Effects of elevated CO2 and temperature on development and consumption rates of Octotoma championi and O. scabripennis feeding on Lantana camara

We carried out a factorial experiment to explore the effect of doubled CO₂ concentration and a 3 °C temperature increase on the development of a complete generation of the beetles Octotoma championi Baly and O. scabripennis Guérin‐Méneville (Coleoptera: Chrysomelidae). These species are biological control agents of Lantana camara L. (Verbenaceae), with a leaf‐mining larval phase and free‐living, leaf‐chewing adults. Plants grown at elevated CO₂ had enhanced above‐ground biomass, thicker leaves, reduced nitrogen concentration, and increased C:N ratios. Under the high temperature treatment, plants grown at ambient CO₂ suffered wilting and premature leaf loss, despite daily watering; this effect was ameliorated at elevated CO₂. The wilting of plants in the ambient CO₂/high temperature treatment reduced the emergence success of the beetles, particularly O. championi. Development time was accelerated by approximately 10–13 days at the higher temperature, but was not affected by CO₂. Neither CO₂ nor temperature affected adult beetle weight. Consumption rates of free‐living beetles were not affected by either CO₂ or temperature. By contrast, in the short‐term trials using excised foliage, beetles given no choice between ambient and elevated CO₂‐grown foliage, consumed more from ambient plants. When beetles were offered a choice between foliage grown at the two CO₂ levels, O. championi did not display a significant preference but O. scabripennis consumed more ambient CO₂‐grown foliage when feeding at the lower temperature. This study indicates that under future conditions of higher temperatures, amelioration of water stress in host plants growing in elevated CO₂ may benefit some endophagous insects by reducing premature leaf loss. Under some circumstances, this benefit may outweigh the deleterious effects of lower leaf nitrogen. Our results also indicate that foliage consumption under elevated CO₂ by mobile, adult insects on whole plants may not be significantly increased, as was previously indicated by short‐term experiments using excised foliage.     

Effects of elevated CO2 and temperature-grown red and sugar maple on gypsy moth performance

Few studies have investigated how tree species grown under elevated CO₂ and elevated temperature alter the performance of leaf‐feeding insects. The indirect effects of an elevated CO₂ concentration and temperature on leaf phytochemistry, along with potential direct effects on insect growth and consumption, may independently or interactively affect insects. To investigate this, we bagged larvae of the gypsy moth on leaves of red and sugar maple growing in open‐top chambers in four CO₂/temperature treatment combinations: (i) ambient temperature, ambient CO₂; (ii) ambient temperature, elevated CO₂ (+ 300 μL L^?1 CO₂); (iii) elevated temperature (+ 3.5°C), ambient CO₂; and (iv) elevated temperature, elevated CO₂. For both tree species, leaves grown at elevated CO₂ concentration were significantly reduced in leaf nitrogen concentration and increased in C: N ratio, while neither temperature nor its interaction with CO₂ concentration had any effect. Depending on the tree species, leaf water content declined (red maple) and carbon‐based phenolics increased (sugar maple) on plants grown in an enriched CO₂ atmosphere. The only observed effect of elevated temperature on leaf phytochemistry was a reduction in leaf water content of sugar maple leaves. Gypsy moth larval responses were dependent on tree species. Larvae feeding on elevated CO₂‐grown red maple leaves had reduced growth, while temperature had no effect on the growth or consumption of larvae. No significant effects of either temperature or CO₂ concentration were observed for larvae feeding on sugar maple leaves. Our data demonstrate strong effects of CO₂ enrichment on leaf phytochemical constituents important to folivorous insects, while an elevated temperature largely has little effect. We conclude that alterations in leaf chemistry due to an elevated CO₂ atmosphere are more important in this plant–folivorous insect system than either the direct short‐term effects of temperature on insect performance or its indirect effects on leaf chemistry.     

