The role of ant-tended extrafloral nectaries in the protection and benefit of a Neotropical rainforest tree

One possible function of extrafloral nectaries is to attract insects, particularly ants, which defend plants from herbivores. We determined whether ants visiting saplings of the tree Stryphnodendronmicrostachyum (Leguminosae) provide protection (decreased plant damage due to ant molestation or killing of herbivores) and benefit (increased plant growth and reproduction associated with ant presence) to the plant. We compared ant and herbivore abundance, herbivore damage and growth of ant-visited plants and ant-excluded plants grown in sun and shade microhabitats of a 6-ha plantation in Costa Rica over a 7-month period. Results show that ants provided protection to plants not by reducing herbivore numbers but by molesting herbivores. Ants also reduced the incidence of pathogen attack on leaves. Protection was greater in the shade than in the sun, probably due to lower herbivore attack in the sun. Protection was also variable within sun and shade habitats, and this variability appeared to be related to variable ant visitation. Results also indicate that ant presence benefits the plant: ant-visited plants grew significantly more in height than ant-excluded plants. The cultivation of ants may serve as an important natural biological control in tropical forestry and agroforestry systems, where increased plant density can otherwise lead to increased herbivore attack. Received: 4 May 1998 / Accepted: 6 October 1998     

Herbivory on Temperate Rainforest Seedlings in Sun and Shade: Resistance,Tolerance and Habitat Distribution

Differential herbivory and/or differential plant resistance or tolerance in sun and shade environments may influence plant distribution along the light gradient. Embothrium coccineum is one of the few light-demanding tree species in the temperate rainforest of southern South America, and seedlings are frequently attacked by insects and snails. Herbivory may contribute to the exclusion of E. coccineum from the shade if 1) herbivory pressure is greater in the shade, which in turn can result from shade plants being less resistant or from habitat preferences of herbivores, and/or 2) consequences of damage are more detrimental in the shade, i.e., shade plants are less tolerant. We tested this in a field study with naturally established seedlings in treefall gaps (sun) and forest understory (shade) in a temperate rainforest of southern Chile. Seedlings growing in the sun sustained nearly 40% more herbivore damage and displayed half of the specific leaf area than those growing in the shade. A palatability test showed that a generalist snail consumed ten times more leaf area when fed on shade leaves compared to sun leaves, i.e., plant resistance was greater in sun-grown seedlings. Herbivore abundance (total biomass) was two-fold greater in treefall gaps compared to the forest understory. Undamaged seedlings survived better and showed a slightly higher growth rate in the sun. Whereas simulated herbivory in the shade decreased seedling survival and growth by 34% and 19%, respectively, damaged and undamaged seedlings showed similar survival and growth in the sun. Leaf tissue lost to herbivores in the shade appears to be too expensive to replace under the limiting light conditions of forest understory. Following evaluations of herbivore abundance and plant resistance and tolerance in contrasting light environments, we have shown how herbivory on a light-demanding tree species may contribute to its exclusion from shade sites. Thus, in the shaded forest understory, where the seedlings of some tree species are close to their physiological tolerance limit, herbivory could play an important role in plant establishment.     

Interactive effects of light environment and herbivory on growth and productivity of an invasive annual vine, Persicaria perfoliata

Plant populations often exist in spatially heterogeneous environments with varying light levels, which can affect plant growth directly through resource availability or indirectly by altering behavior or success of herbivores. The plant vigor hypothesis predicts that herbivores are more likely to attack vigorously growing plants than those that are suppressed, for example in more shaded conditions. Plant tolerance of herbivory can also vary under contrasting resource availability. Observations suggest that damage by Rhinoncomimus latipes Korotyaev (Coleoptera: Curculionidae), introduced into the United States in 2004 as a biological control agent for mile-a-minute weed (Persicaria perfoliata [L.] H. Gross), is greater in the sun than in shade. We compared weevil densities and plant growth in paired plots in full sun or under shade cloth; a second experiment included insecticide-treated plots in sun and shade, to assess the ability of the plant to compensate for herbivore damage. Greater density of weevils and more node damage (indicating internal larval feeding) were found on P. perfoliata plants growing in sun than on those in shade. Nodes were 14% thicker in the sun, which may have provided better larval habitat. Biomass produced by plants without weevils in the sun was about twice that produced in any other treatment. Herbivory had a greater effect on plant growth in the high-light environment than in the shade, apparently because of movement into the sun and increased feeding there by the monophagous herbivore, R. latipes. Results support the plant vigor hypothesis and suggest that high weevil densities in the sunny habitats favored by P. perfoliata can suppress plant growth, negating the resource advantage to plants growing in the sun.     

