Stoichiometry of nutrient excretion by fish: interspecific variation in a hypereutrophic lake

This study investigates how nutrient cycling rates and ratios vary among fish species, with a particular focus on comparing an ecologically dominant detritivore (gizzard shad) to other fishes in a productive lake. We also examined how nutrient cycling rates are mediated by body size (as predicted by allometry theory), and how variation in nutrient cycling is related to body and food nutrient contents (according to predictions of ecological stoichiometry). As predicted by allometry, per capita nitrogen and phosphorus excretion rates increased and mass-specific excretion rates decreased, with increasing mass. Body phosphorus content was correlated with body mass only in one species, bluegill. Contrary to stoichiometric predictions, there was no relationship between body P and mass-normalized P excretion rate, or between body N:P and excreted N:P, when all individuals of all species were considered.
However, at the species level, we observed some support for a body nutrient content effect on excretion as predicted by stoichiometry theory. For example, gizzard shad had lower body P (high body N:P) and also excreted P at higher rates (lower N:P) than bluegill, which had high body P (lower body N:P). We applied the Sterner (1990) homeostatic stoichiometry model to the two most common species in the study – gizzard shad and bluegill and found that food N:P had a greater effect than consumer body N:P on excreted N:P. This indicates that, in terms of variation among these species, nutrient excretion may be more of a function of food nutrient content than the nutrient content of the consumer. These results suggest that stoichiometry can provide a framework for variation among species in nutrient cycling and for evaluating the ecosystem consequences of biodiversity loss.     

Stoichiometry of consumer-driven nutrient recycling across nutrient regimes in streams

Stoichiometric constraints within ecological interactions and their ecosystem consequences may depend on characteristics of the abiotic environment such as background nutrient levels. We assessed whether consumer identity, via differing body stoichiometry, could regulate periphyton stoichiometry across nutrient regimes in open systems. In 60 flow-through artificial streams, we factorially crossed dissolved inorganic nitrogen levels (elevated = 294  μ g L⁻¹, ambient = 26  μ g L⁻¹) with dissolved inorganic phosphorus levels (DIP: elevated = 15  μ g L⁻¹, ambient = 3  μ g L⁻¹) and consumer type [crayfish (body N : P = 18), snails (body N : P = 28) or a control]. At ambient DIP, periphyton in the crayfish treatment had a lower %P and a lower C : P than periphyton in the snail treatment suggesting that consumer identity, probably mediated by differing P-excretion, regulated periphyton P content. At high DIP, consumer identity no longer affected periphyton elemental composition. Therefore, the stoichiometry of consumer-driven nutrient recycling and consumer identity may be less important to ecosystem functioning in environments with elevated nutrient levels.     

Stoichiometry of benthic invertebrate nutrient recycling: interspecific variation and the role of body mass

Ecological stoichiometry (ES) and allometry offer frameworks for predicting how nutrient recycling varies within and among animal species. Despite the importance of benthic-derived nutrients in most aquatic systems, predictions based on ES and allometry have been poorly tested among benthic invertebrate consumers. Here, we show that the rates and ratios at which three freshwater benthic invertebrate species (a crustacean, an insect, and a polychaeta) recycled nitrogen (N) and phosphorus (P) can be partially predicted by ES and allometry depending on whether data are analyzed intra- or interspecifically. Mass-specific N and P excretion rates were negatively correlated with invertebrate body size both among and within taxa, supporting allometric predictions. However, mass-specific N and P excretion rates were positively and negatively correlated to invertebrate body N and P, respectively, but only when data were analyzed intraspecifically. As a corollary, the mass-specific N:P excretion ratio was positively related to body N:P ratio. Such a contrasting pattern on excretion-mediated N and P recycling suggests that stoichiometric constraints regarding consumer-resource imbalances for the three species utilized in this study may be stronger for P than for N. Our results indicate that the variation in nutrient recycling, which is mediated by taxonomic constraints on stoichiometry and allometry, may substantially help us to understand the importance of benthic detritivorous species to the functioning of aquatic ecosystems.     

