We investigate where bottom‐up and top‐down control regulates ecological communities as a mechanism linking ecological gradients to the geography of consumer abundance and biomass. We use standardized surveys of 54 North American grasslands to test alternate hypotheses predicting 100‐fold shifts in the biomass of four common grassland arthropod taxa—Auchenorrhyncha, sucking herbivores, Acrididae, chewing herbivores, Tettigoniidae, omnivores, and Araneae, predators. Bottom‐up models predict that consumer biomass tracks plant quantity (e.g. productivity and standing biomass) and quality (nutrient content) and that ectotherm access to food increases with temperature. Each of the focal trophic groups responded differently to these drivers: the biomass of sucking herbivores and omnivores increased with plant biomass; that of chewing herbivores tracked plant quality; and predator biomass did not depend on plant quality, plant quantity or temperature. The Exploitation Ecosystem Hypothesis is a top‐down hypothesis that predicts a shift from resource limitation of herbivores when plant production is low, to predator limitation when plant production is high. In grasslands where spider biomass was low, herbivore biomass increased with plant biomass, whereas bottom‐up structuring was not evident when spiders were abundant. Furthermore, neither predator biomass nor trophic position (via stable isotope analysis) increased with plant biomass, suggesting predators themselves are top‐down limited. Stable isotope analysis revealed that trophic position of the chewing herbivore and omnivore increased significantly with plant biomass, suggesting these groups increased scavenging and meat consumption in grasslands with higher carbohydrate availability. Taken together, our snapshot sampling documents gradients of food web structure across 54 grasslands, consistent with multiple hypotheses of bottom‐up and top‐down regulation.
The impacts of altered biogeochemical cycles on ecological systems are likely to vary with trophic level. Predicting how these changes will affect ecological food webs is further complicated by human activities, which are simultaneously altering the availability of macronutrients like nitrogen (N) and phosphorus (P), and micronutrients such as sodium (Na). Here we contrast three hypotheses that predict how increasing nutrient availability will shape grassland food webs. We conducted a distributed factorial fertilization experiment (N and P crossed with NaCl) across four North American grasslands, quantifying the responses of aboveground plant biomass and volume, plant tissue and soil elemental concentrations, as well as the abundance of five arthropod functional groups. Fertilization with N and P increased plant biomass and foliar N and P concentrations in grasses but not forbs. Fertilization with Na had no effect on plant biomass but increased foliar Na concentrations. Consistent with the nutrient limitation hypothesis, we found strong evidence of nutrient limitation for insect herbivores across the four sites with sucking (phloem and xylem feeding) herbivores increasing in abundance with NP fertilization and chewing herbivores increasing in response to both Na and NP fertilization, and a trend for increased response of arthropods to lower plant nutrient availability. We found no evidence for an interaction of NaCl and NP on arthropod abundance as predicted by the serial colimitation hypothesis. Finally, consistent with the ecosystem size hypothesis, predator and parasitoid abundances increased with plant volume, but not fertilization. Our results suggest these functional group‐specific responses to changes in plant nutrients and structure are key to predicting the future of grassland food webs in an era with increasing use of N and P fertilizers, and increasing terrestrial inputs of Na from road salt, saline irrigation water, and aerosols due to rising sea levels.
more » « less- NSF-PAR ID:
- 10370208
- Publisher / Repository:
- Wiley Blackwell (John Wiley & Sons)
- Date Published:
- Journal Name:
- Ecology
- Volume:
- 102
- Issue:
- 9
- ISSN:
- 0012-9658
- Format(s):
- Medium: X
- Sponsoring Org:
- National Science Foundation
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One year of our study occurred during an extreme drought. Drought mimics some of the effects of mammalian grazers: decreasing above‐ground plant biomass while increasing plant foliar percentage nitrogen.
We sampled arthropods and nutrient availability on and nearby (“off”) 10 bison‐grazed grazing lawns in a tallgrass prairie in NE Kansas. Total grasshopper abundance was higher on grazing lawns and the magnitude of this difference increased in the wetter year of 2019 compared to 2018, when drought led to high grass foliar nitrogen concentrations on and off grazing lawns. Mixed‐feeding grasshopper abundances were consistently higher on grazing lawns while grass‐feeder and forb‐feeder abundances were higher on lawns only in 2019, the wetter year. In contrast, the abundance of other arthropods (e.g., Hemiptera, Hymenoptera, and Araneae) did not differ on and off lawns, but increased overall in 2019, relative to the drought of 2018.
