Potential impacts of climate change on patterns of insect herbivory on understorey plant species: A transplant experiment

Species‐specific responses to climate change will lead to changes in species interactions across multiple trophic levels. Interactions between plants and their insect herbivores, in particular, may become increasingly disrupted if mobile herbivores respond more rapidly to climatic change than their associated host plants. We present a multispecies transplant experiment aimed at assessing potential climatic impacts on patterns of leaf herbivory. Four shrubby understorey plant species were transplanted outside their native range into a climate 2.5°C warmer in annual mean temperature. After 12 months, we assessed the types and amount of herbivore leaf damage, compared with plants transplanted to a control site within their native range. The overall amount of foliage loss to herbivores ranged from approximately 3–10% across species and sites, a range consistent with most estimates of leaf loss in other studies. The most common types of leaf damage were sucking and chewing and this pattern was consistent for all four plant species at all sites. There were no significant differences in levels and patterns of herbivory between control and warm sites for three out of four plant species. This suggests that with moderate climate warming, most herbivory will continue to be dominated by chewers and suckers, and that the overall level of foliage loss will be similar to that experienced presently.     

Mechanisms for locating resources in space and time: Impacts on the abundance of insect herbivores

Herbivorous insects have the problem both of locating appropriate host plants and ensuring that the plant‐feeding stages of their life cycles are synchronized with the times when those hosts provide a high‐quality food resource. Because the taxonomic range of potential hosts is generally narrow, and the temporal window when those hosts are suitable is often relatively short, developmental (especially diapause) and dispersal mechanisms may be critical factors in determining whether or not a species population is successful in a particular plant community. The present paper considers the impact of diapause and dispersal mechanisms on the ability of insect herbivores to cope with two attributes of their host plants: (i) the diversity of the plant community within which the hosts are located; and (ii) the seasonal predictability of host suitability. Some common dispersal mechanisms used by insect herbivores are much more appropriate to low‐diversity than to high‐diversity plant communities and, similarly, some diapause cues are appropriate only to highly predictable plant phenology. Both agriculture and silviculture characteristically manipulate both these attributes of plant communities, that is, in order to make the human use of plants more efficient, cultivation strategies normally both reduce plant species diversity (often to a condition approaching monoculture) and increase the predictability of plant developmental patterns. Consequently, major pest species in managed systems may not be those that are most common in natural systems, and may be difficult to predict in advance.     

Silica in grasses as a defence against insect herbivores: contrasting effects on folivores and a phloem feeder

1. Silica, deposited as opaline phytoliths in the leaves of grasses, constitutes 2-5% of dry leaf mass, yet its function remains unclear. It has been proposed that silica may act as an antiherbivore defence by increasing the abrasiveness and reducing the digestibility of grass leaves, although there is little direct experimental evidence to support this. 2. We investigated the effects of manipulated silica levels on the abrasiveness of the leaves of five grass species. We also examined the effects of silica levels on the feeding preferences, growth performance and digestion efficiency of two folivorous insects and one phloem-feeding insect. 3. Silica addition resulted in increases to leaf abrasiveness in four of the five grass species studied. Silica addition also deterred feeding by both folivores and reduced their growth rates and digestion efficiency. 4. These effects resulted in lower pupal mass of the lepidopteron larvae Spodoptera exempta and compensatory feeding by the orthopteran, Schistocerca gregaria. In contrast, silica had no effects on the feeding preference or the population growth of the phloem feeder, Sitobion avenae. 5. Our results demonstrate that silica is an effective defence against folivorous insects, both as a feeding deterrent, possibly mediated by increased abrasiveness, and as a digestibility reducer. The effects of silica on pupal mass and development time may impact on herbivore fitness and exposure to natural enemies. 6. These results are the first demonstration of a direct effect of silica on the abrasiveness of grasses and the adverse impact of silica on herbivore preference and performance.     

