Role of inducible defenses in the stability of a tritrophic system

Inducible defenses are a form of phenotypic plasticity that potentially modify direct interactions between various members of an ecological community, generating trait-mediated indirect effects. In this work, the hypothesis that inducible defenses increase the stability of tritrophic chains is tested, through the numerical analysis of a continuous-time model that discriminate between defenses affecting attack rate of predators, and defenses affecting predator handling time. In addition, discrimination between feeding costs of defenses affecting attack rate, and metabolic costs affecting feeding requirement for zero growth are considered. System stability was examined by computing dominant Lyapunov exponents, and through continuation routines of bifurcation points. Background parameter values were taken from two published studies. Our results show that a tritrophic system will generally be stabilized by the incorporation of inducible defenses and by their associated costs, but a number of new outcomes were obtained. Different long-term behavior is predicted if either one or two prey populations exhibit defenses. In the latter case, the defense of the basal prey dominates the dynamics. Handling time based inducible defenses exert a stronger stabilizing effect than attack rate based ones, but also impose a higher extinction risk for top predators. Inducible defenses in particular and trait-mediated indirect effects in general can be important sources of stability in natural systems.     

Induced defenses within food webs: The role of community trade-offs,delayed responses,and defense specificity

In nature, prey and predator species are embedded in complex networks of ecological interactions. As a consequence, organism level reactions such as predator-induced prey defenses will not only influence the dynamics of both the prey exhibiting the response and its inducer predator, but also that of a wider set of populations that interact directly or indirectly with them.In this work our aim is to determine the consequences of community-level side effects, defense specificity, and timing of inducible defenses for the stability of model ecological communities. We shall consider small webs of two and three trophic levels, containing one to three species per level. The model food webs include well-known community motifs that will be studied by means of qualitative analyses of the community matrix. Our results show that side effects that suppress non-focal interactions were able to decrease community stability, particularly when defensive responses were delayed. Conversely, side effects that increase the strength of non-focal interactions stabilized communities. This work also shows that as the defensive response became more specific, it is more likely to obtain a stable community. In general terms, our results revealed that delayed responses decrease the likelihood of system stability. Our results highlight the importance of the underlying biology of species interactions for the definition of the proper topology, and consequently dynamics, of complex ecological networks.     

Inducible defenses,competition and shared predation in planktonic food chains

Ecologists have long debated the role of predation in mediating the coexistence of prey species. Theory has mainly taken a bitrophic perspective that excludes the effects of inducible defenses at different trophic levels. However, inducible defenses could either limit or enhance the effects of predation on coexistence, by means of effects on bottom-up control and population stability. Our aim was to investigate how inducible defenses at different trophic levels affect the possibilities for predator-mediated coexistence, as opposed to competitive exclusion, in replicated experimental plankton communities. In particular, we analyzed how the presence or absence of inducible defenses in algal basal prey affected the outcome of competition between an inducible defended and an undefended herbivore, in the presence or absence of a carnivore. We found the undefended herbivore to be a superior competitor in the absence of predation. This outcome was reversed in the presence of a shared carnivore: populations of the undefended herbivore then strongly declined. The extent of this population decline differed between food webs based on undefended as opposed to inducible defended algal prey. In the former the undefended herbivore became undetectable for most of the duration of the experiment. In the latter the undefended herbivore also crashed to low densities, but it could still be detected during most of the experiment. In food webs based on inducible defended algae, the carnivore failed to reach high densities and exerted weaker top-down control on the two competing herbivores. We conclude that the inducible defense in one of our two competing herbivores allowed the outcome of competition to be reversed when a shared carnivore was added. Inducible defenses in algae did not change this outcome, but they significantly delayed extinction of the undefended herbivore. Predation itself did not promote coexistence in these experimental plankton communities.     

