skip to main content

Title: Energy pathways modulate the resilience of stream invertebrate communities to drought

While climate change is altering ecosystems on a global scale, not all ecosystems are responding in the same way. The resilience of ecological communities may depend on whether food webs are producer‐ or detritus‐based (i.e. ‘green’ or ‘brown’ food webs, respectively), or both (i.e. ‘multi‐channel’ food web).

Food web theory suggests that the presence of multiple energy pathways can enhance community stability and resilience and may modulate the responses of ecological communities to disturbances such as climate change. Despite important advances in food web theory, few studies have empirically investigated the resilience of ecological communities to climate change stressors in ecosystems with different primary energy channels.

We conducted a factorial experiment using outdoor stream mesocosms to investigate the independent and interactive effects of warming and drought on invertebrate communities in food webs with different energy channel configurations. Warming had little effect on invertebrates, but stream drying negatively impacted total invertebrate abundance, biomass, richness and diversity.

Although resistance to drying did not differ among energy channel treatments, recovery and overall resilience were higher in green mesocosms than in mixed and brown mesocosms. Resilience to drying also varied widely among taxa, with larger predatory taxa exhibiting lower resilience.

Our results suggest that the effects of drought on stream communities may vary regionally and depend on whether food webs are fuelled by autochthonous or allochthonous basal resources. Communities inhabiting streams with large amounts of organic matter and more complex substrates that provide refugia may be more resilient to the loss of surface water than communities inhabiting streams with simpler, more homogeneous substrates.

more » « less
Award ID(s):
1802872 1754389
Author(s) / Creator(s):
 ;  ;  ;  
Publisher / Repository:
Date Published:
Journal Name:
Journal of Animal Ecology
Page Range / eLocation ID:
p. 2053-2064
Medium: X
Sponsoring Org:
National Science Foundation
More Like this
  1. Abstract

    Empirical evidence and theory suggest that climate warming and an increase in the frequency and duration of drying events will alter the metabolic balance of freshwater ecosystems. However, the impacts of climate change on ecosystem metabolism may depend on whether energy inputs are of autochthonous or allochthonous origin. To date, few studies have examined how warming and drying may interact to alter stream metabolism, much less how their impacts may depend on the energy‐base of the food web.

    To address this research gap, we conducted a multi‐factorial experiment using outdoor mesocosms to investigate the individual and synergistic effects of warming and drought on metabolic processes in stream mesocosms with green (algal‐based) vs. mixed (algal‐ and detritus‐based) vs. brown (detritus‐based) energy pathways.

    We set up 48 mesocosms with one of three different levels of shade and leaf litter input combinations to create mesocosms with different primary energy channels. In addition, we warmed half of the mesocosms by ~2–3°C. We assessed changes in ecosystem respiration (ER), gross primary production (GPP), net ecosystem production (NEP) and organic matter biomass in warmed and ambient temperature mesocosms before a 24 day drying event and after rewetting.

    Surprisingly, experimental warming had little effect on metabolic processes. Drying, however, led to decreased rates of ER and GPP and led to an overall reduction in NEP. Although the effects of drying were similar across energy channel treatments, reductions in ER and GPP were primarily driven by decreases in biomass of benthic and filamentous algae.

    Overall, we demonstrate that drying led to lower rates of NEP in mesocosms regardless of energy inputs. While warming showed little effect in our study, our results suggest that an increase in the frequency of stream drying events could greatly alter the metabolic balance of many aquatic ecosystems.

    Read the freePlain Language Summaryfor this article on the Journal blog.

    more » « less
  2. 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
  3. Abstract

    Drying intermittent stream networks often have permanent water refuges that are important for recolonisation. These habitats may be hotspots for interactions between fishes and invertebrates as they become isolated, but densities and diversity of fishes in these refuges can be highly variable across time and space.

    Insect emergence from streams provides energy and nutrient subsidies to riparian habitats. The magnitude of such subsidies may be influenced by in‐stream predators such as fishes.

    We examined whether benthic macroinvertebrate communities, emerging adult insects, and algal biomass in permanent grassland stream pools differed among sites with naturally varying densities of fishes. We also manipulated fish densities in a mesocosm experiment to address how fishes might affect colonisation during recovery from hydrologic disturbance.

    Fish biomass had a negative impact on invertebrate abundance, but not biomass or taxa richness, in natural pools. Total fish biomass was not correlated with total insect emergence in natural pools, but orangethroat darter (Etheostoma spectabile) biomass was inversely correlated with emerging Chironomidae biomass and individual midge body size. The interaction in our models between predatory fish biomass and date suggested that fishes may also delay insect emergence from natural pools, altering the timing of aquatic–terrestrial subsidies.

