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

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

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

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

Variable stoichiometric and macronutrient responses to lizard predation in Ozark glade grasshopper communities.

The General Stress Paradigm (GSP) predicts that prey body compositions should shift under chronic predation as prey increase body carbon and decrease body nitrogen content through dietary changes, heightened metabolism, reduced dietary efficiency, and the breakdown of nitrogen rich tissues to make labile carbohydrates available. In our study, we explored how the elemental and macronutrient content along with the morphology of three abundant Ozark glade grasshopper species differed between glades with and without predatory collared lizard (Crotaphytus collaris) populations. Our results indicated that lichen grasshoppers (Trimerotropis saxatilis) increased body C:N ratios in response to predators. Scudder's short-wing grasshoppers (Melanoplus scudderi) increased both body %C and %protein content, while the handsome grasshoppers (Syrbula admirabilis) did not significantly respond to the presence of collared lizards. None of the three grasshopper species showed morphological responses to predation. We also found that elemental and macronutrient content of grasshoppers was not always significantly correlated and was not associated with the same environmental factors, indicating a need to incorporate both perspectives in future research and utilize more accurate macromolecular assays. Overall, we found support for some aspects of the GSP in field-active animals and add to the growing body of evidence that predator-induced changes in prey body composition are more complex than predicted by the original GSP.

Brown meets green: light and nutrients alter detritivore assimilation of microbial nutrients from leaf litter.

In aquatic detrital-based food webs, research suggests that autotroph-heterotroph microbial interactions exert bottom-up controls on energy and nutrient transfer. To address this emerging topic, we investigated microbial responses to nutrient and light treatments during Liriodendron tulipifera litter decomposition and fed litter to the caddisfly larvae Pycnopsyche sp. We measured litter-associated algal, fungal, and bacterial biomass and production. Microbes were also labeled with 14 C and 33 P to trace distinct microbial carbon (C) and phosphorus (P) supporting Pycnopsyche assimilation and incorporation (growth). Litter-associated algal and fungal production rates additively increased with higher nutrient and light availability. Incorporation of microbial P did not differ across diets, except for higher incorporation efficiency of slower-turnover P on low-nutrient, shaded litter. On average, Pycnopsyche assimilated fungal C more efficiently than bacterial or algal C, and Pycnopsyche incorporated bacterial C more efficiently than algal or fungal C. Due to high litter fungal biomass, fungi supported 89.6-93.1% of Pycnopsyche C growth, compared to 0.2% to 3.6% supported by bacteria or algae. Overall, Pycnopsyche incorporated the most C in high nutrient and shaded litter. Our findings affirm others' regarding autotroph-heterotroph microbial interactions and extend into the trophic transfer of microbial energy and nutrients through detrital food webs.

Functional importance and diversity of fungi during standing grass litter decomposition.

Although microbial participation in litter decomposition is widely known within terrestrial soils, the role and significance of microorganisms during the aerial standing litter phase of decomposition remains poorly investigated. We examined the fungi inhabiting standing leaf litter of Schizachyrium scoparium and Schizachyrium tenerum in a Longleaf Pine savanna ecosystem and estimated their contribution to litter decomposition. We identified fungal phylotypes associated with leaf litter and quantified leaf C mass loss, fungal biomass production, and microbial respiration during decomposition. These data were used to construct budgets estimating C flow into and through fungi. Significant losses in S. scoparium (55%) and S. tenerum (67%) leaf C mass were observed during standing decomposition along with concomitant increases in fungal biomass, which reached a maximum of 36 and 33 mgC/g detrital C, respectively. Cumulative fungal production during decomposition totaled 99 ± 6 mgC/g initial detrital C in S. scoparium and 73 ± 5 mgC/g initial detrital C in S. tenerum, indicating that 18 and 11% of the litter C was converted into fungal biomass, respectively. Corresponding estimates of cumulative fungal respiration totaled 106 ± 7 and 174 ± 11 mgC/g initial detrital C in S. scoparium and S. tenerum, respectively. Next generation sequencing identified several fungal phylotypes, with the majority of sequences belonging to the Ascomycota (Dothideomycetes) and Basidiomycota (Agaricomycetes). Fungal phylotypes were similar between litter species and changed over time, showing a successional pattern. These findings extend our understanding of fungal processes to standing litter in terrestrial ecosystems, and highlight the quantitative importance of fungi in C cycling processes.

Filter-feeders have differential bottom-up impacts on green and brown food webs.