The long‐term impacts of predators on prey: inducible defenses,population dynamics,and indirect effects

Despite the amount of research on the inducible defenses of prey against predators, our understanding of the long‐term significance of non‐lethal predators on prey phenotypes, prey population dynamics, and community structure has rarely been explored. Our objectives were to assess the effects of predators on prey defenses, prey population dynamics, and the relative magnitude of density‐ versus trait‐mediated indirect interactions (DMIIs and TMIIs) over multiple prey generations. Using a freshwater snail and three common snail predators, we constructed a series of community treatments with pond mesocosms that manipulated trophic structure, the identity of the top predator, and whether predators were caged or uncaged. We quantified snail phenotypes, snail population size, and resource abundance over multiple snail generations. We found that snails were expressing inducible defenses in our system although the magnitude of the responses varied over time and across predator species. Despite the expression of inducible defenses, caged predators did not reduce snail population size. There also was no evidence of TMIIs throughout the experiment suggesting that TMIIs have a minimal role in the long‐term structure of our communities. The absence of TMIIs was largely driven by the lack of predator‐induced reductions in resource consumption and the lack of consistent reductions in population size with predator cues. In contrast, we detected strong DMIIs associated with lethal predators suggesting that DMIIs are the dominant long‐term mechanism influencing community structure. Our results demonstrate that although predators can have significant effects on prey phenotypes and sometimes cause short‐term TMIIs, there may be few long‐term consequences of these responses on population dynamics and indirect interactions, at least within simple food webs. Research directed towards addressing the long‐term consequences of predator–prey interactions within communities will help to reveal whether the conclusions and predictions generated from short‐term experiments are applicable over ecological and evolutionary timescales.     

Behavioral plasticity in an invaded system: non-native whelks recognize risk from native crabs

Inducible defenses have the potential to affect both invasion success and the structure of invaded communities. However, little is known about the cues used for risk-recognition that influence the expression of inducible defenses in invasive prey, because they involve a novel threat. In laboratory experiments, we investigated behavioral defenses induced by a native crab on two invasive oyster drills (marine whelks Urosalpinx cinerea and Ocinebrina inornata). Both drills hid more often and reduced their feeding rates when they detected predators consuming conspecific prey. Examination of the responses of U. cinerea to specific cue sources (predator kairomones, conspecific alarm cues) indicated that this species had the strongest responses to cues from injured conspecifics, but that it did recognize the novel crab predator. Our observation of native predator (per se) recognition by an invasive marine prey is novel. In addition, we observed that neither species of drill reduced their defensive behavior to reflect predation risk shared by a group of prey. The lack of density dependence in risk-assessment could cause populations of invasive prey to transmit both quantitatively and qualitatively different community effects over the course of an invasion as abundance changes. Together, these findings demonstrate several ways that the risk-assessment strategies could be important in establishment and post-establishment dynamics of invasive prey.     

Linearity in the aggregate effects of multiple predators in a food web

Theory in community ecology often assumes that predator species have similar indirect effects and thus can be treated mathematically as a single functional unit (e.g. guild or trophic level). This assumption is questionable biologically because predator species typically differ in their effects, creating the potential for nonlinearities when they coexist. We evaluated the nature of indirect effects caused by three species of hunting spider predators, singly and in multiple species combinations, on grass and herb plants in experimental old-field food webs. Despite the potential for nonlinearity, indirect effects in different multiple predator combinations consistently did not differ significantly from the respective means of the single species effects. Thus, for this experimental system, the whole was simply the average of the parts. Consequently, models which abstract predator species as single trophic levels would successfully predict indirect effects in this system regardless of the composition of the predator fauna.     

Effect of habitat degradation on competition,carrying capacity,and species assemblage stability

Changes in species’ trophic niches due to habitat degradation can affect intra‐ and interspecific competition, with implications for biodiversity persistence. Difficulties of measuring species’ interactions in the field limit our comprehension of competition outcomes along disturbance gradients. Thus, information on how habitat degradation can destabilize food webs is scarce, hindering predictions regarding responses of multispecies systems to environmental changes. Seagrass ecosystems are undergoing degradation. We address effects of Posidonia oceanica coverage reduction on the trophic organization of a macroinvertebrate community in the Tyrrhenian Sea (Italy), hypothesizing increased trophic generalism, niche overlap among species and thus competition and decreased community stability due to degraded conditions. Census data, isotopic analysis, and Bayesian mixing models were used to quantify the trophic niches of three abundant invertebrate species, and intra‐ and interspecific isotopic and resource‐use similarity across locations differing in seagrass coverage. This allowed the computation of (1) competition strength, with respect to each other and remaining less abundant species and (2) habitat carrying capacity. To explore effects of the spatial scale on the interactions, we considered both individual locations and the entire study area (“‘meadow scale”). We observed that community stability and habitat carrying capacity decreased as P. oceanica coverage declined, whereas niche width, similarity of resource use and interspecific competition strength between species increased. Competition was stronger, and stability lower, at the meadow scale than at the location scale. Indirect effects of competition and the spatial compartmentalization of species interactions increased stability. Results emphasized the importance of trophic niche modifications for understanding effects of habitat loss on biodiversity persistence. Calculation of competition coefficients based on isotopic distances is a promising tool for describing competitive interactions in real communities, potentially extendible to any subset of ecological niche axes for which specimens’ positions and pairwise distances can be obtained.     