Compound effects of induced plant responses on insect herbivores and parasitoids: implications for tritrophic interactions

1. Induced plant responses can affect herbivores either directly, by reducing herbivore development, or indirectly, by affecting the performance of natural enemies. Both the direct and indirect impacts of induction on herbivore and parasitoid success were evaluated in a common experimental system, using clonal poplar trees Populus nigra (Salicales: Salicaceae), the gypsy moth Lymantria dispar (L.) (Lepidoptera: Lymantriidae), and the gregarious parasitoid Glyptapanteles flavicoxis (Marsh) (Hymenoptera: Braconidae). 2. Female parasitoids were attracted to leaf odours from both damaged and undamaged trees, however herbivore‐damaged leaves were three times more attractive to wasps than undamaged leaves. Parasitoids were also attracted to herbivore larvae reared on foliage and to larval frass, but they were not attracted to larvae reared on artificial diet. 3. Prior gypsy moth feeding elicited a systemic plant response that retarded the growth rate, feeding, and survival of gypsy moth larvae, however induction also reduced the developmental success of the parasitoid. 4. The mean number of parasitoid progeny emerging from hosts fed foliage from induced trees was 40% less than from uninduced trees. In addition, the proportion of parasitised larvae that survived long enough to issue any parasitoids was lower on foliage from induced trees. 5. A conceptual and analytical model is provided to describe the net impacts of induced plant responses on parasitoids, and implications for tritrophic interactions and biological control of insect pests are discussed.     

Impacts of urbanization on insect herbivory and plant defences in oak trees

Systematic comparisons of species interactions in urban versus rural environments can improve our understanding of shifts in ecological processes due to urbanization. However, such studies are relatively uncommon and the mechanisms driving urbanization effects on species interactions (e.g. between plants and insect herbivores) remain elusive. Here we investigated the effects of urbanization on leaf herbivory by insect chewers and miners associated with the English oak Quercus robur by sampling trees in rural and urban areas throughout most of the latitudinal distribution of this species. In performing these comparisons, we also controlled for the size of the urban areas (18 cities) and gathered data on CO₂ emissions. In addition, we assessed whether urbanization affected leaf chemical defences (phenolic compounds) and nutritional traits (phosphorus and nitrogen), and whether such changes correlated with herbivory levels. Urbanization significantly reduced leaf chewer damage but did not affect leaf miners. In addition, we found that leaves from urban locations had lower levels of chemical defences (condensed and hydrolysable tannins) and higher levels of nutrients (nitrogen and phosphorus) compared to leaves in rural locations. The magnitude of urbanization effects on herbivory and leaf defences was not contingent upon city size. Importantly, while the effects of urbanization on chemical defences were associated with CO₂ emissions, changes in leaf chewer damage were not associated with either leaf traits or CO₂ levels. These results suggest that effects of urbanization on herbivory occur through mechanisms other than changes in the plant traits measured here. Overall, our simultaneous assessment of insect herbivory, plant traits and abiotic correlates advances our understanding of the main drivers of urbanization effects on plant–herbivore interactions.     

Contrasting effects of insect and molluscan herbivores on plant diversity in a long-term field experiment

The importance of invertebrate herbivores in regulating plant communities remains unclear, due to the absence of long-term exclusion experiments. An experiment in an English grassland involving long-term exclusions of insect and mollusc herbivores, along with rabbit fencing, showed strong, but opposing, effects of the invertebrate herbivores. Plant species richness declined and biomass increased following insect exclusion, due to increased dominance by a grass species, whereas mollusc exclusion led to increased herbs abundance. The two herbivores had a compensatory interaction: molluscs had no effects in the absence of insects and large insect effects depended on the absence of molluscs. The effects of invertebrate exclusion became apparent only after 8 years, and would have been seriously underestimated in shorter studies. Our results suggest that theorists and conservation managers need to shift from their historic focus on vertebrate herbivory, to a recognition that invertebrates can be equally important drivers of plant community structure.     

Fire effects on insect herbivores in an oak savanna: the role of light and nutrients

Abstract.  1. Environmental heterogeneity created by prescribed burning provided the context for testing whether the distribution of an oak specialist (the lace bug, Corythuca arcuata ) could be explained by stoichiometric mismatches between herbivore and host plant composition.
2. Field observations showed that lace bug density was seven-fold higher in frequently burned than in unburned units.
3. Lace bug density did not increase with leaf nutrient concentrations, but was instead associated with higher light levels, higher concentrations of leaf carbon (C), lignin and total phenolics, and lower levels of cellulose. In addition, lace bugs reared on high-light leaves had higher levels of survivorship than those fed on low-light leaves.
4. Sampling restricted to full-sun leaves was used to test whether fire-related changes in leaf nitrogen (N) and phosphorus (P) concentrations have a secondary influence on lace bug success. This sampling provided only limited evidence for nutrient limitation, as decreases in leaf N and P were associated with an increase in lace bug mass but a decrease in density.
5. It is concluded that burning probably promotes lace bug population growth by increasing canopy openness, light penetration, and the availability of C-based metabolites, and thus simple stoichoimetric mismatches between herbivores and host plants are not of primary importance in this system.     

