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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.

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.

Corals thrive in low nutrient environments and the conservation of these globally imperiled ecosystems is largely dependent on mitigating the effects of anthropogenic nutrient enrichment. However, to better understand the implications of anthropogenic nutrients requires a heightened understanding of baseline nutrient dynamics within these ecosystems. Here, we provide a novel perspective on coral reef nutrient dynamics by examining the role of fish communities in the supply and storage of nitrogen (N) and phosphorus (P). We quantified fish-mediated nutrient storage and supply for 144 species and modeled these data onto 172 fish communities (71 729 individual fish), in four types of coral reefs, as well as seagrass and mangrove ecosystems, throughout the Northern Antilles. Fish communities supplied and stored large quantities of nutrients, with rates varying among ecosystem types. The size structure and diversity of the fish communities best predicted N and P supply and storage and N : P supply, suggesting that alterations to fish communities (e.g., overfishing) will have important implications for nutrient dynamics in these systems. The stoichiometric ratio (N : P) for storage in fish mass (~8 : 1) and supply (~20 : 1) was notably consistent across the four coral reef types (but not seagrass or mangrove ecosystems). Published nutrient enrichment studies on corals show that deviations from this N : P supply ratio may be associated with poor coral fitness, providing qualitative support for the hypothesis that corals and their symbionts may be adapted to specific ratios of nutrient supply. Consumer nutrient stoichiometry provides a baseline from which to better understand nutrient dynamics in coral reef and other coastal ecosystems, information that is greatly needed if we are to implement more effective measures to ensure the future health of the world's oceans., (© 2014 John Wiley & Sons Ltd.)

Because nutrient enrichment can increase ecosystem productivity, it may enhance resource flows to adjacent ecosystems as organisms cross ecosystem boundaries and subsidize predators in recipient ecosystems. Here, we quantified the biomass and abundance of aquatic emergence and terrestrial spiders in a reference and treatment stream that had been continuously enriched with nitrogen and phosphorus for 5 years. Because we previously showed that enrichment increased secondary production of stream consumers, we predicted that aquatic emergence flux would be higher in the treatment stream, subsequently increasing the biomass and abundance of terrestrial spiders. Those increases were predicted to be greatest for spiders specializing on aquatic emergence subsidies (e.g., Tetragnathidae). By adding a (15)N stable isotope tracer to both streams, we also quantified nitrogen flow from the stream into the riparian community. Emergence biomass, but not abundance, was higher in the treatment stream. The average body size of emerging adult insects and the relative dominance of Trichoptera adults were also greater in the treatment stream. However, spider biomass did not differ between streams. Spiders also exhibited substantially lower reliance on aquatic emergence nitrogen in the treatment stream. This reduced reliance likely resulted from shifts in the body size distributions and community composition of insect emergence that may have altered predator consumption efficiency in the treatment stream. Despite nutrient enrichment approximately doubling stream productivity and associated cross-ecosystem resource flows, the response of terrestrial predators depended more on the resource subsidy's characteristics that affected the predator's ability to capitalize on such increases.

Responses of detrital pathways to nutrients may differ fundamentally from pathways involving living plants: basal carbon resources can potentially decrease rather than increase with nutrient enrichment. Despite the potential for nutrients to accelerate heterotrophic processes and fluxes of detritus, few studies have examined detritus-nutrient dynamics at whole-ecosystem scales. We quantified organic matter (OM) budgets over three consecutive years in two detritus-based Appalachian (U.S.A.) streams. After the first year, we began enriching one stream with low-level nitrogen and phosphorus inputs. Subsequent effects of nutrients on outputs of different OM compartments were determined using randomized intervention analysis. Nutrient addition did not affect dissolved or coarse particulate OM export but had dramatic effects on fine particulate OM (FPOM) export at all discharges relative to the reference stream. After two years of enrichment, FPOM export was 340% higher in the treatment stream but had decreased by 36% in the reference stream relative to pretreatment export. Ecosystem respiration, the dominant carbon output in these systems, also increased in the treatment stream relative to the reference, but these changes were smaller in magnitude than those in FPOM export. Nutrient enrichment accelerated rates of OM processing, transformation, and export, potentially altering food-web dynamics and ecosystem stability in the long term. The results of our large-scale manipulation of a detrital ecosystem parallel those from analogous studies of soils, in which net loss of organic carbon has often been shown to result from experimental nutrient addition at the plot scale. Streams are useful model systems in which to test the effects of nutrients on ecosystem-scale detrital dynamics because they allow both the tracking of OM conversion along longitudinal continua and the integrated measurement of fluxes of transformed material through downstream sites.

