Density-mediated Indirect Effects Research Articles

The primacy of density‐mediated indirect effects in a community of wolves, elk, and aspen

AbstractThe removal or addition of a predator in an ecosystem can trigger a trophic cascade, whereby the predator indirectly influences plants and/or abiotic processes via direct effects on its herbivore prey. A trophic cascade can operate through a density‐mediated indirect effect (DMIE), where the predator reduces herbivore density via predation, and/or through a trait‐mediated indirect effect (TMIE), where the predator induces an herbivore trait response that modifies the herbivore's effect on plants. Manipulative experiments suggest that TMIEs are an equivalent or more important driver of trophic cascades than are DMIEs. Whether this applies generally in nature is uncertain because few studies have directly compared the magnitudes of TMIEs and DMIEs on natural unmanipulated field patterns. A TMIE is often invoked to explain the textbook trophic cascade involving wolves (Canis lupus), elk (Cervus canadensis), and aspen (Populus tremuloides) in northern Yellowstone National Park. This hypothesis posits that wolves indirectly increase recruitment of young aspen into the overstory primarily through reduced elk browsing in response to spatial variation in wolf predation risk rather than through reduced elk population density. To test this hypothesis, we compared the effects of spatiotemporal variation in wolf predation risk and temporal variation in elk population density on unmanipulated patterns of browsing and recruitment of young aspen across 113 aspen stands over a 21‐year period (1999–2019) in northern Yellowstone National Park. Only 2 of 10 indices of wolf predation risk had statistically meaningful effects on browsing and recruitment of young aspen, and these effects were 8–28 times weaker than the effect of elk density. To the extent that temporal variation in elk density was attributable to wolf predation, our results suggest that the wolf–elk–aspen trophic cascade was primarily density‐mediated rather than trait‐mediated. This aligns with the alternative hypothesis that wolves and other actively hunting predators with broad habitat domains cause DMIEs to dominate whenever prey, such as elk, also have a broad habitat domain. For at least this type of predator–prey community, our study suggests that risk‐induced trait responses can be abstracted or ignored while still achieving an accurate understanding of trophic cascades.

A sit-and-wait predator, but not an active-pursuit predator, alters pollinator-mediated selection on floral traits.

Indirect species interactions are ubiquitous in nature, often outnumbering direct species interactions. Yet despite evidence that indirect interactions have strong ecological effects, relatively little is known about whether they can shape adaptive evolution by altering the strength and/or direction of natural selection. We tested whether indirect interactions affect the strength and direction of pollinator-mediated selection on floral traits of the bumble-bee pollinated wildflower Lobelia siphilitica. We estimated the indirect effects of two pollinator predators with contrasting hunting modes: dragonflies (Aeshnidae and Corduliidae) and ambush bugs (Phymata americana, Reduviidae). Because dragonflies are active pursuit predators, we hypothesized that they would strengthen pollinator-mediated selection by weakening plant-pollinator interactions (i.e., a density-mediated indirect effect). In contrast, because ambush bugs are sit-and-wait predators, we hypothesized that they would weaken or reverse the direction of pollinator-mediated selection by altering pollinator foraging behavior (i.e., a trait-mediated indirect effect). Specifically, if ambush bugs hunt from plants with traits that attract pollinators (i.e., prey), then pollinators will spend less time visiting those plants, weakening or reversing the direction of selection on attractive floral traits. We did not find evidence that high dragonfly abundance strengthened selection on floral traits via a density-mediated indirect effect: neither pollen limitation (a proxy for the strength of plant-pollinator interactions) nor directional selection on floral traits of L. siphilitica differed significantly between high- and low-dragonfly abundance treatments. In contrast, we did find evidence that ambush bug presence affected selection on floral traits via a trait-mediated indirect effect: ambush bugs hunted from L. siphilitica plants with larger daily floral displays, reversing the direction of pollinator-mediated selection on daily display size. These results suggest that indirect species interactions have the potential to shape adaptive evolution by altering natural selection.

Omnivore density affects community structure through multiple trophic cascades.