Gypsy moth feeding in the canopy of a CO2-enriched mature forest

Rising atmospheric carbon dioxide (CO₂) concentration is expected to change plant tissue quality with important implications for plant–insect interactions. Taking advantage of canopy access by a crane and long‐term CO₂ enrichment (530 μ mol mol^?1) of a natural old‐growth forest (web‐free air carbon dioxide enrichment), we studied the responses of a generalist insect herbivore feeding in the canopy of tall trees. We found that relative growth rates (RGR) of gypsy moth (Lymantria dispar) were reduced by 30% in larvae fed on high CO₂‐exposed Quercus petraea, but increased by 29% when fed on high CO₂‐grown Carpinus betulus compared with control trees at ambient CO₂ (370 μ mol mol^?1). In Fagus sylvatica, there was a nonsignificant trend for reduced RGR under elevated CO₂. Tree species‐specific changes in starch to nitrogen ratio, water, and the concentrations of proteins, condensed and hydrolyzable tannins in response to elevated CO₂ were identified to correlate with altered RGR of gypsy moth larvae. Our data suggest that rising atmospheric CO₂ will have strong species‐specific effects on leaf chemical composition of canopy trees in natural forests leading to contrasting responses of herbivores such as those reported here. A future change in host tree preference seems likely with far‐ranging consequences for forest community dynamics.     

Leaf age effects of elevated CO2-grown white oak leaves on spring-feeding lepidopterans

Folivorous insect responses to elevated CO₂-grown tree species may be complicated by phytochemical changes as leaves age. For example, young expanding leaves in tree species may be less affected by enriched CO₂-alterations in leaf phytochemistry than older mature leaves due to shorter exposure times to elevated CO₂ atmospheres. This, in turn, could result in different effects on early vs. late instar larvae of herbivorous insects. To address this, seedlings of white oak (Quercus alba L.), grown in open-top chambers under ambient and elevated CO₂, were fed to two important early spring feeding herbivores; gypsy moth (Lymantria dispar L.), and forest tent caterpillar (Malacosoma disstria Hübner). Young, expanding leaves were presented to early instar larvae, and older fully expanded or mature leaves to late instar larvae. Young leaves had significantly lower leaf nitrogen content and significantly higher total nonstructural carbohydrate:nitrogen ratio as plant CO₂ concentration rose, while nonstructural carbohydrates and total carbon-based phenolics were unaffected by plant CO₂ treatment. These phytochemical changes contributed to a significant reduction in the growth rate of early instar gypsy moth larvae, while growth rates of forest tent caterpillar were unaffected. The differences in insect responses were attributed to an increase in the nitrogen utilization efficiency (NUE) of early instar forest tent caterpillar larvae feeding on elevated CO₂-grown leaves, while early instar gypsy moth larval NUE remained unchanged among the treatments. Later instar larvae of both insect species experienced larger reductions in foliage quality on elevated CO₂-grown leaves than earlier instars, as the carbohydrate:nitrogen ratio of leaves substantially increased. Despite this, neither insect species exhibited changes in growth or consumption rates between CO₂ treatments in the later instar. An increase in NUE was apparently responsible for offsetting reduced foliar nitrogen for the late instar larvae of both species.     

Elevated CO2 levels and herbivore damage alter host plant preferences

Interactions between the moth Spodoptera littoralis and two of its host plants, alfalfa (Medicago sativa) and cotton (Gossypium hirsutum) were examined, using plants grown under ambient (350 ppm) and elevated (700 ppm) CO₂ conditions. To determine strength and effects of herbivore‐induced responses assays were performed with both undamaged (control) and herbivore damaged plants. CO₂ and damage effects on larval host plant preferences were determined through dual‐choice bioassays. In addition, larvae were reared from hatching to pupation on experimental foliage to examine effects on larval growth and development. When undamaged plants were used S. littoralis larvae in consumed more cotton than alfalfa, and CO₂ enrichment caused a reduction in the preference for cotton. With damaged plants larvae consumed equal amounts of the two plant species (ambient CO₂ conditions), but CO₂ enrichment strongly shifted preferences towards cotton, which was then consumed three times more than alfalfa. Complementary assays showed that elevated CO₂ levels had no effect on the herbivore‐induced responses of cotton, whereas those of alfalfa were significantly increased. Larval growth was highest for larvae fed undamaged cotton irrespectively of CO₂ level, and lowest for larvae on damaged alfalfa from the high CO₂ treatment. Development time increased on damaged cotton irrespectively of CO₂ treatment, and on damaged alfalfa in the elevated CO₂ treatment. These results demonstrate that elevated CO₂ levels can cause insect herbivores to alter host plant preferences, and that effects on herbivore‐induced responses may be a key mechanism behind these processes. Furthermore, since the insects were shown to avoid foliage that reduced their physiological performance, our data suggest that behavioural host plant shifts result in partial escape from negative consequences of feeding on high CO₂ foliage. Thus, CO₂ enrichment can alter both physiology and behaviour of important insect herbivores, which in turn may to impact plant biodiversity.     