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]

Streams store nutrients in standing stocks of organic matter (OM) and associated biologically sequestered elements. Unlike standing stocks of autotrophs, detritus is depleted by nutrient enrichment, potentially reducing areal storage of detritus‐associated nutrients. To test effects of nutrient‐loading on storage of nitrogen (N) and phosphorus (P) by autotrophic and detrital‐pool compartments, we quantified the effects of 2 yr of continuous experimental N and P additions on fine benthic organic matter (FBOM), leaves, wood, and biofilms in five forest streams. Our design tested the relative strength of N vs. P on OM nutrient content, areal OM storage, and areal nutrient storage in OM types. Enrichment increased nutrient content of all OM types; %P increased more than %N in leaves, wood, and biofilms, but not FBOM. Biofilm %P and %N increased more than in all detrital types. Areal FBOM and leaf storage declined with nutrient enrichment. Biofilm standing stocks were generally higher with enrichment but were not related to the streamwater N and P gradients. Despite increased OM nutrient content, total areal nutrient storage in leaves and wood decreased due to reduced OM storage. Although annual nutrient storage was stabilized by FBOM, seasonal variation in nutrient storage increased with enrichment. Leaf‐associated nutrient storage was reduced in most seasons, whereas FBOM and biofilm nutrient storage increased in winter and spring, respectively, relative to pretreatment. Overall, the combined responses of all OM types to enrichment resulted in reduced storage and altered seasonal availability of carbon and nutrients, which has implications for consumers and downstream processes. [ABSTRACT FROM AUTHOR]

Tracking carbon (C) flow through ecosystems requires quantification of myriad biophysical processes, including C routing through microbial and metazoan food webs. Yet detailed organic matter budgets are rarely combined with simultaneous measurement of C flows supporting microbial and animal production. Here, we synthesize concurrent data sets on organic matter, microbes, and macroinvertebrates from two detritus‐based stream ecosystems, one of which was subject to experimental nitrogen (N) and phosphorus (P) enrichment. Our synthesis provides new insights into C flow through forest stream ecosystems. Over 3 yr, the reference stream showed a striking balance of inputs and outputs, with a mean surplus of only 7 g C·m−2·yr−1 (~1% of annual inputs), presumably stored in sediments as fine particulate organic matter (FPOM). In contrast, N and P enrichment over 2 yr resulted in severe deficits of C (−576 g C·m−2·yr−1 or ~170% of annual inputs), a shortfall presumably met by stored C. Our data set provides an ecosystem‐based estimate of the fate of forest litter C at ambient nutrient concentrations: 6.2% was leached as dissolved organic C, 40.6% and 8.5% flowed to litter‐associated fungi and bacteria, respectively, 7.5% was consumed by macroinvertebrates, 1.8% was exported as coarse particles, and the remainder (35.4%) was presumably fragmented by biophysical processes. Our calculations also allowed an estimate of inputs into the heterogeneous FPOM pool, which is otherwise difficult to obtain. At naturally low nutrient concentrations, 50.7% was derived from fragmented litter, 39.1% from microbial biomass (mostly fungal), and 10.2% from macroinvertebrate egesta. Nutrient addition drove large changes in C fluxes in the experimental stream, especially in flows of leaf litter to fungi (×1.7 pretreatment) and macroinvertebrates (×2.7), and of FPOM to hydrologic export (×2.6). Our results underscore the key roles of both microbes and metazoans in controlling C flow through detritus‐based ecosystems, as well as how release from persistent nutrient limitation may perturb steady‐state conditions of C inputs vs. outputs. Our analysis also suggests areas for future research, including assessing the relative importance of stored vs. recycled C in fueling detrital food webs subject to altered nutrient regimes and other global‐change drivers. [ABSTRACT FROM AUTHOR]

