Nutrient pollution of freshwater ecosystems results in predictable increases in carbon (C) sequestration by algae. Tests of nutrient enrichment on the fates of terrestrial organic C, which supports riverine food webs and is a source of CO2, are lacking. Using whole-stream nitrogen (N) and phosphorus (P) additions spanning the equivalent of 27 years, we found that average terrestrial organic C residence time was reduced by ~50% as compared to reference conditions as a result of nutrient pollution. Annual inputs of terrestrial organic C were rapidly depleted via release of detrital food webs from N and P co-limitation. This magnitude of terrestrial C loss can potentially exceed predicted algal C gains with nutrient enrichment across large parts of river networks, diminishing associated ecosystem services.

Nutrient-driven perturbations to the resource base of food webs are predicted to attenuate with trophic distance, so it is unclear whether higher-level consumers will generally respond to anthropogenic nutrient loading. Few studies have tested whether nutrient (specifically, nitrogen [N] and phosphorus [P]) enrichment of aquatic ecosystems propagates through multiple trophic levels to affect predators, or whether N vs. P is relatively more important in driving effects on food webs. We conducted two-year whole-stream N and P additions to five streams to generate gradients in N and P concentration and N:P ratio (target N:P = 2, 8, 16, 32, 128). Larval salamanders are vertebrate predators of primary and secondary macroinvertebrate consumers in many heterotrophic headwater streams in which the basal resources are detritus and associated microorganisms. We determined the effects of N and P on the growth rates of caged and free-roaming larval Desmognathus quadramaculatus and the average body size of larval Eurycea wilderae. Growth rates and average body size increased by up to 40% and 60%, respectively, with P concentration and were negatively related to N:P ratio. These findings were consistent across both species of salamanders using different methodologies (cage vs. free-roaming) and at different temporal scales (3 months vs. 2 yr). Nitrogen concentration was not significantly related to increased growth rate or body size of the salamander species tested. Our findings suggest that salamander growth responds to the relaxation of ecosystem-level P limitation and that moderate P enrichment can have relatively large effects on vertebrate predators in detritus-based food webs.

Nutrient enrichment is a key stressor of lakes and streams globally, affecting the relative availability of important basal resources such as algae and detritus. These effects are controlled by responses of autotrophic and heterotrophic microorganisms that subsequently affect primary consumers and higher level predators. Despite the potential for propagation of these bottom‐up effects, few studies have examined how nutrients affect “green” (autotrophic) versus “brown” (heterotrophic) energy pathways to predators via changes in the quantity or type of prey consumed. We studied the pathways by which nutrient enrichment affected two predatory salamander species (Desmognathus quadramaculatus and Eurycea wilderae) using detailed diet analyses before and during 2‐year nutrient additions to five headwater forest streams. The streams were continuously enriched with different concentrations of dissolved nitrogen (N) and phosphorus (P), creating relatively greater N or P concentrations and distinct N:P ratios (2:1, 8:1, 16:1, 32:1 and 128:1) in each stream. Nutrient addition resulted in greater prey number, size and biomass consumed by D. quadramaculatus, an effect driven more by P than by N additions. Some of these effects were greater in the second year of enrichment and were greater for larger individuals. Shifts in the prey composition of D. quadramaculatus included increases in algivores and decreases in detritivores, tracking observed treatment effects on basal resource quantity (e.g. algivore abundance in guts was related to algal biomass, which increased with enrichment, and detritivore abundance in guts was related to detrital standing stocks, which declined with enrichment). For E. wilderae diets, there was limited evidence for increased prey size and number, or for alteration of prey composition with enrichment despite evidence of increased larval growth. We hypothesise that body size differences between the two salamander species partially explain their different dietary responses to enrichment. Our results show that nutrient addition, primarily of P, affected the quantity and composition of predator diets in our nutrient‐poor streams. These effects on diet were consistent with concurrent studies showing that P enrichment resulted in faster growth of salamanders and occurred partly via effects on algal biofilm or “green” food‐web pathways, despite the dominance of detrital or “brown” resources in our heavily shaded forest stream sites. Thus, nutrient enrichment can promote algae‐ versus detritus‐based energy‐flow pathways in nominally light‐limited stream ecosystems, with associated changes in food‐web characteristics and function.