Development of gypsy moth larvae feeding on red maple saplings at elevated CO<Subscript>2</Subscript> and temperature

Predicted increases in atmospheric CO₂ and global mean temperature may alter important plant-insect associations due to the direct effects of temperature on insect development and the indirect effects of elevated temperature and CO₂ enrichment on phytochemicals important for insect success. We investigated the effects of CO₂ and temperature on the interaction between gypsy moth (Lymantria dispar L.) larvae and red maple (Acer rubrum L.) saplings by bagging first instar larvae within open-top chambers at four CO₂/temperature treatments: (1) ambient temperature, ambient CO₂, (2) ambient temperature, elevated CO₂ (+300 l l^-1 CO₂), (3) elevated temperature (+3.5°C), ambient CO₂, and (4) elevated temperature, elevated CO₂. Larvae were reared to pupation and leaf samples taken biweekly to determine levels of total N, water, non-structural carbohydrates, and an estimate of defensive phenolic compounds in three age classes of foliage: (1) immature, (2) mid-mature and (3) mature. Elevated growth temperature marginally reduced (P <0.1) leaf N and significantly reduced (P <0.05) leaf water across CO₂ treatments in mature leaves, whereas leaves grown at elevated CO₂ concentration had a significant decrease in leaf N and a significant increase in the ratio of starch:N and total non-structural carbohydrates:N. Leaf N and water decreased and starch:N and total non-structural carbohydrates:N ratios increased as leaves aged. Phenolics were unaffected by CO₂ or temperature treatment. There were no interactive effects of CO₂ and temperature on any phytochemical measure. Gypsy moth larvae reached pupation earlier at the elevated temperature (female =8 days, P <0.07; male =7.5 days, P <0.03), whereas mortality and pupal fresh weight of insects were unrelated to either CO₂, temperature or their interaction. Our data show that CO₂ or temperature-induced alterations in leaf constituents had no effect on insect performance; instead, the long-term exposure to a 3.5°C increase in temperature shortened insect development but had no effect on pupal weight. It appears that in some tree-herbivorous insect systems the direct effects of an increased global mean temperature may have greater consequences for altering plant-insect interactions than the indirect effects of an increased temperature or CO₂ concentration on leaf constituents.     

Effects of elevated CO2 on development and larval food-plant preference in the butterfly Coenonympha pamphilus (Lepidoptera, Satyridae)

The objective of this study was to determine how increasing atmospheric CO₂ change plant tissue quality in four native grassland grass species (Agrostis stolonifera, Anthoxanthum odoratum, Festuca rubra, Poa pratensis) which are all larval food‐plants of Coenonympha pamphilus (Lepidoptera, Satyridae). We assessed the effect of these changes on the performance and larval food‐plant preference of C. pamphilus in a greenhouse experiment. Furthermore, we tested the interactive effects of elevated CO₂ and soil nutritional availability in F. rubra and its effect an larval development of C. pamphilus. In general, elevated CO₂ decreased leaf water concentration, nitrogen concentration and specific leaf area (SLA), while leaf starch concentration was increased in all grass species. A species‐specific reaction to elevated CO₂ was only found for foliar starch concentration. P. pratensis did not increase its starch concentration under elevated CO₂ conditions, whereas the other three species did. Fertilisation, investigated only for F. rubra, increased leaf nitrogen concentration and amplified the CO₂‐induced decrease in leaf nitrogen. Development time of C. pamphilus was on the average prolonged by two days under elevated CO₂ and the prolongation differed from 0.7 to 5.3 days among food‐plant species. Pupal fresh weight differed marginally between CO₂ treatments. Fertilisation of the larval food‐plant F. rubra shortened development time by one day and significantly increased pupal and adult fresh weights. C. pamphilus larvae showed a clear food‐plant preference among grass species at the age of 36 h or older. Additionally, a change of food‐plant preference under elevated CO₂ was found. Larvae at ambient CO₂ preferred Agrostis stolonifera and F. rubra, while under elevated CO₂Anthoxanthum odoratum and P. pratensis were preferred. The present study demonstrates that larval development of C. pamphilus is affected by food‐plant species and CO₂ induced changes in foliar chemistry. Although we found some species‐specific reactions to elevated CO₂ for foliar chemistry, no such CO₂ by species interaction was found for insect development. The change in food‐plant preference of larvae under elevated CO₂ implies potential changes in selection pressure for grass species and might therefore affect evolutionary processes.     

