Consumer pressure, seed versus safe-site limitation, and plant population dynamics

Plants often suffer reductions in fecundity due to insect herbivory. Whether this loss of seeds has population-level consequences is much debated and often unknown. For many plants, particularly those with long-lived seedbanks, it is frequently asserted that herbivores have minimal impacts on plant abundance because safe-site availability rather than absolute seed number determines the magnitude of future plant recruitment and hence population abundance. However, empirical tests of this assertion are generally lacking and the interplay between herbivory, spatio-temporal variability in seed- or safe-site-limited recruitment, and seedbank dynamics is likely to be complex. Here we use a stochastic simulation model to explore how changes in the spatial and temporal frequency of seed-limited recruitment, the strength of density-dependent seedling survival, and longevity of seeds in the soil influence the population response to herbivory. Model output reveals several surprising results. First, given a seedbank, herbivores can have substantial effects on mean population abundance even if recruitment is primarily safe-site-limited in either time or space. Second, increasing seedbank longevity increases the population effects of herbivory, because annual reductions in seed input due to herbivory are accumulated in the seedbank. Third, population impacts of herbivory are robust even in the face of moderately strong density-dependent seedling mortality. These results imply that the conditions under which herbivores influence plant population dynamics may be more widespread than heretofore expected. Experiments are now needed to test these predictions. Received: 3 November 1999 / Accepted: 15 February 2000     

Consumers limit the abundance and dynamics of a perennial shrub with a seed bank

For nearly 30 years, ecologists have argued that predators of seeds and seedlings seldom have population-level effects on plants with persistent seed banks and density-dependent seedling survival. We parameterized stage-based population models that incorporated density dependence and seed dormancy with data from a 5.5-year experiment that quantified how granivorous mice and herbivorous voles influence bush lupine (Lupinus arboreus) demography. We asked how seed dormancy and density-dependent seedling survival mediate the impacts of these consumers in dune and grassland habitats. In dune habitat, mice reduced analytical lambda (the intrinsic rate of population growth) by 39%, the equilibrium number of aboveground plants by 90%, and the seed bank by 98%; voles had minimal effects. In adjacent grasslands, mice had minimal effects, but seedling herbivory by voles reduced analytical lambda by 15% and reduced both the equilibrium number of aboveground plants and dormant seeds by 63%. A bootstrap analysis demonstrated that these consumer effects were robust to parameter uncertainty. Our results demonstrate that the quantitative strengths of seed dormancy and density-dependent seedling survival--not their mere existence--critically mediate consumer effects. This study suggests that plant population dynamics and distribution may be more strongly influenced by consumers of seeds and seedlings than is currently recognized.     

Long-Term Regional Shifts in Plant Community Composition Are Largely Explained by Local Deer Impact Experiments

The fact that herbivores and predators exert top-down effects to alter community composition and dynamics at lower trophic levels is no longer controversial, yet we still lack evidence of the full nature, extent, and longer-term effects of these impacts. Here, we use results from a set of replicated experiments on the local impacts of white-tailed deer to evaluate the extent to which such impacts could account for half-century shifts in forest plant communities across the upper Midwest, USA. We measured species'' responses to deer at four sites using 10–20 year-old deer exclosures. Among common species, eight were more abundant outside the exclosures, seven were commoner inside, and 16 had similar abundances in- and outside. Deer herbivory greatly increased the abundance of ferns and graminoids and doubled the abundance of exotic plants. In contrast, deer greatly reduced tree regeneration, shrub cover (100–200 fold in two species), plant height, plant reproduction, and the abundance of forbs. None of 36 focal species increased in reproduction or grew taller in the presence of deer, contrary to expectations. We compared these results to data on 50-year regional shifts in species abundances across 62 sites. The effects of herbivory by white-tailed deer accurately account for many of the long-term regional shifts observed in species'' abundances (R² = 0.41). These results support the conjecture that deer impacts have driven many of the regional shifts in forest understory cover and composition observed in recent decades. Our ability to link results from shorter-term, local experiments to regional long-term studies of ecological change strengthens the inferences we can draw from both approaches.     