Stoichiometry and estimates of nutrient standing stocks of larval salamanders in Appalachian headwater streams

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Carbon to nitrogen excretion ratio in lepidopteran larvae: relative importance of ecological stoichiometry and metabolic scaling

The importance of consumers in regulating ecosystem processes has been increasingly recognized. Although insect herbivores have significant impacts on nutrient cycling through excretion in terrestrial systems, few studies have explored how insect species differ in this ecosystem process. Using 130 lepidopteran species, we tested two hypotheses based on ecological stoichiometry and metabolic scaling, respectively, both of which provide a mechanistic framework for consumer‐driven nutrient recycling. Our results highlighted that host plant C:N ratio is the most important determinant of interspecific variation in frass C:N ratio. Insect body mass also partially contributed to the variation in frass C:N ratio. These findings indicate that insect herbivores would play an important role in nutrient recycling with the characteristics of ecological stoichiometry in terrestrial systems.     

Estimating the effects of non‐native species on nutrient recycling using species‐specific and general allometric models

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Ontogeny, diet shifts, and nutrient stoichiometry in fish

Most stoichiometric models do not consider the importance of ontogenetic changes in body nutrient composition and excretion rates. We quantified ontogenetic variation in stoichiometry and diet in gizzard shad, Dorosoma cepedianum , an omnivorous fish with a pronounced ontogenetic diet shift; and zebrafish, Danio rerio, grown in the lab with a constant diet. In both species, body stoichiometry varied considerably along the life cycle. Larval gizzard shad and zebrafish had higher molar C:P and N:P ratios than larger fish. Variation in body nutrient ratios was driven mainly by body P, which increased with size. Gizzard shad body calcium content was highly correlated with P content, indicating that ontogenetic P variation is associated with bone formation. Similar trends in body stoichiometry of zebrafish, grown under constant diet in the laboratory, suggest that ontogeny (e.g. bone formation) and not diet shift is the main factor affecting fish body stoichiometry in larval and juvenile stages. The N:P ratio of nutrient excretion also varied ontogenetically in gizzard shad, but the decline from larvae to juveniles appears to be largely associated with variation in the N:P of alternative food resources (zooplankton vs detritus) rather than by fish body N:P. Furthermore, the N:P ratio of larval gizzard shad excretion appears to be driven more by the N:P ratio at which individuals allocate nutrients to growth, more so than static body N:P, further illustrating the need to consider ontogenetic variation. Our results thus show that fish exhibit considerable ontogenetic variation in body stoichiometry, driven by an inherent increase in the relative allocation of P to bones, whereas ontogenetic variation in excretion N:P ratio of gizzard shad is driven more by variation in food N:P than by body N:P.     

Allometric and trait‐based patterns in parasite stoichiometry

We measured the elemental content (%C, N and P) and ratios (C:N, C:P, N:P) of a diverse assemblage of parasitic helminths to ask whether taxonomy or traits were related to stoichiometric variation among species. We sampled 27 macroparasite taxa, spanning four phyla, infecting vertebrate and invertebrate hosts from freshwater ecosystems in New Jersey. Macroparasites varied widely in elemental content, exhibiting 4.7‐fold variation in %N, 4.6‐fold variation in %P, and 11.5‐fold variation in N:P. Across all species, parasite %P scaled negatively and C:P scaled positively with body size. Similar relationships between parasite P content and body size occurred at the phylum level and within individual species. The allometric scaling of P across species supports the growth rate hypothesis, which predicts that smaller taxa require more P to support relatively higher growth rates. Life cycle stage was related to %N and C:N, with non‐reproductive parasite stages lower in %N and higher in C:N than actively reproducing parasites. Parasite phylum, functional feeding group, and trophic level did not explain elemental variation among species. Organismal stoichiometry is linked to ecological function, and wide variation in macroparasite stoichiometry likely generates diverse patterns in host–parasite nutrient dynamics and variable relationships between parasitism and nutrient cycling.     