Herbivore attack in Casearia nitida influenced by plant ontogenetic variation in foliage quality and plant architecture

Traits influencing plant quality as food and/or shelter for herbivores may change during plant ontogeny, and as a consequence, influence the amount of herbivory that plants receive as they develop. In this study, differences in herbivore density and herbivory were evaluated for two ontogenetic stages of the tropical tree Casearia nitida. To assess plant ontogenetic differences in foliage quality as food for herbivores, nutritional and defensive traits were evaluated in saplings and reproductive trees. Predatory arthropods were quantified and the foraging preferences of a parasitoid wasp of the genus Zacremnops were assessed. In addition, survival rates of lepidopteran herbivores (Geometridae) were evaluated experimentally. Herbivore density was three times higher and herbivory was 66% greater in saplings than in reproductive trees. Accordingly, concentrations of total foliar phenolics were higher in reproductive trees than in saplings, whereas leaf toughness, water and nitrogen concentration did not vary between ontogenetic stages. Survival rates of lepidopteran larvae exposed to natural enemies were equivalent in reproductive trees and saplings. Given the greater herbivore density on saplings, equal survival rates implied a greater foraging effort of predators on reproductive trees. Furthermore, observed foraging of parasitoid wasps was restricted to reproductive trees. I propose that herbivore density, and as a consequence, leaf damage were lower in reproductive trees than in saplings due to both traits influencing food quality, and architectural or unmeasured indirect defensive traits influencing foraging preference of natural enemies of herbivores.     

Herbivores modify selection on plant functional traits in a temperate rainforest understory

There is limited evidence regarding the adaptive value of plant functional traits in contrasting light environments. It has been suggested that changes in these traits in response to light availability can increase herbivore susceptibility. We tested the adaptive value of plant functional traits linked with carbon gain in contrasting light environments and also evaluated whether herbivores can modify selection on these traits in each light environment. In a temperate rainforest, we examined phenotypic selection on functional traits in seedlings of the pioneer tree Aristotelia chilensis growing in sun (canopy gap) and shade (forest understory) and subjected to either natural herbivory or herbivore exclusion. We found differential selection on functional traits depending on light environment. In sun, there was positive directional selection on photosynthetic rate and relative growth rate (RGR), indicating that selection favors competitive ability in a high-resource environment. Seedlings with high specific leaf area (SLA) and intermediate RGR were selected in shade, suggesting that light capture and conservative resource use are favored in the understory. Herbivores reduced the strength of positive directional selection acting on SLA in shade. We provide the first demonstration that natural herbivory rates can change the strength of selection on plant ecophysiological traits, that is, attributes whose main function is resource uptake. Research addressing the evolution of shade tolerance should incorporate the selective role of herbivores.     

Population variation in plant traits associated with ant attraction and herbivory in Chamaecrista fasciculata (Fabaceae)

The benefits of ant–plant–herbivore interactions for the plant depend on the abundance of ants and herbivores and the selective pressures these arthropods exert. In plants bearing extrafloral nectaries (EFN), different mean trait values may be selected for by different populations in response to local herbivore pressure, ultimately resulting in the evolution of differences in plant traits that attract ants as defensive agents against herbivory. To determine if variation in traits that mediate ant–plant interactions reflect herbivore selective pressures, we quantified intra- and inter-population variation in plant traits for eight populations of the EFN-bearing annual Chamaecrista fasciculata (Michx.) (Fabaceae). Censuses in rural and urban areas of Missouri and Illinois (USA) showed population differences in ant attendance and herbivore pressure. Seeds were collected from each population, and plants were grown in a common greenhouse environment to measure sugar production, nectar volume and composition, EFN size and time of emergence, leaf pubescence, and leaf quality throughout plant development. Populations varied mainly in terms of nectary size, sugar production, and nectar volume, but to a lesser degree in leaf pubescence. Populations of C. fasciculata within urban areas (low in insect abundance) had small nectaries and the lowest nectar production. There was a positive correlation across populations between herbivore density and leaf damage by those herbivores on the one hand and sugar production and nectar volume on the other. These results, in conjunction with lack of evidence for maternally based environmental effects, suggest that population differences in herbivore damage have promoted differential evolution of EFN-related traits among populations. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Interactions between willows and insect herbivores under enhanced ultraviolet-B radiation

