N:P, we experimentally enriched five forest streams at different N:P (target molar N:P range 2:1–128:1) for 2 years. We quantified net nutrient exchange (NNE) as the difference between the expected N and P fluxes assuming conservative transport (background concentrations plus experimental inputs) and the observed nutrient fluxes at the downstream end of each experimental stream reach. Supply N:P did not affect the magnitude of NNE for either N or P, but the likelihood of net N and P uptake was greatest at intermediate N:P supply (N:P = 99:1 and 55:1, respectively). Streams appeared to be highly flexible in their N:P uptake and release; the slopes between NNEN and NNEP within each stream increased with supply N:P. Furthermore, slopes comparing supply N:P to uptake and release N:P were near one (0.98 ± 0.06 SE and 0.82 ± 0.13 SE, respectively), indicating a high degree of flexibility. Overall, we found greater stoichiometric flexibility than has been shown in short-term nutrient-addition experiments. We suggest that this flexibility results from changes in nutrient recycling within biofilms or changes in community structure, which may take longer to manifest than the duration of shorter-term experiments. [ABSTRACT FROM AUTHOR]

Diverse global change processes are reshaping the biogeochemistry of stream ecosystems. Nutrient enrichment is a common stressor that can modify flows of biologically important elements such as carbon (C), nitrogen (N), and phosphorus (P) through stream foodwebs by altering the stoichiometric composition of stream organisms. However, enrichment effects on concentrations of other important essential and trace elements in stream taxa are less understood. We investigated shifts in macroinvertebrate ionomes in response to changes in coarse benthic organic matter (CBOM) stoichiometry following N and P enrichment of five detritus-based headwater streams. Concentrations of most elements (17/19) differed among three insect genera (Maccaffertium sp., Pycnopsyche spp., and Tallaperla spp.) prior to enrichment. Genus-specific changes in the body content of: P, magnesium, and sodium (Na) in Tallaperla; P, Na, and cadmium in Pycnopsyche; and P in Maccaffertium were also found across CBOM N:P gradients. These elements increased in Tallaperla but decreased in the other two taxa due to growth dilution at larger body sizes. Multivariate elemental differences were found across all taxa, and ionome-wide shifts with dietary N and P enrichment were also observed in Tallaperla and Pycnopsyche. Our results show that macroinvertebrates exhibit distinct differences in elemental composition beyond C, N, and P and that the ionomic composition of common stream taxa can vary with body size and N and P enrichment. Thus, bottom-up changes in N and P supplies could potentially influence the cycling of lesser studied biologically essential elements in aquatic environments by altering their relative proportions in animal tissues. [ABSTRACT FROM AUTHOR]

Despite being remote, polar and sub-polar regions are increasingly threatened by global ecological change. For instance, South America's sub-Antarctic forest ecoregion is considered one of the world's last wilderness areas and a global reference site for pre-Industrial Revolution nutrient cycles. Nonetheless, the North American beaver (Castor canadensis) was introduced to Tierra del Fuego in 1946 and, as an invasive ecosystem engineer, has transformed the ecology of regional watersheds. Beavers' engineering activities transform forested streams (FS) into beaver ponds (BP), where there is greater light and primary production (allochthonous organic matter) and, consequently, increased basal resource quality. To investigate this, we analyzed algal, diatom, fungal and bacterial fatty acid (FA) biomarkers in three basal resource categories (biofilm, very fine benthic organic matter, coarse benthic organic matter) and benthic consumers from four functional feeding groups (FFG). The amphipod Hyalella spp. was chosen as an indicator species due to its abundance and biomass in both habitats. Hyalella spp. had higher proportions of algal and bacterial FA in BP than FS. In FS, Hyalella spp. (gatherer) and Gigantodax spp. (filterer, Diptera) had greater contributions of higher quality FA (higher in polyunsaturated FA), while Rheochorema magallanicum (predator, Trichoptera) and Meridialaris spp. (scraper, Ephemeroptera) showed lower quality monounsaturated and saturated FA. All FFGs showed evidence of microbial FA and had higher levels of autochthonous FA biomarkers than their food resources. Scrapers had the greatest proportion of autochthonous FA. These data provide new insights into the utilization of basal resources by stream consumers in sub-Antarctic streams and how beavers modify these ecosystems. [ABSTRACT FROM AUTHOR]

