Nitrogen and phosphorus enrichment cause declines in invertebrate populations: a global meta-analysis

Human-driven changes in nitrogen (N) and phosphorus (P) inputs are modifying biogeochemical cycles and the trophic state of many habitats worldwide. These alterations are predicted to continue to increase, with the potential for a wide range of impacts on invertebrates, key players in ecosystem-level processes. Here, we present a meta-analysis of 1679 cases from 207 studies reporting the effects of N, P, and combined N + P enrichment on the abundance, biomass, and richness of aquatic and terrestrial invertebrates. Nitrogen and phosphorus additions decreased invertebrate abundance in terrestrial and aquatic ecosystems, with stronger impacts under combined N + P additions. Likewise, N and N + P additions had stronger negative impacts on the abundance of tropical than temperate invertebrates. Overall, the effects of nutrient enrichment did not differ significantly among major invertebrate taxonomic groups, suggesting that changes in biogeochemical cycles are a pervasive threat to invertebrate populations across ecosystems. The effects of N and P additions differed significantly among invertebrate trophic groups but N + P addition had a consistent negative effect on invertebrates. Nutrient additions had weaker or inconclusive impacts on invertebrate biomass and richness, possibly due to the low number of case studies for these community responses. Our findings suggest that N and P enrichment affect invertebrate community structure mainly by decreasing invertebrate abundance, and these effects are dependent on the habitat and trophic identity of the invertebrates. These results highlight the important effects of human-driven nutrient enrichment on ecological systems and suggest a potential driver for the global invertebrate decline documented in recent years.     

Signatures of nutrient limitation and co‐limitation: responses of autotroph internal nutrient concentrations to nitrogen and phosphorus additions

Humans are modifying the availability of nutrients such as nitrogen (N) and phosphorus (P), and it is therefore important to understand how these nutrients, independently or in combination, influence the growth and nutrient content of primary producers. Using meta‐analysis of 118 field and laboratory experiments in freshwater, marine and terrestrial ecosystems, we tested hypotheses about co‐limitation of N and P by comparing the effects of adding N alone, P alone, and both N and P together on internal N (e.g. %N, C:N) and P (e.g. %P, C:P) concentrations in autotroph communities. In particular, we tested the following predictions. First, if only one nutrient was limiting, addition of that nutrient should decrease the concentration of the other nutrient, but addition of the non‐limiting nutrient would have no effect on the internal concentration of the limiting nutrient. If community co‐limitation was occurring then addition of either nutrient should result in a decrease in the internal concentration of the other nutrient. Community co‐limitation could also result in no change – or even an increase – in N concentrations in response to P addition if P stimulated growth of N fixers. Finally, if biochemically dependent co‐limitation was occurring, addition of a limiting nutrient would not decrease, and could even increase, the concentration of the other, co‐limited nutrient. We found no general evidence for the decrease in the internal concentration of one nutrient due to addition of another nutrient. The one exception to this overall pattern was marine systems, where N addition decreased internal P concentrations. In contrast, P addition increased internal N concentrations across all experiments, consistent with co‐limitation. These results have important implications for understanding the roles that N and P play in controlling producer growth and internal nutrient accumulation as well as for managing the effects of nutrient enrichment in ecosystems. Synthesis On a global scale, humans have doubled nitrogen (N) inputs and quadrupled phosphorus (P) inputs relative to pre‐industrial levels. N and P fertilization influences autotroph internal nutrient concentrations and ratios and thereby affects a variety of community and ecosystem processes, including decomposition and consumer population dynamics. It is therefore critical to understand the effects of nutrient additions on the growth and nutrient concentrations of primary producers. We used meta‐analysis to evaluate the responses of autotroph internal N and P concentrations to additions of N, P, and N+P and make inferences about limitation and co‐limitation of N and P across marine, terrestrial, and freshwater ecosystems. We found little evidence for single‐nutrient limitation, highlighting the fact that multiple nutrients generally limit primary production.     

Plant species loss from European semi‐natural grasslands following nutrient enrichment – is it nitrogen or is it phosphorus?

