The role of animals in modulating nutrient cycling [hereafter, consumer-driven nutrient dynamics (CND)] has been accepted as an important influence on both community structure and ecosystem function in aquatic systems. Yet there is great variability in the influence of CND across species and ecosystems, and the causes of this variation are not well understood. Here, we review and synthesize the mechanisms behind CND in fresh waters. We reviewed 131 articles on CND published between 1973 and 1 June 2015. The rate of new publications in CND has increased from 1.4 papers per year during 1973-2002 to 7.3 per year during 2003-2015. The majority of investigations are in North America with many concentrating on fish. More recent studies have focused on animal-mediated nutrient excretion rates relative to nutrient demand and indirect impacts (e.g. decomposition). We identified several mechanisms that influence CND across levels of biological organization. Factors affecting the stoichiometric plasticity of consumers, including body size, feeding history and ontogeny, play an important role in determining the impact of individual consumers on nutrient dynamics and underlie the stoichiometry of CND across time and space. The abiotic characteristics of an ecosystem affect the net impact of consumers on ecosystem processes by influencing consumer metabolic processes (e.g. consumption and excretion/egestion rates), non-CND supply of nutrients and ecosystem nutrient demand. Furthermore, the transformation and transport of elements by populations and communities of consumers also influences the flow of energy and nutrients across ecosystem boundaries. This review highlights that shifts in community composition or biomass of consumers and eco-evolutionary underpinnings can have strong effects on the functional role of consumers in ecosystem processes, yet these are relatively unexplored aspects of CND. Future research should evaluate the value of using species traits and abiotic conditions to predict and understand the effects of consumers on ecosystem-level nutrient dynamics across temporal and spatial scales. Moreover, new work in CND should strive to integrate knowledge from disparate fields of ecology and environmental science, such as physiology and ecosystem ecology, to develop a comprehensive and mechanistic understanding of the functional role of consumers. Comparative and experimental studies that develop testable hypotheses to challenge the current assumptions of CND, including consumer stoichiometric homeostasis, are needed to assess the significance of CND among species and across freshwater ecosystems.
OnlineOpen: This article is available free online at www.blackwell-synergy.com SUMMARY 1. Pacific salmon and steelhead once contributed large amounts of marine-derived carbon, nitrogen and phosphorus to freshwater ecosystems in the Pacific Northwest of the United States of America (California, Oregon, Washington and Idaho). Declines in historically abundant anadromous salmonid populations represent a significant loss of returning nutrients across a large spatial scale. Recently, a manufactured salmon carcass analogue was developed and tested as a safe and effective method of delivering nutrients to freshwater and linked riparian ecosystems where marine-derived nutrients have been reduced or eliminated. 2. We compared four streams: two reference and two treatment streams using salmon carcass analogue(s) (SCA) as a treatment. Response variables measured included: surface streamwater chemistry; nutrient limitation status; carbon and nitrogen stable isotopes; periphyton chlorophyll a and ash-free dry mass (AFDM); macroinvertebrate density and biomass; and leaf litter decomposition rates. Within each stream, upstream reference and downstream treatment reaches were sampled 1 year before, during, and 1 year after the addition of SCA. 3. Periphyton chlorophyll a and AFDM and macroinvertebrate biomass were significantly higher in stream reaches treated with SCA. Enriched stable isotope (d 15 N) signatures were observed in periphyton and macroinvertebrate samples collected from treatment reaches in both treatment streams, indicating trophic transfer from SCA to consumers. Densities of Ephemerellidae, Elmidae and Brachycentridae were significantly higher in treatment reaches. Macroinvertebrate community composition and structure, as measured by taxonomic richness and diversity, did not appear to respond significantly to SCA treatment. Leaf breakdown rates were variable among treatment streams: significantly higher in one stream treatment reach but not the other. Salmon carcass analogue treatments had no detectable effect on measured water chemistry variables. 4. Our results suggest that SCA addition successfully increased periphyton and macroinvertebrate biomass with no detectable response in streamwater nutrient concentrations. Correspondingly, no change in nutrient limitation status was detected based on dissolved inorganic nitrogen to soluble reactive phosphorus ratios (DIN/SRP) and nutrient-diffusing substrata experiments. Salmon carcass analogues appear to increase freshwater productivity. 5. Salmon carcass analogues represent a pathogen-free nutrient enhancement tool that mimics natural trophic transfer pathways, can be manufactured using recycled fish