N : P stoichiometry and protein : RNA ratios in vascular plants: an evaluation of the growth-rate hypothesis

The growth-rate hypothesis states that fast-growing organisms need relatively more phosphorus-rich RNA to support rapid rates of protein synthesis, and therefore predicts, within and among taxa, increases in RNA and phosphorus content (relative to protein and nitrogen content) with increased growth rate. Here, we present a test of this hypothesis in vascular plants. We determined nitrogen : phosphorus ratios and protein : RNA ratios in pines growing at different rates due to nutrient conditions. In general, when comparing leaves of the same species at low and high growth rates, the faster-growing plants had higher RNA content, higher %N and %P, and lower protein : RNA ratios, but not consistently lower N : P ratios. We found no link between growth rate and foliar N : P or protein : RNA when comparing multiple species of different inherent growth rates. We conclude that plants adjust the balance of protein and RNA to favour either speed or efficiency of protein synthesis, but this balance does not alone dictate leaf stoichiometry.     

Tissue nutrient concentrations in freshwater aquatic macrophytes: high inter-taxon differences and low phenotypic response to nutrient supply

1. The elemental composition and stoichiometry of aquatic plants has often been suggested to reflect the nutrient enrichment of aquatic habitats. However, the relationship is often weak. Moreover, uncertainties remain in the relevance of laboratory derived critical plant tissue nutrient concentrations to maximum yield or growth rates in the field.
2. Aquatic vascular plants and bryophytes, overlying water and sediment samples were collected to test whether freshwater aquatic macrophytes: (i) show tissue nutrient deficiencies when growing in oligotrophic freshwater habitats, and (ii) have strict homeostatic stoichiometry.
3. Plant nutrient concentrations were significantly related to total inorganic nitrogen (or nitrate), total dissolved phosphorus and sediment total phosphorus. However, these relationships were weak. Virtually all the variance in plant tissue nutrient concentrations, however, could be explained by species (taxon) identity.
4. Critical tissue nutrient concentrations for 95% maximum yield or 95% maximum growth rate in aquatic angiosperms, determined from laboratory bioassays, suggested that nutrients should not limit yield in wild aquatic macrophytes. However, there were a substantial number of samples where potential growth rate limitation was possible, particularly due to phosphorus.
5. Strict C : N : P stoichiometric ratios were found for both vascular plants and bryophytes, suggesting little scope for plants as indicators of nutrient enrichment, but provide robust stoichiometric data for studies on ecosystem metabolism and nutrient cycling.     

Feedbacks of consumer nutrient recycling on producer biomass and stoichiometry: separating direct and indirect effects

Herbivores can have both direct (consumptive) and indirect (nutrient‐mediated) effects on primary producer biomass and nutrient stoichiometry. Ecological stoichiometry theory predicts that herbivores of contrasting body stoichiometry will differentially remineralize nutrients, resulting in feedbacks on producer stoichiometry. We experimentally separated direct and indirect effects of aquatic vertebrate grazers on periphyton by manipulating grazer abundance and identity in mesocosms, and using grazer exclusion cages to expose periphyton to recycled nutrients in the absence of direct grazing. In experiment 1, we used a catfish with high body phosphorus (low body N:P), Ancistrus triradiatus, to assess consumptive versus nutrient‐mediated effects of grazer density on periphyton. In experiment 2, we compared the nutrient‐mediated effects of grazing by Ancistrus triradiatus and Rana palmipes, a tadpole with low body phosphorus and high body N:P. In experiment 1, we found that increasing catfish density led to lower biomass and particulate nutrients in periphyton through direct consumptive effects, but that nutrient‐mediated indirect effects enhanced periphyton biomass when grazers were experimentally separated from direct contact with periphyton. As predicted by stoichiometry theory, nutrient recycling by this P‐rich grazer tended to increase algal C:P and N:P (although effects were not statistically significant), while their consumptive effects reduced algal C:P and N:P. In experiment 2, grazer identity had strong effects on dissolved water nutrient concentrations, N recycling (measured with a ¹⁵N tracer), and periphyton stoichiometry. In accordance with stoichiometry theory, catfish increased N concentrations and recycling rates leading to higher periphyton N:P, while tadpoles had greater effects on P availability leading to lower periphyton N:P. Our experiments elucidate the importance of both the density and identity of grazers in controlling periphyton biomass and stoichiometry through consumptive and nutrient‐mediated effects, and support the power of ecological stoichiometry theory to predict feedbacks on producer stroichiometry arising from consumer stoichiometry through nutrient recycling.     

