Bottom-up and top-down effects on insect herbivores do not vary among sites of different salinity

It has been suggested, but rarely tested, that the relative strength of top-down and bottom-up factors in communities varies along an environmental stress gradient. We compared the strength of bottom-up and top-down effects on the densities of insect herbivores along a range of sites of different salinities in west-central Florida. We used a 2 x 2 factorial design with plots divided into four treatments: (1) bottom-up manipulation, where fertilizer was applied to increase plant quality; (2) top-down manipulation, where sticky traps were used to reduce the effects of natural enemies (parasitoids); (3) bottom-up and top-down manipulation, where fertilizer was applied and sticky traps were used; and (4) control plots. These plots were established along a range of salinities among seven different sites containing the salt marsh plant Borrichia frutescens. In each plot, we determined the parasitism levels and abundances of the sap sucker Pissonotus quadripustulatus, the gall maker Asphondylia borrichiae, and the lepidopteran stem borer Argyresthia spp. Gall density, Pissonotus density, and stem borer density were significantly higher in lower salinity sites, suggesting a strong effect of environmental stress. There was a significant increase of galls and Pissonotus and a marginally significant increase of bored stems on fertilized plots but not on trapped plots. There was a significant interaction of site and fertilizer on gall parasitism. There were no interactions of either treatment with salinity on herbivore densities. The general lack of interaction between salinity level and other treatments on herbivore densities contrasts with our previous result where treatment effects did vary with salinity level on a large experimentally generated salinity gradient at one site. Thus, the results of the present paper suggest that, while environmental stress can modify top-down and bottom-up effects on herbivores at single sites, variation in site-to-site factors, possibly including clonal identity of plant, affects herbivore densities so much as to swamp out any observable interaction between environmental stress and top-down or bottom-up factors.     

Top-down control inhibits spatial self-organization of a patterned landscape

Regular, self-organized spatial patterns in primary producers have been described in a wide range of ecosystems and are predicted to affect community production and resilience. Although consumers are abundant in most systems, the effect of trophic interactions on pattern formation in primary producers remains unstudied. We studied the effects of top-down control by herbivores on a self-organized landscape of regularly spaced, diatom-covered hummocks alternating with water-filled hollows on an intertidal mudflat in The Netherlands. Spatial patterns developed during spring but were followed by a rapid collapse in summer, leading to a flat landscape with low diatom densities and little variation in sediment bed level. This dramatic decline co-occurred with a gradual increase of benthic herbivores. A manipulative field experiment, where benthic herbivores were removed from the sediment, revealed that both diatom growth and hummock formation were inhibited by the activity of benthic herbivores. Our study provides clear evidence of top-down control of spatial self-organized patterns by benthic herbivores within a biological-geomorphic landscape.     

Rainfall and soils modify plant community response to grazing in Serengeti National Park

Terrestrial plant community responses to herbivory depend on resource availability, but the separate influences of different resources are difficult to study because they often correlate across natural environmental gradients. We studied the effects of excluding ungulate herbivores on plant species richness and composition, as well as available soil nitrogen (N) and phosphorus (P), across eight grassland sites in Serengeti National Park (SNP), Tanzania. These sites varied independently in rainfall and available soil N and P. Excluding herbivores decreased plant species richness at all sites and by an average of 5.4 species across all plots. Although plant species richness was a unimodal function of rainfall in both grazed and ungrazed plots, fences caused a greater decrease in plant species richness at sites of intermediate rainfall compared to sites of high or low rainfall. In terms of the relative or proportional decreases in plant species richness, excluding herbivores caused the strongest relative decreases at lower rainfall and where exclusion of herbivores increased available soil P. Herbivore exclusion increased among-plot heterogeneity in species composition but decreased coexistence of congeneric grasses. Compositional similarity between grazed and ungrazed treatments decreased with increasing rainfall due to greater forb richness in exclosures and greater sedge richness outside exclosures and was not related to effects of excluding herbivores on soil nutrients. Our results show that plant resources, especially water and P, appear to modulate the effects of herbivores on tropical grassland plant diversity and composition. We show that herbivore effects on soil P may be an important and previously unappreciated mechanism by which herbivores influence plant diversity, at least in tropical grasslands.     