Impact of elevated CO2 and increased temperature on Japanese beetle herbivory

To examine how the major elements of global change affect herbivory in agroecosystems, a multifactorial experiment was conducted where soybeans were grown at two levels of carbon dioxide and temperature, including those predicted for 2050, under otherwise normal field conditions. Japanese beetles (Popillia japonica Newman) were enclosed on foliage for 24 h, after which the beetle survivorship, total and per capita leaf consumption, and leaf protease inhibitor activity were measured. The direct effect of temperature on beetle consumption and survivorship also was measured under controlled environmental conditions. No differences in total foliage consumption were observed; however, beetles forced to feed at elevated temperature in the field demonstrated greater per capita consumption and reduced survivorship compared to beetles feeding at ambient temperature. Survivorship was also greater for beetles that consumed foliage grown under elevated CO₂, but there were no interactive effects of CO₂ and temperature, and no differences in leaf chemistry were resolved. Leaf consumption by beetles increased strongly with increasing temperature up to ～37° C, above which increased mortality caused a precipitous decrease in consumption. An empirical model based on the temperature dependence of leaf consumption and flight suggests that the 3.5°C increase in temperature predicted for 2050 will increase the optimal feeding window for the Japanese beetle by 290%. Elevated temperature and CO₂ operating independently have the potential to greatly increase foliage damage to soybean by chewing insects, such as Popillia japonica, potentially affecting crop yields.     

Effects of elevated CO2 on feeding preference and performance of the gypsy moth (Lymantria dispar) larvae

Multiple‐choice assays were conducted in the laboratory to examine the effect of CO₂‐induced changes in leaf quality on growth, nutritional indices and preferences of the gypsy moth (Lymantria dispar) larvae. The tested tree species, one‐year‐old aspen (Populus pseudo‐simonii Kitag.), two‐year‐old birch (Betula platyphylla) and three‐year‐old oak (Quercus mongolica Fisch.) were transplanted to open‐top chambers at ambient or elevated CO₂ (650 ppm) concentrations in May 2005. The present study was conducted in 2006. Leaves from the upper and lower crowns of each tree species were sampled and analysed, respectively. On both measurement dates (24 July 2006 and 16 August 2006), there were significant CO₂ effects on levels of condensed tannin, starch, nitrogen and C : N ratio. But only on the latter date, were the concentrations of total phenolics, soluble sugar, carbon and total non‐structural carbohydrates significantly affected by elevated CO₂. Leaf dry weight content and specific leaf weight were almost completely unaffected by CO₂ enrichment. Gypsy moth larvae exhibited a clear selectivity for tested leaf types (leaves from the upper and lower crowns of each tree species) even in their early instar stage, with the upper leaves of P. pseudo‐simonii being the most preferable and the lower leaves of Q. mongolica being the least preferred. The changes in leaf quality significantly reduced larval growth and altered partial indices of insect performance (e.g., relative growth rate and relative consumption rate). However, at least in this short‐term choice feeding assay (13 days), the CO₂‐induced changes in leaf quality had no significant effects on food preference behaviour of the gypsy moth larvae, neither within the limited range of host plants nor within the leaves at different canopy heights of the same tree species.     

CO2 and O3 effects on host plant preferences of the forest tent caterpillar (Malacosoma disstria)

Elevated levels of CO₂ and O₃ affect plant growth and phytochemistry, which in turn can alter physiological performance of associated herbivores. Little is known, however, about how generalist insect herbivores respond behaviorally to CO₂‐ and O₃‐mediated changes in their host plants. This research examined the effects of elevated CO₂ and O₃ levels on host plant preferences and consumption of forest tent caterpillar (FTC, Malacosoma disstria Hbn.) larvae. Dual choice feeding assays were performed with foliage from birch (Betula papyrifera Marsh.) and aspen (Populus tremuloides Michx., genotypes 216 and 259). Trees were grown at the Aspen Free Air CO₂ Enrichment (FACE) facility near Rhinelander, WI, USA, and had been exposed to ambient or elevated concentrations of CO₂ and/or O₃. Levels of nutritional and secondary compounds were quantified through phytochemical analyses. The results showed that elevated O₃ levels increased FTC larval preferences for birch compared with aspen, whereas elevated CO₂ levels had the opposite effect. In assays with the two aspen genotypes, addition of both CO₂ and O₃ caused a shift in feeding preferences from genotype 259 to genotype 216. Consumption was unaffected by experimental treatments in assays comparing aspen and birch, but were increased for larvae given high O₃ foliage in the aspen genotype assays. Elevated levels of CO₂ and O₃ altered tree phytochemistry, but did not explain shifts in feeding preferences. The results demonstrate that increased levels of CO₂ and O₃ can alter insect host plant preferences both between and within tree species. Also, consequences of altered host quality (e.g., compensatory consumption) may be buffered by partial host shifts in situations when alternative plant species are available. Environmentally induced changes in host plant preferences may have the potential to alter the distribution of herbivory across plant genotypes and species, as well as competitive interactions among them.     