Herbivory and plant resource competition: a review of two interacting interactions

This review discusses the prevalence and potential for interactive effects between herbivory and competition on plant growth and biomass, and it is apparent that such effects typically arise when there is a mismatch between the spatial scale of herbivore behaviour (food or patch choice) and the spatial heterogeneity of the plant community. Historically, such interactive effects have been examined using two approaches. Studies using the first approach have excluded plant neighbors and herbivores in a factorial experiment, and scored effects on plant biomass. Studies using the second approach have observed herbivore abundance or herbivory on plants with or without plant neighbors, and have identified a large number of mechanisms underlying such interactive effects. The two types of studies have produced somewhat conflicting results, where interactive effects have been commonly observed in studies using the second approach and only rarely in studies using the first approach. This is most likely a consequence of a biased choice of study systems, where studies using the first approach have primarily studied mammalian herbivory while studies using the second approach have been more focussed on insect herbivory. Moreover, studies using the first approach have typically been very small-scale manipulations and this probably precludes most possible interactive effects in systems with mammalian herbivory. This points to the fact that studies examining interactive effects of herbivory and plant competition should more carefully consider the behaviour and life history of herbivores included in the study prior to the design of removal experiments.     

Insect-plant interactions on a planet of weeds

Two conflicting views confront ecologists and evolutionary biologists on the degree of symmetry in interactions between plants and phytophagous insects. The symmetrical view holds that insects and plants have strong effects on one another's evolutionary and ecological dynamics. Thus, herbivores are regarded as a major influence on plant distribution and abundance in contemporary ecosystems, and coevolution is commonly invoked to explain adaptive radiation in plants and insects, host specialization in insects, as well as much of the morphological and chemical variety observed in plants. The asymmetrical view acknowledges that plants have major effects on insects, but claims that insects seldom impose significant effects on plants. Proponents of the asymmetric view tend to ignore or discount insect-plant interactions in communities and ecosystems altered by human impacts. If we recognize the scope and scale of human impacts, and ways in which these impacts change insect-plant interactions, then our views about symmetry or asymmetry in insect-plant interactions will change. To understand, predict, and manage insect herbivory we need to study it in all its manifestations. In particular, the study of interactions involving alien species is both an urgent priority for environmental management and potentially a source of ecological insights on the role of herbivores in plant population and community dynamics. A complete theory of insect/host plant interactions must explain and predict interactions both within and beyond the native range. Such a theory might guide efforts to deal with environmental problems stemming from rapid rates of extinction and homogenization of the world's biota.     

Simultaneous effects of food limitation and inducible resistance on herbivore population dynamics

Many herbivore populations fluctuate temporally, but the causes of those fluctuations remain unclear. Plant inducible resistance can theoretically cause herbivore population fluctuations, because herbivory may induce plant changes that reduce the survival or reproduction of later-feeding herbivores. Herbivory can also simply reduce the quantity of food available for later feeders and this, too, can cause population fluctuations. Inducible resistance and food limitation often occur simultaneously, yet whether they jointly facilitate or suppress herbivore fluctuations remains largely unexplored. We present models that suggest that food limitation and inducible resistance may have synergistic effects on herbivore population dynamics. The population-level response of the food plant to herbivory and the details of how inducible resistance affects herbivore performance both influence the resulting herbivore dynamics. Our results identify some biological properties of plant-herbivore systems that might determine whether or not cycles occur, and suggest that future empirical and theoretical population dynamics studies should account for the effects of both food limitation and inducible resistance.     