Asymmetry in community regulation: effects of predators and productivity

Eutrophication and predator additions and extinctions are occurring in ecosystems worldwide. Although theory predicts that both will strongly alter the distribution of biomass in whole communities, empirical evidence has not been consolidated to quantitatively determine whether these theoretical predictions are generally borne out in real ecosystems. Here we analyze data from two types of trophic cascade studies, predator removals in factorial combination with fertilization and observed productivity gradients, to assess the role of top-down and bottom-up forces in structuring multi-trophic communities and compare results from these analyses to those from an extensive database of trophic cascade studies. We find that herbivore biomass declines and plant biomass increases in the presence of predators, regardless of system productivity. In contrast, while plants are increased by fertilization, this effect does not significantly increase herbivores in either the presence or absence of predators. These patterns are consistent among marine, freshwater, and terrestrial ecosystems and are largely independent of study size and duration. Thus, top-down effects of predation are transferred through more trophic levels than are bottom-up effects of eutrophication, showing strong asymmetry in the direction of control of biomass distribution in communities.     

Warming modifies trophic cascades and eutrophication in experimental freshwater communities

Climate warming is occurring in concert with other anthropogenic changes to ecosystems. However, it is unknown whether and how warming alters the importance of top-down vs. bottom-up control over community productivity and variability. We performed a 16-month factorial experimental manipulation of warming, nutrient enrichment, and predator presence in replicated freshwater pond mesocosms to test their independent and interactive impacts. Warming strengthened trophic cascades from fish to primary producers, and it decreased the impact of eutrophication on the mean and temporal variation of phytoplankton biomass. These impacts varied seasonally, with higher temperatures leading to stronger trophic cascades in winter and weaker algae blooms under eutrophication in summer. Our results suggest that higher temperatures may shift the control of primary production in freshwater ponds toward stronger top-down and weaker bottom-up effects. The dampened temporal variability of algal biomass under eutrophication at higher temperatures suggests that warming may stabilize some ecosystem processes.     

Induced defenses in herbivores and plants differentially modulate a trophic cascade

Inducible defenses are dynamic traits that modulate the strength of both plant-herbivore and herbivore-carnivore interactions. Surprisingly few studies have considered the relative contributions of induced plant and herbivore defenses to the overall balance of bottom-up and top-down control. Here we compare trophic cascade strengths using replicated two-level and three-level plankton communities in which we systematically varied the presence or absence of induced defenses at the plant and/or herbivore levels. Our results show that a trophic cascade, i.e., significantly higher plant biomass in three-level than in two-level food chains, occurred whenever herbivores were undefended against carnivores. Trophic cascades did not occur when herbivores exhibited an induced defense. This pattern was obtained irrespective of the presence or absence of induced defenses at the plant level. We thus found that herbivore defenses, not plant defenses, had an overriding effect on cascade strength. We discuss these results in relation to variation in cascade strengths in natural communities.     

Turning up the heat: temperature influences the relative importance of top-down and bottom-up effects

Understanding how communities respond to changes in temperature is a major challenge for community ecology. Temperature influences the relative degree to which top-down and bottom-up forces structure ecological communities. In greenhouse experiments using the aquatic community found in pitcher plants (Sarracenia purpurea), I tested how temperature affected the relative importance of top-down (mosquito predation) and bottom-up (ant carcasses) forces on protozoa and bacteria populations. While bottom-up effects did not vary consistently with temperature, the top-down effects of predators on protozoa increased at higher temperatures. These results suggest that temperature could change the relative importance of top-down and bottom-up effects in ecological communities. Specifically, higher temperature may increase the strength of top-down effects by raising predator metabolic rate and concomitant processes (e.g., activity, foraging, digestion, growth) relative to cooler temperatures. These findings apply broadly to an understanding of trophic interactions in a variable environment and are especially relevant in the context of ongoing climate change.     