Context dependency of insect and mammalian herbivore effects on tall thistle (Cirsium altissimum) populations

Aims Identifying factors that drive variation in herbivore effects on plant populations can provide insight for explaining plant distributions and for limiting weeds. Abiotic resource availability to plants is a key explanation for variation in herbivore effects on individual plants, but the role of resources in determining herbivore effects on plant populations is largely unexplored. We tested the hypothesis that soil nutrient availability drives variation in insect and mammal herbivore effects on tall thistle (Cirsium altissimum) population growth.     

Responses of insect herbivores and herbivory to habitat fragmentation: a hierarchical meta‐analysis

Loss and fragmentation of natural habitats can lead to alterations of plant–animal interactions and ecosystems functioning. Insect herbivory, an important antagonistic interaction is expected to be influenced by habitat fragmentation through direct negative effects on herbivore community richness and indirect positive effects due to losses of natural enemies. Plant community changes with habitat fragmentation added to the indirect effects but with little predictable impact. Here, we evaluated habitat fragmentation effects on both herbivory and herbivore diversity, using novel hierarchical meta‐analyses. Across 89 studies, we found a negative effect of habitat fragmentation on abundance and species richness of herbivores, but only a non‐significant trend on herbivory. Reduced area and increased isolation of remaining fragments yielded the strongest effect on abundance and species richness, while specialist herbivores were the most vulnerable to habitat fragmentation. These fragmentation effects were more pronounced in studies with large spatial extent. The strong reduction in herbivore diversity, but not herbivory, indicates how important common generalist species can be in maintaining herbivory as a major ecosystem process.     

Ultrastructure and the importance of wear in the dentition of the halfbeak (Pisces: Hemiramphidae) pharyngeal mill

To assess how tooth microstructure and composition might facilitate the pharyngeal mill mechanism of halfbeaks, apatite structure and iron content were determined by scanning electron microscopy and energy dispersive X‐ray analysis for Hyporhamphus regularis ardelio, Arrhamphus sclerolepis krefftii, and Hemiramphus robustus. Iron was present in developing teeth and was concentrated along the shearing edge of spatulate incisiform teeth, which dominate the occlusive wear zone in all three species. A model based on tooth structure and wear rate is proposed to explain how halfbeaks maintain a fully functional occlusion zone throughout growth and consequent tooth addition and replacement. Replacement teeth erupt and wear rapidly so that a constant occlusion plane is always present. Iron within the tooth tissue reduces the wear rate of the cutting edge while simultaneously maintaining its sharpness and efficiency. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

The potential interplay of posture, digestive anatomy, density of ingesta and gravity in mammalian herbivores: why sloths do not rest upside down

1. The interaction between the density of ingesta and gravity observed in the digestive systems of ruminant herbivores should receive attention in other non‐ruminant herbivorous mammals. The resting postures adopted by non‐ruminants are of particular interest. 2. A new interpretation of established findings regarding the digestive tract of sloths illustrates that the interplay of posture, anatomy, the density of ingesta and gravity can provide a novel explanation of behavioural and morphological adaptations in herbivores, as the average particle size and dry‐matter content increases within their forestomach from its caudal towards its cranial portion. In sloths, this could be indicative of a stratification of ingesta occurring in the upright sitting posture adopted while resting, as opposed to their characteristic upside down posture when moving. 3. The sitting resting posture of sloths could therefore be an adaptation to exploit the tendency of the forestomach contents to stratify in order to pass larger, more difficult‐to‐digest particles faster from the fermentation chamber.     