Human activities have dramatically altered global patterns of nitrogen (N) and phosphorus (P) availability. This pervasive nutrient pollution is changing basal resource quality in food webs, thereby affecting rates of biological productivity and the pathways of energy and material flow to higher trophic levels.Here, we investigate how the stoichiometric quality of basal resources modulates patterns of material flow through food webs by characterizing the effects of experimental N and P enrichment on the trophic basis of macroinvertebrate production and flows of dominant food resources to consumers in five detritus‐based stream food webs.After a pre‐treatment year, each stream received N and P at different concentrations for 2 years, resulting in a unique dissolved N:P ratio (target range from 128:1 to 2:1) for each stream. We combined estimates of secondary production and gut contents analysis to calculate rates of material flow from basal resources to macroinvertebrate consumers in all five streams, during all 3 years of study.Nutrient enrichment resulted in a 1.5× increase in basal resource flows to primary consumers, with the greatest increases from biofilms and wood. Flows of most basal resources were negatively related to resource C:P, indicating widespread P limitation in these detritus‐based food webs. Nutrient enrichment resulted in a greater proportion of leaf litter, the dominant resource flow‐pathway, being consumed by macroinvertebrates, with that proportion increasing with decreasing leaf litter C:P. However, the increase in efficiency with which basal resources were channelled into metazoan food webs was not propagated to macroinvertebrate predators, as flows of prey did not systematically increase following enrichment and were unrelated to basal resource flows.This study suggests that ongoing global increases in N and P supply will increase organic matter flows to metazoan food webs in detritus‐based ecosystems by reducing stoichiometric constraints at basal trophic levels. However, the extent to which those flows are propagated to the highest trophic levels likely depends on responses of individual prey taxa and their relative susceptibility to predation. [ABSTRACT FROM AUTHOR]

Elevated streamwater nitrogen (N) and phosphorus (P) concentrations can stimulate microbial activity on detrital C and accelerate its breakdown in stream ecosystems. Our study evaluated whether nutrient–detrital relationships are robust across a moderately altered land-use gradient and can be used to identify functional impairment of stream ecosystems. We tested the relative importance of N vs P as likely drivers of these responses, whether responses differed for labile or recalcitrant standardized substrates, and whether responses were detectable across streams with other stressors, which can potentially mask nutrient effects. Two studies were conducted in 23 sites in southeastern US streams. These streams differed in land use and exhibited low-to-moderate gradients in N and P. In study 1, we used 9 sites to compare the relationships between nutrient (N and P) concentrations and microbial respiration and breakdown of 2 standardized C substrates: recalcitrant oak wood veneer and labile cellulose sponge. Both of these substrates are low in nutrient content but differ structurally. In the best supported models, respiration and breakdown rates were positively related to streamwater P, but not N, after 4 wk of stream incubation. Microbial respiration increased 4.2 and 1.2× and breakdown increased 1.8 and 2.3× on cellulose and wood, respectively, across the P gradient. Temperature (+) and specific conductivity (−) were also in top models for wood respiration. Respiration and breakdown were highly correlated for both substrates, indicating the importance of microbial processing in driving breakdown rates. In study 2, we used 23 sites to test for association between landscape nutrient (N and P) gradients and wood veneer breakdown and whether detrital stoichiometry was a better predictor of breakdown than streamwater nutrient concentrations. Wood breakdown was related to P, but not N, and increased 4.1× in 12 wk across the P gradient. Wood nutrient content (increased %N and %P, reduced C∶N and C∶P) was also related to streamwater P and better predicted breakdown (C∶P r 2 = 0.75, C∶N r 2 = 0.87) than streamwater nutrient concentrations. Streamwater P concentrations appeared to stimulate breakdown to a degree that indicates impaired stream function. Our study showed that standardized detrital substrates responses to nutrients 1) were greater to streamwater P than N concentration gradients, 2) occurred on both labile and recalcitrant substrates, and 3) were detectable across landscape gradients with other stressors (e.g., temperature, specific conductivity). These responses likely reflect effects of excess nutrients on diverse C resources in these streams. Wood veneers integrated streamwater nutrient effects, were resistant to physical abrasion, and exhibited significant mass loss even when detritivores were excluded, indicating their value in stream functional assessments under a wide range of stream conditions. [ABSTRACT FROM AUTHOR]