Omnivores can dampen trophic cascades by feeding at multiple trophic levels, yet few studies have evaluated how intraspecific variation of omnivores influences community structure. The speckled dace (Rhinichthys osculus) is a common and omnivorous minnow that consumes algae and invertebrates. We studied effects of size and size structure on top-down control by dace and how effects scaled with density. Dace were manipulated in a mesocosm experiment and changes in invertebrate and algal communities and ecosystem function were monitored. Omnivores affected experimental communities via two distinct trophic pathways (benthic and pelagic). In the benthic pathway, dace reduced macroinvertebrate biomass, thereby causing density-mediated indirect effects that led to increased benthic algal biomass. Dace also reduced pelagic predatory macroinvertebrate biomass (hemipterans), thereby increasing the abundance of emerging insects. The effect of dace and hemipterans on emerging insects was mediated by a non-linear response to dace with peak emergence at intermediate dace density. In contrast with recent studies, omnivore size and size structure had no clear effect, indicating that small and large dace in our experiment shared similar functional roles. Our results support that the degree to which omnivores dampen trophic cascades depends on their relative effect on multiple trophic levels, such that the more omnivorous a predator is, the more likely cascades will be dampened. Availability of abundant macroinvertebrates, and the absence of top predators, may have shifted dace diets from primary to secondary consumption, strengthening density-dependent trophic cascades. Both omnivore density and dietary shifts are important factors influencing omnivore-mediated communities.

Environmental stress gradients regulate the relative importance of predator density- and trait-mediated indirect effects in oyster reef communities.

Predators affect community structure by influencing prey density and traits, but the importance of these effects often is difficult to predict. We measured the strength of blue crab predator effects on mud crab prey consumption of juvenile oysters across a flow gradient that inflicts both physical and sensory stress to determine how the relative importance of top predator density‐mediated indirect effects (DMIEs) and trait‐mediated indirect effects (TMIEs) change within systems. Overall, TMIEs dominated in relatively benign flow conditions where blue crab predator cues increased oyster survivorship by reducing mud crab–oyster consumption. Blue crab DMIEs became more important in high sensory stress conditions, which impaired mud crab perception of blue crab chemical cues. At high physical stress, the environment benefitted oyster survival by physically constraining mud crabs. Thus, factors that structure communities may be predicted based on an understanding of how physical and sensory performances change across environmental stress gradients.

Novel parasite invasion leads to rapid demographic compensation and recovery in an experimental population of guppies

The global movement of pathogens is altering populations and communities through a variety of direct and indirect ecological pathways. The direct effect of a pathogen on a host is reduced survival, which can lead to decreased population densities. However, theory also suggests that increased mortality can lead to no change or even increases in the density of the host. This paradoxical result can occur in a regulated population when the pathogen's negative effect on survival is countered by increased reproduction at the lower density. Here, we analyze data from a long-term capture-mark-recapture experiment of Trinidadian guppies (Poecilia reticulata) that were recently infected with a nematode parasite (Camallanus cotti). By comparing the newly infected population with a control population that was not infected, we show that decreases in the density of the infected guppy population were transient. The guppy population compensated for the decreased survival by a density-dependent increase in recruitment of new individuals into the population, without any change in the underlying recruitment function. Increased recruitment was related to an increase in the somatic growth of uninfected fish. Twenty months into the new invasion, the population had fully recovered to preinvasion densities even though the prevalence of infection of fish in the population remained high (72%). These results show that density-mediated indirect effects of novel parasites can be positive, not negative, which makes it difficult to extrapolate to how pathogens will affect species interactions in communities. We discuss possible hypotheses for the rapid recovery.

Specific parasites indirectly influence niche occupation of non-hosts community members.