Carbon dioxide‐ and temperature‐mediated changes in plant defensive compounds alter food utilization of herbivores

Although the impact of elevated carbon dioxide and rising temperature on plants and animals has been extensively documented recently, only limited understanding exists regarding their combined effects. The objective of this research was to address the consequences of using combinations of elevated CO₂ and elevated temperature on a plant's defensive chemistry, and subsequent utilization of the plant as insect food. Our results indicated that elevated CO₂ and increased temperature, for the most part, act independently on the production of defensive compounds in broccoli leaves (Brassica oleracea L. var. italica). CO₂ concentrations had significant effects on the foliar water content, total phenolic compounds, polyphenol oxidase and trypsin inhibitor concentrations. The herbivore Spodoptera litura (Fabricius; Lepidoptera: Noctuidae) responded to changes in the plant secondary chemistry, with larvae consuming more plant materials that had been exposed to elevated CO₂. The food utilization efficiencies of second‐instar larvae were more sensitive to CO₂‐treated foliage than those of the third‐ and fourth‐instar larvae. Temperature did exert a significant effect on food utilization (ECD) by the larvae. Our study will provide important information in future predictions on plant–insect interactions as a result of climate change. The study also demonstrated that since various larval stages might respond differently to climate change, this possibility needs to be considered in future forecasting and monitoring.     

Response of quaking aspen genotypes to enriched CO2: foliar chemistry and tussock moth performance

Abstract

• 1 Genetic variation in the phytochemical responses of plants to CO₂ enrichment is likely to alter trophic dynamics, and to shift intraspecific selection pressures on plant populations. We evaluated the independent and interactive effects of atmospheric CO₂ and quaking aspen (Populus tremuloides Michx.) genotype on chemical composition of foliage and performance of the whitemarked tussock moth (Orgyia leucostigma J. E. Sm.).
• 2 This research was conducted at the Aspen FACE (Free Air CO₂ Enrichment) site in northern Wisconsin, U.S.A. Leaf samples were collected periodically from each of three genetically variable aspen genotypes growing under ambient and elevated CO₂, and analysed for levels of primary and secondary metabolites. Tussock moth larvae were reared in situ on experimental trees, and development times and pupal masses were recorded.
• 3 Foliar chemical composition varied among aspen genotypes and in response to CO₂ enrichment. However, chemical responses of trees to elevated CO₂ were generally consistent across genotypes.
• 4 Larval development times varied among host genotypes and increased slightly for insects on high‐CO₂ plants. Enriched CO₂ tended to reduce insect pupal masses, particularly for females on one of the three aspen genotypes.
• 5 CO₂ × genotype interactions observed for plant chemistry and insect performance in this study with a small number of genotypes are probably too few, and too weak, to shift selection pressures in aspen populations. These results differ, however, from earlier work in which more substantial CO₂ × genotype interactions were observed for plant chemistry.

     

Elevated atmospheric CO2: effects on phytochemistry, insect performance and insect-parasitoid interactions

This study was conducted to examine the effects of CO₂-mediated changes in tree chemistry on the performance of the gypsy moth ((Lymantria dispar L.) and the parasitold Cotesia melanoscela (Ratz.). We used carbon-nutrient balance theory to develop hypotheses regarding changes in tree chemistry and the performance of both insects under elevated CO₂. As predicted, levels of foliar nitrogen declined and concentrations of carbon-based compounds (e.g. starch and phenolics) increased under elevated CO₂. Gypsy moth performance (e.g. growth, development) was altered by CO₂-mediated changes in foliar chemistry, but the magnitude was small and varied across tree species. Larvae feeding on high CO₂ aspen exhibited the largest reduction in performance, relative to larvae feeding on birch, oak, or maple. Parasitism by C. melanoscela significantly prolonged gypsy moth development and reduced growth rates. Overall, the effect of parasitism on gypsy moth performance did not differ between CO₂ treatments. Altered gypsy moth performance on high CO₂ foliage in turn affected parasitoid performance, but the response was variable: parasitoid mortality increased and adult female size declined slightly under high CO₂, while development time and adult male size were unaffected. Our results suggest that CO₂-induced changes in plant chemistry were buffered to the extent that effects on third trophic level interactions were weak to non-existent for the system examined in this study.     