Stoichiometric ratios of resources and consumers have been used to predict nutrient limitation across diverse terrestrial and aquatic ecosystems. In forested headwater streams, coarse and fine benthic organic matter (CBOM, FBOM) are primary basal resources for the food web, and the distribution and quality of these organic matter resources may therefore influence patterns of secondary production and nutrient cycling within stream networks or among biomes. We measured carbon (C), nitrogen (N), and phosphorus (P) content of CBOM and FBOM and calculated their stoichiometric ratios (C/N, C/P, N/P) from first- to fourth-order streams from tropical montane, temperate deciduous, and boreal forests, and tallgrass prairie, to compare the magnitude and variability of these resource types among biomes. We then used the ratios to predict nutritional limitations for consumers of each resource type. Across biomes, CBOM had consistently higher %C and %N, and higher and more variable C/N and C/P than FBOM, suggesting that microbial processing results in more tightly constrained elemental composition in FBOM than in CBOM. Biome-specific differences were observed in %P and N/P between the two resource pools; CBOM was lower in %P but higher in N/P than FBOM in the tropical montane and temperate deciduous forest biomes, while CBOM was higher in %P but similar in N/P than FBOM in the grassland and boreal forest biomes. Stable 13C isotopes suggest that FBOM likely derives from CBOM in tropical and temperate deciduous forest, but that additional non-detrital components may contribute to FBOM in boreal forests and grasslands. Comparisons of stoichiometric ratios of CBOM and FBOM to estimated needs of aquatic detritivores suggest that shredders feeding on CBOM are more likely to experience nutrient (N and/or P) than C limitation, whereas collector-gatherers consuming FBOM are more likely to experience C than N and/or P limitation. Our results suggest that differences in basal resource elemental content and stoichiometric ratios have the potential to affect consumer production and ecosystem rates of C, N, and P cycling in relatively consistent ways across diverse biomes. [ABSTRACT FROM AUTHOR]

North American beavers ( Castor canadensis) were introduced to Tierra del Fuego Island in 1946 for their fur, and have since spread across the archipelago and onto the South American mainland. We assessed the impact of invasive beavers on streams of these forested watersheds by quantifying the trophic basis of production (TBP) and consumptive organic matter flows of benthic macroinvertebrate assemblages. TBP was determined in two streams: clear- and black-water. Stable isotopes were used across four streams to further elucidate food web structure and dominant pathways. TBP and stable isotopes showed that terrestrially derived organic matter (amorphous detritus, leaves, and wood) supported a majority of secondary production in the benthic food webs at all sites (forested reaches, beaver ponds, and sections downstream of ponds with foraged riparian zones). The magnitude of these flows was enhanced in beaver-modified sites compared with forested habitats (4.0–5.3× increase g AFDM m−2 year−1 in pond habitats, 1.1–2.1× increase in downstream habitats). Diatoms were the only autochthonous resource identified in macroinvertebrate guts, but their contribution to secondary production was small. Consumptive flows mirrored trends in TBP (i.e., dominance of terrestrial sources and greater magnitude in beaver ponds). Collector–gatherer consumption of amorphous detrital material dominated food web flows in all habitats, but was higher in beaver ponds relative to other habitats. Food web structure was simplified in beaver ponds; only two of the five possible functional groups contributed >1% of total organic matter flow in ponds (collector–gatherers and predators). Consumptive flows to predators increased in ponds, and stable isotopes of nitrogen and carbon (δ15N and δ13C) corroborated a relatively greater importance of predators (greater trophic distance), as well as less diversity of basal resources (less variation in δ13C) in ponds. Our findings indicate that invasive beaver’s engineering activities resulted in greater flows of terrestrial organic matter subsidies to in-stream food webs, which had a relatively greater change in the clear-water than in the black-water stream. Owing to the fact that these streams were naturally dependent on allochthonous resources for a majority of production and material flows, changes wrought by beavers to streams in forested environments are probably less than in watersheds with inherently greater dependence on autochthonous production such as the adjacent steppe biome. [ABSTRACT FROM AUTHOR]

Species invasions are of global significance, but predicting their impacts can be difficult. Introduced ecosystem engineers, however, provide an opportunity to test the underlying mechanisms that may be common to all invasive engineers and link relationships between changes in diversity and ecosystem function, thereby providing explanatory power for observed ecological patterns. Here we test specific predictions for an invasive ecosystem engineer by quantifying the impacts of habitat and resource modifications caused by North American beavers ( Castor canadensis) on aquatic macroinvertebrate community structure and stream ecosystem function in the Cape Horn Biosphere Reserve, Chile. We compared responses to beavers in three habitat types: (1) forested (unimpacted) stream reaches, (2) beaver ponds, and (3) sites immediately downstream of beaver dams in four streams. We found that beaver engineering in ponds created taxonomically simplified, but more productive, benthic macroinvertebrate assemblages. Specifically, macroinvertebrate richness, diversity and number of functional feeding groups were reduced by half, while abundance, biomass and secondary production increased three- to fivefold in beaver ponds compared to forested sites. Reaches downstream of beaver ponds were very similar to natural forested sections. Beaver invasion effects on both community and ecosystem parameters occurred predominantly via increased retention of fine particulate organic matter, which was associated with reduced macroinvertebrate richness and diversity (via homogenization of benthic microhabitat) and increased macroinvertebrate biomass and production (via greater food availability). Beaver modifications to macroinvertebrate community structure were largely confined to ponds, but increased benthic production in beaver-modified habitats adds to energy retention and flow for the entire stream ecosystem. Furthermore, the effects of beavers on taxa richness (negative) and measures of macroinvertebrate biomass (positive) were inversely related. Thus, while a generally positive relationship between diversity and ecosystem function has been found in a variety of systems, this work shows how they can be decoupled by responding to alterative mechanisms. [ABSTRACT FROM AUTHOR]