Aim Although many studies support the prevailing paradigm of nitrogen (N)‐driven biodiversity loss, some have argued that phosphorus (P) may be the main culprit. This questions the generality of the global threat through N enrichment. The major objective here was to quantify the relative importance of soil N and P in explaining patterns of plant species richness, under different levels of N and P limitation. Location North‐western Europe. Methods We collected soil, productivity and plant species data from 132 semi‐natural grasslands located along a gradient of nutrient availability and atmospheric N deposition. We used linear mixed models to investigate the relation between soil nutrients, acidity, limitation and productivity on one side, and indices for plant species richness on the other. Results Mixed models explained between 38 and 50% of the total variation in species numbers, forbs and endangered species. Soil P was significantly negatively related to total species number, forbs and endangered species. Soil N was only significantly negatively related to number of forbs and endangered species. Compared with soil P, the explained variation attributed to soil N was between five‐ and twenty‐fold lower. P‐limited grasslands exhibited higher species richness, numbers of forbs and endangered species. Species richness and number of forbs decreased with lower soil acidity. N deposition was negatively related to the number of forbs and endangered species, as well as to soil acidity. Productivity was weakly positively related to soil P and negatively to species and forb numbers. We found no interaction factors between the explanatory variables. Main conclusions P enrichment can present a greater threat to biodiversity than N enrichment in at least some terrestrial ecosystems. However, as N‐ and P‐driven species loss appeared independent, our results suggest that simultaneously reducing N and P inputs is a prerequisite for maintaining maximum plant diversity.     

Variation in Detrital Resource Stoichiometry Signals Differential Carbon to Nutrient Limitation for Stream Consumers Across Biomes

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.     

Changes in stoichiometric constraints on epilithon and benthic macroinvertebrates in response to slight nutrient enrichment of mountain rivers

1. To assess changes in stoichiometric constraints on stream benthos, we measured elemental composition of epilithon and benthic macroinvertebrates in intrinsically P‐limited mountain rivers, upstream and downstream of low‐level anthropogenic nutrient enrichment by effluents of municipal wastewater treatment plants. 2. While there was a broad range in the elemental composition of epilithon (C : P ratios of 200–16 500, C : N ratios of 8–280, N : P ratios of 8–535) and heptageniid mayfly scrapers (C : P ratios of 125–300, C : N ratios of 5.1–7.2, N : P ratios of 20–60), the average C : P ratio of epilithon was 10‐fold lower and the average C : N ratio twofold lower at more nutrient‐rich downstream sites. Nutrient ratios in benthic macroinvertebrates were lower than in epilithon and varied little between relatively nutrient‐poor and nutrient‐rich sites. 3. We modified the existing definition of producer‐consumer elemental imbalance to allow for variation in consumer nutrient content. We defined this ‘non‐homeostatic’ imbalance as the perpendicular distance between the producer and consumer C : P, C : N, or N : P ratios, and the 1 : 1 line. 4. At P‐limited sites, the estimated mayfly N : P recycling ratio was higher than the N : P ratio in epilithon, suggesting nutrient recycling by consumers could accentuate P‐limitation of epilithon. 5. Measuring the degree of producer–consumer nutrient imbalance may be important in predicting the magnitude of effects from nutrient enrichment and can help elucidate the causes and consequences of ecological patterns and processes in rivers.     

Species richness loss after nutrient addition as affected by N:C ratios and phytohormone GA3 contents in an alpine meadow community

Aims From the light-competition hypothesis, competition for light is asymmetric and the observed increases in plant-size variability with increasingly denser canopies are primarily due to competition for light. Greater plant height provides pre-emptive access to light and produces increased height differences among species. The question is what produces these differences in plant height or height growth response among species in response to fertilization.Methods In 2009, a field experiment of N, P and N + P enrichments at three levels each was initiated in an alpine meadow on the northeast Qinghai-Tibet Plateau. Effects of fertilization on species richness, aboveground net primary production (ANPP), relative light intensity and plant height of different plant functional groups were determined. Festuca ovina (grass), Kobresia humilis (sedge), Oxytropis ochrocephala (legume), Taraxacum lugubre (rosette forb) and Geranium pylzowianum (upright forb) were selected as exemplars of each of the indicated functional groups. The N:C ratios in aboveground biomass, gibberellic acid (GA 3) concentrations in leaves, plant heights and height relative growth rate (RGR) of these exemplar species were analyzed in detail.Important findings Species richness of grasses significantly increased with increasing N + P levels. Species richness of legumes and upright forbs decreased after N and N + P additions. P addition had no significant effect on species richness. The effects of N + P addition on species richness and ANPP were consistently stronger than those of the single N or P fertilization. Reductions in species richness caused by nutrient addition paralleled the increases in ANPP and decreases in light intensity under the canopies, indicating indirect effect of nutrient addition on species richness via ANPP-induced light competition. The exemplar species that responded most positively to fertilization in height and RGR also displayed stronger increases in their GA 3 content and N:C ratios. GA 3 concentrations and N:C ratios were positively correlated with height RGR when the data were pooled for all species. The tallest and the fastest-growing grass, F. ovina, had the largest increase in N:C ratios and the highest leaf GA 3 concentrations after nutrient addition. These results indicated that differential responses of GA 3 concentrations and N:C ratios to fertilization were related to the inequality in plant heights among species.     