Widespread foliage δ15N depletion under elevated CO2: inferences for the nitrogen cycle

Leaf ¹⁵N signature is a powerful tool that can provide an integrated assessment of the nitrogen (N) cycle and whether it is influenced by rising atmospheric CO₂ concentration. We tested the hypothesis that elevated CO₂ significantly changes foliage δ¹⁵N in a wide range of plant species and ecosystem types. This objective was achieved by determining the δ¹⁵N of foliage of 27 field‐grown plant species from six free‐air CO₂ enrichment (FACE) experiments representing desert, temperate forest, Mediterranean‐type, grassland prairie, and agricultural ecosystems. We found that within species, the δ¹⁵N of foliage produced under elevated CO₂ was significantly lower (P<0.038) compared with that of foliage grown under ambient conditions. Further analysis of foliage δ¹⁵N by life form and growth habit revealed that the CO₂ effect was consistent across all functional groups tested. The examination of two chaparral shrubs grown for 6 years under a wide range of CO₂ concentrations (25–75 Pa) also showed a significant and negative correlation between growth CO₂ and leaf δ¹⁵N. In a select number of species, we measured bulk soil δ¹⁵N at a depth of 10 cm, and found that the observed depletion of foliage δ¹⁵N in response to elevated CO₂ was unrelated to changes in the soil δ¹⁵N. While the data suggest a strong influence of elevated CO₂ on the N cycle in diverse ecosystems, the exact site(s) at which elevated CO₂ alters fractionating processes of the N cycle remains unclear. We cannot rule out the fact that the pattern of foliage δ¹⁵N responses to elevated CO₂ reported here resulted from a general drop in δ¹⁵N of the source N, caused by soil‐driven processes. There is a stronger possibility, however, that the general depletion of foliage δ¹⁵N under high CO₂ may have resulted from changes in the fractionating processes within the plant/mycorrhizal system.     

Seeing the forest for the trees: long-term exposure to elevated CO2 increases some herbivore densities

The effects of elevated CO₂ on plant growth and insect herbivory have been frequently investigated over the past 20 years. Most studies have shown an increase in plant growth, a decrease in plant nitrogen concentration, an increase in plant secondary metabolites and a decrease in herbivory. However, such studies have generally overlooked the fact that increases in plant production could cause increases of herbivores per unit area of habitat. Our study investigated leaf production, herbivory levels and herbivore abundance per unit area of leaf litter in a scrub‐oak system at Kennedy Space Center, Florida, under conditions of ambient and elevated CO₂, over an 11‐year period, from 1996 to 2007. In every year, herbivory, that is leafminer and leaftier abundance per 200 leaves, was lower under elevated CO₂ than ambient CO₂ for each of three species of oaks, Quercus myrtifolia, Quercus chapmanii and Quercus geminata. However, leaf litter production per 0.1143 m² was greater under elevated CO₂ than ambient CO₂ for Q. myrtifolia and Q. chapmanii, and this difference increased over the 11 years of the study. Leaf production of Q. geminata under elevated CO₂ did not increase. Leafminer densities per 0.1143 m² of litterfall for Q. myrtifolia and Q. chapmanii were initially lower under elevated CO₂. However, shortly after canopy closure in 2001, leafminer densities per 0.1143 m² of litter fall became higher under elevated CO₂ and remained higher for the remainder of the experiment. Leaftier densities per 0.1143 m² were also higher under elevated CO₂ for Q. myrtifolia and Q. chapmanii over the last 6 years of the experiment. There were no differences in leafminer or leaftier densities per 0.1143 m² of litter for Q. geminata. These results show three phenomena. First, they show that elevated CO₂ decreases herbivory on all oak species in the Florida scrub‐oak system. Second, despite lower numbers of herbivores per 200 leaves in elevated CO₂, increased leaf production resulted in higher herbivore densities per unit area of leaf litter for two oak species. Third, they corroborate other studies which suggest that the effects of elevated CO₂ on herbivores are species specific, meaning they depend on the particular plant species involved. Two oak species showed increases in leaf production and herbivore densities per 0.1143 m² in elevated CO₂ over time while another oak species did not. Our results point to a future world of elevated CO₂ where, despite lower plant herbivory, some insect herbivores may become more common.     