Nutrient recycling by consumers can strongly impact nutrient availability for autotrophic and heterotrophic microbes, thus impacting functions such as primary production and decomposition. Filter-feeding freshwater mussels form dense, multispecies assemblages in aquatic ecosystems and have been shown to play a critical role in nutrient cycling. Mussel excretion can enhance benthic primary production and influence algal species composition. However, the role of mussels in brown or detritus-based food webs and species-specific differences has received considerably less attention. Here, using mesocosm experiments, we assessed how three species of freshwater mussels that occupy three different phylogenetic tribes influenced benthic algal accrual, ecosystem metabolism, cotton strip decomposition, leaf litter (Acer saccharum) decomposition, and litter-associated fungal biomass measured as ergosterol. Additionally, we measured mussel excretion and biodeposition rates and assessed the stoichiometry (C:N, C:P, and N:P) of the benthic algae, cotton strips, and leaf litter. In comparison to controls without mussels, generally, mussel treatments had higher benthic algal biomass composed of more diatoms, higher gross primary productivity and net ecosystem production rates, and higher cotton strip tensile strength loss, but there was not a difference in ecosystem respiration rates, leaf litter decomposition rates, or fungal biomass. Benthic algae had lower C:N and higher N:P in mussel treatment tanks and cotton strip C:N was lower in mesocosms with mussels. Our results suggest that nutrient regeneration by mussels most strongly regulates green food webs, with some impacts to brown food webs, suggesting that consumers have interactive effects on microbial functioning in freshwaters.

Periphytic algae decouple fungal activity from leaf litter decomposition via negative priming.

1. Well-documented in terrestrial settings, priming effects describe stimulated heterotrophic microbial activity and decomposition of recalcitrant carbon by additions of labile carbon. In aquatic settings, algae produce labile exudates which may elicit priming during organic matter decomposition, yet the directions and mechanisms of aquatic priming effects remain poorly tested. 2. We tested algal-induced priming during decomposition of two leaf species of contrasting recalcitrance, Liriodendron tulipifera and Quercus nigra, in experimental streams under light or dark conditions. We measured litter-associated algal, bacterial, and fungal biomass and activity, stoichiometry, and litter decomposition rates over 43 days. 3. Light increased algal biomass and production rates and increased bacterial abundance 141-733% and fungal production rates 20-157%. Incubations with a photosynthesis inhibitor established that algal activity directly stimulated fungal production rates in the short-term. 4. Algal-stimulated fungal production rates on both leaf species were not coupled to long-term increases in fungal biomass accrual or litter decomposition rates, which were 154-157% and 164-455% greater in the dark, respectively. The similar patterns on fast- vs. slow-decomposing L. tulipifera and Q. nigra, respectively, indicated that substrate recalcitrance may not mediate priming strength or direction. 5. In this example of negative priming, periphytic algae decoupled fungal activity from decomposition, likely by providing labile carbon invested toward greater fungal growth and reproduction instead of recalcitrant carbon degradation. If common, algal-induced negative priming could stimulate heterotrophy reliant on labile carbon yet suppress decomposition of recalcitrant carbon, modifying energy and nutrients available to upper trophic levels and enhancing organic carbon storage or export in well-lit aquatic habitats.

Interspecific homeostatic regulation and growth across aquatic invertebrate detritivores: a test of ecological stoichiometry theory.

Across resource quality gradients, primary consumers must regulate homeostasis and release of nutrients to optimize growth and fitness. Based primarily on internal body composition, the ecological stoichiometry theory (EST) offers a framework to generalize interspecific patterns of these responses, yet the predictions and underlying assumptions of EST remain poorly tested across many species. We used controlled laboratory feeding experiments to measure homeostasis, nutrient release, and growth across seven field-collected aquatic invertebrate detritivore taxa fed wide resource carbon:nitrogen (C:N) and carbon:phosphorus (C:P) gradients. We found that most invertebrates exhibited strict stoichiometric homeostasis (average 1/H = - 0.018 and 0.026 for C:N and C:P, respectively), supporting assumptions of EST. However, the stoichiometry of new tissue production during growth intervals (growth stoichiometry) deviated - 30 to + 54% and - 145 to + 74% from initial body C:N and C:P, respectively, and across species, growth stoichiometry was not correlated with initial body stoichiometry. Notably, smaller non- and hemimetabolous invertebrates exhibited low, decreasing growth C:N and C:P, whereas larger holometabolous invertebrates exhibited high, often increasing growth C:N and C:P. Despite predictions of EST, interspecific sensitivity of egestion stoichiometry and growth rates to the resource gradient were weakly related to internal body composition across species. While the sensitivity of these patterns differed across taxa, such differences carried a weak phylogenetic signal and were not well predicted by EST. Our findings suggest that traits beyond internal body composition, such as feeding behavior, selective assimilation, and ontogeny, are needed to generalize interspecific patterns in consumer growth and nutrient release across resource quality gradients.

Quantity and quality limit detritivore growth: mechanisms revealed by ecological stoichiometry and co-limitation theory.

