Nutrient and stoichiometric time series measurements of decomposing coarse detritus in freshwaters.

Decomposition of coarse detritus (e.g., dead organic matter larger than ~1 mm such as leaf litter or animal carcasses) in freshwater ecosystems is well described in terms of mass loss, particularly as rates that compress mass loss into one number (e.g., a first-order decay coefficient, or breakdown rate, "k"); less described are temporal changes in the elemental composition of these materials during decomposition, with important implications for elemental cycling from microbes to ecosystems. This stands in contrast with work in the terrestrial realm, where a focus on detrital elemental cycling has provided a sharper mechanistic understanding of decomposition, especially with specific processes such as immobilization and mineralization. Notably, freshwater ecologists often measure carbon (C), nitrogen (N), and phosphorus (P), and their stoichiometric ratios in decomposing coarse materials, including carcasses, wood, leaf litter, and more, but these measurements remain piecemeal. These detrital nutrients are measurements of the entire detrital-microbial complex and are integrative of numerous processes, especially nutrient immobilization and mineralization, and associated microbial growth and death. Thus, data relevant to an elemental, mechanistically focused decomposition ecology are available in freshwaters, but have not been fully applied to that purpose. We synthesized published detrital nutrient and stoichiometry measurements at a global scale, yielding 4038 observations comprising 810 decomposition time series (i.e., measurements within a defined cohort of decomposing material through time) to build a basis for understanding the temporality of elemental content in freshwater detritus. Specifically, the dataset focuses on temporally and ontogenetically (mass loss) explicit measurements of N, P, and stoichiometry (C:N, C:P, N:P). We also collected ancillary data, including detrital characteristics (e.g., species, lignin content), water physiochemistry, geographic location, incubation system type, and methodological variables (e.g., bag mesh size). These measurements are important to unlocking mechanistic insights into detrital ontogeny (the temporal trajectory of decomposing materials) that can provide a deeper understanding of heterotroph-driven C and nutrient cycling in freshwaters. Moreover, these data can help to bridge aquatic and terrestrial decomposition ecology, across plant or animal origin. By focusing on temporal trajectories of elements, this dataset facilitates cross-ecosystem comparisons of fundamental decomposition controls on elemental fluxes. It provides a strong starting point (e.g., via modeling efforts) for comparing processes such as immobilization and mineralization that are understudied in freshwaters. Time series from decomposing leaf litter, particularly in streams, are common in the dataset, but we also synthesized ontogenies of animal-based detritus, which tend to decompose rapidly compared with plant-based detritus that contains high concentrations of structural compounds such as lignin and cellulose. Although animal-based data were rare, comprising only three time series, their inclusion in this dataset underscores the opportunities to develop an understanding of decomposition that encompasses all detrital types, from carrion to leaf litter. There are no copyright or proprietary restrictions on the dataset; please cite this data paper when reusing these materials.

Nutrient and stoichiometry dynamics of decomposing litter in stream ecosystems: A global synthesis.

Decomposing organic matter forms a substantial resource base, fueling the biogeochemical function and secondary production of most aquatic ecosystems. However, detrital N (nitrogen) and P (phosphorus) dynamics remain relatively unexplored in aquatic ecosystems relative to terrestrial ecosystems, despite fundamentally linking microbial processes to ecosystem function across broad spatial scales. We synthesized 217 published time series of detrital carbon (C), N, P, and their stoichiometric ratios (C:N, C:P, N:P) from stream ecosystems to analyze the temporal nutrient dynamics of decomposing litter using generalized additive models. Model results indicated that detritus was a net source of N (irrespective of inorganic or organic form) to the environment, regardless of initial N content. In contrast, P sink/source dynamics were more strongly influenced by the initial P content, in which P-poor litters were sinks for nutrients until these shifted to net P mineralization after ~40% mass loss. However, large variations surrounded both the N and P predictions, suggesting the importance of nonmicrobial factors such as fragmentation by invertebrates. Detrital C:N ratios converged and became more similar toward the end of the decomposition, suggesting predictable microbial functional effects throughout detrital ontogeny. C:P and N:P ratios also converged to some degree, but these model predictions were less robust than for C:N, due in part to the lower number of published detrital C:P time series. The explorations of environmental covariate effects were frequently limited by a few coincident covariate measurements across studies, but temperature, N availability, and P tended to accelerate the existing ontogenetic patterns in C:N. Our analysis helps to unite organic matter decomposition across aquatic-terrestrial boundaries by describing the basic patterns of elemental flows catalyzed by decomposition in streams, and points to a research agenda with which to continue addressing gaps in our knowledge of detrital nutrient dynamics across ecosystems.

Transport of N and P in U.S. streams and rivers differs with land use and between dissolved and particulate forms.

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.

Nutrients and temperature additively increase stream microbial respiration.