We used a recently published, open‐access data set of U.S. streamwater nitrogen (N) and phosphorus (P) concentrations to test whether watershed land use differentially influences N and P concentrations, including the relative availability of dissolved and particulate nutrient fractions. We tested the hypothesis that N and P concentrations and molar ratios in streams and rivers of the United States reflect differing nutrient inputs from three dominant land‐use types (agricultural, urban and forested). We also tested for differences between dissolved inorganic nutrients and suspended particulate nutrient fractions to infer sources and potential processing mechanisms across spatial and temporal scales. Observed total N and P concentrations often exceeded reported thresholds for structural changes to benthic algae (58, 57% of reported values, respectively), macroinvertebrates (39% for TN and TP), and fish (41, 37%, respectively). The majority of dissolved N and P concentrations exceeded threshold concentrations known to stimulate benthic algal growth (85, 87%, respectively), and organic matter breakdown rates (94, 58%, respectively). Concentrations of both N and P, and total and dissolved N:P ratios, were higher in streams and rivers with more agricultural and urban than forested land cover. The pattern of elevated nutrient concentrations with agricultural and urban land use was weaker for particulate fractions. The % N contained in particles decreased slightly with higher agriculture and urbanization, whereas % P in particles was unrelated to land use. Particulate N:P was relatively constant (interquartile range = 2–7) and independent of variation in DIN:DIP (interquartile range = 22–152). Dissolved, but not particulate, N:P ratios were temporally variable. Constant particulate N:P across steep DIN:DIP gradients in both space and time suggests that the stoichiometry of particulates across U.S. watersheds is most likely controlled either by external or by physicochemical instream factors, rather than by biological processing within streams. Our findings suggest that most U.S. streams and rivers have concentrations of N and P exceeding those considered protective of ecological integrity, retain dissolved N less efficiently than P, which is retained proportionally more in particles, and thus transport and export high N:P streamwater to downstream ecosystems on a continental scale.

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

Free-flowing river segments provide refuges for many imperiled aquatic biota that have been extirpated elsewhere in their native ranges. These biodiversity refuges are also foci of conservation concerns because species persisting within isolated habitat fragments may be particularly vulnerable to local environmental change. We have analyzed long-term (14-and 20-y) survey data to assess evidence of fish species declines in two southeastern U.S. rivers where managers and stakeholders have identified potentially detrimental impacts of current and future land uses. The Conasauga River (Georgia and Tennessee) and the Etowah River (Georgia) form free-flowing headwaters of the extensively dammed Coosa River system. These rivers are valued in part because they harbor multiple species of conservation concern, including three federally endangered and two federally threatened fishes. We used data sets comprising annual surveys for fish species at multiple, fixed sites located at river shoals to analyze occupancy dynamics and temporal changes in species richness. Our analyses incorporated repeated site-specific surveys in some years to estimate and account for incomplete species detection, and test for species-specific (rarity, mainstem-restriction) and year-specific (elevated frequencies of low-or high-flow days) covariates on occupancy dynamics. In the Conasauga River, analysis of 26 species at 13 sites showed evidence of temporal declines in colonization rates for nearly all taxa, accompanied by declining species richness. Four taxa (including one federally endangered species) had reduced occupancy across the Conasauga study sites, with three of these taxa apparently absent for at least the last 5 y of the study. In contrast, a similar fauna of 28 taxa at 10 sites in the Etowah River showed no trends in species persistence, colonization, or occupancy. None of the tested covariates showed strong effects on persistence or colonization rates in either river. Previous studies and observations identified contaminants, nutrient loading, or changes in benthic habitat as possible causes for fish species declines in the Conasauga River. Our analysis provides baseline information that could be used to assess effectiveness of future management actions in the Conasauga or Etowah rivers, and illustrates the use of dynamic occupancy models to evaluate evidence of faunal decline from time-series data.

The majority of terrestrial net primary production decomposes, fueling detrital food webs and converting dead plant carbon to atmospheric CO2. There is considerable interest in determining the sensitivity of this process to climate warming. A common approach has been to use spatial gradients in temperature (i.e., latitude or elevation) to estimate temperature sensitivity. However, these studies typically relate decomposition rates to average temperatures at each site along such gradients, ignoring within‐site temperature variation. To evaluate the potential effects of temperature variation on estimates of temperature sensitivity, we simulated plant litter decomposition using both randomly generated and real time series of temperature. This simulation approach illustrated how temperature variation leads to higher decomposition rates at a given mean temperature than is predicted from simulations in which temperature is held constant. Increases in decomposition rate were most evident at cooler sites, where temporal variation in temperature tends to be greater than at warmer sites. This unbalanced effect of temperature variation shifted the slope of the relationships between average temperature and decomposition rate, resulting in lower estimated temperature sensitivities than were used to simulate decomposition. For example, estimates of activation energy (Ea) were as much as 0.15 eV lower than the true Ea when decomposition was simulated with the true Ea set to the canonical respiration value of 0.65 eV. We found that the estimated Ea was lower than the true Ea for surface, soil, and air temperatures, but not for stream temperatures, for which there was only a weak relationship between temperature variation and mean temperature. Our results suggest that commonly used methods may underestimate the temperature dependence of litter decomposition, particularly in terrestrial environments. We encourage publication of temperature data that include variation estimates and suggest an alternative method for calculating temperature sensitivity that accounts for variation in temperature.