The coexistence of phylogenetically related species is an attractive topic because of the potentially intense interspecific competition. The most often investigated mechanisms mediating coexistence of these species are environmental filtering and niche partitioning. However, the role of other factors, such as species-specific parasites, is still poorly understood. Along the riparian understory of a tropical forest, we explored niche occupation and coexistence between Chrysso intervales and Helvibis longicauda, two related syntopic web-building spiders. We also investigated the effect of H. longicauda mortality induced by a specific fungus parasite, Gibellula pulchra, on the dynamic of C. intervales spatial distribution. Coexistence was mediated mainly by a fine-scale horizontal spatial segregation. H. longicauda built webs almost exclusively close to the river, while C. intervales occupied adjacent areas (10-20m away from margins). We also found differentiation in other niche dimensions that might allow coexistence, such as in plants occupied, height of web placement, width of leaves used for thread attachment and phenology. H. longicauda mortality caused by fungi was higher during winter than in summer. Consequently, the abundance of C. intervales increased at distances close to the river, indicating competitive release through a density-mediated indirect effect. This demonstrates how non-competitive specific-antagonists can indirectly affect other non-hosts competing community members and influence their spatial distribution in fine-scale ranges. We suggest that environmental filtering restricts H. longicauda to cooler regions; niche partitioning leads populations to occupy different vertical strata and competitive exclusion precludes C. intervales to reach river margins, generating an unusual horizontal zonation pattern.

Density-mediated indirect effects from active predators and narrow habitat domain prey.

The hunting-mode-habitat-domain-range framework suggests that the mechanism driving trophic cascades (i.e., trait-mediated indirect interactions [TMIIs] vs. density-mediated indirect interactions [DMIIs]) should depend upon the functional traits of predators and prey. For example, trophic cascades containing active, broad habitat domain range (BHDR) predators interacting with narrow habitat domain range (NHDR) prey are predicted to arise primarily via TMIIs, because these prey should reduce their conspicuous activity in the presence of these predators. Unfortunately, this hypothesis is difficult to test given the strong bias against studies assessing trophic cascades containing NHDR prey. Furthermore, this hypothesis ignores evidence that (1) active predators can have high consumption rates on prey, (2) continuously responding to active predators foraging across broad areas is energetically costly for prey, and (3) cues from active, BHDR predators may not influence prey density. We examined the TMIIs and total indirect interaction (TII) produced during interactions between an active, BHDR ladybeetle predator (Naemia seriata) and its NHDR prey (scale insects). We exposed scale insects to nonlethal and lethal ladybeetle predators in laboratory mesocosms for 15 weeks. We measured the growth of the scale insect's host plant (cordgrass) and the population density of scale insects. Contrary to theory, nonlethal ladybeetles did not induce TMIIs. However, lethal ladybeetles increased cordgrass total and root dry biomass by 36% and 44%, respectively, suggesting the presence of strong DMIIs. Additionally, both lethal and nonlethal ladybeetles reduced scale insect population density. Our findings suggest that DMIIs, rather than TMIIs, can result from interactions between active BHDR predators and NHDR prey.