We studied the effects of elevated ultraviolet-B radiation on interactions between insect herbivores and their host plants by exposing two species of phytochemically different willows, Salix myrsinifolia and S. phylicifolia, to a modulated increase in ultraviolet radiation in an outdoor experiment and monitoring the colonisation of insect herbivores on these willows. We examined the effect of increased ultraviolet-B (UV-B) radiation on (1) the quality of willow leaves, (2) the distribution and abundance of insect herbivores feeding on these willows, (3) the resulting amount of damage, and (4) the performance of insect larvae feeding on the exposed plant tissue. Six clones of each of the two willow species were grown in eight blocks for 12 weeks in the UV-B irradiation field. The clones were exposed to a constant 50% increase in UV-B radiation (simulating 20-25% ozone depletion), to a small increase in UV-A radiation or to ambient solar irradiation. We allowed colonisation on the willows by naturally occurring insects, but also introduced adults of a leaf beetle, Phratora vitellinae, a specialist herbivore on S. myrsinifolia. Increased UV-B radiation did not affect any of the measured indices of plant quality. However, numbers of P. vitellinae on S. myrsinifolia were higher in plants with UV-B treatment compared with UV-A and shade controls. In laboratory tests, growth of the second-instar larva of P. vitellinae was not affected by UV-B treatment of S. myrsinifolia, but was retarded on UV-B treated leaves of S. phylicifolia. In addition, naturally occurring insect herbivores were more abundant on willows exposed to elevated UV-B radiation compared to those grown under control treatments. In spite of the increased abundance of insect herbivores, willows treated with elevated UV-B did not suffer more herbivore damage than willows exposed to ambient solar radiation (shade control). The observed effects of UV-B on herbivore abundance, feeding and growth varied significantly due to spatial variation in environment quality, as indicated by the UV-treatment x block interaction. The results suggest that (1) environmental variation modifies the effects of UV-B radiation on plant-insect interactions and (2) specialist herbivores might be more sensitive to chemical changes in their secondary host plants (S. phylicifolia) than to changes in their primary hosts (S. myrsinifolia).     

Influence of previous frost damage on tree growth and insect herbivory of Eucalyptus globulus globulus

The plant stress hypothesis suggests that some herbivores favour stressed plants, whereas the plant vigour hypothesis proposes that other herbivores prefer vigorous plants. The effects of a prior stress, that of frost damage, were examined on the subsequent growth of Eucalyptus globulus globulus and on the response of insect herbivores. Frost damage affected tree growth by reducing new leaf area and increasing specific leaf area (SLA). However, herbivore abundance was not affected by prior frost damage. Two feeding trials using Anoplognathus chloropyrus and Hyalarcta huebneri and a morphometric study of Ctenarytaina eucalypti were conducted to assess the performance of herbivores on trees that had suffered more or less frost damage. Consumption by A. chloropyrus and H. huebneri was unaffected by foliage origin (damaged versus healthy). Hyalarcta huebneri grew faster when fed leaves from previously damaged trees, and C. eucalypti from previously damaged trees were larger than those from healthy trees. Enhanced insect performance on frost damaged plants may have resulted from the high specific leaf area (most likely thinner) leaves. The herbivore abundance data did not support the hypothesis that previously frost damaged plants are preferred by insects. However, increased growth of H. huebneri and larger body size of C. eucalypti on damaged trees indicates that previously stressed trees may produce leaves of higher nutritional value.     