Rising temperatures and nutrient enrichment are co-occurring global-change drivers that stimulate microbial respiration of detrital carbon, but nutrient effects on the temperature dependence of respiration in aquatic ecosystems remain uncertain. We measured respiration rates associated with leaf litter, wood, and fine benthic organic matter (FBOM) across seasonal temperature gradients before (PRE) and after (ENR1, ENR2) experimental nutrient (nitrogen [N] and phosphorus [P]) additions to five forest streams. Nitrogen and phosphorus were added at different N:P ratios using increasing concentrations of N (~80-650 µg/L) and corresponding decreasing concentrations of P (~90-11 µg/L). We assessed the temperature dependence, and microbial (i.e., fungal) drivers of detrital mass-specific respiration rates using the metabolic theory of ecology, before vs. after nutrient enrichment, and across N and P concentrations. Detrital mass-specific respiration rates increased with temperature, exhibiting comparable activation energies (E, electronvolts [eV]) for all substrates (FBOM E = 0.43 [95% CI = 0.18-0.69] eV, leaf litter E = 0.30 [95% CI = 0.072-0.54] eV, wood E = 0.41 [95% CI = 0.18-0.64] eV) close to predicted MTE values. There was evidence that temperature-driven increased respiration occurred via increased fungal biomass (wood) or increased fungal biomass-specific respiration (leaf litter). Respiration rates increased under nutrient-enriched conditions on leaves (1.32×) and wood (1.38×), but not FBOM. Respiration rates responded weakly to gradients in N or P concentrations, except for positive effects of P on wood respiration. The temperature dependence of respiration was comparable among years and across N or P concentration for all substrates. Responses of leaf litter and wood respiration to temperature and the combined effects of N and P were similar in magnitude. Our data suggest that the temperature dependence of stream microbial respiration is unchanged by nutrient enrichment, and that increased temperature and N + P availability have additive and comparable effects on microbial respiration rates.

Seasonally distinct taxonomic and functional shifts in macroinvertebrate communities following dam removal.

Dam removal is an increasingly popular restoration tool, but our understanding of ecological responses to dam removal over time is still in the early stages. We quantified seasonal benthic macroinvertebrate density, taxonomic composition, and functional traits for three years after lowhead dam removal in three reaches of the Olentangy River (Ohio, USA): two upstream of former dam (one restored, one unrestored), and one downstream of former dam. Macroinvertebrate community density, generic richness, and Shannon-Wiener diversity decreased between ∼9 and ∼15 months after dam removal; all three variables consistently increased thereafter. These threshold responses were dependent on reach location: density and richness increased ∼15 months after removal in upstream reaches versus ∼19 months downstream of the former dam. Initial macroinvertebrate density declines were likely related to seasonality or life-history characteristics, but density increased up to 2.27נfrom year to year in three out of four seasons (late autumn, early spring, summer) across all reaches. Macroinvertebrate community composition was similar among the three reaches, but differed seasonally based on non-metric multidimensional scaling (NMDS) and analysis of similarity (ANOSIM). Seasonal differences among communities tended to decrease after dam removal. We detected community-wide shifts in functional traits such as multivoltinism, depositional habitat use, burrowing, and collector-gatherer feeding mode. We observed that these traits were expressed most strongly with Chironomidae, which was the most abundant family. Our results suggest that seasonal environmental conditions can play a role in the response and recovery of macroinvertebrate communities-often used to monitor ecosystem condition-following dam removal. In particular, macroinvertebrate density and diversity can show recovery after dam removal, especially in seasons when macroinvertebrate density is typically lowest, with concomitant changes to functional trait abundance. Thus, we recommend scientists and managers consider responses to dam removal throughout the year. Further, similar density, generic richness, and functional traits among reaches suggest that channel restoration after dam removal may initially have equivocal effects on invertebrate communities.

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.

Convergence of detrital stoichiometry predicts thresholds of nutrient-stimulated breakdown in streams.

Nutrient enrichment of detritus-based streams increases detrital resource quality for consumers and stimulates breakdown rates of particulate organic carbon (C). The relative importance of dissolved inorganic nitrogen (N) vs. phosphorus (P) for detrital quality and their effects on microbial- vs. detritivore-mediated detrital breakdown are poorly understood. We tested effects of experimental N and P additions on detrital stoichiometry (C:N, C:P) and total and microbial breakdown (i.e., with and without detritivorous shredders, respectively) of five detritus types (four leaf litter species and wood) with different initial C : nutrient content. We enriched five headwater streams continuously for two years at different relative availabilities of N and P and compared breakdown rates and detrital stoichiometry to pretreatment conditions. Total breakdown rates increased with nutrient enrichment and were predicted by altered detrital stoichiometry. Streamwater N and P, fungal biomass, and their interactions affected stoichiometry of detritus. Streamwater N and P decreased detrital C:N, whereas streamwater P had stronger negative effects on detrital C:P. Nutrient addition and fungal biomass reduced C:N by 70% and C:P by 83% on average after conditioning, compared to only 26% for C:N and 10% for C:P under pretreatment conditions. Detritus with lowest initial nutrient content changed the most and had greatest increases in total breakdown rates. Detrital stoichiometry was reduced and differences among detritus types were homogenized by nutrient enrichment. With enrichment, detrital nutrient content approached detritivore nutritional requirements and stimulated greater detritivore vs. microbial litter breakdown. We used breakpoint regression to estimate values of detrital stoichiometry that can potentially be used to indicate elevated breakdown rates. Breakpoint ratios for total breakdown were 41 (C:N) and 1518 (C:P), coinciding with total breakdown rates that were ~1.9 times higher when C:N or C:P fell below these breakpoints. Microbial and shredder-mediated breakdown rates both increased when C:N and C:P were reduced, suggesting that detrital stoichiometry is useful for predicting litter breakdown dominated by either microbial or shredder activity. Our results show strong effects of nutrient enrichment on detrital stoichiometry and offer a robust link between a potential holistic nutrient loading metric (decreased and homogenized detrital stoichiometry) and increased C loss from stream ecosystems.

