skip to main content

Title: Experimental N and P additions alter stream macroinvertebrate community composition via taxon‐level responses to shifts in detrital resource stoichiometry

Increases in nitrogen (N) and phosphorus (P) availability are changing animal communities, partly by altering stoichiometric imbalances between consumers and their food. Testing relationships between resource stoichiometry and consumer assemblage structure requires ecosystem‐level manipulations that have been lacking to date.

We analysed patterns of macroinvertebrate community composition in five detritus‐based headwater streams subject to experimental whole‐stream N and P additions that spanned a steep gradient in dissolved N:P ratio (2:1, 8:1, 16:1, 32:1, 128:1) over 2 years, following a 1‐year pre‐treatment period.

We predicted that shifts in leaf litter stoichiometry would drive overall patterns of community composition via greater responses of shredders to enrichment than other taxa, as shredders dominate primary consumer biomass and experience larger consumer–resource elemental imbalances than other functional groups in stream ecosystems. Specifically, we expected litter C:P to be a significant predictor of shredder biomass given the greater relative imbalances between shredder and litter C:P than C:N. Finally, we tested whether shredder responses to enrichment were related to other taxon‐level traits, including body size and stoichiometry, larval life span and growth rate.

Whole‐community composition shifted similarly across the five streams after enrichment, largely driven by increased shredder and predator biomass. These shifts were limited to the autumn/winter seasons and related to decreased leaf litter C:P, highlighting important links between the quality of seasonal litter subsidies and community phenology.

Among 10 taxa that drove structural shifts, two declined while other taxa from the same functional/taxonomic groups responded positively, suggesting that specific life‐history traits may determine sensitivity to enrichment.

Increases in total shredder biomass, and in biomass of several common shredders, were associated with lower litter C:P. Body C:P did not predict shredder response to enrichment. However, weak negative relationships between shredder response and body size, and larval life span, suggest that small‐bodied and short‐lived taxa may be more responsive to shifting resource stoichiometry.

Moderate anthropogenic increases in N and P availability affect resource stoichiometry and can alter animal communities, influencing additional food web and ecosystem properties. We provide support for ecological stoichiometry as a framework for predicting such outcomes based on changes in the elemental composition of resource pools.

Aplain language summaryis available for this article.

more » « less
Award ID(s):
Author(s) / Creator(s):
 ;  ;  ;  ;
Publisher / Repository:
Date Published:
Journal Name:
Functional Ecology
Page Range / eLocation ID:
p. 855-867
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    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.

    more » « less
  2. Abstract

    Herbivorous ectothermic vertebrates are more diverse and abundant at lower latitudes. While thermal constraints may drive this pattern, its underlying cause remains unclear. We hypothesized that this constraint stems from an inability to meet the elevated phosphorus demands of bony vertebrates feeding on P‐poor plant material at cooler temperatures because low gross growth efficiency at warmer temperatures facilitates higher P ingestion rates. We predicted that dietary carbon:phosphorus (C:P) should exceed the threshold elemental ratio between carbon and P‐limited growth (TERC:P) for herbivores feeding at cooler temperatures, thereby limiting the range of herbivorous ectothermic vertebrates facing P‐limited growth.

    We tested this hypothesis using the Andean suckermouth catfishesAstroblepusandChaetostoma.Astroblepusare invertivores that inhabit relatively cool, high‐elevation streams whileChaetostomaare grazers that inhabit relatively warm, low‐elevation streams. We calculated TERC:Pfor each genus across its elevational range and compared these values to measured values of food quality over an elevational gradient in the Andes. We also broadly summarized measurements of TERC:Pacross diverse groups of fishes.

    Supporting our hypothesis, we found that dietary C:P was predicted to exceed the TERC:Pfor the grazerChaetostomanear the highest elevation where this genus has been recorded. Conversely, the TERC:Pfor the invertivoreAstroblepuswas consistently higher than that ofChaetostomaand thus its dietary C:P never approached the TERC:P. We found that, among all fishes, omnivores had higher average TERC:Pthan invertivores, and TERC:Pdid not vary with temperature.