长期施肥黄土旱塬农田土壤-微生物-植物系统碳氮磷生态化学计量特征

探讨长期不同施肥制度对农田土壤、植物生态系统的碳(C)、氮(N)、磷(P)含量及其生态化学计量比的影响,可为揭示该系统能量平衡和养分循环,实现农业生态系统元素平衡及可持续发展提供参考意义。以位于黄土高原半干旱地区的长武国家黄土高原农业生态实验站长期施肥试验为研究对象,选取不施肥(CK)、单施氮肥(N)、单施磷肥(P)、施氮磷肥(NP)、单施有机肥(M)、氮肥配施有机肥(NM)、磷肥配施有机肥(PM)、氮磷肥配施有机肥(NPM)8个处理,分析了黄土旱塬典型农田土壤-微生物-植物生态系统中C、N、P含量及其生态化学计量变化规律。研究结果表明:1)长期单施有机肥和化肥配施有机肥处理可显著提高土壤和有机质C、N、P含量。2)氮、磷肥的输入显著降低了土壤和小麦C∶N、N∶P,施P显著降低了有机态C∶P和小麦C∶P;有机肥配施对微生物生物量和小麦C∶N∶P的影响更为明显。3)长期有机肥配施条件下土壤养分和小麦化学计量比存在较强的相关关系。微生物生物量碳与有机C、N、P呈显著正相关,土壤微生物生物量氮与土壤N、P总量呈显著正相关,微生物生物量磷与土壤C、N、P总量含量呈显著负相关;植株碳含量与微生物...     

Non‐additive effects of litter mixing are suppressed in a nutrient‐enriched stream

Investigations of how species compositional changes interact with other aspects of global change, such as nutrient mobilization, to affect ecosystem processes are currently lacking. Many studies have shown that mixed species plant litters exhibit non‐additive effects on ecosystem functions in terrestrial and aquatic systems. Using a full‐factorial design of three leaf litter species with distinct initial chemistries (carbon:nitrogen; C:N) and breakdown rates (Liriodendron tulipifera, Acer rubrum and Rhododendron maximum), we tested for additive and non‐additive effects of litter species mixing on breakdown in southeastern US streams with and without added nutrients (N and phosphorus). We found a non‐additive (antagonistic) effect of litter mixing on breakdown rates under reference conditions but not when nutrient levels were elevated. Differential responses among single‐species litters to nutrient enrichment contributed to this result. Antagonistic litter mixing effects on breakdown were consistent with trends in litter C:N, which were higher for mixtures than for single species, suggesting lower microbial colonization on mixtures. Nutrient enrichment lowered C:N and had the greatest effect on the lowest‐ (R. maximum) and the least effect on the highest‐quality litter species (L. tulipifera), resulting in lower interspecific variation in C:N. Detritivore abundance was correlated with litter C:N in the reference stream, potentially contributing to variation in breakdown rates. In the nutrient‐enriched stream, detritivore abundance was higher for all litter and was unrelated to C:N. Thus, non‐additive effects of litter mixing were suppressed by elevated streamwater nutrients, which increased nutrient content of all litter, reduced variation in C:N among litter species and increased detritivore abundance. Nutrients reduced interspecific variation among plant litters, the base of important food web pathways in aquatic ecosystems, affecting predicted mixed‐species breakdown rates. More generally, world‐wide mobilization of nutrients may similarly modify other effects of biodiversity on ecosystem processes.     