生态系统基于性状调节的物种间接作用:特征、成因及后果

PDF HTML阅读 XML下载 导出引用 引用提醒 生态系统基于性状调节的物种间接作用:特征、成因及后果 DOI: 10.5846/stxb201610011986 作者: 作者单位: 作者简介: 通讯作者: 中图分类号: 基金项目: 国家自然科学基金项目（31230012）；国家科技基础性工作专项（2012FY111900）；教育部创新团队发展计划（IRT-16R11）；中国博士后面上基金（2016M590246） Characteristics, causes, and consequences of trait-mediated indirect interactions in ecosystems Author: Affiliation: Fund Project: 摘要 | 图/表 | 访问统计 | 参考文献 | 相似文献 | 引证文献 | 资源附件 | 文章评论 摘要:物种之间的间接作用关系是维持生物群落结构以及生态系统功能的关键因素。目前的理论模型和实验性研究均认为，物种间接作用的传递主要是由物种的密度变化所引起。但大量的实验证据表明，生物个体在形态、生理、行为和生活史性状的适应性变化，是物种间形成间接互作关系的另一个重要机制。生态学家把这种基于物种性状调控的种间关系称为性状调节的间接作用。深入了解性状调节的间接作用类型，发生机制和作用途径，有助于阐释自然界中多物种种间关系的复杂性和多样性形成机制。对性状调节的概念和性状的分类进行系统的总结，指出性状可塑性和物种特异性是物种间性状调节关系形成的两个重要机制。与其他类型的种间互作一样，性状调节的种间作用是驱动物种进化的重要力量。此外，分析了性状调节在自然群落中传递的几个重要途径，并强调这些间接作用在影响和调节生态系统功能和过程中的重要作用。最后，就性状调节种间关系的研究对象、研究尺度、以及研究方法等问题提出若干建议，为今后对这一问题的研究提供相关参考。 Abstract:Indirect species interactions play a critical role in shaping community structure and the functioning of ecosystems. Previous empirical and theoretical studies commonly assumed that indirect interactions were caused in general by changes in population density of interacting species, which is referred to as ‘density-mediated indirect interactions.’ However, there is growing evidence that changes in traits of the individual intermediate species can also be an important mechanism that regulates indirect interactions among species, and ecologists term these species interactions ‘trait-mediated indirect interactions’ (TMⅡs). TMⅡs are prevalent in both aquatic and terrestrial systems, and their effects are often as strong as, or stronger than, those of density-mediated indirect effects. Studies on the characteristics, causes, and consequences of TMⅡs are essential to understand the complexity and diversity of multispecies interactions in natural ecosystems fully. In this paper, we summarize the concept and classification of TMⅡs, and suggest that trait plasticity and species specificity are the two intrinsic mechanisms for the occurrence of TMⅡs. We also review the direct evidence for TMⅡs in various permutations of interaction webs that can have both trophic and non-trophic links, and then highlight the profound effects of TMⅡs on regulation of ecological functions and processes in a wide range of ecological systems. Finally, we propose future developments regarding the effects of TMⅡs to provide a reference framework for related research. 参考文献 相似文献 引证文献

Disease Ecology Meets Ecosystem Science

Growing evidence indicates that parasites—when considered—can play influential roles in ecosystem structure and function, highlighting the need to integrate disease ecology and ecosystem science. To strengthen links between these traditionally disparate fields, we identified mechanisms through which parasites can affect ecosystems and used empirical literature searches to explore how commonly such mechanisms have been documented, the ecosystem properties affected, and the types of ecosystems in which they occur. Our results indicate that ecosystem-disease research has remained consistently rare, comprising less than 2% of disease ecology publications. Existing studies from terrestrial, freshwater, and marine systems, however, demonstrate that parasites can strongly affect (1) biogeochemical cycles of water, carbon, nutrients, and trace elements, (2) fluxes of biomass and energy, and (3) temporal ecosystem dynamics including disturbance, succession, and stability. Mechanistically, most studies have demonstrated density-mediated indirect effects, rather than trait-mediated effects, or direct effects of parasites, although whether this is representative remains unclear. Looking forward, we highlight the importance of applying traits-based approaches to predict when parasites are most likely to exert ecosystem-level effects. Future research should include efforts to extend host–parasite studies across levels of ecological organization, large-scale manipulations to experimentally quantify ecosystem roles of parasites, and the integration of parasites and disease into models of ecosystem functioning.

Top predators and habitat complexity alter an intraguild predation module in pond communities.

Predator diversity and habitat complexity frequently influence species interactions at lower trophic levels, yet their joint investigation has been performed infrequently despite the demonstrated importance of each individual factor. We investigated how different top predators and varying habitat complexity influence the function of an intraguild predation module consisting of two larval salamanders, intraguild predator Ambystoma annulatum and intraguild prey A. maculatum. We manipulated predator food webs and habitat complexity in outdoor mesocosms. Top predators significantly influenced body condition and survival of A. annulatum, but habitat complexity had minimal effects on either response. A three-way interaction among the covariates top predator identity, habitat complexity and A. annulatum survival influenced body condition and survival of A. maculatum via a density-mediated indirect effect. Different top predator combinations had variable effects in different habitat complexity treatments on intraguild predator (A. annulatum) survival that subsequently influenced intraguild prey (A. maculatum) body condition and survival. Future work should consider how different top predators influence other food web components, which should vary due to predator attributes and the physical environments in which they co-occur.