Resource quantity and quality are fundamental bottom-up constraints on consumers. Best understood in autotroph-based systems, co-occurrence of these constraints may be common but remains poorly studied in detrital-based systems. Here, we used a laboratory growth experiment to test limitation of the detritivorous caddisfly larvae Pycnopsyche lepida across a concurrent gradient of oak litter quantity (food supply) and quality (phosphorus : carbon [P:C ratios]). Growth increased simultaneously with quantity and quality, indicating co-limitation across the resource gradients. We merged approaches of ecological stoichiometry and co-limitation theory, showing how co-limitation reflected shifts in C and P acquisition throughout homeostatic regulation. Increased growth was best explained by elevated consumption rates and improved P assimilation, which both increased with elevated quantity and quality. Notably, C assimilation efficiencies remained unchanged and achieved maximum 18% at low quantity despite pronounced C limitation. Detrital C recalcitrance and substantive post-assimilatory C losses probably set a minimum quantity threshold to achieve positive C balance. Above this threshold, greater quality enhanced larval growth probably by improving P assimilation toward P-intensive growth. We suggest this interplay of C and P acquisition contributes to detritivore co-limitation, highlighting quantity and quality as potential simultaneous bottom-up controls in detrital-based ecosystems, including under anthropogenic change like nutrient enrichment.

Comparing the Ecological Stoichiometry in Green and Brown Food Webs - A Review and Meta-analysis of Freshwater Food Webs.

The framework of ecological stoichiometry was developed primarily within the context of "green" autotroph-based food webs. While stoichiometric principles also apply in "brown" detritus-based systems, these systems have been historically understudied and differ from green ones in several important aspects including carbon (C) quality and the nutrient [nitrogen (N) and phosphorus (P)] contents of food resources for consumers. In this paper, we review work over the last decade that has advanced the application of ecological stoichiometry from green to brown food webs, focusing on freshwater ecosystems. We first review three focal areas where green and brown food webs differ: (1) bottom-up controls by light and nutrient availability, (2) stoichiometric constraints on consumer growth and nutritional regulation, and (3) patterns in consumer-driven nutrient dynamics. Our review highlights the need for further study of how light and nutrient availability affect autotroph-heterotroph interactions on detritus and the subsequent effects on consumer feeding and growth. To complement this conceptual review, we formally quantified differences in stoichiometric principles between green and brown food webs using a meta-analysis across feeding studies of freshwater benthic invertebrates. From 257 datasets collated across 46 publications and several unpublished studies, we compared effect sizes (Pearson's r) of resource N:C and P:C on growth, consumption, excretion, and egestion between herbivorous and detritivorous consumers. The meta-analysis revealed that both herbivore and detritivore growth are limited by resource N:C and P:C contents, but effect sizes only among detritivores were significantly above zero. Consumption effect sizes were negative among herbivores but positive for detritivores in the case of both N:C and P:C, indicating distinct compensatory feeding responses across resource stoichiometry gradients. Herbivore P excretion rates responded significantly positively to resource P:C, whereas detritivore N and P excretion did not respond; detritivore N and P egestion responded positively to resource N:C and P:C, respectively. Our meta-analysis highlights resource N and P contents as broadly limiting in brown and green benthic food webs, but indicates contrasting mechanisms of limitation owing to differing consumer regulation. We suggest that green and brown food webs share fundamental stoichiometric principles, while identifying specific differences toward applying ecological stoichiometry across ecosystems.

Dietary and taxonomic controls on incorporation of microbial carbon and phosphorus by detritivorous caddisflies.

Heterotrophic microbes on detritus play critical roles in the nutrition of detritivorous animals, yet few studies have examined factors controlling the acquisition of microbial nutrients toward detritivore growth, which is termed "incorporation". Here, we assessed effects of detrital substrate identity (leaf type), background nutrients, and detritivore species identity on detritivore incorporation of microbial carbon (C) and phosphorus (P) in leaf litter diets. We fed oak and maple litter conditioned under two nutrient concentrations (50 or 500 µg P L(-1)) to the detritivorous caddisfly larvae Ironoquia spp., Lepidostoma spp., and Pycnopsyche lepida and used the radioisotopes 14C as glucose and 33P as phosphate to dually trace incorporation of microbial C and P by caddisflies. Incorporation efficiencies of microbial C (mean ± SE = 12.3 ± 1.3%) were one order of magnitude higher than gross growth efficiencies for bulk detrital C from recent studies (1.05 ± 0.08%). Litter type did not affect incorporation of microbial nutrients; however, caddisflies incorporated microbial P 11 % less efficiently when fed litter from the higher P concentration. Two lower body C:P species (Pycnopsyche and Ironoquia) exhibited 9.9 and 7.1% greater microbial C and 19.0 and 17.7% greater microbial P incorporation efficiencies, respectively, than the higher body C:P species (Lepidostoma). Our findings support ecological stoichiometry theory on post-ingestive regulation that animals fed lower C:P diets should reduce P incorporation efficiency due to excess diet P or alleviation of P-limited growth, and that lower C:P species must incorporate dietary C and P more efficiently to support fast growth of P-rich tissues.