Caterpillar assemblages on Chusquea bamboos in southern Ecuador: abundance,guild structure,and the influence of host plant quality

1. Information on the guild structure of foliage‐associated tropical insects is scarce, especially as caterpillars are mostly considered only as herbivores feeding on living leaves. However, many caterpillar species display alternative trophic associations, feeding on dead or withered leaves or epiphylls (‘non‐herbivores’). 2. To determine the contribution of these non‐herbivores, caterpillar communities associated with Chusquea Kunth (Poaceae) in the Andes of southern Ecuador were investigated. Caterpillars were collected at two elevation levels (montane rainforest ～2000 m and elfin forest at ～3000 m a.s.l.) and assigned to three feeding guilds (strict herbivores, non‐herbivores, and switchers) based on feeding trials. Foliage quality and leaf area were recorded to test for their influence on guild composition and caterpillar density. 3. Three hundred and eighty‐nine individuals belonging to 175 Lepidoptera species associated with Chusquea bamboos were found. The species richness of caterpillars was similarly high at both elevation levels but varied between feeding guilds. Approximately half (46.5%) displayed an alternative feeding association, i.e. were non‐herbivores (31.1%) or switchers (15.4%). 4. Caterpillar density was nearly two‐fold higher in the elfin forest, but only strict herbivores and switchers increased significantly with elevation. Leaf area positively influenced the density of strict herbivores and switchers; foliage quality only affected strict herbivores. The density of non‐herbivores did not differ significantly between the forest types and was not related to leaf area or foliage quality. 5. The present study underpins that non‐herbivores make up a considerable fraction of caterpillar communities in tropical mountain ecosystems and demonstrates that elevation, foliage quality and available plant biomass further shape feeding guild composition.     

Avian pest control in tea plantations of sub-Himalayan plains of Northeast India: Mixed-species foraging flock matters

This study reports on the biocontrol role birds play in caterpillar pest control of tea plantations of Northeast India. In this area large tracts of tea plantations have been extensively defoliated by the recent invasion of two forest-dwelling geometrid looper caterpillars, Hyposidra spp. and a lymantriid hairy caterpillar, Arctornis submarginata. This exacerbated tea herbivory by two resident pest caterpillars, Biston suppressaria and Eterusia magnifera. Currently there are no identified resident insect predators for any life stage of Hyposidra spp. and A. submarginata. Larvae of these pests drop from tea bushes using salivary thread, allowing caterpillars to escape from insect predators. The study identified 38 native insectivorous bird species in tea plantations, of which four species (Asian-pied starling, Chestnut-tailed starling, Jungle Myna, Red-vented Bulbul) could be potential control agents of looper and hairy caterpillar pests. These species had high population densities. Their cumulative abundances represented a major proportion of the total bird community during both the infested (86.44%) and non-infested phase (75.34%). They foraged in mixed-species flocks in both tea foliage and on the ground. This behavior is suited to capture foliage-living and dropped caterpillars that were flushed from tea bushes by foraging birds. Abundance and species richness of overall tea layer-foraging birds were higher in infested phase when compared to non-infested phase. The predation rate of four bird species of the foraging flock varied significantly. These results suggest that birds should be considered as important biological control agent of caterpillar pests of tea and considered in pest management plans.     

Bird rookeries have different effects on different feeding guilds of herbivores and alter the feeding behavior of a common caterpillar

Aggregations of nesting birds are common in many landscapes and have a broad impact on their surrounding habitat through the nutrient input and disturbance of their guano depositions. Ecological theory makes specific predictions about how disturbances or nutrient pulses will affect the interactions between different trophic levels of organisms. This study dissects the effects of a multi-species bird rookery on plant-herbivore interactions on coast live oak trees (Quercus agrifolia). I found that different feeding guilds of herbivores were affected in opposing directions by the rookery. I observed less chewing damage within the rookery than outside, but more damage from piercing/sucking and galling herbivores within the rookery than outside. To understand why chewing guild herbivores are negatively affected by the environmental impacts of the rookery, I explored the behavioral response of a specialist caterpillar (Phryganidia californica) to various rookery conditions. Bird guano had a direct negative impact on P. californica foraging. P. californica movement was impaired by guano addition to twigs, and caterpillars preferred to eat leaves without guano on them. The rookery also had an indirect negative effect on P. californica foraging, as P. californica preferred clean leaves from forest areas outside of the rookery over clean leaves from within the rookery. This study suggests that while it may be possible to make accurate predictions about the effect of large events (such as rookery formation) on different trophic levels, understanding the response of individual species within that trophic level requires an understanding of aspects of their natural history—such as feeding mode and behavior.     

Nucleopolyhedroviruses of forest and western tent caterpillars: cross-infectivity and evidence for activation of latent virus in high-density field populations

Abstract. 1. Cyclic population dynamics of forest caterpillars are often associated with epizootics of nucleopolyhedrovirus, but it is not known how these viruses persist between generations or through the fluctuations in host population density. 2. To explore the question of virus persistence at different phases of the population cycle, the nucleopolyhedroviruses of two species of tent caterpillar that co‐occur in British Columbia, Canada, Malacosoma californicum pluviale (western tent caterpillar) and Malacosoma disstria (forest tent caterpillar), were characterised. The cross‐infectivity of the viruses in these two host species was investigated to determine whether there might be a route for virus persistence via the alternative host species. Any virus produced in the cross‐infections was characterised to confirm true cross‐infection or to ascertain whether cross‐inoculation triggered latent virus persisting within the population. 3. The virus associated with forest tent caterpillars (MadiNPV) did not infect western tent caterpillars from low‐density populations, nor did it trigger a latent virus infection; however, inoculation of forest tent caterpillars from high‐density populations with virus from western tent caterpillars (McplNPV) resulted in viral infection, but without a dose–response relationship. 4. Analysis of DNA profiles of virus resulting from cross‐infection of the forest tent caterpillar with McplNPV, revealed that 88% of these infections were caused by MadiNPV rather than McplNPV; however the virus from all 44 infected individuals was identical and differed in DNA profile from the stock MadiNPV used for cross‐infection. This suggests strongly that forest tent caterpillars from high‐density field populations harbour a latent, persistent, or sublethal form of MadiNPV that was triggered by exposure to nucleopolyhedrovirus from the western tent caterpillar. 5. Virus was not activated in western tent caterpillars collected over 2 years of late population decline and the first year of population increase.     