Mechanisms underlying plant sexual dimorphism in multi‐trophic arthropod communities

A growing body of research documents the importance of plant genetic effects on arthropod community structure. However, the mechanisms underlying these effects are often unclear. Additionally, plant genetic effects have largely been quantified in common gardens, thus inflating the estimates of their importance by minimizing levels of natural variation. Using Valeriana edulis, a dioecious plant with genetically based sex determination, we conducted surveys and experiments on wild-grown individuals to document field patterns of arthropod association between the sexes and the mechanisms underlying these plant genetic effects. Three years of surveys revealed strong and consistent sex-biased arthropod association in wild-grown plants: female plants supported 4-fold, 1.5-fold, and 4-fold higher densities of aphids, aphid predators, and aphid-tending ants, respectively, compared to males. There was mixed evidence that the female bias for aphids was due to higher plant quality, while we found no difference between plant sexes in aphid preference or the top-down effects of predators and tending ants. Female bias for ants was due to both the greater attractiveness of female plants (direct effect mediated by floral nectar) and an independent, weaker effect of higher aphid abundance on females (density-mediated indirect effect). Conversely, the female bias for predators was driven solely by the greater attractiveness of female plants. We did not find interaction modification, i.e., ant-aphid and predator-aphid interactions were equivalent between plant sexes. Plant sex explained 0.24%, 2.28%, and 4.42% of the variance in aphids, predators, and ants, respectively, values comparable to but slightly weaker than those previously reported from common-garden studies. In contrast to the prediction of diminished plant genetic effects with increasing trophic level, we show how weak indirect effects on predators and parasitoids (via herbivores) can be complemented by strong direct effects via common plant traits (floral resources). In summary, we document direct and indirect effects of genetically based sex on a multi-trophic arthropod community that were expressed in wild-grown plants across multiple years.

Restricting Prey Dispersal Can Overestimate the Importance of Predation in Trophic Cascades

Predators can affect prey populations and, via trophic cascades, predators can indirectly impact resource populations (2 trophic levels below the predator) through consumption of prey (density-mediated indirect effects; DMIEs) and by inducing predator-avoidance behavior in prey (trait-mediated indirect effects; TMIEs). Prey often employ multiple predator-avoidance behaviors, such as dispersal or reduced foraging activity, but estimates of TMIEs are usually on individual behaviors. We assessed direct and indirect predator effects in a mesocosm experiment using a marine food chain consisting of a predator (toadfish – Opsanus tau), prey (mud crab - Panopeus herbstii) and resource (ribbed mussel – Geukensia demissa). We measured dispersal and foraging activity of prey separately by manipulating both the presence and absence of the predator, and whether prey could or could not disperse into a predator-free area. Consumption of prey was 9 times greater when prey could not disperse, probably because mesocosm boundaries increased predator capture success. Although predator presence did not significantly affect the number of crabs that emigrated, the presence of a predator decreased resource consumption by prey, which resulted in fewer resources consumed for each prey that emigrated in the presence of a predator, and reduced the overall TMIE. When prey were unable to disperse, TMIEs on mussel survival were 3 times higher than the DMIEs. When prey were allowed to disperse, the TMIEs on resource survival increased to 11-times the DMIEs. We found that restricting the ability of prey to disperse, or focusing on only one predator-avoidance behavior, may be underestimating TMIEs. Our results indicate that the relative contribution of behavior and consumption in food chain dynamics will depend on which predator-avoidance behaviors are allowed to occur and measured.

Density-Dependent Effects of Amphibian Prey on the Growth and Survival of an Endangered Giant Water Bug.