Increases in nitrogen (N) and phosphorus (P) availability are changing animal communities, partly by altering stoichiometric imbalances between consumers and their food. Testing relationships between resource stoichiometry and consumer assemblage structure requires ecosystem‐level manipulations that have been lacking to date.We analysed patterns of macroinvertebrate community composition in five detritus‐based headwater streams subject to experimental whole‐stream N and P additions that spanned a steep gradient in dissolved N:P ratio (2:1, 8:1, 16:1, 32:1, 128:1) over 2 years, following a 1‐year pre‐treatment period.We predicted that shifts in leaf litter stoichiometry would drive overall patterns of community composition via greater responses of shredders to enrichment than other taxa, as shredders dominate primary consumer biomass and experience larger consumer–resource elemental imbalances than other functional groups in stream ecosystems. Specifically, we expected litter C:P to be a significant predictor of shredder biomass given the greater relative imbalances between shredder and litter C:P than C:N. Finally, we tested whether shredder responses to enrichment were related to other taxon‐level traits, including body size and stoichiometry, larval life span and growth rate.Whole‐community composition shifted similarly across the five streams after enrichment, largely driven by increased shredder and predator biomass. These shifts were limited to the autumn/winter seasons and related to decreased leaf litter C:P, highlighting important links between the quality of seasonal litter subsidies and community phenology.Among 10 taxa that drove structural shifts, two declined while other taxa from the same functional/taxonomic groups responded positively, suggesting that specific life‐history traits may determine sensitivity to enrichment.Increases in total shredder biomass, and in biomass of several common shredders, were associated with lower litter C:P. Body C:P did not predict shredder response to enrichment. However, weak negative relationships between shredder response and body size, and larval life span, suggest that small‐bodied and short‐lived taxa may be more responsive to shifting resource stoichiometry.Moderate anthropogenic increases in N and P availability affect resource stoichiometry and can alter animal communities, influencing additional food web and ecosystem properties. We provide support for ecological stoichiometry as a framework for predicting such outcomes based on changes in the elemental composition of resource pools. A plain language summary is available for this article. [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]

Abstract: Humans are altering nutrient dynamics through myriad pathways globally. Concurrent with the addition of nutrients via municipal, industrial, and agricultural sources, widespread consumer exploitation is changing consumer‐mediated nutrient dynamics drastically. Thus, altered nutrient dynamics can occur through changes in the supply of multiple nutrients, as well as through changes in the sources of these nutrients. Seagrass ecosystems are heavily impacted by human activities, with highly altered nutrient dynamics from multiple causes. We simulate scenarios of altered nutrient supply and ratios, nitrogen:phosphorus (N:P), from two nutrient sources in seagrass ecosystems: anthropogenic fertilizer and fish excretion. In doing so we tested expectations rooted in ecological theory that suggest the importance of resource dynamics for predicting primary producer dynamics. Ecosystem functions were strongly altered by artificial fertilizer (e.g., seagrass growth increased by as much as 140%), whereas plant/algae community structure was most affected by fish‐mediated nutrients or the interaction of both treatments (e.g., evenness increased by ~140% under conditions of low fish nutrients and high anthropogenic nutrients). Interactions between the nutrient sources were found for only two of six response variables, and the ratio of nutrient supply was the best predictor for only one response. These findings show that seagrass structure and function are well predicted by supply of a single nutrient (either N or P). Importantly, no single nutrient best explained the majority of responses—measures of community structure were best explained by the primary limiting nutrient to this system (P), whereas measures of growth and density of the dominant producer in the system were best explained by N. Thus, while our findings support aspects of theoretical expectations, the complexity of producer community responses belies broad generalities, underscoring the need to manage for multiple simultaneous nutrients in these imperiled coastal ecosystems. [ABSTRACT FROM AUTHOR]