Environmental and stoichiometric controls on microbial carbon-use efficiency in soils

Carbon (C) metabolism is at the core of ecosystem function. Decomposers play a critical role in this metabolism as they drive soil C cycle by mineralizing organic matter to CO(2) . Their growth depends on the carbon-use efficiency (CUE), defined as the ratio of growth over C uptake. By definition, high CUE promotes growth and possibly C stabilization in soils, while low CUE favors respiration. Despite the importance of this variable, flexibility in CUE for terrestrial decomposers is still poorly characterized and is not represented in most biogeochemical models. Here, we synthesize the theoretical and empirical basis of changes in CUE across aquatic and terrestrial ecosystems, highlighting common patterns and hypothesizing changes in CUE under future climates. Both theoretical considerations and empirical evidence from aquatic organisms indicate that CUE decreases as temperature increases and nutrient availability decreases. More limited evidence shows a similar sensitivity of CUE to temperature and nutrient availability in terrestrial decomposers. Increasing CUE with improved nutrient availability might explain observed declines in respiration from fertilized stands, while decreased CUE with increasing temperature and plant C?:?N ratios might decrease soil C storage. Current biogeochemical models could be improved by accounting for these CUE responses along environmental and stoichiometric gradients.     

Importance of Terrestrial Arthropods as Subsidies in Lowland Neotropical Rain Forest Stream Ecosystems

The importance of terrestrial arthropods has been documented in temperate stream ecosystems, but little is known about the magnitude of these inputs in tropical streams. Terrestrial arthropods falling from the canopy of tropical forests may be an important subsidy to tropical stream food webs and could also represent an important flux of nitrogen (N) and phosphorus (P) in nutrient‐poor headwater streams. We quantified input rates of terrestrial insects in eight streams draining lowland tropical wet forest in Costa Rica. In two focal headwater streams, we also measured capture efficiency by the fish assemblage and quantified terrestrially derived N‐ and P‐excretion relative to stream nutrient uptake rates. Average input rates of terrestrial insects ranged from 5 to 41 mg dry mass/m²/d, exceeding previous measurements of aquatic invertebrate secondary production in these study streams, and were relatively consistent year‐round, in contrast to values reported in temperate streams. Terrestrial insects accounted for half of the diet of the dominant fish species, Priapicthys annectens. Although terrestrially derived fish excretion was found to be a small flux relative to measured nutrient uptake rates in the focal streams, the efficient capture and processing of terrestrial arthropods by fish made these nutrients available to the local stream ecosystem. This aquatic‐terrestrial linkage is likely being decoupled by deforestation in many tropical regions, with largely unknown but potentially important ecological consequences.     

Consumer-resource stoichiometry in detritus-based streams

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.     

Applications of ecological stoichiometry for sustainable acquisition of ecosystem services

Human activities have differentially altered biogeochemical cycling at local, regional and global scales. We propose that a stoichiometric approach, examining the fluxes of multiple elements and the ratio between them, may be a useful tool for better understanding human effects on ecosystem processes and services. The different scale of impacts of the elements carbon, nitrogen and phosphorus and the different nature of their biogeochemical cycles, imply a large variation of their stoichiometric ratios in space and time and thus divergent impacts on biota.
In this paper, we examine the effects of anthropogenic perturbations on nutrient ratios in ecosystems in two examples and one case study. Altered stoichiometry in agricultural systems (example 1) can affect not only crop yield and quality but also the interactions between plants and their pollinators, pests and pathogens. Human activities have also altered stoichiometry in coastal ecosystems (example 2). Increased N loading has especially lead to increased N:P and reduced Si:N ratios, with detrimental effects on ecosystem services derived from coastal pelagic food webs, such as fish yield and water quality. The terrestrial–aquatic linkage in stoichiometric alterations is illustrated with a case study, the Mississippi River watershed, where anthropogenic activities have caused stoichiometric changes that have propagated through the watershed into the northern Gulf of Mexico.
Coupled with altered stoichiometric nutrient inputs are the inherent differences in variation and sensitivity of different ecosystems to anthropogenic disturbance. Furthermore, the connections among the components of a watershed may result in downstream cascades of disrupted functioning. Applying a multiple element perspective to understanding and addressing societal needs is a new direction for both ecological stoichiometry and sustainability.     