Amphibian predator-insect prey relationships are common in terrestrial habitats, but amphibian larvae are preyed upon by a variety of aquatic hemipterans in aquatic habitats. This paper suggests that the survival of the nymphs of the endangered aquatic hemipteran Kirkaldyia (=Lethocerus) deyrolli (Belostomatidae: Heteroptera) is directly and indirectly affected by the abundance of their amphibian larval prey (tadpoles). Young nymphs of K. deyrolli mainly feed on tadpoles, regardless of differences in prey availability. Nymphs provided with tadpoles grow faster than nymphs provided with invertebrate prey. Therefore, tadpole consumption seems to be required to allow the nymphs to complete their larval development. In addition, the survival of K. deyrolli nymphs was greater during the period of highest tadpole density (June) than during a period of low tadpole density (July). Higher tadpole density moderates predation pressure from the water scorpion Laccotrephes japonensis (Nepidae: Heteroptera) on K. deyrolli nymphs; i.e., it has a density-mediated indirect effect. These results suggest that an abundance of tadpoles in June provides food for K. deyrolli nymphs (a direct bottom-up effect) and moderates the predation pressure from L. japonensis (an indirect bottom-up effect). An abundance of amphibian prey is indispensable for the conservation of this endangered giant water bug species.

A tale of two pesticides: how common insecticides affect aquatic communities

Summary 1. Recent ecotoxicology studies show that pesticide exposure can alter community composition, structure and function. Generally, community responses to pesticides are driven by trait- and density-mediated indirect effects resulting from sublethal and lethal effects of pesticide exposure on vulnerable taxa. These effects depend upon the concentration of the pesticide and the frequency of exposure. 2. While more research is needed to understand community-level responses to pesticide exposure, testing the effects of multitudes of registered chemicals on ecologically relevant communities is overwhelming. Recent reviews suggest that contaminants with similar modes of action should produce comparable community-level responses because they have similar direct effects and, as a result, similar indirect effects; this hypothesis remains largely untested. 3. We subjected pond communities [containing zooplankton, phytoplankton, periphyton and leopard frog tadpoles (Rana pipiens)] to several applications (single applications of medium or high concentrations or weekly applications of a lower concentration) of two acetylcholine esterase inhibiting insecticides, malathion and carbaryl that have comparable toxicity for aquatic organisms. 4. We found that both insecticides cause comparable trophic cascades that affect zooplankton and phytoplankton abundances; however, their effects on amphibians diverged, especially when exposed to higher concentrations of insecticides. Malathion caused a trophic cascade beginning with a decline in cladocerans followed by increases in phytoplankton. At a medium concentration, this cascade also caused a subsequent decrease in periphyton. Carbaryl caused a similar trophic cascade with the highest application, a weak trophic cascade with the medium application and no cascade with smallest application. Malathion directly reduced tadpole survival at all concentrations. Survivors in the two higher treatments were larger at metamorphosis while survivors in the lowest treatments were smaller and developed slowly. In contrast, carbaryl was not directly toxic to tadpoles, but indirectly reduced survival because slow growth and development prevented some tadpoles from metamorphosing before the mesocosms dried at medium and low applications. 5. These results suggest that these common pesticides, which share the same mode of action, have similar effects on zooplankton and algae, but differences in the strength and timing of their effects on tadpoles reduce the generality of responses at higher trophic levels. Overall, general predictive models of contaminant effects could be improved by incorporating the relative timing of direct and indirect effects of exposure.

THE IMPACT OF CANNIBALISM IN THE PREY ON PREDATOR-PREY SYSTEMS.

Cannibalism is ubiquitous in natural communities and has the potential to alter the functional relationship of predator-prey interactions. Although cannibalistic species are frequently subject to predation, the consequences of cannibalism in the prey for predator-prey interactions are poorly understood. Using a dragonfly larvae system, I provide the first experimental evidence that cannibalism in the prey creates behavior- and density-mediated indirect effects that result in nonlinear predator-prey interactions. As a consequence, cannibalism in the prey altered the functional relationship of the predator and its prey and reduced the impact of the predator on prey mortality by 47%. By parameterizing a mechanistic predation model, I show that the nonlethal interaction between cannibals and predators reduced cannibalism rates, which explained almost two times more of the observed mortality reduction than the consumption of cannibals. However, only a model that accounted for both behavioral interactions and the consumption of cannibals could predict ∼100% of the observed mortality. Using the mechanistic model, I discuss the long-term effects of cannibalism on community dynamics and how they can differ from effects of simple density-dependent mortality. In general, these results demonstrate the importance of accounting for the trophic structure in cannibalistic populations and the resulting nonlinear interactions to predict predator-prey dynamics.