Pelagic-benthic coupling of the microbial food web modifies nutrient cycles along a cascade-dammed river

• Structure of multi-trophic microbial groups were analyzed using DNA metabarcoding. • Discontinuity and trophic interactions were observed along the dam-fragmented river. • C, N and P cycles are driven by top-down and bottom-up forces of microbial food web. • Pelagic-benthic coupling may intensify nutrient accumulation in the river system. Cascade dams disrupt the river continuum, altering hydrology, biodiversity and nutrient flux. Describing the diversity of multi-trophic microbiota and assessing microbial contributions to the ecosystem processes are prerequisites for the restoration of these aquatic systems. This study investigated the microbial food web structure along a cascade-dammed river, paying special attention to the multi-trophic relationships and the potential role of pelagic-benthic coupling in nutrient cycles. Our results revealed the discontinuity in bacterial and eukaryotic community composition, functional group proportion, as well as α-diversity due to fragmentation by damming. The high microbial dissimilarity along the river, with the total multi-trophic β-diversity was 0.84, was almost completely caused by species replacement. Synchronization among trophic levels suggests potential interactions of the pelagic and the benthic groups, of which the β-diversities were primarily influenced by geographic and environmental factors, respectively. Dam-induced environmental variations, especially hydrological and nutrient variables, potentially influence the microbial food web via both top-down and bottom-up forces. We proposed that the cycles of carbon, nitrogen and phosphorus are influenced by multi-trophic groups through autotrophic and heterotrophic processes, predator–prey relationships, as well as the release of nutrients mainly by microfauna. Our results advance the notion that pelagic-benthic trophic coupling may intensify the accumulation of organic carbon, ammonium and inorganic phosphorus, thereby changing the biogeochemical patterns along river systems. As a consequence, researchers should pay more attention to the multi-trophic studies when assessing the environmental impacts, and to provide the necessary guidance for the ecological conservation and restoration of the dam-regulated systems.     

Pulses of dead periodical cicadas increase herbivory of American bellflowers

Resource pulses can have both direct bottom-up and indirect top-down effects on their consumers, but comparatively few studies have investigated the top-down effects of naturally occurring resource pulses on plants. This study describes two years of field experiments conducted to determine the indirect effects of 17-year periodical cicadas (Magicicada spp.) on herbivory in American bellflowers (Campanulastrum americanum). In 2004, the area of damaged leaves on cicada-supplemented plants was 78% greater than the area of damaged leaves on control plants. In 2005, cicada-supplemented plants were more likely to experience herbivory by mammalian herbivores than control plants. When large herbivores were excluded, similar patterns of leaf herbivory were observed, but these differences were not statistically significant. These results suggest that the pulsed input of dead periodical cicada bodies increased rates of herbivory on bellflowers, and that this effect was largely mediated by the selective foraging of large mammalian herbivores. More broadly, this study suggests that pulses of limiting resources can have both positive direct effects on plants and negative indirect effects due to selective herbivory, and that the net effects of pulsed resources on plants may depend on the composition and behavior of the surrounding herbivore community.     

Bottom-up regulation of plant community structure in an aridland ecosystem

We conducted a long-term rodent exclosure experiment in native grass- and shrub-dominated vegetation to evaluate the importance of top-down and bottom-up controls on plant community structure in a low-productivity aridland ecosystem. Using multiple regressions and analysis of covariance, we assessed how bottom-up precipitation pulses cascade through vegetation to affect rodent populations, how rodent populations affect plant community structure, and how rodents alter rates of plant community change over time. Our findings showed that bottom-up pulses cascade through the system, increasing the abundances of plants and rodents, and that rodents exerted no control on plant community structure and rate of change in grass-dominated vegetation, and only limited control in shrub-dominated vegetation. These results were discussed in the context of top-down effects on plant communities across broad gradients of primary productivity. We conclude that bottom-up regulation maintains this ecosystem in a state of low primary productivity that constrains the abundance of consumers such that they exert limited influence on plant community structure and dynamics.     

Defoliation synchronizes aboveground growth of co-occurring C4 grass species

The aboveground net primary productivity (ANPP) of grass communities in grasslands and savannas is primarily determined by precipitation quantity. Recent research, motivated by predictions of changes in the distribution of rainfall events by global climate change models, indicates that ANPP may be affected by rainfall distribution as much as by annual totals. Grazing and community composition are also known to affect grassland ANPP. The manner in which interactions between rainfall distribution, grazing, and community composition affect the relationship between precipitation and ANPP represents a critical knowledge gap. The effects of community composition and grazing on aboveground growth responses to intraseasonal variation in water availability were investigated at seven grassland sites with a nonselective clipping experiment. The aboveground growth of the dominant C4 species at each site was measured at regular intervals for 2-3 growing seasons in the presence or absence of regular defoliation. In the absence of defoliation, there was a general lack of synchrony of intraseasonal growth among co-occurring species. Variation in growth rates was high and was only partially explained by variation in rainfall. Regular defoliation increased growth synchrony at all sites, but changes in growth responses to rainfall varied between sites. These results suggest that community composition will be important in determining ANPP-precipitation relationships under conditions of altered rainfall distribution. However this effect appears to be a result of species responding differently to soil water or other resources rather than to rainfall per se. Grazing may override the effects of community composition by reducing differences in growth patterns between species and has the potential to weaken precipitation controls on ANPP.     