Ecosystem development in short‐term postglacial chronosequences: N and P limitation in glacier forelands from Santa Inés Island,Magellan Strait

Glacier foreland moraines provide an ideal model to examine the patterns of ecosystem development and the evolution of nitrogen and phosphorous limitation over successional time. In this paper, we focus on a 400‐year soil chronosequence in the glacier forelands of Santa Inés Island in the Magellan Strait, southern Chile by examining forest development on phosphorus (P)‐poor substrates in a uniquely unpolluted region of the world. Results show a steady increase in tree basal area and a humped trend in tree species richness over four centuries of stand development. The increase in basal area suggests that the late successional tree species were more efficient nutrient users than earlier successional ones. Total contents of carbon (C) and nitrogen (N) in soils increased during the chronosequence, reaching an asymptote in late succession. The net increases in soil C : N, C : P and N : P ratios observed over successional time suggest that nutrient limitation is maximal in 400‐year‐old substrates. Foliar C : N and C : P ratios also increased over time to reach an asymptote in old‐growth stages, following soil stoichiometric relationships; however the foliar N‐to‐P ratio remained constant throughout the chronosequence. Biological N fixation was greater in early postglacial succession, associated with the presence of the symbiotic N‐fixer Gunnera magellanica. Declining trends of δ¹⁵N in surface soils through the 400‐year chronosequence are evidence of decreasing N losses in old‐growth forests. In synthesis, glacier foreland chronosequences at this high South American latitude provide evidence for increasing efficiency of N and P use in the ecosystem, with the replacement of shade‐intolerant pioneers by more efficient, shade‐tolerant tree species. This pattern of ecosystem development produces a constant foliar N : P ratio, regardless of variation in soil N‐to‐P ratio over four centuries.     

Response of a Stoichiometrically Imbalanced Ecosystem to Manipulation of Nutrient Supplies and Ratios

Cuatro Ciénegas Basin (CCB) is a desert ecosystem that hosts a large diversity of water bodies. Many surface waters in this basin have imbalanced nitrogen (N) to phosphorus (P) stoichiometry (total N:P > 100 by atoms), where P is likely to be a limiting nutrient. To investigate the effects of nutrient stoichiometry on planktonic and sediment ecosystem components and processes, we conducted a replicated in situ mesocosm experiment in Lagunita, a shallow pond located in the southwest region of the basin. Inorganic N and P were periodically added to mesocosms under three different N:P regimes (P only, N:P = 16 and N:P = 75) while the control mesocosms were left unamended. After three weeks of fertilization, more than two thirds of the applied P was immobilized into seston or sediment. The rapid uptake of P significantly decreased biomass C:P and N:P ratios, supporting the hypothesis that Lagunita is P-limited. Meanwhile, simultaneous N and P enrichment significantly enhanced planktonic growth, increasing total planktonic biomass by more than 2-fold compared to the unenriched control. With up to 76% of added N sequestered into the seston, it is suspected that the Lagunita microbial community also experienced strong N-limitation. However, when N and P were applied at N:P = 75, the microbes remained in a P-limitation state as in the untreated control. Two weeks after the last fertilizer application, seston C:P and N:P ratios returned to initial levels but chlorophyll a and seston C concentrations remained elevated. Additionally, no P release from the sediment was observed in the fertilized mesocosms. Overall, this study provides evidence that Lagunita is highly sensitive to nutrient perturbation because the biota is primarily P-limited and experiences a secondary N-limitation despite its high TN:TP ratio. This study serves as a strong basis to justify the need for protection of CCB ecosystems and other low-nutrient microbe-dominated systems from anthropogenic inputs of both N and P.     

Relationship between soil nutrient availability and plant species richness in a tropical semi‐arid environment

Question: What is the relationship between soil fertility and plant species richness in the ‘fertile islands’ occurring beneath two species of legume (Cercidium praecox and Prosopis laevigata)? Location: Tehuacán‐Cuicatlán region, central Mexico. Methods: Plant richness was measured in three micro‐environments (below canopies of C. praecox, below canopies of P. laevigata and in areas without canopies). The concentration of soil nutrients (C, N and P), C and N in the microbiota, and processes of ecosystem functioning (net C mineralization rate and N mineralization) were measured. The relationship between soil variables and plant richness were assessed with ANCOVAs. Results: Soil nutrients and species richness increases markedly under fertility islands. There were higher concentrations of C and N in the soil, faster rates of C mineralization, and higher species richness under P. laevigata canopies. The relationship between soil fertility and species richness was always positive except for total N, ammonium and net C mineralization rate under C. praecox, and for available P under P. laevigata. Conclusions: The sign of the relationship between soil fertility and species richness varies according to the nutrient and the micro‐environment. Positive relationships could result from between species complementarity and facilitation. Negative relationships could be explained by a specific limitation threshold for some soil resources (P and N for plants and C for the soil microbiota) which eliminate the possibilities of between species complementarity and facilitation above that threshold. As in all observational studies, these relationships should be considered only correlational.     