昆虫取食诱导的植物防御反应

植物被昆虫取食后可产生直接防御或间接防御。直接防御通过增加有毒的次生代谢产物或防御蛋白对昆虫生理代谢产生不利的影响,但对植物的消耗较大。间接防御通过释放挥发性化合物吸引天敌昆虫,并以此控制植食性昆虫。特异性的昆虫激发子(insect specific elicitors)能够诱导挥发性化合物的释放。多种信号途径参与昆虫取食诱导的植物防御反应,它们之间的相互作用协同或拮抗。了解昆虫取食诱导的植物防御反应,对于害虫综合治理策略的完善具有重要的意义。     

Autochthonous or allochthonous resources determine the characteristic population dynamics of ecosystem engineers and their impacts

Ecosystem engineering, or the modification of physical environments by organisms, can influence trophic interactions and thus food web dynamics. Although existing theory exclusively considers engineers using autochthonous resources, many empirical studies show that they often depend on allochthonous resources. By developing a simple mathematical model involving an ecosystem engineer that modifies the physical environment through its activities, its resource, and physical environment modified by the engineer, we compare the effects of autochthonous and allochthonous resources on the dynamics and stability of community with ecosystem engineers. To represent a variety of real situations, we consider engineers that alter either resource productivity, engineer feeding rate on the resource, or engineer mortality, and incorporate time-lagged responses of the physical environment. Our model shows that the effects of ecosystem engineering on community dynamics depend greatly on resource types. When the engineer consumes autochthonous resources, the community can exhibit oscillatory dynamics if the engineered environment affects engineer’s feeding rate or mortality. These cyclic behaviors are, however, stabilized by a slowly responding physical environment. When allochthonous resources are supplied as donor-controlled, on the other hand, the engineer population is unlikely to oscillate but instead can undergo unbounded growth if the engineered environment affects resource productivity or engineer mortality. This finding suggests that ecosystem engineers utilizing allochthonous resources may be more likely to reach high abundance and cause strong impacts on ecosystems. Our results highlight that community-based, compounding effects of trophic and physical biotic interactions of ecosystem engineers depend crucially on whether the engineers utilize autochthonous or allochthonous resources.     

Tri‐trophic interactions: bridging species,communities and ecosystems

A vast body of research demonstrates that many ecological and evolutionary processes can only be understood from a tri‐trophic viewpoint, that is, one that moves beyond the pairwise interactions of neighbouring trophic levels to consider the emergent features of interactions among multiple trophic levels. Despite its unifying potential, tri‐trophic research has been fragmented, following two distinct paths. One has focused on the population biology and evolutionary ecology of simple food chains of interacting species. The other has focused on bottom‐up and top‐down controls over the distribution of biomass across trophic levels and other ecosystem‐level variables. Here, we propose pathways to bridge these two long‐standing perspectives. We argue that an expanded theory of tri‐trophic interactions (TTIs) can unify our understanding of biological processes across scales and levels of organisation, ranging from species evolution and pairwise interactions to community structure and ecosystem function. To do so requires addressing how community structure and ecosystem function arise as emergent properties of component TTIs, and, in turn, how species traits and TTIs are shaped by the ecosystem processes and the abiotic environment in which they are embedded. We conclude that novel insights will come from applying tri‐trophic theory systematically across all levels of biological organisation.     

Inducible defences and the paradox of enrichment

In order to evaluate the effects of inducible defences on community stability and persistence, we analyzed models of bitrophic and tritrophic food chains that incorporate consumer-induced polymorphisms. These models predict that intra-specific heterogeneity in defence levels resolves the paradox of enrichment for a range of top-down effects that affect consumer death rates and for all possible levels of primary productivity. We show analytically that this stability can be understood in terms of differences in handling times on the different prey types. Our predictions still hold when defences also affect consumer attack rates. The predicted stability occurs in both bitrophic and tritrophic food chains.
Inducible defences may promote population persistence in tritrophic food chains. Here the minimum densities of cycling populations remain bound away from zero, thus decreasing the risk of population extinctions. However, the reverse can be true for the equivalent bitrophic predator–prey model. This shows that theoretical extrapolations from simple to complex communities should be made with caution. Our results show that inducible defences are among the ecological factors that promote stability in multitrophic communities.     