Stoichiometric traits of stickleback: Effects of genetic background,rearing environment,and ontogeny

Phenotypes can both evolve in response to, and affect, ecosystem change, but few examples of diverging ecosystem‐effect traits have been investigated. Bony armor traits of fish are good candidates for this because they evolve rapidly in some freshwater fish populations, and bone is phosphorus rich and likely to affect nutrient recycling in aquatic ecosystems. Here, we explore how ontogeny, rearing environment, and bone allocation among body parts affect the stoichiometric phenotype (i.e., stoichiometric composition of bodies and excretion) of threespine stickleback. We use two populations from distinct freshwater lineages with contrasting lateral plating phenotypes (full vs. low plating) and their hybrids, which are mostly fully plated. We found that ontogeny, rearing environment, and body condition were the most important predictors of organismal stoichiometry. Although elemental composition was similar between both populations and their hybrids, we found significant divergence in phosphorus allocation among body parts and in phosphorus excretion rates. Overall, body armor differences did not explain variation in whole body phosphorus, phosphorus allocation, or phosphorus excretion. Evolutionary divergence between these lineages in both allocation and excretion is likely to have important direct consequences for ecosystems, but may be mediated by evolution of multiple morphological or physiological traits beyond plating phenotype.     

Soil acidity, ecological stoichiometry and allometric scaling in grassland food webs

The factors regulating the structure of food webs are a central focus of community and ecosystem ecology, as trophic interactions among species have important impacts on nutrient storage and cycling in many ecosystems. For soil invertebrates in grassland ecosystems in the Netherlands, the site-specific slopes of the faunal biomass to organism body mass relationships reflected basic biochemical and biogeochemical processes associated with soil acidity and soil C : N : P stoichiometry. That is, the higher the phosphorus availability in the soil, the higher, on average, the slope of the faunal biomass size spectrum (i.e., the higher the biomass of large-bodied invertebrates relative to the biomass of small invertebrates). While other factors may also be involved, these results are consistent with the growth rate hypothesis from biological stoichiometry that relates phosphorus demands to ribosomal RNA and protein production. Thus our data represent the first time that ecosystem phosphorus availability has been associated with allometry in soil food webs (supporting information available online). Our results have broad implications, as soil invertebrates of different size have different effects on soil processes.     

Biogeographic bases for a shift in crop C : N : P stoichiometries during domestication

We lack both a theoretical framework and solid empirical data to understand domestication impacts on plant chemistry. We hypothesised that domestication increased leaf N and P to support high plant production rates, but biogeographic and climate patterns further influenced the magnitude and direction of changes in specific aspects of chemistry and stoichiometry. To test these hypotheses, we used a data set of leaf C, N and P from 21 herbaceous crops and their wild progenitors. Domestication increased leaf N and/or P for 57% of the crops. Moreover, the latitude of the domestication sites (negatively related to temperature) modulated the domestication effects on P (+), C (?), N : P (?) and C : P (?) ratios. Further results from a litter decomposition assay showed that domestication effects on litter chemistry affected the availability of soil N and P. Our findings draw attention to evolutionary effects of domestication legacies on plant and soil stoichiometry and related ecosystem services (e.g. plant yield and soil fertility).     