The effect of disturbance on an ant–plant mutualism

Protective ant–plant mutualisms—where plants provide food or shelter to ants and ants protect the plants from herbivores—are a common feature in many ecological communities, but few studies have examined the effect of disturbance on these interactions. Disturbance may affect the relationship between plants and their associated ant mutualists by increasing the plants’ susceptibility to herbivores, changing the amount of reward provided for the ants, and altering the abundance of ants and other predators. Pruning was used to simulate the damage to buttonwood mangrove (Conocarpus erectus) caused by hurricanes. Pruned plants grew faster than unpruned plants, produced lower levels of physical anti-herbivore defenses (trichomes, toughness), and higher levels of chemical defenses (tannins) and extrafloral nectaries. Thus, simulated hurricane damage increased plant growth and the amount of reward provided to ant mutualists, but did not have consistent effects on other anti-herbivore defenses. Both herbivores and ants increased in abundance on pruned plants, indicating that the effects of simulated hurricane damage on plant traits were propagated to higher trophic levels. Ant-exclusion led to higher leaf damage on both pruned and upruned plants. The effect of ant-exclusion did not differ between pruned and unpruned plants, despite the fact that pruned plants had higher ant and herbivore densities, produced more extrafloral nectaries, and had fewer physical defenses. Another common predator, clubionid spiders, increased in abundance on pruned plants from which ants had been excluded. I suggest that compensatory predation by these spiders diminished the effect of ant-exclusion on pruned plants.     

Herbivore-induced responses and patch heterogeneity affect abundance of arthropods on plants

Abstract.  1. Plants respond to herbivore damage by inducing defences that can affect the abundance of herbivores and predators. These tritrophic interactions may be influenced by heterogeneity in plant neighbourhood.
2. In the present study, the effects of induced responses on the abundance of herbivores (flea beetles and aphids), omnivores (pirate bugs and thrips), and predators (lady beetles and spiders) on individual plants and their neighbours between and within patches composed of three tomato plants was investigated.
3. Herbivore damage was manipulated to create homogeneous patches where either all or none of the plants had defences induced by herbivore damage, and heterogeneous patches where only one of the plants was induced.
4. Arthropod abundance on plants at different scales was compared by testing between patch effects (patch level), for neighbourhood effects at the plant phenotype level (neighbourhood level), and between near and far plants (within patch position).
5. At the patch level , plants in homogeneously induced patches contained fewer flea beetles and pirate bugs, but more lady beetles, compared with homogeneously non-induced patches. There was no effect of patch type on the abundance of aphids, thrips, and spiders on plants.
6. At the neighbourhood level , induced plants in heterogeneous patches contained more flea beetles and pirate bugs compared with induced plants in homogeneous patches, indicating that the abundance of some herbivores and omnivores on induced plants varied depending on the phenotype of the other plants within the patch. Within patch position, there was no evidence that the abundance of herbivores or predators on non-induced plants was affected by proximity to an induced plant.
7. Therefore, variation in plant neighbourhood generated by induced plant responses affected the abundance of three arthropods from three feeding guilds.     

Contrasting cascades: insectivorous birds increase pine but not parasitic mistletoe growth

1. Intraguild predation occurs when top predators feed upon both intermediate predators and herbivores. Intraguild predators may thus have little net impact on herbivore abundance. Variation among communities in the strength of trophic cascades (the indirect effects of predators on plants) may be due to differing frequencies of intraguild predation. Less is known about the influence of variation within communities in predator-predator interactions upon trophic cascade strength. 2. We compared the effects of a single predator community between two sympatric plants and two herbivore guilds. We excluded insectivorous birds with cages from ponderosa pine Pinus ponderosa trees parasitized by dwarf mistletoe Arceuthobium vaginatum. For 3 years we monitored caged and control trees for predatory arthropods that moved between the two plants, foliage-feeding caterpillars and sap-feeding hemipterans that were host-specific, and plant damage and growth. 3. Excluding birds increased the abundance of ant-tended aphids on pine and resulted in an 11% reduction in pine woody growth. Mutualist ants protected pine-feeding aphids from predatory arthropods, allowing aphid populations to burgeon in cages even though predatory arthropods also increased in cages. By protecting pine-feeding aphids from predatory arthropods but not birds, mutualist ants created a three-tiered linear food chain where bird effects cascaded to pine growth via aphids. 4. In contrast to the results for tended aphids on pine, bird exclusion had no net effects on untended pine herbivores, the proportion of pine foliage damaged by pine-feeding caterpillars, or the proportion of mistletoe plants damaged by mistletoe-feeding caterpillars. These results suggest that arthropod predators, which were more abundant in cages as compared with control trees, compensated for bird predation of untended pine and mistletoe herbivores. 5. These contrasting effects of bird exclusion support food web theory: where birds were connected to pine by a linear food chain, a trophic cascade occurred. Where birds fed as intraguild predators, the reticulate food webs linking birds to pine and mistletoe resulted in no net effects on herbivores or plant biomass. Our study shows that this variation in food web structure occurred between sympatric plants and within plants between differing herbivore guilds.     