Analysis of trophic interactions reveals highly plastic response to climate change in a tri-trophic High-Arctic ecosystem

As a response to current climate changes, individual species have changed various biological traits, illustrating an inherent phenotypic plasticity. However, as species are embedded in an ecological network characterised by multiple consumer–resource interactions, ecological mismatches are likely to arise when interacting species do not respond homogeneously. The approach of biological networks analysis calls for the use of structural equation modelling (SEM), a multidimensional analytical setup that has proven particularly useful for analysing multiple interactions across trophic levels. Here we apply SEM to a long-term dataset from a High-Arctic ecosystem to analyse how phenological responses across three trophic levels are coupled to snowmelt patterns and how changes may cascade through consumer–resource interactions. Specifically, the model included the effect of snowmelt on a High-Arctic tri-trophic system of flowers, insects and waders (Charadriiformes), with latent factors representing phenology (timing of life history events) and performance (abundance or reproduction success) for each trophic level. The effects derived from the model demonstrated that the time of snowmelt directly affected plant and arthropod phenology as well as the performance of all included trophic levels. Additionally, timing of snowmelt appeared to indirectly influence wader phenology as well as plant, arthropod and wader performance through effects on adjacent trophic levels and lagged effects. The results from the tri-trophic community presented here emphasise that effects of climate on species in consumer–resource systems may propagate through trophic levels.     

Modelling food web complexity: The consequences of individual-based, spatially explicit behavioural ecology on trophic interactions

We present a prototype simulator that enables one to explore the influence of individual behaviour on the dynamics and structural complexity of food webs. In the simulations, individuals act according to simple, biologically plausible rules in a spatially explicit setting. We present the results of a series of simulation experiments on artificial, tri-trophic level food chains used to calibrate the simulator against real-world systems and to demonstrate the simulators promise for ecological modelling. Our primary objective was to discover the biological features leading to stability of artificial food chains over ecological time and under different conditions of trophic efficiency. This involved a qualitative analysis of food chains comprised of a plant, a herbivore and a carnivore species. We explored the consequences of allowing individual heterotrophs to make active choices about resource selection (perception and intentional behaviour) under high and low degrees of trophic efficiency. We found that individuals had to adopt realistic behavioural ecological strategies, such as active resource selection, for systems to persist, especially under conditions in which trophic efficiencies were of the magnitude observed in real systems (e.g. 10%). Our results reaffirm previous convictions that a better understanding of food web interactions in real-world systems will require approaches that blend animal behavioural ecology with population and community ecology. However, the evidence comes from a new mathematical perspective.     

Amphibian defenses against ultraviolet-B radiation

As part of an overall decline in biodiversity, amphibian populations throughout the world are disappearing. There are a number of potential causes for these declines, including those related to environmental changes such as increasing ultraviolet-B (UV-B) radiation due to stratospheric ozone depletion. UV-B radiation can kill amphibian embryos or can cause sublethal effects that can harm amphibians in later life stages. However, amphibians have defenses against UV-B damage that can limit damage or repair it after exposure to UV-B radiation. These include behavioral, physiological, and molecular defenses. These defenses differ interspecifically, with some species more able to cope with exposure to UV-B than others. Unfortunately, the defense mechanisms of many species may not be effective against increasing persistent levels of UV-B radiation that have only been present for the past several decades due to human-induced environmental damage. Moreover, we predict that persistent UV-B-induced mortality and sublethal damage in species without adequate defenses could lead to changes in community structure. In this article we review the effects of UV-B radiation on amphibians and the defenses they use to avoid solar radiation and make some predictions regarding community structure in light of interspecific differences in UV-B tolerance.     