    There was an increase over time in algal biomass (chlorophyll‐a) in mesocosms, but this did not differ among fish density treatments. Regardless, fish presence in mesocosms reduced the abundance of colonising insects and total invertebrate biomass. Mesocosm invertebrate communities in treatments without fishes were characterised by more Chironomidae, Culicidae, and Corduliidae.

    Results suggest that fishes influence invertebrates in habitats that represent important refuges during hydrologic disturbance, hot spots for subsidy exports to riparian food webs, and source areas for colonists during recovery from hydrologic disturbance. Fish effects in these systems include decreasing invertebrate abundance, shifting community structure, and altering patterns of invertebrate emergence and colonisation.

    more » « less
  4. Abstract

    Animals exert both direct and indirect controls over elemental cycles, linking primary producer‐based (green) and decomposer‐based (brown) food webs through top‐down trophic interactions and bottom‐up element regeneration. Where animals are aggregated at high biomass, they create hotspots of elemental cycling. The relative importance of animal control on elemental cycling depends on animal biomass, species functional traits (i.e. feeding mode and stoichiometry), and their overlap.

    We evaluated how animal community complexity affects the mechanisms regulating energy flow to the brown food web. We conducted a mesocosm experiment where we varied the biomass and overlap of animals with different life history and stoichiometric traits (stream fish and mussels) and measured how this influenced the quantity and fraction of labile carbon available to microbes. We used linear models and structural equation modelling to evaluate direct (excretion) and indirect (herbivory, nutrient availability, and nutrient stoichiometry) effects of animals on bioavailable dissolved organic carbon (BDOC) concentration.

    In experimental stream mesocosms, we found support for both direct (DOC excretion) and indirect (grazing) animal influences on BDOC concentration. Although we found that snail, fish, and mussel biomass increased nutrient concentrations, neither nutrient concentration nor stoichiometry had a significant effect on BDOC concentration. This has been due to the high background nutrient concentration context of our stream mesocosm water. Snails, probably due to their high biomass and small body size, exerted a significant positive direct control on BDOC concentration. Fish and mussels exerted a significant negative indirect control on BDOC via their effects (grazing and bioturbation) on algal biomass.

    Our results imply that primary consumers with different feeding strategies provide a key mechanism regulating the flow of DOC into the brown food web through direct (excretion) and indirect (grazing) controls on primary producers. This highlights that animals can provide important controls on the production of bioavailable organic energy supporting microbes in aquatic ecosystems, but the importance of these controls depends on the nutrient context and the distribution of primary producer and animal biomasses.

    more » « less
  5. Abstract

    Biogeochemical cycling has often been characterized by physical and microbial processes, yet animals can be essential mediators of energy and nutrients in ecosystems. Excretion by aggregated animals can be an important local source of inorganic nutrients in green food webs; however, whether animals are a source of dissolved energy that can support brown food webs is understudied.

    We tested whether animal aggregations are a substantial flux of bioavailable dissolved organic matter (DOM) by studying spatially stable, biogeochemical hotspots formed by filter‐feeding freshwater mussels. We used parallel‐factor analysis to quantify DOM fluorescent components composition of mussel excretion and expected digestive breakdown of particulate food sources would lead to excretion of labile DOM. Next, we combined measured excretion rates of DOM, ammonium (, N) and phosphorous (SRP; P) for 22 species with biomass estimates for 14 aggregations to quantify contributions of DOM, N and P to local availability. Because mussels occupy distinct stoichiometric niches, we anticipated that differences in species biomass and assemblage structure would elicit different flux and stoichiometries of aggregate excretion.

    Aggregate dissolved organic carbon (DOC) excretion was minor (1%–11%) compared to N (12%–2,860%) and P (1%–97%), yet generalities across assemblages emerged regarding organic matter transformation by mussels towards labile protein‐like compounds compared to abundant aromatic, humic compounds in ambient water.

    Aggregate excretion of labile DOM was a substantial pool of bioavailable energy, contributing 2%–114% of local labile DOM. Spatial differences in assemblage structure led to strong differences in aggregate flux and stoichiometry driven by biomass and stoichiometric trait expression of species with contrasting dominance patterns.

    Under the nutrient conditions of our study (high C:nutrient), biogeochemical hotspots associated with low‐trophic position animal biomass may indirectly control energy flow to the brown food web by shifting C:nutrient stoichiometry available to microbes or directly by increasing the flux of microbially available DOM. Collectively, our results highlight a potentially substantial flux of labile energy and nutrients to microbial communities through the transformation of ingested organic matter by aggregations of animals and emphasize that shared functional trait classification may not translate into shared ecological function.

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

    more » « less