Influence of plant genotype and environment on oviposition preference and offspring survival in a gallmaking herbivore

Summary Plant resistance to insect herbivores may derive from traits influencing herbivore preference, traits influencing the suitability of the plant as a host, or both. However, the plant traits influencing host-plant selection by ovipositing insect herbivores may not completely overlap those traits that affect larval survival, and distinct traits may exhibit different levels of genetic vs. environmental control. Therefore, resource supply to the host plant could affect oviposition preference and larval performance differently in different plant genotypes. To test this hypothesis, the effects of resistance level, plant genotype, and resource supply to the host plant on oviposition preference and larval performance of a gallmaking herbivore, and on various plant traits that could influence these, were examined. Replicates of four genotypes of Solidago altissima, grown under low, medium, or high levels of nutrient supply in full sun or with medium levels of nutrients in shade, were exposed to mass-released Eurosta solidaginis. The number of plants ovipunctured was significantly affected by plant genotype and the interaction between genotype and nutrient supply to the host plant: one susceptible and one resistant genotype were more preferred, and preference tended to increase with nutrient supply in the more-preferred genotypes. The growth rate of ovipunctured plants during the oviposition period was significantly greater than that of unpunctured plants. Bud diameter (which was strongly correlated with plant growth rate), leaf area, and leaf water content were significant determinants of the percentage of plants ovipunctured, explaining 74% of the variance. The number of surviving larvae was significantly affected by plant genotype, but no effect of nutrient or light supply to the host plant was detected. The ratio of bud diameter to bud length was positively related to the percentage of ovipunctured plants that formed galls, suggesting that the accurate placement of eggs near the apical meristem by ovipositing females may be easier in short, thick buds. No significant correlation was observed between oviposition preference and larval survival at the population level. These results suggest that the plant traits affecting oviposition preference may exhibit different magnitudes of phenotypic plasticity than those affecting larval survival, and that the degree of phenotypic plasticity in plant traits affecting oviposition preference may differ among genotypes within a species.     

Plant quality and predation risk mediated by plant ontogeny: consequences for herbivores and plants

Bottom‐up and top‐down impacts on herbivores can be influenced by plant productivity, structural complexity, vigor and size. Although these traits are likely to vary with plant development, the influence of plant ontogeny on the relative importance of plant quality (i.e. bottom‐up forces) and predation risk (i.e. top‐down forces) has been the focus of little previous investigation. We evaluated the role of plant ontogeny for the relative importance of bottom‐up and top‐down forces on insect herbivore abundance, species richness, and species diversity attacking the tropical tree Casearia nitida. We also quantified the cascading effects on herbivory, growth and reproduction of this plant species. Plant quality traits (nitrogen and phenolic compounds) were assessed in saplings and reproductive trees. Bottom‐up forces were manipulated by fertilizing plants from both ontogenetic stages. Top‐down forces were manipulated by excluding insectivorous birds from saplings and reproductive trees. Plant ontogeny influenced foliage quality in terms of total phenolics, which were in greater concentration in reproductive trees than in saplings; however, it did not influence bottom‐up forces as modified by fertilization. Bird exclusion increased herbivore density with the same magnitude on both stages. Ontogeny influenced species diversity, which was greater in reproductive trees than in saplings, and also influenced treatment impacts on species richness and diversity. Although top‐down forces increased herbivory equally on plants of each ontogenetic stage, the two stages showed different overcompensation responses to increased damage: caged saplings produced greater leaf biomass than non‐caged saplings, whereas caged trees increased in height proportionally more than non‐caged trees. In sum, plant ontogeny influenced the impact of bird predation on herbivore density, species richness, and species diversity, and the growth variables affected by increased damage in caged plants. We suggest that plant ontogeny can contribute to some extent to the influence of plant quality and the third trophic level on herbivores in this system.     