Temporal Variation in the Importance of a Dominant Consumer to Stream Nutrient Cycling

Animal excretion can be a significant nutrient flux within ecosystems, where it supports primary production and facilitates microbial decomposition of organic matter. The effects of excretory products on nutrient cycling have been documented for various species and ecosystems, but temporal variation in these processes is poorly understood. We examined variation in excretion rates of a dominant grazing snail, Elimia clavaeformis, and its contribution to nutrient cycling, over the course of 14 months in a well-studied, low-nutrient stream (Walker Branch, east Tennessee, USA). Biomass-specific excretion rates of ammonium varied over twofold during the study, coinciding with seasonal changes in food availability (measured as gross primary production) and water temperature (multiple linear regression, R ² = 0.57, P = 0.053). The contribution of ammonium excretion to nutrient cycling varied with seasonal changes in both biological (that is, nutrient uptake rate) and physical (that is, stream flow) variables. On average, ammonium excretion accounted for 58% of stream water ammonium concentrations, 26% of whole-stream nitrogen demand, and 66% of autotrophic nitrogen uptake. Phosphorus excretion by Elimia was contrastingly low throughout the year, supplying only 1% of total dissolved phosphorus concentrations. The high average N:P ratio (89:1) of snail excretion likely exacerbated phosphorus limitation in Walker Branch. To fully characterize animal excretion rates and effects on ecosystem processes, multiple measurements through time are necessary, especially in ecosystems that experience strong seasonality.     

Exploring patterns and mechanisms of interspecific and intraspecific variation in body elemental composition of desert consumers

Key processes such as trophic interactions and nutrient cycling are often influenced by the element content of organisms. Previous analyses have led to some preliminary understanding of the relative importance of evolutionary and ecological factors determining animal stoichiometry. However, to date, the patterns and underlying mechanisms of consumer stoichiometry at interspecific and intraspecific levels within natural ecosystems remain poorly investigated. Here, we examine the association between phylogeny, trophic level, body size, and ontogeny and the elemental composition of 22 arthropod as well as two lizard species from the coastal zone of the Atacama Desert in Chile. We found that, in general, whole‐body P content was more variable than body N content both among and within species. Body P content showed a significant phylogenetic signal; however, phylogeny explained only 4% of the variation in body P content across arthropod species. We also found a significant association between trophic level and the element content of arthropods, with carnivores having 15% greater N and 70% greater P contents than herbivores. Elemental scaling relationships across species were only significant for body P content, and even the P content scaling relationship was not significant after controlling for phylogeny. P content did decrease significantly with body size within most arthropod species, which may reflect the size dependence of RNA content in invertebrates. In contrast, larger lizards had higher P contents and lower N:P ratios than smaller lizards, which may be explained by size‐associated differences in bone and scale investments. Our results suggests that structural differences in material allocation, trophic level and phylogeny can all contribute to variation in the stoichiometry of desert consumers, and they indicate that the elemental composition of animals can be useful information for identifying broad‐scale linkages between nutrient cycling and trophic interactions in terrestrial food webs.     

Nutrient recycling by fish in streams along a gradient of agricultural land use

Nutrient recycling is an essential ecosystem process provided by animals. In many aquatic systems, fish have been identified as important in ecosystem nutrient recycling; however, this importance can vary widely between systems. The factors controlling intersystem variation in animal‐mediated nutrient cycling have rarely been examined and as such it remains unclear what impact human landscape changes will have upon these processes. Here we examined rates of nutrient recycling for temperate stream fish assemblages along a gradient of agricultural land use (proportion cropland in the watershed: 1–59%). We quantified nutrient excretion rates of both ammonium–N (NH₄⁺–N) and phosphate (as soluble reactive phosphate: SRP) for fish assemblages at eight streams in southern Ontario, Canada with species‐specific excretion measurements and quantitative assemblage sampling. For both nutrients, total assemblage excretion exhibited a strong positive relationship with riparian cropland. The distance required for fish assemblages to turn over ambient nutrient pools was shorter for cropland systems, indicating that the relative importance of excreted nutrients was higher in these systems. Based on measured uptake rates of NH₄⁺–N in two streams (one higher cropland and one low cropland) and on modeled uptake rates for all streams, the proportion of ecosystem demand that can be satisfied by excretion is generally higher in the more agricultural streams. These patterns appear to be driven largely by disproportionate increases in fish assemblage biomass with increasing stream nutrient concentrations.     