    Our results suggest that, at least for Andean suckermouth catfishes, cool temperatures constrain herbivory at higher elevations. Increased gross growth efficiency at cooler temperatures evidently restricts the ability of P‐limited consumers to meet P demand. However, our survey of fish TERC:Pestimates suggests that some fishes are able to circumvent this constraint through behavioural and life‐history adaptations that reduce P demand or increase P use efficiency.

    The physiological trade‐offs underlying these functional shifts reveal that geographic dietary patterns can be predicted by stoichiometric theory, but variation in food quality and consumer traits that reduce P demand and/or increase P efficiency can create exceptions to these patterns.

    Aplain language summaryis available for this article.

    more » « less
  3. Abstract

    Climate change is increasing the frequency, severity, and extent of wildfires and drought in many parts of the world, with numerous repercussions for the physical, chemical, and biological characteristics of streams. However, information on how these perturbations affect top predators and their impacts on lower trophic levels in streams is limited.

    The top aquatic predator in southern California streams is nativeOncorhynchus mykiss, the endangered southern California steelhead trout (trout). To examine relationships among the distribution of trout, environmental factors, and stream invertebrate resources and assemblages, we sampled pools in 25 stream reaches that differed in the presence (nine reaches) or absence (16 reaches) of trout over 12 years, including eight reaches where trout were extirpated during the study period by drought or post‐fire flood disturbances.

    Trout were present in deep pools with high water and habitat quality. Invertebrate communities in trout pools were dominated by a variety of medium‐sized collector–gatherer and shredder invertebrate taxa with non‐seasonal life cycles, whereas tadpoles and large, predatory invertebrates (Odonata, Coleoptera, Hemiptera [OCH]), often with atmospheric breather traits, were more abundant in troutless than trout pools.

    Structural equation modelling of the algal‐based food web indicated a trophic cascade from trout to predatory invertebrates to collector–gatherer taxa and weaker direct negative trout effects on grazers; however, both grazers and collector–gatherers also were positively related to macroalgal biomass. Structural equation modelling also suggested that bottom‐up interactions and abiotic factors drove the detritus‐based food web, with shredder abundance being positively related to leaf litter (coarse particulate organic matter) levels, which, in turn, were positively related to canopy cover and negatively related to flow. These results emphasise the context dependency of trout effects on prey communities and of the relative importance of top‐down versus bottom‐up interactions on food webs, contingent on environmental conditions (flow, light, nutrients, disturbances) and the abundances and traits of component taxa.

    Invertebrate assemblage structure changed from a trout to a troutless configuration within a year or two after trout were lost owing to post‐fire scouring flows or drought. Increases in OCH abundance after trout were lost were much more variable after drought than after fire. The reappearance of trout in one stream resulted in quick, severe reductions in OCH abundance.

    These results indicate that climate‐change induced disturbances can result in the extirpation of a top predator, with cascading repercussions for stream communities and food webs. This study also emphasises the importance of preserving or restoring refuge habitats, such as deep, shaded, perennial, cool stream pools with high habitat and water quality, to prevent the extirpation of sensitive species and preserve native biodiversity during a time of climate change.

    more » « less
  4. Abstract

    The effects of nutrient pollution on coral reef ecosystems are multifaceted. Numerous experiments have sought to identify the physiological effects of nutrient enrichment on reef‐building corals, but the results have been variable and sensitive to choices of nutrient quantity, chemical composition and exposure duration.

    To test the effects of chronic, ecologically relevant nutrient enrichment on coral growth and photophysiology, we conducted a 5‐week continuous dosing experiment on two Hawaiian coral species,Porites compressaandPocillopora acuta. We acclimated coral fragments to five nutrient concentrations (0.1–7 µMand 0.06–2.24 µM) with constant stoichiometry 2.5:1 nitrate to phosphate) bracketing in situ observations from reefs throughout the Pacific.