Delayed induced responses of birch glandular trichomes and leaf surface lipophilic compounds to mechanical defoliation and simulated winter browsing

Changes in morphology and chemistry of leaf surface in response to herbivore damage may increase plant resistance to subsequent herbivore attack; however, there is lack of studies on induced responses of glandular trichomes and their exudates in woody plants and on effects of these changes on herbivores. We studied delayed induced responses in leaf surface traits of five clones of silver birch (Betula pendula Roth) subjected to various types of mechanical defoliation and simulated winter browsing. Glandular trichome density and concentrations of the majority of surface lipophilic compounds increased in trees defoliated during the previous summer. This induced response was systemic, since control branches in branch defoliated trees responded to the treatments similarly to defoliated branches, but differently from control trees. In contrast to defoliation treatments, simulated winter browsing reduced glandular trichome density on the following summer and had fewer effects on individual surface lipophilic compounds. Moreover, constitutive density of glandular trichomes was negatively correlated with induced total amount of lipophilic compounds per trichome, indicating a trade-off between constitutive and induced resistance in silver birch. Induced changes in leaf surface traits had no significant effect on leaf damage by chewers, miners and gall mites, but increased susceptibility of birch trees to aphids. However, leaf damage by chewers, miners and gall mites in defoliated (but not in control) trees was correlated with concentrations of some fatty acids and triterpenoids, although the direction of relationships varied among herbivore species. This indicates that induction of surface lipophilic compounds may influence birch resistance to herbivores. Our study thus demonstrated both specificity of elicitation of induced responses of birch leaf surface traits by different types of damage and specificity of the effects of these responses on different types of herbivores.Electronic Supplementary Material Supplementary material is available in the online version of this article at and is accessible for authorized users.     

Clumped distribution of oak leaf miners between and within plants

Leaf miners typically show non-random distributions both between and within plants. We tested the hypothesis that leaf miners on two oak species were clumped on individual host trees and individual branches and addressed whether clumping was influenced by aspects of plant quality and how clumping and/or interactions with other oak herbivores affected leaf-miner survivorship. Null models were used to test whether oak herbivores and different herbivore guilds co-occur at the plant scale. Twenty individual Quercus geminata plants and 20 Quercus laevis plants were followed over the season for the appearance of leaf miners and other herbivores, and foliar nitrogen, tannin concentration, leaf toughness and leaf water content were evaluated monthly for each individual tree. The survivorship of the most common leaf miners was evaluated by following the fate of marked mines in several combinations that involved intra- and inter-specific associations. We observed that all leaf miners studied were clumped at the plant and branch scale, and the abundance of most leaf-miner species was influenced by plant quality traits. Mines that occurred singly on leaves exhibited significantly higher survivorship than double and triple mines and leaves that contained a mine or a leaf gall and a mine and damage by chewers exhibited lowest survivorship. Although leaf miners were clumped at individual host trees, null model analyses indicated that oak herbivores do not co-occur significantly less than expected by chance and there was no evidence for biological mechanisms such as inter-specific competition determining community structure at the plant scale. Thus, despite co-occurrence resulting in reduced survivorship at the leaf scale, such competition was not strong enough to structure separation of these oak herbivore communities.     

Attracting pollinators and avoiding herbivores: insects influence plant traits within and across years

Perennial plants interact with herbivores and pollinators across multiple growing seasons, and thus may respond to herbivores and pollinators both within and across years. Joint effects of herbivores and pollinators influence plant traits, but while some of the potential interactions among herbivory, pollination, plant size, and plant reproductive traits have been well studied, others are poorly understood. This is particularly true for perennial plants where effects of herbivores and pollinators may manifest across years. Here, we describe two experiments addressing the reciprocal interactions of plant traits with herbivore damage and pollination across 2 years using the perennial plant Chamerion angustifolium. We measured (1) plant responses to manipulation of damage and pollination in the year of treatment and the subsequent season, (2) damage and pollination responses to manipulation of plant size and flowering traits in the year of treatment, and (3) plant-mediated indirect interactions between herbivores and pollinators. We found that plant traits had little effect on damage and pollination, but damage and pollination affected plant traits in both the treatment year and the subsequent year. We found evidence of indirect effects between leaf herbivores and pollinators in both directions; indirect effects of pollinators on leaf herbivores have not been previously demonstrated. Our results indicate that pollen receipt results in shorter plants with fewer stems but does not change flower number, while leaf herbivory results in taller plants with fewer flowers. Together, herbivory and pollination may contribute to intermediate plant height and plants with fewer stems and flowers in our system.     