Experimental Manipulation of Grassland Plant Diversity Induces Complex Shifts in Aboveground Arthropod Diversity

Changes in producer diversity cause multiple changes in consumer communities through various mechanisms. However, past analyses investigating the relationship between plant diversity and arthropod consumers focused only on few aspects of arthropod diversity, e.g. species richness and abundance. Yet, shifts in understudied facets of arthropod diversity like relative abundances or species dominance may have strong effects on arthropod-mediated ecosystem functions. Here we analyze the relationship between plant species richness and arthropod diversity using four complementary diversity indices, namely: abundance, species richness, evenness (equitability of the abundance distribution) and dominance (relative abundance of the dominant species). Along an experimental gradient of plant species richness (1, 2, 4, 8, 16 and 60 plant species), we sampled herbivorous and carnivorous arthropods using pitfall traps and suction sampling during a whole vegetation period. We tested whether plant species richness affects consumer diversity directly (i), or indirectly through increased productivity (ii). Further, we tested the impact of plant community composition on arthropod diversity by testing for the effects of plant functional groups (iii). Abundance and species richness of both herbivores and carnivores increased with increasing plant species richness, but the underlying mechanisms differed between the two trophic groups. While higher species richness in herbivores was caused by an increase in resource diversity, carnivore richness was driven by plant productivity. Evenness of herbivore communities did not change along the gradient in plant species richness, whereas evenness of carnivores declined. The abundance of dominant herbivore species showed no response to changes in plant species richness, but the dominant carnivores were more abundant in species-rich plant communities. The functional composition of plant communities had small impacts on herbivore communities, whereas carnivore communities were affected by forbs of small stature, grasses and legumes. Contrasting patterns in the abundance of dominant species imply different levels of resource specialization for dominant herbivores (narrow food spectrum) and carnivores (broad food spectrum). That in turn could heavily affect ecosystem functions mediated by herbivorous and carnivorous arthropods, such as herbivory or biological pest control.     

Plant Diversity Impacts Decomposition and Herbivory via Changes in Aboveground Arthropods

Loss of plant diversity influences essential ecosystem processes as aboveground productivity, and can have cascading effects on the arthropod communities in adjacent trophic levels. However, few studies have examined how those changes in arthropod communities can have additional impacts on ecosystem processes caused by them (e.g. pollination, bioturbation, predation, decomposition, herbivory). Therefore, including arthropod effects in predictions of the impact of plant diversity loss on such ecosystem processes is an important but little studied piece of information. In a grassland biodiversity experiment, we addressed this gap by assessing aboveground decomposer and herbivore communities and linking their abundance and diversity to rates of decomposition and herbivory. Path analyses showed that increasing plant diversity led to higher abundance and diversity of decomposing arthropods through higher plant biomass. Higher species richness of decomposers, in turn, enhanced decomposition. Similarly, species-rich plant communities hosted a higher abundance and diversity of herbivores through elevated plant biomass and C:N ratio, leading to higher herbivory rates. Integrating trophic interactions into the study of biodiversity effects is required to understand the multiple pathways by which biodiversity affects ecosystem functioning.     

Impacts of invasive plants on resident animals across ecosystems,taxa, and feeding types: a global assessment

As drivers of global change, biological invasions have fundamental ecological consequences. However, it remains unclear how invasive plant effects on resident animals vary across ecosystems, animal classes, and functional groups. We performed a comprehensive meta‐analysis covering 198 field and laboratory studies reporting a total of 3624 observations of invasive plant effects on animals. Invasive plants had reducing (56%) or neutral (44%) effects on animal abundance, diversity, fitness, and ecosystem function across different ecosystems, animal classes, and feeding types while we could not find any increasing effect. Most importantly, we found that invasive plants reduced overall animal abundance, diversity and fitness. However, this significant overall effect was contingent on ecosystems, taxa, and feeding types of animals. Decreasing effects of invasive plants were most evident in riparian ecosystems, possibly because frequent disturbance facilitates more intense plant invasions compared to other ecosystem types. In accordance with their immediate reliance on plants for food, invasive plant effects were strongest on herbivores. Regarding taxonomic groups, birds and insects were most strongly affected. In insects, this may be explained by their high frequency of herbivory, while birds demonstrate that invasive plant effects can also cascade up to secondary consumers. Since data on impacts of invasive plants are rather limited for many animal groups in most ecosystems, we argue for overcoming gaps in knowledge and for a more differentiated discussion on effects of invasive plant on native fauna.     