Understanding these local scale patterns of abundances and community composition improves predictability of arthropod responses to ongoing habitat change.
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Evidence for global insect declines mounts, increasing our need to understand underlying mechanisms. We test the nutrient dilution (ND) hypothesis—the decreasing concentration of essential dietary minerals with increasing plant productivity—that particularly targets insect herbivores. Nutrient dilution can result from increased plant biomass due to climate or CO2enrichment. Additionally, when considering long-term trends driven by climate, one must account for large-scale oscillations including El Niño Southern Oscillation (ENSO), North Atlantic Oscillation (NAO), and Pacific Decadal Oscillation (PDO). We combine long-term datasets of grasshopper abundance, climate, plant biomass, and end-of-season foliar elemental content to examine potential drivers of abundance cycles and trends of this dominant herbivore. Annual grasshopper abundances in 16- and 22-y time series from a Kansas prairie revealed both 5-y cycles and declines of 2.1–2.7%/y. Climate cycle indices of spring ENSO, summer NAO, and winter or spring PDO accounted for 40–54% of the variation in grasshopper abundance, mediated by effects of weather and host plants. Consistent with ND, grass biomass doubled and foliar concentrations of N, P, K, and Na—nutrients which limit grasshopper abundance—declined over the same period. The decline in plant nutrients accounted for 25% of the variation in grasshopper abundance over two decades. Thus a warming, wetter, more CO2-enriched world will likely contribute to declines in insect herbivores by depleting nutrients from their already nutrient-poor diet. Unlike other potential drivers of insect declines—habitat loss, light and chemical pollution—ND may be widespread in remaining natural areas.
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Aim: Ongoing alterations to Earth’s biogeochemical cycles (e.g., via fertilization, burning of fossil fuels, and pollution) are expected to impact plants, plant consumers and all subsequent trophic levels. While fertilization experiments often reveal arthropod nutrient limitation by nitrogen and phosphorus via effects on plant nutrient density and biomass, these macronutrients are only two of many nutrients important to arthropod fitness. Micronutrients are key to osmoregulation and enzyme function and can interact synergistically with macronutrients to shape the geography of arthropod abundance. We examine arthropod response to macro- and micronutrient fertilization as a function of nutrient type, application amount, duration, frequency, and plant responses to fertilization with the goal of addressing how ongoing alterations to biogeochemical cycles will shape future grassland food webs.more » « less
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Abstract Aim Multiple hypotheses predict how gradients of nutrient availability, plant biomass, and temperature shape trophic pyramids. We aim to disentangle the simultaneous influence of those factors and their indirect effects on trophic structure and individual trophic levels.
Location United States.
Time period 2017.
Major taxa studied Invertebrates.
Methods To examine differences in trophic pyramid shape and abundance within trophic levels and across ecological gradients, we conducted 54 standardized surveys of invertebrate communities in North American grasslands. We tested for the direct and indirect effects of plant biomass, temperature, sodium (Na), other essential elements (e.g. N, P, and K), and toxic heavy metals, (e.g. Ar and Pb) in plant tissue on both individual trophic levels, and trophic pyramid shape, estimated as the community trophic mean (CTM).
Results Plant sodium increased CTM, indicating that high plant sodium concentrations are associated with top‐heavy invertebrate trophic pyramids. Sites with higher plant biomass had higher proportions of herbivores compared to higher trophic levels. Finally, increasing temperature resulted in more top‐heavy trophic pyramids. Overall, plant biomass, temperature, and plant chemistry directly and indirectly affected the abundances within different trophic levels, highlighting the complexity of factors regulating trophic structure.
Main conclusions Trophic structure of grassland invertebrate communities is strongly influenced by plant sodium, plant biomass, and to a lesser extent, temperature. Grasslands occupy 30% of the Earth’s terrestrial surface and are an imperiled ecosystem due to conversion to row crop agriculture. As biogeochemistry and temperature in the Anthropocene are increasingly modified, our results have considerable implications for the trophic structure of future grassland communities.