Nutrients stimulate leaf breakdown rates and detritivore biomass: bottom-up effects via heterotrophic pathways

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. Electronic supplementary material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

An experimental test of the hypothesis of non‐homeostatic consumer stoichiometry in a plant litter–microbe system

Stoichiometric homeostasis of heterotrophs is a common, but not always well‐examined premise in ecological stoichiometry. We experimentally evaluated the relationship between substrate (plant litter) and consumer (microorganisms) stoichiometry for a tropical terrestrial decomposer system. Variation in microbial C : P and N : P ratios tracked that of the soluble litter fraction, but not that of bulk leaf litter material. Microbial N and P were not isometrically related, suggesting higher rates of P than N sequestration in microbial biomass. Shifts in microbial stoichiometry were related to changes in microbial community structure. Our results indicate that P in dissolved form is a major driver of terrestrial microbial stoichiometry, similar to aquatic environments. The demonstrated relative plasticity in microbial C : P and N : P and the critical role of P have important implications for theoretical modelling and contribute to a process‐based understanding of stoichiometric relationships and the flow of elements across trophic levels in decomposer systems.     

Effects of phosphorus enrichment and grazing snails on modern stromatolitic microbial communities

1. The effects of phosphorus enrichment and grazing snails on a benthic microbial community that builds stromatolic oncolites were examined in an experiment at Rio Mesquites, Cuatro Ciénegas, Mexico. Chemical analyses of stream water samples indicated that overall atomic ratios of total nitrogen (N) to total phosphorus (P) were approximately 110, indicating a strong potential for P‐limitation of microbial growth. 2. Phosphorus enrichment involved addition of 5 μmol Na₂HPO₄ L^?1 to streamside microcosms receiving intermittent inputs of stream water while grazer manipulation involved removal of the dominant grazer, the snail Mexithauma quadripaludium. After 7 weeks, we examined responses in organic matter content, C : N : P ratios, metabolism (P removal, primary production, dark respiration, and calcification), and microbial community structure using molecular fingerprinting of 16S rRNA genes. 3. Manipulation of snails did not affect response variables measured in these treatments (organic matter, C : P ratio, P removal rate). However, P enrichment significantly decreased the C : P and N : P ratios of surficial materials in the oncolites (organic matter content was unchanged), increased net and gross photosynthesis (oxygen consumption in the dark was unchanged), increased rates of calcification, and increased diatoms relative to cyanobacteria. Heterotrophic Eubacteria and Archaea were only modestly affected. Thus, our results indicate weak grazing effects but strong impacts of P in this benthic system. 4. We hypothesise that a state of severe P‐limitation is imposed on autotrophic production in this food web due, at least in part, to co‐precipitation of phosphate during calcite deposition. This produces severe P‐limitation of the benthic algae and cyanobacteria, resulting in high C : P ratio of microbial mats relative to the biomass of photoautotrophs (phytoplankton, terrestrial foliage) in other ecosystems. In turn, this high C : P ratio is likely to generate severe stoichiometric constraints on the herbivores, thus limiting their populations and resulting in weak overall grazing impacts.     

Development of a diverse epiphyte community in response to phosphorus fertilization

The role of terrestrial soil nutrient supply in determining the composition and productivity of epiphyte communities has been little investigated. In a montane Hawaiian rainforest, we documented dramatic increases in the abundance and species richness of canopy epiphytes in a forest that had been fertilized annually with phosphorus (P) for 15 years; there was no response in forest that had been fertilized with nitrogen (N) or other nutrients. The response of N-fixing lichens to P fertilization was particularly strong, although mosses and non-N-fixing lichens also increased in abundance and diversity. We show that enhancement of canopy P availability is the most likely factor driving the bloom in epiphytes. These results provide strong evidence that terrestrial soil fertility may structure epiphyte communities, and in particular that the abundance of N-fixing lichens – a functionally important epiphyte group – may be particularly sensitive to ecosystem P availability.