Effects of plant traits and the relative abundance of common woody species on seedling herbivory in the Thousand Island Lake region

Aims Plant-herbivore interaction is a hot topic in the study of biodiversity and ecosystem functions. Herbivores can negatively affect seedling growth and therefore can alter the dynamics of plant recruitment. However, previous studies do not fully reveal the relative importance of different plant functional traits on herbivory intensity and rarely link herbivory to the relative abundance of plant species.Methods Here, we measured 11 plant functional traits and the relative abundance of seedlings of 16 common woody species in the subtropical forests on 29 islands in Thousand Island Lake, East China. We then used multivariate regression and variance partitioning to test the contribution of functional traits and the relative abundance to interspecific differences of insect herbivory intensity.Important findings Our study found that both plant functional traits (e.g. carbon nitrogen ratio, leaf thickness) and the relative abundance of woody species played important roles in herbivory intensity, and they jointly contributed 54% of the variance of the interspecific differences. Among these factors, species with higher defensive ability, lower nutrient content and higher relative abundance had lower herbivory intensity. We suggest to consider both individual level traits (functional traits) and community level attributes (the relative abundance) in future herbivory studies.     

Herbivory and population dynamics of invasive and native <Emphasis Type="Italic">Lespedeza</Emphasis>

Some exotic plants are able to invade habitats and attain higher fitness than native species, even when the native species are closely related. One explanation for successful plant invasion is that exotic invasive plant species receive less herbivory or other enemy damage than native species, and this allows them to achieve rapid population growth. Despite many studies comparing herbivory and fitness of native and invasive congeners, none have quantified population growth rates. Here, we examined the contribution of herbivory to the population dynamics of the invasive species, Lespedeza cuneata, and its native congener, L. virginica, using an herbivory reduction experiment. We found that invasive L. cuneata experienced less herbivory than L. virginica. Further, in ambient conditions, the population growth rate of L. cuneata (λ = 20.4) was dramatically larger than L. virginica (λ = 1.7). Reducing herbivory significantly increased fitness of only the largest L. virginica plants, and this resulted in a small but significant increase in its population growth rate. Elasticity analysis showed that the growth rate of these species is most sensitive to changes in the seed production of small plants, a vital rate that is relatively unaffected by herbivory. In all, these species show dramatic differences in their population growth rates, and only 2% of that difference can be explained by their differences in herbivory incidence. Our results demonstrate that to understand the importance of consumers in explaining the relative success of invasive and native species, studies must determine how consumer effects on fitness components translate into population-level consequences. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Forage nutritive quality in the Serengeti ecosystem: the roles of fire and herbivory

Fire and herbivory are important determinants of nutrient availability in savanna ecosystems. Fire and herbivory effects on the nutritive quality of savanna vegetation can occur directly, independent of changes in the plant community, or indirectly, via effects on the plant community. Indirect effects can be further subdivided into those occurring because of changes in plant species composition or plant abundance (i.e., quality versus quantity). We studied relationships between fire, herbivory, rainfall, soil fertility, and leaf nitrogen (N), phosphorus (P), and sodium (Na) at 30 sites inside and outside of Serengeti National Park. Using structural equation modeling, we asked whether fire and herbivory influences were largely direct or indirect and how their signs and strengths differed within the context of natural savanna processes. Herbivory was associated with enhanced leaf N and P through changes in plant biomass and community composition. Fire was associated with reduced leaf nutrient concentrations through changes in plant community composition. Additionally, fire had direct positive effects on Na and nonlinear direct effects on P that partially mitigated the indirect negative effects. Key mechanisms by which fire reduced plant nutritive quality were through reductions of Na-rich grasses and increased abundance of Themeda triandra, which had below-average leaf nutrients.     