Aim Neotropical highland streams have shown diminished ecosystem functioning after amphibian extirpation infected by the chytrid fungus Batrachochytrium dendrobatidis. The loss of amphibians could affect communities of aquatic insects co-occurring in these streams in various ways. We examined patterns of species and genetic diversity of these communities and their evolutionary history along the chytrid expansion gradient to elucidate potential community responses.Location Six streams over a 320-km transect in Panama affected by chytrid expansion from west to east for up to 14 years, and two apparently chytrid-free streams in the east.Methods Patterns of a-and b-diversity were investigated at three hierarchical levels: genus, species and haplotypes. Genus identification was based on morphology, and putative species were inferred by grouping the DNA barcodes (749 cox1 sequences) with the GMYC method on all collected individuals of Ephemeroptera, Trichoptera, Coleoptera and Plecoptera.Results A total of 96 genera in 43 families (9 orders) of insects were encountered. Genus-level a-diversity was higher in the easternmost streams, possibly due to a separate biogeographical history, whereas b-diversity was constant along the chytrid expansion gradient. Community DNA barcoding resulted in 426 cox1 haplotypes and 154 putative species, most of them limited to single sites. High b-diversity along the gradient at both species and haplotype levels argues against community homogenization by migration in the wake of amphibian declines. In contrast, phylo-b-diversity was low, indicating community similarity at deep levels.Main conclusions Aquatic insect communities in this region are influenced by long-term limited dispersion that generated high endemicity. The pattern persists mostly unperturbed after disease-driven amphibian declines; hence, if indeed insects fill the niches vacated by tadpoles, they would originate from local communities rather than immigration. Given the unique evolutionary history and physical isolation of local assemblages, the ecosystem deterioration carries the risk of losing unique diversity.
Summary 1. Relationships between biodiversity and ecosystem function are of increasing interest, particularly in freshwater ecosystems where species losses are occurring at unprecedented rates. Amphibian declines have been associated with a loss of ecosystem function in neotropical streams, but little is known of the potential roles of stream‐dwelling tadpoles in leaf decomposition. Leaf litter is an important energy source to streams, and the breakdown of this material to fine particulate organic matter (FPOM) is a key ecosystem function. 2. We used mesocosms in a natural stream setting to quantify the effects of grazing tadpoles, shredding macroinvertebrates and a combination of the two on leaf decomposition and associated microbial activity. We measured respiration rates of decomposing leaves, particulate organic matter (POM) and leaf biofilm biomass and C : N : P ratios, and leaf area loss in 4 treatments: Control, tadpole only (TP), tadpole and shredding macroinvertebrates (TP + INV) and shredding macroinvertebrates only (INV). We hypothesised that tadpoles would enhance leaf decomposition by changing nutrient availability and stimulating microbial activity. 3. Respiration rates ranged from 3.1 to 6.0 mg O2 dry mass−1 h−1 and were significantly higher in the TP and TP + INV treatments than in the control. The TP + INV treatment had significantly higher POM in chambers than the control and INV treatments. The TP treatment had significantly lower leaf biofilm biomass than the control and INV treatments. 4. Tadpoles influenced the elemental balance of C and N in POM and leaf biofilm. In contrast to our prediction, molar C : N ratios were higher in the TP + INV treatment than in the control. Mean molar N : P ratios in POM were higher in the TP + INV treatment than in any other treatment. Leaf biofilm followed a similar pattern, but both TP and TP + INV had significantly higher N : P ratios than the control and INV treatments. Leaf area loss was greatest when tadpoles and invertebrates were together (TP + INV = 0.6% leaf area loss per mg organism) than separate (TP = 0.1%, INV = 3%), indicating facilitation. 5. Tadpoles indirectly affected leaf decomposition by influencing microbial communities and macroinvertebrate feeding. As such, ongoing amphibian declines may adversely affect a critical ecosystem function in freshwater habitats.
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