The structure and stability of model ecosystems assembled in a variable environment

To explore how environmental variability may create non‐random community structure, we simulated the assembly of model communities under varying levels of environmental variability. We assembled communities by creating a large pool of randomly constructed species, and then added species from this pool sequentially, allowing extinctions of invading and resident species to occur until the community became saturated. Because much current research on community structure focuses on single trophic levels, we constructed species pools consisting only of competitors. To compare with more realistic communities, we also created species pools with multiple trophic levels. For both types of communities, following assembly we calculated a variety of metrics of community structure, and five measures of community stability. Communities assembled under high environmental variability had fewer species, fewer and weaker interactions among species, and greater evenness in abundance of persisting species. For single trophic‐level communities, community size was dictated primarily by competitive exclusion. In contrast, for multiple trophic‐level communities, community size was increasingly limited by dynamical instabilities as environmental variability increased. Differences in community structure resulting from assembly under high environmental variability led to differences in community stability. According to two measures of stability related to population variability – the characteristic return rate to equilibrium and the coefficient of variation in individual species densities – stability increased for communities assembled under high environmental variability. In contrast, three additional measures of stability that are not directly related to population variability showed a variety of patterns, either increasing, decreasing, or remaining constant. Thus, communities assembled in highly variable environments are not necessarily generically more stable. Our results demonstrate that environmental variability can structure communities and affect their stability properties in non‐trivial ways. Thus, when making predictions about the response of communities to future extinctions or environmental degradation, account should be given to the forces responsible for community structure.     

The ecology and evolution of inducible defenses

Inducible defenses are responses activated through a previous encounter with a consumer or competitor that confer some degree of resistance to subsequent attacks. While the importance of inducible resistance has long been known in host-parasite interactions, it is only recently that its importance has emerged in other natural systems. Although the structural defenses produced by invertebrates to their competitors and predators are by no means the same as an immune response triggered by parasites, these responses all share the properties of (1) specificity, (2) amplification and (3) memory. This review discusses the following ecological consequences and evolutionary causes of inducible defenses: (1) Inducible defenses render historical factors important in biological interactions and can affect the probability of individual survival and growth, as well as affect population dynamics of consumers in some circumstances. (2) Although the benefits of inducible defenses are often balanced by fitness costs, including reduced growth, reproductive output and survivorship, the role of costs and benefits in the evolution of inducible defenses is by no means clear. A more integrated approach would involve a multivariate analysis of the role of natural selection on the inducible characters of interest, their norms of reaction and correlated fitness characters. (3) The disproportionate representation of inducible, morphological defenses among clonal organisms may be due to both a higher rate of origination and enhanced selection to maintain these defenses in clonal taxa. (4) Inducible defenses should be most common when reliable cues are available, attacks by biological agents are unpredictable, and the fitness gains of defenses are balanced by the costs. An integrated approach to studying inducible defenses would thus combine mechanistic estimates of costs, population-level estimates of defense effectiveness, and genetic estimates of correlations between fitness and inducible characters. This will allow us to estimate rates of evolution in these phenotypically plastic threshold characters.     

Short-term analysis of the phytoplankton structure and dynamics in two ponds with distinct trophic states from Cierva Point (maritime Antarctica)

Phytoplankton communities dominating Musgos and Papúa ponds with differing trophic states were sampled over 3 days enabling the detection of the physiological and population responses of microalgae to short-scale changes in biotic and abiotic factors, rather than frequently analyzed changes in community composition responses to long-scale environmental changes. We hypothesized that both environments undergoing diel changes would be dominated by phytoplankton with generalist strategies, while community structure would be mostly dictated by the trophic state of each water body. The phytoplankton biovolumes of both ponds were strongly dominated by euplanktonic nanoflagellated Chlorophyta, while phycocyanin-rich picocyanobacteria dominated the picophytoplankton. Parallel diel cycles of air and water temperatures were more pronounced on a sunny, warm day which prompted algal photosynthesis, revealed by strong increases in dissolved oxygen and pH. Nutrient and phytoplanktonic chlorophyll a confirmed the hypertrophic condition of Papúa pond. This accounted for the distinct community composition encountered in each pond, which remained stable throughout the study, as revealed by the SIMI index. The inverse relationship between the chl a/abundance ratio and the abundances of dominant species together with varying net growth rates (k′) showed algal reproduction, yet densities remained rather stable in both cases. In Musgos pond, fluctuations in k′ for small and median ciliates shadowed those of pico- and nanophytoplankton, respectively, strongly suggesting that they can control algal growth in these 2-level trophic chains.     