Small mammal herbivores mediate the effects of soil nitrogen and invertebrate herbivores on grassland diversity

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Plant diversity effects on arthropods and arthropod-dependent ecosystem functions in a biodiversity experiment

Biodiversity-ecosystem function experiments test how species diversity influences fundamental ecosystem processes. Historically, arthropod driven functions, such as herbivory and pest-control, have been thought to be influenced by direct and indirect associations among species. Although a number of studies have evaluated how plant diversity affects arthropod communities and arthropod-mediated ecosystem processes, it remains unclear whether diversity effects on arthropods are sufficiently consistent over time such that observed responses can be adequately predicted by classical hypotheses based on associational effects. By combining existing results from a long-term grassland biodiversity experiment (Jena Experiment) with new analyses, we evaluate the consistency of consumer responses within and across taxonomic, trophic, and trait-based (i.e. vertical stratification) groupings, and we consider which changes in arthropod community composition are associated with changes in consumer-mediated ecosystem functions.Overall, higher plant species richness supported more diverse and complex arthropod communities and this pattern was consistent across multiple years. Vegetation-associated arthropods responded more strongly to changes in plant species richness than ground-dwelling arthropods. Additionally, increases in plant species richness were associated with shifts in the species-abundance distributions for many, but not all taxa. For example, highly specialized consumers showed a decrease in dominance and an increase in the number of rare species with increasing plant species richness. Most ecosystem processes investigated responded to increases in plant species richness in the same way as the trophic group mediating the process, e.g. both herbivory and herbivore diversity increase with increasing plant species richness. In the Jena Experiment and other studies, inconsistencies between predictions based on classic hypotheses of associational effects and observed relationships between plant species richness and arthropod diversity likely reflect the influence of multi-trophic community dynamics and species functional trait distributions. Future research should focus on testing a broader array of mechanisms to unravel the biological processes underlying the biodiversity-ecosystem functioning relationships.     

Ecological Restoration Treatments Increase Butterfly Richness and Abundance: Mechanisms of Response

Few ecosystem restoration studies evaluate whether arthropods are important components of ecosystem recovery. We tested the hypothesis that ponderosa pine restoration treatments would increase adult butterfly species richness and abundance as a direct result of increased understory diversity and abundance. To examine mechanisms that potentially affect adult butterfly distribution, we quantified host plant frequency, nectar plant abundance, and insolation (light intensity) in restoration treatment and control forests. This study is unique, because this is the first invertebrate monitoring in ponderosa pine forest restoration treatments in the U.S. Southwest and also because these treatments are the first replicated ponderosa pine restoration treatments at a landscape scale. Three patterns emerged: (1) butterfly species richness and abundance were 2 and 3 times greater, respectively, in restoration treatment units than in paired control forests 1 year after treatment, and 1.5 and 3.5 times greater, respectively, 2 years after treatment, ordination of control and treatment sampling units using butterfly assemblages showed significant separation of control and restoration treatment units after restoration treatment; (2) host plant and nectar plant species richness showed little difference between treated and control forests even 2 years after treatment; and (3) insolation (light intensity) was significantly greater in treated forests after restoration. We suggest that changes in the butterfly assemblage may occur due to light intensity effects before plant community changes occur or can be detected. Butterfly assemblage differences will have additional cascading effects on the ecosystem as prey for higher trophic levels and through plant interactions including herbivory and pollination.     

How does herbivory affect individuals and populations of the perennial herb Paeonia officinalis?

In this study we quantified variability in foliage herbivory and pre-dispersal seed predation and its effects on plant performance and demography in populations of a rare and protected perennial herb, Paeonia officinalis. An individual-based survey was performed during four years in four populations, which contained plants in both open habitat and woodland. We detected marked spatial and temporal variation among and within populations in foliage herbivory (by insects) and pre-dispersal seed predation (by insects, rodents and Roe Deer). Foliage herbivory decreased with plant demographic stages in open habitats, from seedlings to reproductive individuals, but no significant trend was detected in woodland habitats. This may be due to different demographic origin of larger vegetative plants in this habitat. Depending on demographic stage, herbivory was higher in open habitats or not significantly different between habitats. This suggests differences in herbivore abundance in different habitats within sites. Pre-dispersal seed predation remained weak and did not depend on habitat. We did not detect any consequence of foliage herbivory on seedling mortality and individual growth in our study. Our results illustrate the need to investigate plant-herbivore interactions over several years in distinct populations in order to more accurately evaluate herbivore impact on plant population dynamics.     