DEFINING AND MEASURING THE IMPACT OF DYNAMIC TRAITS ON INTERSPECIFIC INTERACTIONS

Trait- and density-mediated indirect effects describe different pathways by which indirect interactions in food webs are propagated from one species to another, through changes in intermediate species. A series of articles in Ecology has progressively altered the original definitions of "trait-mediated" to the point where understanding is being impeded. The most recent of these articles are two meta-analyses that use "trait-mediated" to describe the demographic costs to a prey species of employing anti-predator defenses. These same articles introduce a companion term, "density-mediated interaction", apparently to describe direct and indirect interactions that only involve changes in population density due to consumption by predators. This new terminology has many disadvantages, including (1) using a general term for a relatively narrow group of processes; (2) using "mediated" in a manner inconsistent with existing terminology; (3) confusing the accepted definitions of different types of indirect effects; and (4) providing a highly incomplete measure of the impact of behavior on the predator-prey interaction. Solutions to these problems and the meaning of the meta-analyses are discussed.

Effects of experimental duration and density of Japanese dace Tribolodon hakonensis on the strength of trophic cascades on benthic algae

To analyze density-mediated indirect effects through trophic cascades caused by Japanese dace Tribolodon hakonensis on the algal biomass and growth of ayu Plecoglossus altivelis altivelis, (freshwater fish belonging to Salmoniformes), I introduced zero, five, ten, and 20 dace with and without ayu into experimental pools. Ayu fed predominantly on benthic algae, whereas dace fed on both invertebrates and algae. The percentage of algae in the food content of dace was correlated with the number of dace in a pool. In the experiment lasting 8 days, trophic cascades on algae were not clear. In contrast, when the experimental duration was extended to 20 days, the number of gastropods and ephemeropteran nymphs was negatively correlated with the number of dace, consequently increasing the algal biomass and ayu growth through trophic cascades. Compared with pools with five dace, the presence of ten and 20 dace had similar gross effects on the number of algivorous invertebrates and algal biomass and greater gross effects on the ayu growth. The significant increase in ayu growth in pools with ten and 20 dace strongly suggests that small positive effects on algal growth by dace accumulated and extended to ayu during a 20-day experimental period. These results indicate the importance of analyzing interactions between the density and behavior of the top predator and the experimental duration on the intensity of trophic cascades.

LETHAL AND NONLETHAL PREDATOR EFFECTS ON AN HERBIVORE GUILD MEDIATED BY SYSTEM PRODUCTIVITY

Indirect effects propagated through intervening species in a food web have important effects on community properties. Traditionally, these indirect effects have been conceptualized as mediated through density changes of the intervening species, but it is becoming increasingly apparent that those mediated through trait (phenotypic) responses also can be very important. Because density- and trait-mediated indirect effects have different properties, it is critical that we understand the mechanisms of transmission in order to predict how they will interact, and when or where they will be important. In this study, we examined the mechanisms and consequences of the lethal (density-mediated) and nonlethal (trait-mediated) effects of a larval odonate predator on a guild of four herbivore species (a larval anuran and three species of snails) and their resources. We also manipulated system productivity in order to explore the effects of environmental context on the transmission of these two types of indirect effects. We show that trait-mediated effects arising from the predator can be very strong relative to density-mediated effects on both the competing herbivores and the species composition and production of their resources. A number of these indirect effects are shown to be contingent on productivity of the system. We further present evidence that trait- and density-mediated indirect effects originating from a predator may be transmitted independently through different routes in a food web, particularly when spatial responses of the transmitting prey are involved. Finally, effects on prey growth due to trait responses to the predator varied from negative to positive in predictable ways as a function of time and indirect effects on the larger food web. These results indicate the important role that trait-mediated indirect effects can play in trophic cascades and keystone predator interactions, and we discuss how the mechanisms involved can be incorporated in theory.

