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1. Aggregated filter‐feeders govern the flux and stoichiometry of locally available energy and nutrients in rivers

   https://doi.org/10.1111/1365-2435.13778  

   Hopper, Garrett W. ; Chen, Shuo ; Sánchez González, Irene ; Bucholz, Jamie R. ; Lu, YueHan ; Atkinson, Carla L. ; Gonzalez, ed., Angélica ( March 2021 , Functional Ecology)

   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.

    

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2. Do mobile consumers homogenize the distribution of resources in stream food webs? A test with overlapping fish and mussel aggregations

   https://doi.org/10.1111/fwb.13872  

   Vaughn, Caryn C. ; Parr, Thomas B. ; DuBose, Traci P. ; Gates, Kiza K. ; Hopper, Garrett W. ; Gido, Keith B. ( December 2021 , Freshwater Biology)

   In streams, unionoid mussels and fish form aggregations that exert bottom‐up and top‐down effects on food webs, but the magnitude and spatial extent of their effects are controlled by species traits. Sedentary mussels live burrowed in the sediment in patchily distributed dense aggregations (mussel beds) where they filter seston and provide a local, relatively constant nutrient subsidy. In contrast, fish move on and off mussel beds, and thus comprise a transient nutrient subsidy.

   We asked how overlap between fish and mussels influences nutrient recycling and resource distribution in streams. We conducted an 8‐week study in experimental streams where we created mussel beds (comprised of two species,Actinonaias ligamentinaandAmblema plicata), manipulated the occurrence of a grazing minnow (Campostoma anomalum), and tracked nutrient (nitrogen and phosphorus) and resource (algae, detritus, and chironomids) abundance up and downstream of the mussel beds.

   In general, neither consumer had strong effects on the concentration or spatial distribution of nutrients. Water turnover time in our experimental streams may have diluted fish and mussel nutrient excretion effects, making it difficult to detect spatial patterns during a given sampling period.

   Fish controlled the abundance and productivity of algae. In treatments without fish, large mats of filamentous algae formed early in the experiment. These algae senesced, decomposed, and were not replaced. When fish were present, algae consisted of attached biofilms with consistent biomass and spatial distribution over time.

   Although previous work has shown that mussels can have strong, seasonal bottom‐up effects on both primary and secondary production, our results suggested that adding grazing mobile fishes, led to a more consistent and homogenous supply of algal resources. Because mussels rarely occur in the absence of fish, considering their combined influence on ecosystem dynamics is likely to be important.

    

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3. Carnivore identity mediates the effects of light and nutrients on aquatic food‐chain efficiency

   https://doi.org/10.1111/fwb.12790  

   Rock, Amber M. ; Hall, Mia R. ; Vanni, Michael J. ; González, María J. ( June 2016 , Freshwater Biology)

   Food chain efficiency (FCE), the proportion of primary production converted to production of the top trophic level, can influence several ecosystem services as well as the biodiversity and productivity of each trophic level. AquaticFCEis affected by light and nutrient supply, largely via effects on primary producer stoichiometry that propagate to herbivores and then carnivores. Here, we test the hypothesis that the identity of the top carnivore mediatesFCEresponses to changes in light and nutrient supply.

   We conducted a large‐scale, 6‐week mesocosm experiment in which we manipulated light and nutrient (nitrogen and phosphorus) supply and the identity of the carnivore in a 2 × 2 × 2 factorial design. We quantified the response ofFCEand the biomass and productivity of each trophic level (phytoplankton, zooplankton, and carnivore). We used an invertebrate,Chaoborus americanus, and a vertebrate, bluegill sunfish (Lepomis macrochirus), as the two carnivores in this study because of the large difference in phosphorus requirements between these taxa.

   We predicted that bluegill would be more likely to experience P‐limitation due to higher P requirements, and hence thatFCEwould be lower in the bluegill treatments than in theChaoborustreatments. We also expected the interactive effect of light and nutrients to be stronger in the bluegill treatments. Within a carnivore treatment, we predicted highestFCEunder low light and high nutrient supply, as these conditions would produce high‐quality (low C:nutrient) algal resources. In contrast, if food quantity had a stronger effect on carnivore production than food quality, carnivore production would increase proportionally with primary production, thusFCEwould be similar across light and nutrient treatments.

   Carnivore identity mediated the effects of light and nutrients onFCE, and as predictedFCEwas higher in food chains withChaoborusthan with bluegill. Also as predicted,FCEinChaoborustreatments was higher under low light. However,FCEin bluegill treatments was higher at high light supply, opposite to our predictions. In addition, bluegill production increased proportionally with primary production, whileChaoborusproduction was not correlated with primary production, suggesting that bluegill responded more strongly to food quantity than to food quality. These carnivore taxa differ in traits other than body stoichiometry, for example, feeding selectivity, which may have contributed to the observed differences inFCEbetween carnivores.

   Comparison of our results with those from previous experiments showed thatFCEresponds similarly to light and nutrients in food chains withChaoborusand larval fish (gizzard shad: Clupeidae), but very differently in food chains with bluegill. These findings warrant further investigation into the mechanisms related to carnivore identity (e.g., developmental stage, feeding selectivity) underlying these responses, and highlight the importance of considering both top‐down and bottom‐up effects when evaluating food chain responses to changing light and nutrient conditions.

    

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

   https://doi.org/10.1111/1365-2435.13289  

   Demi, Lee M. ; Benstead, Jonathan P. ; Rosemond, Amy D. ; Maerz, John C. ; El‐Sabaawi, ed., Rana ( February 2019 , Functional Ecology)

   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.

    

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5. Influence of fishes on macroinvertebrate communities and insect emergence production in intermittent stream refuges

   https://doi.org/10.1111/fwb.13509  

   Bonjour, Sophia M. ; Whiles, Matt R. ; Gido, Keith B. ( April 2020 , Freshwater Biology)

   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.

    

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