Atmospheric change alters performance of an invasive forest insect

Atmospheric change and species invasions are arguably two of the most important factors affecting the long‐term sustainability of natural ecosystems. We examined the independent and interactive effects of atmospheric carbon dioxide (CO₂) and tropospheric ozone (O₃) on the foliar quality of two host species and performance of an invasive folivorous insect. Trembling aspen (Populus tremuloides) and paper birch (Betula papyrifera) were grown at the Aspen FACE research site in northern Wisconsin, USA, under all combinations of ambient and elevated CO₂ and O₃. We measured the effects of elevated CO₂ and O₃ on aspen and birch phytochemistry and on the survivorship, development time, growth, and fecundity of the gypsy moth (Lymantria dispar). Elevated CO₂ had little effect on, whereas elevated O₃ altered, the composite phytochemical profiles of aspen and birch. Nutritional quality in aspen and birch leaves was marginally affected by elevated CO₂ and reduced by elevated O₃. Both gases increased concentrations of phenolic and structural compounds in aspen and birch. Elevated CO₂ offset reduced foliar quality under elevated O₃, but only in aspen, and to a greater extent later than earlier in spring. Elevated CO₂ generally had beneficial effects on, while elevated O₃ detrimentally affected, gypsy moth performance. Elevated CO₂ ameliorated most of the reductions in gypsy moth performance under elevated O₃. Our findings suggest that atmospheric change can alter foliar quality in gypsy moth hosts sufficiently to influence gypsy moth performance, but that these responses will depend on interactions among CO₂, O₃, and tree species. Our findings also contrast with those of earlier studies at Aspen FACE, indicating that foliar quality responses to environmental change are likely influenced by tree stand age and longevity of exposure to pollutants to the extent that they affect plant‐herbivore interactions differently over decadal time spans.     

Effects of elevated CO2 and O3 on leaf damage and insect abundance in a soybean agroecosystem

By altering myriad aspects of leaf chemistry, increasing concentrations of CO₂ and O₃ in the atmosphere derived from human activities may fundamentally alter the relationships between insect herbivores and plants. Because exposure to elevated CO₂ can alter the nutritional value of leaves, some herbivores may increase consumption rates to compensate. The effects of O₃ on leaf nutritional quality are less clear; however, increased senescence may also reduce leaf quality for insect herbivores. Additionally, changes in secondary chemistry and the microclimate of leaves may render plants more susceptible to herbivory in elevated CO₂ and O₃. Damage to soybean (Glycine max L.) leaves and the size and composition of the insect community in the plant canopy were examined in large intact plots exposed to elevated CO₂ (~550 μmol mol⁻¹) and elevated O₃ (1.2*ambient) in a fully factorial design with a Soybean Free Air Concentration Enrichment system (SoyFACE). Leaf area removed by folivorous insects was estimated by digital photography and insect surveys were conducted during two consecutive growing seasons, 2003 and 2004. Elevated CO₂ alone and in combination with O₃ increased the number of insects and the amount of leaf area removed by insect herbivores across feeding guilds. Exposure to elevated CO₂ significantly increased the number of western corn rootworm (Diabrotica virgifera) adults (foliage chewer) and soybean aphids (Aphis glycines; phloem feeder). No consistent effect of elevated O₃ on herbivory or insect population size was detected. Increased loss of leaf area to herbivores was associated with increased carbon-to-nitrogen ratio and leaf surface temperature. Soybean aphids are invasive pests in North America and new to this ecosystem. Higher concentrations of CO₂ in the atmosphere may increase herbivory in the soybean agroecosystem, particularly by recently introduced insect herbivores. Handling editor: Gary Felton.     