Most nutrient enrichment studies in aquatic systems have focused on autotrophic food webs in systems where primary producers dominate the resource base. We tested the heterotrophic response to long-term nutrient enrichment in a forested, headwater stream. Our study design consisted of 2 years of pretreatment data in a reference and treatment stream and 2 years of continuous nitrogen (N) + phosphorus addition to the treatment stream. Studies were conducted with two leaf species that differed in initial C:N, Rhododendron maximum (rhododendron) and Acer rubrum (red maple). We determined the effects of nutrient addition on detrital resources (leaf breakdown rates, litter C:N and microbial activity) and tested whether nutrient enrichment affected macroinvertebrate consumers via increased biomass. Leaf breakdown rates were ca. 1.5 and 3× faster during the first and second years of enrichment, respectively, in the treatment stream for both leaf types. Microbial respiration rates of both leaf types were 3× higher with enrichment, and macroinvertebrate biomass associated with leaves increased ca. 2–3× with enrichment. The mass of N in macroinvertebrate biomass relative to leaves tended to increase with enrichment up to 6× for red maple and up to 44× for rhododendron leaves. Lower quality (higher C:N) rhododendron leaves exhibited greater changes in leaf nutrient content and macroinvertebrate response to nutrient enrichment than red maple leaves, suggesting a unique response by different leaf species to nutrient enrichment. Nutrient concentrations used in this study were moderate and equivalent to those in streams draining watersheds with altered land use. Thus, our results suggest that similarly moderate levels of enrichment may affect detrital resource quality and subsequently lead to altered energy and nutrient flow in detrital food webs. [ABSTRACT FROM AUTHOR]

While many studies have measured effects of nutrient enrichment on higher trophic levels in grazing food webs, few such studies exist for detritus-based systems. We measured effects of nitrogen and phosphorus addition on growth of larval Eurycea wilderae in a heterotrophic headwater stream using a repeated mark-recapture design. Growth estimates for 208 recaptured larvae (control stream n = 92; treatment stream n = 116) resulted in a growth rate of 0.0027 d−1 in each stream prior to enrichment, whereas during enrichment treatment growth rates ( g = 0.0069 d−1 [±0.0019, 95% C.I.]) were significantly higher than control ( g = 0.0043 d−1 [±0.0007, 95% C.I.]). Results indicate that E. wilderae growth is tightly linked to the detrital resource and that growth may be indirectly affected by both quantity and quality of detritus. This study provides some of the first evidence that nutrient enrichment of detritus-based systems can influence multiple trophic levels in ways similar to autotrophic systems. [ABSTRACT FROM AUTHOR]

Pristine wilderness is a scarce global resource, but exotic species are so common that they constitute a principal component of worldwide ecological change. The relationship between these two topics, invasion and remoteness, was the impetus behind five years of fieldwork aimed at identifying the assemblage and range of exotic vertebrates in Cape Horn, Chile, identified as one of the world's most pristine wilderness areas. While the archipelago has extremely low human population density and vast tracts of undisturbed land, we discovered that several terrestrial vertebrate groups were dominated by exotic species. Native birds were diverse (approx. 154 spp), and only 1.3% (or two spp.) were introduced. In contrast, exotic terrestrial mammals (12 spp.) and freshwater fish (three spp.) outnumbered their native counterparts, constituting 55% and 75% of the assemblages. Using GIS, we found that not all areas were impacted equally, largely due to intensity of human occupation. Human settled islands (Navarino and Tierra del Fuego) hosted the greatest number of exotics, but humans alone did not explain observed patterns. Remote islands also had introduced species. North American beavers (Castor canadensis), American minks (Mustela vison) and feral domestic dogs and cats were particularly widespread, and their range in isolated parts of the study area raised important ecological and management questions. In conclusion, the Cape Horn Archipelago retained areas free of exotic vertebrates, particularly parts of the Cape Horn and Alberto D'Agostini National Parks, but at many sites introduced species were overwhelming native biota and altering these previously remote natural ecosystems. [ABSTRACT FROM AUTHOR]