Atmospheric nitrogen-deposition may intensify phosphorus limitation of shallow epilithic periphyton in unproductive lakes

1. We tested whether increasing atmospheric nitrogen (N) deposition along a north–south gradient intensifies epilithic phosphorus (P) limitation in oligotrophic Swedish lakes from the north to the south. We examined the epilithic community at a shallow depth from seven northern and six southern Swedish lakes, and also compared the results with a lake located geographically between the two groups. We determined lake nutrient state, epilithic nutrient ratios and epilithic algal composition, as well as grazer N : P ratios, grazer-epilithon N : P imbalance, and N : P cycling ratios.
2. Epilithic communities appear to be generally more N-limited in the northern lakes and more P-limited in the southern lakes. Lake water total N (Tot-N) and epilithic N : P ratios were lower in northern than in southern lakes and the proportion of N₂-fixing cyanobacteria was higher in northern than in southern lakes.
3. Gastropod grazers had lower N : P imbalances and cycled less N relative to P in northern than in southern lakes.
4. Atmospheric N-deposition showed a strong positive correlation with lake water Tot-N and a much weaker positive correlation with epilithon N : P ratios. Atmospheric N-deposition also correlated negatively with the proportion of N₂-fixing cyanobacteria.
5. There are indications that increased atmospheric N-deposition towards the south might intensify P-limitation of epilithic algae and invertebrate grazers, although more studies are needed to show the strength and generality of our findings.     

Inter‐specific Variation in Foliar Nutrients and Resorption of Nine Canopy‐tree Species in a Secondary Neotropical Rain Forest

The influence of environmental gradients on the foliar nutrient economy of forests has been well documented; however, we have little understanding of what drives variability among individuals within a single forest stand, especially tropical forests. We evaluated inter‐ and intra‐specific variation in nutrient resorption, foliar nutrient concentrations and physical leaf traits of nine canopy tree species within a 1‐ha secondary tropical rain forest in northeastern Costa Rica. Both nitrogen (N) and phosphorus (P) resorption efficiency (RE) and proficiency of the nine tree species varied significantly among species, but not within. Both N and P RE were significantly negatively related to leaf specific strength. Green leaf N and P concentrations were strongly negatively related to leaf mass per area, and senesced leaf nutrient concentrations were significantly positively related to green leaf nutrient concentrations. This study reveals a strong influence of physical leaf traits on foliar nutrient and resorption traits of co‐occurring species in a secondary wet tropical forest stand.     

Resource composition mediates the effects of intraspecific variability in nutrient recycling on ecosystem processes

Despite the growing evidence for individual variation in trophic niche within populations, its potential indirect effects on ecosystem processes remains poorly understood. In particular, few studies have investigated how intraspecific trophic variability can modulate the effects of consumers on ecosystems through potential changes in nutrient excretion rates. Here, we first quantified the level of intraspecific trophic variability in 11 wild populations of the omnivorous fish Lepomis gibbosus. Outputs from stomach content and stable isotope analyses revealed that the degree of trophic specialization and trophic positions were highly variable between and within these wild populations. There was intrapopulation variation in trophic position of more than one trophic level, suggesting that individuals consumed a range of plant and animal resources. We then experimentally manipulated intraspecific trophic variability to assess how it can modulate consumer‐mediated nutrient effects on relevant processes of ecosystem functioning. Specifically, three food sources varying in nutrient quality (e.g. plant material, macro‐invertebrate and fish meat) were used individually or in combination to simulate seven diet treatments. Results indicated that intraspecific variability in growth and nitrogen excretion rates were more related to the composition of the diet rather than the degree of specialization, and increased with the trophic position of the diet consumed. We subsequently used microcosms and showed that critical ecosystem functions, such as primary production and community respiration, were affected by the variability in excretory products, and this effect was biomass‐dependent. These results highlight the importance of considering variation within species to better assess the effects of individuals on ecosystems and, more specifically, the effects of consumer‐mediated nutrient recycling because the body size and the trophic ecology of individuals are affected by a large spectrum of natural and human‐induced environmental changes.