    Nutrient enrichment linearly increased photophysiological performance of both species within 3 weeks. The effect of nutrients onP. acutaphotochemical efficiency increased through time while a consistent response inP. compressaindicated acclimation to elevated nutrients within 5 weeks. Endosymbiont densities and total chlorophyll concentrations also increased proportionally with nutrient enrichment inP. acuta, but not inP. compressa, revealing contrasting patterns of host–symbiont acclimatization.

    The two species also exhibited contrasting effects of nutrient enrichment on skeletal growth. Calcification was enhanced at low nutrient enrichment (1 µM) inP. acuta, but comparable to the control at higher concentrations, whereas calcification was reduced inP. compressa(30%–35%) above 3 µM.

    Stable isotope analysis revealed species‐specific nitrogen uptake dynamics in the coral–algal symbiosis. The endosymbionts ofP. acutaexhibited increased nitrogen uptake (decreased δ15N) and incorporation (19%–31% decrease in C:N ratios) across treatments. In contrast,P. compressaendosymbionts maintained constant δ15N values and low levels of nitrogen incorporation (9%–11% decrease in C:N ratios). The inability ofP. acutato regulate endosymbiont nutrient uptake may indicate an emerging destabilization in the coral–algal symbiosis under nutrient enrichment that could compromise resistance to additional environmental stressors.

    Our results highlight species‐specific differences in the coral–algal symbiosis, which influence responses to chronic nutrient enrichment. These findings showcase how symbioses can vary among closely related taxa and underscore the importance of considering how life‐history traits modify species response to environmental change.

    A freePlain Language Summarycan be found within the Supporting Information of this article.

    more » « less
  5. Abstract

    Understanding how nutrient limitation affects algal biomass and production is a long‐standing interest in aquatic ecology. Nutrients can influence these whole‐community characteristics through several mechanisms, including shifting community composition. Therefore, incorporating the joint responses of biomass, taxonomic composition, and production of algal communities, and relationships among them, is important for understanding effects of nutrient enrichment.

    In shallow subarctic Lake Mývatn, Iceland, benthic algae compose a majority of whole‐lake primary production, support high secondary production, and influence nutrient cycling. Given the importance of these ecosystem processes, the factors that limit benthic algae have a large effect on the function and dynamics of the Mývatn system.

    In a 33‐day nutrient enrichment experiment conducted in Lake Mývatn, we measured the joint responses of benthic algal biomass, primary production, and composition to nitrogen (N) and phosphorus (P) supplementation. We enriched N and P using nutrient‐diffusing agar overlain by sediment, with three levels of N and P that were crossed in a factorial design.

    We found little evidence of community‐wide nutrient limitation, as chlorophyll‐aconcentrations showed a negligible response to nutrients. Gross primary production (GPP) was unaffected by P and inhibited by N enrichment after 10 days, although the inhibitory effect of N diminished by day 33.

    In contrast to biomass and primary production, community composition was strongly affected by N and marginally affected by P, with some algal groups increasing and others decreasing with enrichment. The taxa with the most negative and positive responses to N enrichment were Fragilariaceae andScenedesmus, respectively.

    The abundances of particular algal groups, based on standardised cell counts, were related toGPPmeasured at the end of the experiment.Oocystiswas negatively associated withGPPbut was unaffected by N or P, while Fragilariaceae andScenedesmuswere positively associated withGPPbut had opposite responses to N. As a result, nutrient‐induced compositional shifts did not alterGPP.

    Overall, our results show that nutrient enrichment can have large effects on algal community composition while having little effect on total biomass and primary production. Our study suggests that nutrient‐driven compositional shifts may not alter the overall ecological function of algal communities if (1) taxa have contrasting responses to nutrient enrichment but have similar effects on ecological processes, and/or (2) taxa that have strong influences on ecological function are not strongly affected by nutrients.

    more » « less