Herbivory and the evolution of divaricate plants: Structural defences lost on an offshore island

Many island plants are characterized by unique morphology. For example, the high branching angles and small leaves of divaricate plants are a common feature of the New Zealand flora. The divaricate growth form may be an adaptation to deter browsing by extinct avian herbivores (moa); alternatively aspects of the insular climate may be responsible. However, our understanding of the selective pressures responsible for the high branching angles and small leaves of divaricate plants is incomplete. Here, I tested for differences in traits associated with the divaricate growth form between plants from Chatham Island and the New Zealand mainland. Moa never reached the Chatham Islands and its flora is derived from plants on mainland New Zealand. Therefore, I predicted Chatham Island plants to have lost morphological adaptations that may have deterred moa herbivory. Traits were quantified on 316 individuals in the field, allowing for 12 island‐mainland taxonomic comparisons. Chatham Island plants consistently produced smaller branching angles, larger leaves, shorter internodes and larger stems than related mainland plants. Results are therefore consistent with the hypothesis that selection for small leaves and high angled branching may be relaxed on the Chatham Islands due to an absence of moa. Smaller branching angles and larger leaves may offer a competitive advantage to Chatham Island plants.     

Effects of insect herbivory on induced chemical defences and compensation during early plant development in Penstemon virgatus

Background and Aims

The lack of studies assessing the simultaneous expression of tolerance and resistance traits during seedling development and overall seedling defences as compared with adult plants, in general, constitutes a significant research need that can greatly improve our understanding of overall investment in defences during plant ontogeny.

Methods

Using two seedling and two juvenile stages of the perennial herb Penstemon virgatus (Plantaginaceae) evaluations were made of (a) patterns of investment in constitutive chemical defences [i.e. iridoid glycosides (IGs)], and (b) simultaneous variation in the short-term ability of seedling and juvenile stages to induce resistance traits, measured as induced chemical defences, or tolerance traits, measured as compensatory re-growth following moderate levels of damage by a specialist insect herbivore.

Key Results

Plants were highly defended during most of their transition from seedling to early juvenile stages, reaching a constant approx. 20 % dry weight total IGs. Furthermore, following 30 % above-ground tissue damage, seedlings and juvenile stages were equally able to induce resistance, by raising their IG concentration by approx. 8 %, whereas compensatory re-growth was only achieved at young juvenile but not seedling stages.

Conclusions

Two major trends emerged from this study: (1) in contrast to expected and previously observed trends, in this perennial plant species, seedlings seem to be one of the most well-defended stages as compared with adult ones; (2) high levels of constitutive defences did not limit the ability of young developmental stages to induce resistance following damage, although this response may come with a cost (i.e. decreased compensation) in young seedling stages. Hence, as has been previously demonstrated in few other systems, these results points towards an indirect evidence for a trade-off between tolerance and resistance traits at some, but not all, developmental stages; making them often difficult to detect.     

Nutrient composition of soil and plants may predict the distribution and abundance of specialist insect herbivores: implications for agent selection in weed biological control

Abstract  Refining the process of selecting specialist herbivores that are used as biological control agents to maximise effects on the targets can diminish the risk to non-target species. While biotic factors (e.g. plant demography and agent–host interactions) have been explored for clues to improve the way we make agent selection decisions, recent ecological research indicates that abiotic factors related to the habitat (e.g. plant and soil nutrient composition, and soil characteristics) are important predictors of insect herbivore community composition. In this paper we explore the relevance of plant nutrient composition to aid in selecting agents for the invasive Eurasian perennial Euphorbia esula (leafy spurge) in Illinois, USA. We propose that an approach that compares such abiotic factors across the native and invaded ranges of plants in conjunction with the community composition of specialist herbivores in the native range may yield valuable clues in selecting agents that are most likely to establish and regulate populations of the target weed.     