APPARENT COMPETITION, QUANTITATIVE FOOD WEBS, AND THE STRUCTURE OF PHYTOPHAGOUS INSECT COMMUNITIES

Phytophagous insects and their natural enemies make up one of the largest and most diverse groups of organisms on earth. Ecological processes, in particular negative indirect effects mediated by shared natural enemies (apparent competition), may be important in structuring phytophagous insect communities. The potential for indirect interactions can be assessed by analyzing the trophic structure of insect communities, and we claim that quantitative food webs are particularly well suited for this task. We review the experimental evidence for both short-term and long-term apparent competition in phytophagous insect communities and discuss the possible interactions between apparent competition and intraguild predation or shared mutualists. There is increasing evidence for the importance of trait-mediated as well as density-mediated indirect effects. We conclude that there is a need for large-scale experiments manipulating communities in their entirety and a greater integration of community and chemical ecology.

Simulation of trait- and density-mediated indirect effects induced by piscivorous predators

The study demonstrates how to use a spatially explicit individual-based model (IBM) to tackle the problem of resolving density-mediated (DMII) and trait-mediated indirect interactions (TMII) when manipulative experimentation is difficult on the scale of community-level interactions in the real word. As an example we simulated predator-induced behavioural changes in a roach ( Rutilus rutilus (L.)) population before and after stocking a lake with an additional pelagic predator ( Sander lucioperca (L.)). By this it was possible to calculate bioenergetic gains and costs of trait changes as well as the relative proportions of DMIIs and TMIIs in roach consumption on littoral and pelagic resources in the lake food web. Despite higher net activity costs of 64%, roach migrating horizontally over the diel cycle had a net benefit (higher energy intake of 75%) compared to roach that were restricted to the littoral as a behavioural response to the new predator. By separating the components of the indirect predator effects, we demonstrate that a predator-induced modification in prey traits (behaviour, feeding activity) could contribute substantially to the net indirect effect of a predator even when there are strong density effects. When considering the predation pressure by roach on pelagic resources, the net TMIIs due to the behavioural adaptation of the consumer in response to the predator were 13–14 times stronger than the DMIIs alone. When comparing the different effects on littoral resources it can be shown that the TMIIs approximately compensate the DMIIs thus resulting in a nearly zero net effect. In der vorliegenden Arbeit wurde ein räumlich explizites individuenbasiertes Modell (IBM) verwendet, um dichteabhängige (DMII) und eigenschaftsabhängige (TMII) indirekte Interaktionen zu vergleichen. Da sich dies auf der Ebene der Lebensgemeinschaft unter realen experimentellen Bedingungen als besonders schwierig erwiesen hat, wurden räuberinduzierte Verhaltensänderungen einer Plötzenpopulation ( Rutilus rutilus (L.)) vor und nach dem Besatz mit einem zusätzlichen pelagischen Räuber, dem Zander ( Sander lucioperca (L.)), simuliert. Dadurch war es möglich, sowohl die energetischen Kosten und Nutzen der räuberinduzierten Verhaltensänderung als auch die relativen Anteile von DMIIs and TMIIs an der Plötzenkonsumtion auf litorale und pelagische Ressourcen innerhalb des Nahrungsnetzes zu kalkulieren. Verglichen mit den Plötzen, welche sich aufgrund des neuen Räubers vor allem im Litoral aufhielten, hatten die täglich (horizontal) wandernden Plötzen trotz höherer Nettoaktivitätskosten von 64% eine größere Energieaufnahme von 75%. Durch die Auflösung der verschiedenen indirekten Räubereffekte konnten wir demonstrieren, dass räuberinduzierte Modifikationen von Beuteeigenschaften (Verhalten, Fraßaktivität) erheblich zu den indirekten Nettoeffekten eines Räubers beitragen können, auch wenn starke dichteabhängige Effekte vorhanden sind. Wenn man die Plötzenkonsumtion auf pelagische Ressourcen betrachtet, so ergeben sich 13-14 fach stärkere TMIIs als DMIIs. Wenn man die verschiedenen Effekte auf litorale Ressourcen vergleicht, zeigt es sich, dass TMIIs die DMIIs kompensieren, wodurch der indirekte Nettoeffekt fast null beträgt.