Herbivory mediates grass-endophyte relationships

Endophytic fungi are plant symbionts living asymptomatically within plant tissues. Neotyphodium spp., which are asexual vertically transmitted systemic fungal endophytes of cool-season grasses, are predicted to be plant mutualists. These endophytes increase host plant resistance to environmental stresses and/or increase the production of alkaloid-based herbivore deterrents. The ubiquity of this defense mutualism is unclear, and a variety of alternative mechanisms may explain the observed variation in infection rates, levels of deterrence, and the maintenance of asexual endophytes in grass populations. We found that grass-endophyte interactions are variable and ordered along an herbivory gradient in an undisturbed subarctic alpine ecosystem. Native grass populations in grazed sites had significantly greater frequency of Neotyphodium infection compared to ungrazed sites. Tillers from grazed sites had significantly higher hyphal densities compared to ungrazed sites. The ability of grass-Neotyphodium constituents to deter vertebrate herbivory in natural systems is thought to be rare. In grazed meadows, we showed that endophyte infection resulted in the deterrence of grazing by native vertebrate herbivores. However, the same herbivores did not distinguish between infected and uninfected grass harvested from ungrazed areas. These results demonstrate that the relationship between vertically transmitted endophytes and grasses in the alpine tundra vary greatly within populations. This may be based in part on defense mutualism and is consistent, under varying levels of herbivory, with the predictions of optimal defense theory.     

Above- and belowground responses of Arctic tundra ecosystems to altered soil nutrients and mammalian herbivory

Theory and observation indicate that changes in the rate of primary production can alter the balance between the bottom-up influences of plants and resources and the top-down regulation of herbivores and predators on ecosystem structure and function. The exploitation ecosystem hypothesis (EEH) posited that as aboveground net primary productivity (ANPP) increases, the additional biomass should support higher trophic levels. We developed an extension of EEH to include the impacts of increases in ANPP on belowground consumers in a similar manner as aboveground, but indirectly through changes in the allocation of photosynthate to roots. We tested our predictions for plants aboveground and for phytophagous nematodes and their predators belowground in two common arctic tundra plant communities subjected to 11 years of increased soil nutrient availability and/or exclusion of mammalian herbivores. The less productive dry heath (DH) community met the predictions of EEH aboveground, with the greatest ANPP and plant biomass in the fertilized plots protected from herbivory. A palatable grass increased in fertilized plots while dwarf evergreen shrubs and lichens declined. Belowground, phytophagous nematodes also responded as predicted, achieving greater biomass in the higher ANPP plots, whereas predator biomass tended to be lower in those same plots (although not significantly). In the higher productivity moist acidic tussock (MAT) community, aboveground responses were quite different. Herbivores stimulated ANPP and biomass in both ambient and enriched soil nutrient plots; maximum ANPP occurred in fertilized plots exposed to herbivory. Fertilized plots became dominated by dwarf birch (a deciduous shrub) and cloudberry (a perennial forb); under ambient conditions these two species coexist with sedges, evergreen dwarf shrubs, and Sphagnum mosses. Phytophagous nematodes did not respond significantly to changes in ANPP, although predator biomass was greatest in control plots. The contrasting results of these two arctic tundra plant communities suggest that the predictions of EEH may hold for very low ANPP communities, but that other factors, including competition and shifts in vegetation composition toward less palatable species, may confound predicted responses to changes in productivity in higher ANPP communities such as the MAT studied here.     

