Species-specific traits predict whole-assemblage detritus processing by pond invertebrates.

Functional trait diversity determines if ecosystem processes are sensitive to shifts in species abundances or composition. For example, trait variation suggests detritivores process detritus at different rates and make different contributions to whole-assemblage processing, which could be sensitive to compositional shifts. Here, we used a series of microcosm experiments to quantify species-specific coarse and fine particulate organic matter (CPOM and FPOM) processing for ten larval caddisfly species and three non-caddisfly species in high-elevation wetlands. We then compared trait-based models including life history, dietary, and extrinsic traits to determine which traits explained interspecific variation in detritus processing. Finally, we compared processing by mixed caddisfly assemblages in microcosms and natural ponds to additive predictions based on species-specific processing to determine if single-species effects are additive in multi-species assemblages. We found considerable interspecific variation in biomass-specific CPOM (13-fold differences) and FPOM (8-fold differences) processing. Furthermore, on a mass-specific basis, amphipods, chironomids, and caddisflies processed similar amounts of detritus, suggesting non-shredder taxa could process more than previously recognized. Trait models including dietary percent detritus, development rate, body size, and wetland hydroperiod explained 81 and 57% of interspecific variation in CPOM and FPOM processing, respectively. Finally, species-specific additive predictions were strikingly similar to mixed-assemblage processing in microcosms and natural ponds, with the largest difference being a 15% overestimate. Thus, additivity of species-specific processing suggests single-species rates may be useful for understanding functional consequences of shifting assemblages, and a trait-based approach to predicting species-specific processing could support generating additive predictions of whole-assemblage processing.

Understanding the evolution of nutritive taste in animals: Insights from biological stoichiometry and nutritional geometry.

A major conceptual gap in taste biology is the lack of a general framework for understanding the evolution of different taste modalities among animal species. We turn to two complementary nutritional frameworks, biological stoichiometry theory and nutritional geometry, to develop hypotheses for the evolution of different taste modalities in animals. We describe how the attractive tastes of Na-, Ca-, P-, N-, and C-containing compounds are consistent with principles of both frameworks based on their shared focus on nutritional imbalances and consumer homeostasis. Specifically, we suggest that the evolution of multiple nutritive taste modalities can be predicted by identifying individual elements that are typically more concentrated in the tissues of animals than plants. Additionally, we discuss how consumer homeostasis can inform our understanding of why some taste compounds (i.e., Na, Ca, and P salts) can be either attractive or aversive depending on concentration. We also discuss how these complementary frameworks can help to explain the evolutionary history of different taste modalities and improve our understanding of the mechanisms that lead to loss of taste capabilities in some animal lineages. The ideas presented here will stimulate research that bridges the fields of evolutionary biology, sensory biology, and ecology.

Differential responses of macroinvertebrate ionomes across experimental N:P gradients in detritus-based headwater streams.

Diverse global change processes are reshaping the biogeochemistry of stream ecosystems. Nutrient enrichment is a common stressor that can modify flows of biologically important elements such as carbon (C), nitrogen (N), and phosphorus (P) through stream foodwebs by altering the stoichiometric composition of stream organisms. However, enrichment effects on concentrations of other important essential and trace elements in stream taxa are less understood. We investigated shifts in macroinvertebrate ionomes in response to changes in coarse benthic organic matter (CBOM) stoichiometry following N and P enrichment of five detritus-based headwater streams. Concentrations of most elements (17/19) differed among three insect genera (Maccaffertium sp., Pycnopsyche spp., and Tallaperla spp.) prior to enrichment. Genus-specific changes in the body content of: P, magnesium, and sodium (Na) in Tallaperla; P, Na, and cadmium in Pycnopsyche; and P in Maccaffertium were also found across CBOM N:P gradients. These elements increased in Tallaperla but decreased in the other two taxa due to growth dilution at larger body sizes. Multivariate elemental differences were found across all taxa, and ionome-wide shifts with dietary N and P enrichment were also observed in Tallaperla and Pycnopsyche. Our results show that macroinvertebrates exhibit distinct differences in elemental composition beyond C, N, and P and that the ionomic composition of common stream taxa can vary with body size and N and P enrichment. Thus, bottom-up changes in N and P supplies could potentially influence the cycling of lesser studied biologically essential elements in aquatic environments by altering their relative proportions in animal tissues.

Experimental N and P additions relieve stoichiometric constraints on organic matter flows through five stream food webs.

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.

Litter P content drives consumer production in detritus-based streams spanning an experimental N:P gradient.

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.