Growth responses of gypsy moth larvae to elevated CO2: the influence of methods of insect rearing

Abstract The effects of elevated CO₂ on foliar chemistry of two tree species (Populus pseudo‐simonii Kitag. and Betula platyphylla) and on growth of gypsy moth (Lymantria dispar L.) larvae were examined. Furthermore, we focused on the comparison of results on the growth responses of larvae obtained from two methods of insect rearing, the no‐choice feeding trial performed in the laboratory or in situ in open‐top chambers. On the whole, both primary and secondary metabolites in the leaves of the two tree species were significantly affected by main effects of time (sampling date), CO₂ and species. Elevated CO₂ significantly increased the C : N ratio and concentrations of the soluble sugar, starch, total nonstructural carbohydrates, total phenolics and condensed tannins, but significantly decreased the concentration of nitrogen. Higher contents of total phenolics and condensed tannins were detected in the frass of larvae reared in elevated CO₂ treatments. Overall, the growth of gypsy moth larvae were significantly inhibited by elevated CO₂ and CO₂‐induced changes in leaf quality. Our study did not indicate the two methods of insect rearing could influence the direction of effects of elevated CO₂ on the growth of individual insects; however, the magnitude of negative effects of elevated CO₂ on larval growth did differ between the two insect rearing methods, and it seems that the response magnitude was also mediated by larval age and host plant species.     

Acclimation of photosynthesis to elevated CO2 in onion (Allium cepa) grown at a range of temperatures

Onion (Allium cepa) was grown in the field within temperature gradient tunnels (providing about ‐2.5°C to +2.5°C from outside temperatures) maintained at either 374 or 532 μmol mol^?1 CO₂. Plant leaf area was determined non‐destructively at 7 day intervals until the time of bulbing in 12 combinations of temperature and CO₂ concentration. Gas exchange was measured in each plot at the time of bulbing, and the carbohydrate content of the leaf (source) and bulb (sink) was determined. Maximum rate of leaf area expansion increased with mean temperature. Leaf area duration and maximum rate of leaf area expansion were not significantly affected by CO₂. The light‐saturated rates of leaf photosynthesis (A_sat) were greater in plants grown at normal than at elevated CO₂ concentrations at the same measurement CO₂ concentration. Acclimation of photosynthesis decreased with an increase in growth temperature, and with an increase in leaf nitrogen content at elevated CO₂. The ratio of intercellular to atmospheric CO₂ (C₁/C₃ ratio) was 7.4% less for plants grown at elevated compared with normal CO₂. A_sat in plants grown at elevated CO₂ was less than in plants grown at normal CO₂ when compared at the same C₁. Hence, acclimation of photosynthesis was due both to stomatal acclimation and to limitations to biochemical CO₂ fixation. Carbohydrate content of the onion bulbs was greater at elevated than at normal CO₂. In contrast, carbohydrate content was less at elevated compared with normal CO₂ in the leaf sections in which CO₂ exchange was measured at the same developmental stage. Therefore, acclimation of photosynthesis in fully expanded onion leaves was detected despite the absence of localised carbohydrate accumulation in these field‐grown crops.     

Interactive effects of pre-industrial, current and future [CO2] and temperature on an insect herbivore of Eucalyptus

Both atmospheric [CO₂] and average surface temperatures are predicted to increase with potentially different, additive or opposing, effects on leaf quality and insect herbivore activity. Few studies have directly measured the interactive effects of concurrent changes in [CO₂] and temperature on insect herbivores. None have done so over the entire developmental period of a tree-feeding insect, and none have compared responses to low pre-industrial [CO₂] and present day [CO₂] to estimate responses to future increases. Eucalypt herbivores may be particularly sensitive to climate-driven shifts in plant chemistry, as eucalypt foliage is naturally low in [N]. In this study, we assessed the development of the eucalypt herbivore Doratifera quadriguttata exposed concurrently to variable [CO₂] (290, 400, 650 μmol mol^?1) and temperature (ambient, ambient +4 °C) on glasshouse-grown Eucalyptus tereticornis. Overall, insects performed best on foliage grown at pre-industrial [CO₂], indicating that modern insect herbivores have already experienced nutritional shifts since industrialisation. Rising [CO₂] increased specific leaf mass and leaf carbohydrate concentration, subsequently reducing leaf [N]. Lower leaf [N] induced compensatory feeding and impeded insect performance, particularly by prolonging larval development. Importantly, elevated temperature dampened the negative effects of rising [CO₂] on larval performance. Therefore, rising [CO₂] over the past 200 years may have reduced forage quality for eucalypt insects, but concurrent temperature increases may have partially compensated for this, and may continue to do so in the future. These results highlight the importance of assessing plant–insect interactions within the context of multiple climate-change factors because of the interactive and potentially opposing effects of different factors within and between trophic levels.