Ontogenetic niche shift, food-web coupling, and alternative stable states

Several recent studies have shown that food web coupling by ontogenetic niche shifts can generate alternative stable states (ASS). However, these studies mainly considered cases where juvenile and adult stages are the top level consumers. The conditions under which ASS occur in more structurally diverse food web configurations have not been explored. In this study, I examine the influence of food-chain length and the trophic positions of juveniles and adults on the existence of ASS. Comprehensive model analysis showed that if both juveniles and adults are top predators, ASS are possible irrespective of their trophic level, because of overcompensation in reproduction and maturation due to strong density dependence, as previously predicted. However, the following potential food-web effects were found: ASS potential (1) disappears if either or both the juveniles and adults have a predator and (2) is once again observed if another predator is added on the stage-specific predator. These mechanisms involve (1) top–down control that relaxes intrastage food competition and (2) top–down cascade that intensifies the intrastage competition, respectively. Furthermore, it was illustrated that the environmental conditions under which ASS occurred varied in complex ways with the coupled food-web configurations. My results provide a novel concept that anthropogenic changes in local community structure (e.g., species extinction and invasion) propagate through space and may cause or prevent regime shifts in broad-scale community structure by altering the resilience to environmental perturbations.     

Poor phenotypic integration of blue mussel inducible defenses in environments with multiple predators

Aquatic prey encounter an array of threat cues from multiple predators and killed conspecifics, yet the vast majority of induced defenses are investigated using cues from single predator species. In most cases, it is unclear if odors from multiple predators will disrupt defenses observed in single-predator induction experiments. We experimentally compared the inducible defenses of the common marine mussel Mytilus edulis to waterborne odor from pairwise combinations of three predators representing two attack strategies. Predators included the sea star, Asterias vulgaris (= Asterias rubens ), and the crabs Carcinus maenas and Cancer irroratus . The mussels increased adductor muscle mass in response to cues from unfed Asterias (a predatory seastar that pulls mussel shells open) and increased shell thickness in response to unfed Carcinus, a predatory crab that crushes or peels shells. However, the mussels did not express either predator specific response when exposed to the combined cues of Asterias and Carcinus , and mussels did not increase shell thickness when exposed to cues from Cancer alone or any pairwise combination of the three predators. Shell closure or 'clamming up' did not occur in response to any predator combination. These results suggest that predator-specific responses to the Asterias and Carcinus are poorly integrated and cannot be expressed simultaneously. Simultaneous cues from multiple predators affect the integration of predator specific defenses and predator odors from functionally similar predators do not necessarily initiate similar defenses. Ultimately, the degree that prey can integrate potentially disparate defenses in a multiple predator environment may have ecological ramifications and represent a seldom explored facet of the evolution of inducible defenses.     

Positive Effects of Plant Genotypic and Species Diversity on Anti-Herbivore Defenses in a Tropical Tree Species

Despite increasing evidence that plant intra- and inter-specific diversity increases primary productivity, and that such effect may in turn cascade up to influence herbivores, there is little information about plant diversity effects on plant anti-herbivore defenses, the relative importance of different sources of plant diversity, and the mechanisms for such effects. For example, increased plant growth at high diversity may lead to reduced investment in defenses via growth-defense trade-offs. Alternatively, positive effects of plant diversity on plant growth may lead to increased herbivore abundance which in turn leads to a greater investment in plant defenses. The magnitude of trait variation underlying diversity effects is usually greater among species than among genotypes within a given species, so plant species diversity effects on resource use by producers as well as on higher trophic levels should be stronger than genotypic diversity effects. Here we compared the relative importance of plant genotypic and species diversity on anti-herbivore defenses and whether such effects are mediated indirectly via diversity effects on plant growth and/or herbivore damage. To this end, we performed a large-scale field experiment where we manipulated genotypic diversity of big-leaf mahogany (Swietenia macrophylla) and tree species diversity, and measured effects on mahogany growth, damage by the stem-boring specialist caterpillar Hypsipyla grandella, and defensive traits (polyphenolics and condensed tannins in stem and leaves). We found that both forms of plant diversity had positive effects on stem (but not leaf) defenses. However, neither source of diversity influenced mahogany growth, and diversity effects on defenses were not mediated by either growth-defense trade-offs or changes in stem-borer damage. Although the mechanism(s) of diversity effects on plant defenses are yet to be determined, our study is one of the few to test for and show producer diversity effects on plant chemical defenses.