Responses of insect herbivores to sharing a host plant with a hemiparasite: impacts on preference and performance differ with feeding guild

1. Root hemiparasites are common components of many ecosystems and can affect both the biomass and the nutritional quality of the plants they infect. The consequences of these modifications for the preference and performance of three herbivore feeding guilds sharing a host with the hemi‐parasite were examined. 2. It was predicted that as the hemiparasite increased in biomass its impact on the host would increase, as would the indirect impacts on the herbivores. It was also predicted that herbivores from different feeding guilds would respond differently to the presence of the hemiparasite, reflecting the extent to which they utilise resources disrupted by the parasite and hence are in competition with it. 3. The preference and performance of phloem‐feeding aphids, xylem‐feeding spittle bugs, and leaf‐feeding grasshoppers were measured on the host grass species, Holcus lanatus L. (Poaceae), with and without attachment from the hemi‐parasite, Rhinanthus minor L. (Orobanchaceae). 4. The effects of R. minor on the host were dependent on the hemiparasite's stage of growth, being most pronounced when it was at peak biomass. At this stage it caused a significant reduction in the biomass, water content, and total nitrogen content of the host plants. 5. Overall, herbivores benefited from, or preferred, shared host plants more than uninfected plants. The aphid benefited from sharing a host with R. minor, showing increased population growth on, and preference for, parasitised plants. The spittle bug also showed a preference for parasitised plants. The grasshopper, Chorthippus brunneus Thunberg (Orthoptera: Acrididae), did not show a preference for, or a performance response to, parasitised hosts, but it consumed significantly more plant material when caged on parasitised plants. 6. These data support the prediction that invertebrate herbivores responded to changes in host plant traits driven by the hemiparasite, and strongly suggest that these indirect interactions could impact on population and community processes within natural communities.     

No detectable impacts of frequent burning on foliar C and N or insect herbivory in an Australian eucalypt forest

Question: What is the effect of frequent low intensity prescribed fire on foliar nutrients and insect herbivory in an Australian eucalypt forest? Location: Lorne State Forest (Bulls Ground Frequent Burning Study), mid‐north coast, New South Wales, Australia. Methods: Eighteen independent sites were studied representing three experimental fire regimes: fire exclusion (at least 45 years), frequently burnt (every 3 years for 35 years) and fire exclusion followed by the recent introduction of frequent burning (two fires in 6 years). Mature leaves were collected from the canopy of Eucalyptus pilularis trees at each site and analysed for nutrients and damage by invertebrate herbivores. Results: Almost 75% of all leaves showed some signs of leaf damage. The frequency of past fires had no effect on carbon and nitrogen content of canopy leaves. These results were consistent with assessments of herbivore damage where no significant differences were found in the amount of invertebrate herbivory damage to leaves across fire treatments. Conclusions: This eucalypt forest displayed a high degree of resilience to both frequent burning and fire exclusion as determined by foliar nutrients and damage by insect herbivores. Fire frequency had no detectable ecological impact on this aspect of forest health.     

Top-down effects of insect herbivores during early succession: influence on biomass and plant dominance

We tested the hypothesis that phytophagous insects would have a strong top-down effect on early successional plant communities and would thus alter the course of succession. To test this hypothesis, we suppressed above-ground insects at regular intervals with a broad-spectrum insecticide through the first 3 years of old-field succession at three widely scattered locations in central New York State. Insect herbivory substantially reduced total plant biomass to a similar degree at all three sites by reducing the abundance of meadow goldenrod, Solidago altissima. As a result, Euthamia graminifolia dominated control plots whereas S. altissima dominated insecticide-treated plots by the third year of succession. S. altissima is the dominant old-field herbaceous species in this region but typically requires at least 5 years to become dominant. Past explanations for this delay have implicated colonization limitation whereas our data demonstrate that insect herbivory is a likely alternative explanation. A widespread, highly polyphagous insect, the xylem-tapping spittlebug, Philaenus spumarius, appeared to be the herbivore responsible for the reduction in standing crop biomass at all three sites. Insect herbivory typically caused little direct leaf tissue loss for the ten plant species we examined, including S. altissima. Consequently, the amount of leaf area removed was not a reliable indicator of the influence of insect herbivory on standing crop biomass or on early succession. Overall, we found a strong top-down effect of insect herbivores on biomass at several sites, so our results may be broadly applicable. These findings run counter to generalizations that top-down effects of herbivores, particularly insects, are weak in terrestrial systems. These generalizations may not apply to insects, such as spittlebugs, that can potentially mount an effective defense (i.e., spittle) against predators and subsequently reach relatively high abundance on common plant species. Our results suggest that insect herbivory may play an important but often overlooked role during early old-field succession. Received: 26 December 1998 / Accepted: 3 April 1999