Detrital stoichiometry as a critical nexus for the effects of streamwater nutrients on leaf litter breakdown rates.

Nitrogen (N) and phosphorus (P) concentrations are elevated in many freshwater systems, stimulating breakdown rates of terrestrially derived plant litter; however, the relative importance of N and P in driving litter breakdown via microbial and detritivore processing are not fully understood. Here, we determined breakdown rates of two litter species, Acer rubrum (maple) and Rhododendron maximum (rhododendron), before (PRE) and during two years (YR1, YR2) of experimental N and P additions to five streams, and quantified the relative importance of hypothesized factors contributing to breakdown. Treatment streams received a gradient of P additions (low to high soluble reactive phosphorus [SRP]; ~10-85 µg/L) crossed with a gradient of N additions (high to low dissolved inorganic nitrogen [DIN]; ~472-96 µg/L) to achieve target molar N:P ratios ranging from 128 to 2. Litter breakdown rates increased above pre-treatment levels by an average of 1.1-2.2x for maple, and 2.7-4.9x for rhododendron in YR1 and YR2. We used path analysis to compare fungal biomass, shredder biomass, litter stoichiometry (nutrient content as C:N or C:P), discharge, and streamwater temperature as predictors of breakdown rates and compared models containing streamwater N, P or N + P and litter C:N or C:P using model selection criteria. Litter breakdown rates were predicted equally with either streamwater N or P (R2 = 0.57). In models with N or P, fungal biomass, litter stoichiometry, discharge, and shredder biomass predicted breakdown rates; litter stoichiometry and fungal biomass were most important for model fit. However, N and P effects may have occurred via subtly different pathways. Litter N content increased with fungal biomass (N-driven effects) and litter P content increased with streamwater P availability (P-driven effects), presumably via P storage in fungal biomass. In either case, the effects of N and P through these pathways were associated with higher shredder biomass and breakdown rates. Our results suggest that N and P stimulate litter breakdown rates via mechanisms in which litter stoichiometry is an important nexus for associated microbial and detritivore effects.

Low-to-moderate nitrogen and phosphorus concentrations accelerate microbially driven litter breakdown rates.

Particulate organic matter (POM) processing is an important driver of aquatic ecosystem productivity that is sensitive to nutrient enrichment and.drives ecosystem carbon (C) loss. Although studies of single concentrations of nitrogen (N) or phosphorus (P) have shown effects at relatively low concentrations, responses of litter breakdown rates along gradients of low-to-moderate N and P concentrations are needed to establish likely interdependent effects of dual N and P enrichment on baseline activity in stream ecosystems. We established 25 combinations of dissolved inorganic N (DIN; 55-545 µg/L) and soluble reactive P (SRP; 4-86 µg/L) concentrations with corresponding N:P molar ratios of 2-127 in experimental stream channels. We excluded macroinvertebrates, focusing on microbially driven breakdown of maple (Acer rubrum L.) and rhododendron (Rhododendron maximum L.) leaf litter. Breakdown rates, k, per day (d-1) and per degree-day (dd-l), increased by up to 6X for maple and 12× for rhododendron over our N and P enrichment gradient compared to rates at low ambient N and P concentrations. The best models of k (d- and dd-1) included litter species identity and N and P concentrations; there was evidence for both additive and interactive effects of N and P. Models explaining variation in k dd-1 were supported by N and P for both maple and rhododendron (R =0.67 and 0.33, respectively). Residuals in the relationship between k dd-1 and N concentration were largely explained by P, but residuals for k dd-1 and P. concentration were less adequately explained by N. Breakdown rates were more closely related to nutrient concentrations than variables associated with measurements of two mechanistic parameters associated with C loss (fungal biomass and microbial respiration rate). We also determined the effects of nutrient addition on litter C: nutrient stoichiometry and found reductions in litter C:N and C:P along our experimental nutrient gradient. Our results indicate that microbially driven litter processing rates increase across low-to-moderate nutrient gradients that are now common throughout human-modified landscapes.

Freshwater ecology. Experimental nutrient additions accelerate terrestrial carbon loss from stream ecosystems.

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.