Savanna tree density, herbivores, and the herbaceous community: bottom-up vs. top-down effects

Herbivores choose their habitats both to maximize forage intake and to minimize their risk of predation. For African savanna herbivores, the available habitats range in woody cover from open areas with few trees to dense, almost-closed woodlands. This variation in woody cover or density can have a number of consequences for herbaceous species composition, cover, and productivity, as well as for ease of predator detection and avoidance. Here, we consider two alternative possibilities: first, that tree density affects the herbaceous vegetation, with concomitant "bottom-up" effects on herbivore habitat preferences; or, second, that tree density affects predator visibility, mediating "top-down" effects of predators on herbivore habitat preferences. We sampled sites spanning a 10-fold range of tree densities in an Acacia drepanolobium-dominated savanna in Laikipia, Kenya, for variation in (1) herbaceous cover, composition, and species richness; (2) wild and domestic herbivore use; and (3) degree of visibility obstruction by the tree layer. We then used structural equation modeling to consider the potential influences that tree density may have on herbivores and herbaceous community properties. Tree density was associated with substantial variation in herbaceous species composition and richness. Cattle exhibited a fairly uniform use of the landscape, whereas wild herbivores, with the exception of elephants, exhibited a strong preference for areas of low tree density. Model results suggest that this was not a response to variation in herbaceous-community characteristics, but rather a response to the greater visibility associated with more open places. Elephants, in contrast, preferred areas with higher densities of trees, apparently because of greater forage availability. These results suggest that, for all but the largest species, top-down behavioral effects of predator avoidance on herbivores are mediated by tree density. This, in turn, appears to have cascading effects on the herbaceous vegetation. These results shed light on one of the major features of the "landscape of fear" in which African savanna herbivores exist.     

喀斯特山地草地植物群落结构与相似性特征

选择贵州典型喀斯特山地草地植物群落为研究对象,采用标准样地调查法,对不同海拔、坡向的植物群落结构、物种分布、种群习性等展开研究,结果表明,（1）喀斯特山地草地植物群落结构随海拔的升高,多年生草本植物种类比例逐渐降低,一年生草本植物和灌木植物没有明显的变化规律;喀斯特山地草地植物群落是以多年生草本为主,一年生草本和灌木为辅的植物群落结构类型。（2）喀斯特山地草地植物群落中,斑茅（Saccharum arundinaceum）、紫茎泽兰（Eupatorium adenophorum）、扭黄茅（Heteropogon contortus）、莠竹（Microstegium nudum）种群分别为不同海拔样点的优势种群,优势种群和次优种群以多年生草本植物主要。（3）喀斯特山地草地不同海拔间的植物群落相似性系数均在0~0.25之间,属于极不相似水平,海拔梯度间群落结构差异较大。喀斯特山地草地植物群落物种多样性较高,同时坡向对于植物群落相似性的影响较小,海拔是影响群落间差异的主要因素。     

Mercury bioaccumulation in arthropods from typical community habitats in a zinc-smelting area

This study assessed the enrichment of mercury in the food web from the different community habitats in a zinc-smelting area of China. We used a nitrogen stable isotope technique to analyze trophic level relationships among arthropods and found that the first trophic level consisted of plants in the different community habitats, the second trophic level consisted of herbivores such as locusts and grasshoppers (primary consumers), and the third trophic level included spiders and mantes (secondary consumers). Mercury enrichment in the primary consumers was not evident, but enrichment in arthropods of the third trophic level was significant. The average of enrichment coefficients in spiders and mantes was greater than 1. The δ¹⁵N values indicated that mercury concentrations accumulated from primary producers to top carnivorous arthropods increased. In this zinc-smelting area, the biological amplification of mercury in the food web is significant. It is reasonable to assume that humans, located at the top of the food chain, are exposed to biomagnified levels of mercury.     

Climate warming and predation risk during herbivore ontogeny

Phenological effects of climate change are expected to differ among species, altering interactions within ecological communities. However, the nature and strength of these effects can vary during ontogeny, so the net community-level effects will be the result of integration over an individual's lifetime. I resolved the mechanism driving the effects of warming and spider predation risk on a generalist grasshopper herbivore at each ontogenetic stage and quantified the treatment effects on a measure of reproductive fitness. Spiders caused nymphal grasshoppers to increase the proportion of herbs in their diet, thus having a positive indirect effect on grasses and a negative indirect effect on herbs. Warming strengthened the top-down effect by affecting spiders and grasshoppers differently. In cooler, ambient conditions, grasshoppers and spiders had a high degree of spatial overlap within the plant canopy. Grasshopper position was unaffected by temperature, but spiders moved lower in the canopy in response to warming. This decreased the spatial overlap between predator and prey, allowing nymphal grasshoppers to increase daily feeding time. While spiders decreased grasshopper growth and reproductive fitness in ambient conditions, spiders had no effect on grasshopper fitness in warmed treatments. The study demonstrates the importance of considering the ontogeny of behavior when examining the effects of climate change on trophic interactions.     