Abstract: Ecological stoichiometry theory (EST) is a key framework for predicting how variation in N:P supply ratios influences biological processes, at molecular to ecosystem scales, by altering the availability of C, N, and P relative to organismal requirements. We tested EST predictions by fertilizing five forest streams at different dissolved molar N:P ratios (2, 8, 16, 32, 128) for two years and tracking responses of macroinvertebrate consumers to the resulting steep experimental gradient in basal resource stoichiometry (leaf litter %N, %P, and N:P). Nitrogen and P content of leaf litter, the dominant basal resource, increased in all five streams following enrichment, with steepest responses in litter %P and N:P ratio. Additionally, increases in primary consumer biomass and production occurred in all five streams following N and P enrichment (averages across all streams: biomass by 1.2×, production by 1.6×). Patterns of both biomass and production were best predicted by leaf litter N:P and %P and were unrelated to leaf litter %N. Primary consumer production increased most in streams where decreases in leaf litter N:P were largest. Macroinvertebrate predator biomass and production were also strongly positively related to litter %P, providing robust experimental evidence for the primacy of P limitation at multiple trophic levels in these ecosystems. However, production of predatory macroinvertebrates was not related directly to primary consumer production, suggesting the importance of additional controls for macroinvertebrates at upper trophic positions. Our results reveal potential drivers of animal production in detritus‐based ecosystems, including the relative importance of resource quality vs. quantity. Our study also sheds light on the more general impacts of variation in N:P supply ratio on nutrient‐poor ecosystems, providing strong empirical support for predictions that nutrient enrichment increases food web productivity whenever large elemental imbalances between basal resources and consumer demand are reduced. [ABSTRACT FROM AUTHOR]

Abstract: Nitrogen (N) and phosphorus (P) enrichment reduces organic carbon (C) storage in detritus‐based stream ecosystems, but the relative effects of N and P concentrations and ratios on stream metabolic rates have not previously been tested. We tracked changes in whole‐stream ecosystem respiration (ER) and gross primary productivity (GPP), particulate organic matter (POM) standing stocks, fungal biomass, and POM‐specific respiration rates before and during 2 yr of experimental N and P enrichment in five forest streams. Nutrient additions (∼ 96 μg N L−1 to ∼ 472 μg N L−1 and ∼ 10 μg P L−1 to ∼ 85 μg P L−1) targeted dissolved N : P molar ratios of 2, 8, 16, 32, and 128. Whole‐stream ER was positively related to standing stock of wood, a seasonably stable POM compartment that varied by up to 2× among streams. Nutrient enrichment generally increased ER but had no effect on low‐level GPP. Prior to nutrient enrichment, ER was higher at lower N : P, but during enrichment ER increased with increasing N : P. Respiration rates on leaf litter and wood increased with enrichment but decreased with increasing P, and the quantity of leaf litter generally declined with increasing N. Respiration rates on fine benthic organic matter (FBOM) were higher with increasing N : P, and FBOM standing stocks decreased with increasing N. Fungal biomass did not change with nutrient enrichment. Compared to pre‐enrichment conditions, nutrients increased seasonal variation in leaf litter standing stocks and whole‐stream respiration rates. Our results demonstrate how nutrient‐stimulated loss of C from detritus‐based ecosystems occurs through the maintenance of enhanced respiration rates on detrital resources that are particularly sensitive to N inputs. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