Winter predation by insectivorous birds and consequences for arthropods and plants in summer

Top-down effects of predators can have important consequences for ecosystems. Insectivorous birds frequently have strong predation effects on herbivores and other arthropods, as well as indirect effects on herbivores’ host plants. Diet studies have shown that birds in temperate ecosystems consume arthropods in winter as well as in summer, but experimental studies of bird predation effects have not attempted to quantitatively separate winter predation impacts from those in summer. To understand if winter foraging by insectivorous birds has consequences for arthropods or plants, we performed a meta-analysis of published bird exclusion studies in temperate forest and shrubland habitats. We categorized 85 studies from 41 publications by whether birds were excluded year-round or only in summer, and analyzed arthropod and plant response variables. We also performed a manipulative field experiment in which we used a factorial design to exclude birds from Quercus velutina Lam. saplings in winter and summer, and censused arthropods and herbivore damage in the following growing season. In the meta-analysis, birds had stronger negative effects on herbivores in studies that included winter exclusion, and this effect was not due to study duration. However, this greater predation effect did not translate to a greater impact on plant damage or growth. In the field experiment, winter exclusion did not influence herbivore abundance or their impacts on plants. We have shown that winter feeding by temperate insectivorous birds can have important consequences for insect herbivore populations, but the strength of these effects may vary considerably among ecosystems. A full understanding of the ecological roles of insectivorous birds will require explicit consideration of their foraging in the non-growing season, and we make recommendations for how future studies can address this.     

Leaf damage and functional traits along a successional gradient in Brazilian tropical dry forests

Herbivory has significant impacts on individual plants and plant communities, both at ecological and evolutionary time scales. In this context, this study aims to evaluate herbivore damage and its relationship with leaf chemical and structural traits, nutritional status, and forest structural complexity along a successional gradient. We predicted that trees in early successional stages support conservative traits related to drought tolerance (high specific leaf mass and phenolics), whereas trees in light-limited, late successional stages tend to enhance light acquisition strategies (high nitrogen content). We sampled 261 trees from 26 species in 15 plots (50 × 20 m; five per successional stage). From each tree, twenty leaves were collected for leaf trait measures. Phenolic content increased whereas specific leaf mass and nitrogen content decreased from early to late stages. However, leaf damage did not differ among successional stages. Our results partially corroborate the hypothesis that early successional plants in tropical dry forests exhibit leaf traits involved in the conservative use of water. The unexpected decrease in nitrogen content along the chronosequence is likely related to the fact that thinner leaves with low specific leaf mass could have less nitrogen-containing mesophyll per unit area. Mechanisms affecting herbivory intensity varied across scales: at the species level, leaf damage was negatively correlated with tannin concentration and specific leaf mass; at the plot level, leaf damage was positively affected by forest structural complexity. Herbivory patterns in tropical forests are difficult to detect because abiotic factors and multiple top-down and bottom-up forces directly and indirectly affect herbivores.     

Differential induced response to generalist and specialist herbivores by Lindera benzoin (Lauraceae) in sun and shade

Theoretically, induced defenses should be prevalent within low resource environments like the forest understory where constitutive defenses would be costly. Also, the induced response should be stronger when the herbivore is a generalist rather than a specialist, which often have mechanisms to avoid or overcome plant defenses. These ideas have been previously tested for herbaceous species, and we examined these predictions in Lindera benzoin (spicebush), a common woody shrub of the eastern deciduous forest. Lindera benzoin plants in contrasting light environments served as control plants or were subjected to one of four treatments: application of jasmonic acid, clipping, herbivory by the specialist Epimecis hortaria (tulip tree beauty) and herbivory by the generalist Spodoptera exigua (beet armyworm). Following treatment, we assessed induced responses by measuring leaf chemistry (C/N ratio, protein content, and peroxidase activity), and by using insect bioassays with E. hortaria larvae. We found no difference in peroxidase activity between light environments in controls, plants treated with clipping or jasmonic acid. In plants subject to insect herbivory, peroxidase activity was greater in shade plants than in sun plants. The magnitude of this increase in the shade varied between the herbivores, with a 32 fold increase in plants exposed to the generalist S. exigua and a 9 fold increase in plants exposed to the specialist E. hortaria . Leaves from shade plants had more protein and lower C/N ratios than leaves from sun plants, regardless of induction treatment. In control plants, E. hortaria larvae consumed more leaf biomass and achieved greater final weights in the sun than in the shade, but these differences disappeared with induction treatments were applied. These results are among the first to show rapid induction in a woody plant, and different levels of induction with light environments and with specialist versus generalist herbivores.     

Plant neighbours mediate bird predation effects on arthropod abundance and herbivory

相似文献