The ecology of predispersal insect herbivory on tree reproductive structures in natural forest ecosystems

Plant-insect interactions are key model systems to assess how some species affect the distribution, the abundance, and the evolution of others. Tree reproductive structures represent a critical resource for many insect species, which can be likely drivers of demography, spatial distribution, and trait diversification of plants. In this review, we present the ecological implications of predispersal herbivory on tree reproductive structures by insects (PIHR) in forest ecosystems. Both insect's and tree's perspectives are addressed with an emphasis on how spatiotemporal variation and unpredictability in seed availability can shape such particular plant-animal interactions. Reproductive structure insects show strong trophic specialization and guild diversification. Insects evolved host selection and spatiotemporal dispersal strategies in response to variable and unpredictable abundance of reproductive structures in both space and time. If PIHR patterns have been well documented in numerous systems, evidences of the subsequent demographic and evolutionary impacts on tree populations are still constrained by time-scale challenges of experimenting on such long-lived organisms, and modeling approaches of tree dynamics rarely consider PIHR when including biotic interactions in their processes. We suggest that spatially explicit and mechanistic approaches of the interactions between individual tree fecundity and in sect dynamics will clarify predictions of the demogenetic implications of PIHR in tree populations. In a global change context, further experimental and theoretical contributions to the likelihood of life-cycle disruptions between plants and their specialized herbivores, and to how these changes may gen erate novel dynamic patterns in each partner of the interaction are increasingly critical.     

Equivalence in the strength of deer herbivory on above and below ground communities

Herbivores exert a strong influence on the species composition and richness of plant communities, but the magnitude of their effect on belowground communities remains poorly understood. While an increasing number of studies acknowledge the importance of documenting belowground effects of herbivores, very few of these evaluate variation in the strength of the response from aboveground to belowground communities. Our study documents the long-term consequences of sustained deer herbivory for plant and arthropod communities adjacent to 15 exclosures that have been in place since 1996. We hypothesized that herbivory would alter the composition and diversity of communities, but the strength of the effects of herbivory would weaken from plants, to leaf-litter invertebrates, and to belowground microarthropod communities. First, we found that herbivory negatively impacted plant seedling and sapling abundance and performance, reduced the abundance of ants and the taxonomic richness of arthropods in the litter layer and reduced the richness of soil microarthropod communities. Second, in contrast to our hypothesis, the magnitude of effect size did not vary among trophic levels, indicating that effects of deer herbivory cascade from plants to the leaf-litter and soil arthropod communities with equal strength. While much recent research has focused on how specific traits of plants may mediate the effects of herbivory on associated species, our results suggest that indirect effects of herbivory might influence many components of belowground communities.     

Field evidence for indirect interactions between foliar‐feeding insect and root‐feeding nematode communities on Nicotiana tabacum

Abstract 1. As herbivory often elicits systemic changes in plant traits, indirect interactions via induced plant responses may be a pervasive feature structuring herbivore communities. Although the importance of this phenomenon has been emphasised for herbivorous insects, it is unknown if and how induced responses contribute to the organisation of other major phytoparasitic taxa. 2. Survey and experimental field studies were used to investigate the role of plants in linking the dynamics of foliar‐feeding insects and root‐feeding nematodes on tobacco, Nicotiana tabacum. 3. Plant‐mediated interactions between insects and nematodes could largely be differentiated by insect feeding guild, with positive insect–nematode interactions predominating with leaf‐chewing insects (caterpillars) and negative interactions occurring with sap‐feeding insects (aphids). For example, insect defoliation was positively correlated with the abundance of root‐feeding nematodes, but aphids and nematodes were negatively correlated. Experimental field manipulations of foliar insect and nematode root herbivory also tended to support this outcome. 4. Overall, these results suggest that plants indirectly link the dynamics of divergent consumer taxa in spatially distinct ecosystems. This lends support to the growing perception that plants play a critical role in propagating indirect effects among a diverse assemblage of consumers.