We assessed the key role of aquatic fungi in modifying coarse particulate organic matter ( CPOM) by affecting its breakdown rate, nutrient concentration and conversion to fine particulate organic matter ( FPOM). Overall, we hypothesised that fungal-mediated conditioning and breakdown of CPOM would be accelerated when nutrient concentrations are increased and tested the degree to which fungi were critical to CPOM processing and to FPOM production by an invertebrate consumer., We manipulated the presence and absence of fungi, exogenous nutrients [nitrogen (N) and phosphorus (P)] and an invertebrate consumer in a full-factorial laboratory experiment and quantified their effects on CPOM mass loss and nutrient concentration, and the quantity and nutrient concentration of FPOM produced during leaf breakdown., Mass of CPOM lost and the quantity of FPOM produced were highest in nutrient-enriched treatments containing fungal decomposers. Across all treatments, FPOM produced was a constant proportion of CPOM lost (67%). Loss of CPOM due to shredders was highest at fungal biomass values >16 mg g dry mass−1, which occurred with nutrient enrichment., Nitrogen concentration in CPOM increased in treatments with nutrients and fungi, while CPOM P concentration was primarily affected by nutrients. In contrast, FPOM P concentration declined in treatments with fungi and nutrients, suggesting either sequestration via CPOM-associated fungi or preferential assimilation by shredders. Nitrogen in FPOM increased with nutrients, but was unaffected by fungi., Our results indicate that aquatic fungi play a critical role in facilitating energy and nutrient flow through detrital pathways and that their ability to mediate organic matter transformations is significantly influenced by nutrient enrichment. [ABSTRACT FROM AUTHOR]

Small-scale experiments and theory suggest that ecological functions provided by communities become more stable with increased species richness. Whether these patterns manifest at regional spatial scales and within species-rich communities (e.g., coral reefs) is largely unknown. We quantified five biogeochemical processes, and an aggregate measure of multifunctionality, in species-rich coastal fish communities to test three questions: (1) Do previously predicted biodiversity-ecosystem-function relationships hold across large spatial scales and in highly diverse communities? (2) Can additional covariates of community structure improve these relationships? (3) What is the role of community biomass and functional group diversity in maintaining biogeochemical processes under various scenarios of species loss across ecosystem types? These questions were tested across a large regional gradient of coral reef, mangrove and seagrass ecosystems. Statistical models demonstrated that species richness and the mean maximum body size per species strongly predicted biogeochemical processes in all ecosystem types, but functional group diversity was only a weak predictor. Simulating three scenarios of species loss demonstrated that conserving community biomass alone increased the ability for communities to maintain ecosystem processes. Multifunctionality of biogeochemical processes was maintained least in simulations that conserved biomass and community structure, underscoring the relative lack of importance of community structure in maintaining multiple simultaneous ecosystem functions in this system. Findings suggest that conserving community biomass alone may be sufficient to sustain certain biogeochemical processes, but when considering conservation of multiple simultaneous biogeochemical processes, management efforts should focus first on species richness. [ABSTRACT FROM AUTHOR]

Abstract: The traditional model of nutrient availability in coastal estuarine ecosystems is based on predictable inputs of nitrogen (N) and phosphorus (P) via riverine and oceanic sources, respectively. But coastlines with low nutrient input from these sources may not fit into this simple framework. Here we use observational (seagrass nutrient content) and experimental (nutrient enrichment assays) data for assessing nutrient availability and limitation for primary producers along a spatial transect extending from the mouth (nearest to the ocean) to the terminal portion (boundary with the terrestrial ecosystem) of three coastal mangrove-lined tidal creeks in The Bahamas. Compiling seagrass nutrient content from all sites showed a negative relationship between seagrass nutrient limitation (either N or P) and distance from mouth, but this pattern differed across sites with respect to which nutrient was more limiting. Our experimental results demonstrated patterns of decreased response by microalgae to dual nutrient enrichment in one site with distance from the creek mouth, and increased response to single nutrient enrichment in another, with the third showing no trend along this gradient. Our findings show that Bahamian mangrove wetlands are extremely nutrient-limited ecosystems, and that the most limiting nutrient varied among sites. In general, these ecosystems deviate from the typical paradigm of spatial nutrient limitation patterns in estuaries. We suggest that various site-specific biological and physical factors may be more important than large-scale hydrologic factors in driving trends of nutrient availability in coastal ecosystems under strong nutrient constraints, such as in The Bahamas. Our findings suggest that even minor changes in nutrient loading rates can have significant implications for primary production in subtropical oligotrophic systems. [Copyright &y& Elsevier]

The article presents a study which examines the impact of overfishing and habitat fragmentation in the decline of population-level nutrient recycling of gray snapper in Bahamas tidal creeks. The study uses excretion measures and field surveys of fish abundance to evaluate the population-level excretion rates of gray snapper (Lutjanus griseus). Result shows that the human disturbances have resulted in the reductions of recycling rates.