The top-down mechanism for body-mass-abundance scaling

Scaling relationships between mean body masses and abundances of species in multitrophic communities continue to be a subject of intense research and debate. The top-down mechanism explored in this paper explains the frequently observed inverse linear relationship between body mass and abundance (i.e., constant biomass) in terms of a balancing of resource biomasses by behaviorally and evolutionarily adapting foragers, and the evolutionary response of resources to this foraging pressure. The mechanism is tested using an allometric, multitrophic community model with a complex food web structure. It is a statistical model describing the evolutionary and population dynamics of tens to hundreds of species in a uniform way. Particularities of the model are the detailed representation of the evolution and interaction of trophic traits to reproduce topological food web patterns, prey switching behavior modeled after experimental observations, and the evolutionary adaptation of attack rates. Model structure and design are discussed. For model states comparable to natural communities, we find that (1) the body-mass abundance scaling does not depend on the allometric scaling exponent of physiological rates in the form expected from the energetic equivalence rule or other bottom-up theories; (2) the scaling exponent of abundance as a function of body mass is approximately -1, independent of the allometric exponent for physiological rates assumed; (3) removal of top-down control destroys this pattern, and energetic equivalence is recovered. We conclude that the top-down mechanism is active in the model, and that it is a viable alternative to bottom-up mechanisms for controlling body-mass-abundance relations in natural communities.     

Disentangling trophic relationships in a High Arctic tundra ecosystem through food web modeling

Determining the manner in which food webs will respond to environmental changes is difficult because the relative importance of top-down vs. bottom-up forces in controlling ecosystems is still debated. This is especially true in the Arctic tundra where, despite relatively simple food webs, it is still unclear which forces dominate in this ecosystem. Our primary goal was to assess the extent to which a tundra food web was dominated by plant-herbivore or predator-prey interactions. Based on a 17-year (1993-2009) study of terrestrial wildlife on Bylot Island, Nunavut, Canada, we developed trophic mass balance models to address this question. Snow Geese were the dominant herbivores in this ecosystem, followed by two sympatric lemming species (brown and collared lemmings). Arctic foxes, weasels, and several species of birds of prey were the dominant predators. Results of our trophic models encompassing 19 functional groups showed that <10% of the annual primary production was consumed by herbivores in most years despite the presence of a large Snow Goose colony, but that 20-100% of the annual herbivore production was consumed by predators. The impact of herbivores on vegetation has also weakened over time, probably due to an increase in primary production. The impact of predators was highest on lemmings, intermediate on passerines, and lowest on geese and shorebirds, but it varied with lemming abundance. Predation of collared lemmings exceeded production in most years and may explain why this species remained at low density. In contrast, the predation rate on brown lemmings varied with prey density and may have contributed to the high-amplitude, periodic fluctuations in the abundance of this species. Our analysis provided little evidence that herbivores are limited by primary production on Bylot Island. In contrast, we measured strong predator-prey interactions, which supports the hypothesis that this food web is primarily controlled by top-down forces. The presence of allochthonous resources subsidizing top predators and the absence of large herbivores may partly explain the predominant role of predation in this low-productivity ecosystem.     

Trophic levels and trophic tangles: the prevalence of omnivory in real food webs

The concept of trophic levels is one of the oldest in ecology and informs our understanding of energy flow and top-down control within food webs, but it has been criticized for ignoring omnivory. We tested whether trophic levels were apparent in 58 real food webs in four habitat types by examining patterns of trophic position. A large proportion of taxa (64.4%) occupied integer trophic positions, suggesting that discrete trophic levels do exist. Importantly however, the majority of those trophic positions were aggregated around integer values of 0 and 1, representing plants and herbivores. For the majority of the real food webs considered here, secondary consumers were no more likely to occupy an integer trophic position than in randomized food webs. This means that, above the herbivore trophic level, food webs are better characterized as a tangled web of omnivores. Omnivory was most common in marine systems, rarest in streams, and intermediate in lakes and terrestrial food webs. Trophic-level-based concepts such as trophic cascades may apply to systems with short food chains, but they become less valid as food chains lengthen.