1. We determined the effects of nutrient enrichment on wood decomposition rates and microbial activity during a 3-year study in two head water streams at Coweeta Hydrologic Laboratory, NC, U.S.A. After a 1-year pretreatment period, one of the streams was continuously enriched with inorganic nutrients (nitrogen and phosphorus) for 2 years while the other stream served as a reference. We determined the effects of enrichment on both wood veneers and sticks, which have similar carbon quality but differ in physical characteristics (e.g. surface area to volume ratios, presence of bark) that potentially affect microbial colonisation and activity. 2. Oak wood veneers (0.5 mm thick) were placed in streams monthly and allowed to decompose for approximately 90 days. Nutrient addition stimulated ash-free dry mass loss and increased mean nitrogen content, fungal biomass and microbial respiration on veneers in the treatment stream compared with the reference. The magnitude of the response to enrichment was great, with mass loss 6.1 times, and per cent N, fungal biomass and microbial respiration approximately four times greater in the treatment versus reference stream. 3. Decomposition rate and nitrogen content of maple sticks (ca. 1–2 cm diameter) also increased; however, the effect was less pronounced than for veneers. Wood response overall was greater than that determined for leaves in a comparable study, supporting the hypothesis that response to enrichment may be greater for lower quality organic matter (high C : N) than for higher quality (low C : N) substrates. 4. Our results show that moderate nutrient enrichment can profoundly affect decomposition rate and microbial activity on wood in streams. Thus, the timing and availability of wood that provides retention, structure, attachment sites and food in stream ecosystems may be affected by nutrient concentrations raised by human activities. [ABSTRACT FROM AUTHOR]

Abstract Stoichiometric relationships between consumers and resources in detritus-based ecosystems have received little attention, despite the importance of detritus in most food webs. We analysed carbon (C), nitrogen (N), and phosphorus (P) content of invertebrate consumers, and basal food resources in two forested headwater streams (one reference and the other nutrient-enriched). We found large elemental imbalances between consumers and food resources compared with living plant-based systems, particularly in regard to P content, which were reduced with enrichment. Enrichment significantly increased nutrient content of food resources (consistent with uptake of N and P by detritus-associated microbes). P content of some invertebrates also increased in the enriched vs. reference stream, suggesting deviation from strict homeostasis. Nutrient content varied significantly among invertebrate functional feeding groups, orders and, to some extent, size classes. Future application of stoichiometric theory to detritus-based systems should consider the potential for relatively large consumer-resource elemental imbalances and P storage by insect consumers. [ABSTRACT FROM AUTHOR]

We examined benthic algal response to nutrient enrichment by nitrogen (N), phosphorus (P), and N + P in mangrove wetlands of The Bahamas, to test the hypothesis that human impacts (fragmentation) on these ecosystems altered nutrient limitation, thereby changing the frequency and/or magnitude at which ecological synergies occurred. Fragmentation occurred due to road construction, resulting in reduced hydrological connectivity between the wetlands and marine environment. Strong, persistent, and synergistic nutrient colimitation occurred in both pristine and fragmented estuaries. Ecosystem fragmentation did not alter the biomass response to dual nutrient enrichment, but did alter the relative magnitude of the nonadditive response. That is, synergistic responses were less extreme in fragmented systems. This was supported by the strong, negative relationship between ambient algal biomass (a surrogate for background productivity) and the strength of synergistic responses (R² = 0.69 and 0.79, year 1 and year 2, respectively). Bahamian coastal ecosystems exhibited the greatest synergistic responses reported for a marine ecosystem, suggesting that the benthic algal communities associated with Bahamian wetlands are among the most nutrient-limited marine ecosystems. Our findings provide a case study illustrating how altered nutrient dynamics associated with land-use change may decrease the frequency and/or magnitude of synergistic responses to nutrients in aquatic ecosystems. [ABSTRACT FROM AUTHOR]