Density dependence in insect performance within individual plants: induced resistance to Spodoptera exigua in tomato

Net intraspecific density dependence experienced by insect herbivores at the scale of single plants can be a function both of induced resistance in the plant and other interactions among individual herbivores. Theory suggests that non‐linearity in the form of this density dependence can influence the effects of plants on herbivore population dynamics. This study examined both net density dependence at the scale of single plants, and changes in plant quality with herbivore density for Spodoptera exigua caterpillars on tomato plants. One experiment measured the growth of caterpillars moving freely about the plant at different densities, the distribution of damage by these caterpillars, and the quality of the plant as food for caterpillars (growth of caterpillars on undamaged leaf tissue excised from the plant). A second experiment measured plant quality for plants with different amounts of damage by caterpillars confined to particular leaves in mesh bags. Growth of S. exigua caterpillars was found to be negatively density dependent, and this was in part due to decreases in plant quality both as herbivore density increased and as the amount of damage increased. The response of plant quality to herbivores was found to have non‐linear features; there was both a threshold below which no significant decreases in quality (as measured by herbivore growth) occurred, and the decrease in herbivore performance saturated at the highest damage levels. In addition, it was found that caterpillar damage was significantly more aggregated than expected when multiple caterpillars occupy a single plant. This study confirms that host plants have the potential to be a source of density dependence that affects herbivore performance.     

Foraging behaviour of caterpillars given a choice of plant genotypes in the presence of insect predators

1. To examine the effects of predators and plant genotype on the behaviour, patterns of herbivory, growth, and survivorship of caterpillars, an experiment was conducted under semi‐natural conditions, with two host plant genotypes (low vs. high iridoid glycosides) of plantain Plantago lanceolata, two kinds of herbivores (noncryptic specialist Junonia coenia vs. cryptic generalist Pyrrharctia isabella), and two levels of caterpillar predation (with and without Podisus maculiventris stinkbugs). 2. Choice tests conducted in the laboratory showed that in three out of four cases, caterpillars preferred leaves from the low iridoid glycoside genotype. 3. In the field experiment, the presence of predators affected the amount of leaf material eaten per plot and the proportion of leaf material eaten by the caterpillars as expected, but it did not affect the use of plant genotypes by Junonia caterpillars within the plots. In contrast, the caterpillar density–predation treatments affected the proportion of leaves eaten by Pyrrharctia caterpillars that were of the low iridoid glycoside genotype, but not entirely as expected. The caterpillars used both genotypes equally when predators were present but the high iridoid glycoside genotype was used more by caterpillars at low density and without predators. 4. The message then is that on the scale that caterpillars could be choosy about intake of iridoid glycosides they were not choosy among plant genotypes; however they were choosy between leaves, which can differ in iridoid glycoside concentration via plant genotype and leaf age.     

Temperature and population density: interactional effects of environmental factors on phenotypic plasticity,immune defenses,and disease resistance in an insect pest

Temperature and crowding are key environmental factors mediating the transmission and epizooty of infectious disease in ectotherm animals. The host physiology may be altered in a temperature‐dependent manner and thus affects the pathogen development and course of diseases within an individual and host population, or the transmission rates (or infectivity) of pathogens shift linearly with the host population density. To our understanding, the knowledge of interactive and synergistic effects of temperature and population density on the host–pathogen system is limited. Here, we tested the interactional effects of these environmental factors on phenotypic plasticity, immune defenses, and disease resistance in the velvetbean caterpillar Anticarsia gemmatalis. Upon egg hatching, caterpillars were reared in thermostat‐controlled chambers in a 2 × 4 factorial design: density (1 or 8 caterpillars/pot) and temperature (20, 24, 28, or 32°C). Of the immune defenses assessed, encapsulation response was directly affected by none of the environmental factors; capsule melanization increased with temperature in both lone‐ and group‐reared caterpillars, although the lone‐reared ones presented the most evident response, and hemocyte numbers decreased with temperature regardless of the population density. Temperature, but not population density, affected considerably the time from inoculation to death of velvetbean caterpillar. Thus, velvetbean caterpillars succumbed to Anticarsia gemmatalis multiple nucleopolyhedrovirus (AgMNPV) more quickly at higher temperatures than at lower temperatures. As hypothesized, temperature likely affected caterpillars' movement rates, and thus the contact between conspecifics, which in turn affected the phenotypic expression of group‐reared caterpillars. Our results suggest that environmental factors, mainly temperature, strongly affect both the course of disease in velvetbean caterpillar population and its defenses against pathogens. As a soybean pest, velvetbean caterpillar may increase its damage on soybean fields under a scenario of global warming as caterpillars may reach the developmental resistance faster, and thus decrease their susceptibility to biological control by AgMNPV.     

The role of leaf defensive traits in oaks on the preference and performance of a polyphagous herbivore,Orgyia vetusta

1. Leaves possess traits that mediate the preference and performance of herbivores. Most evidence for the importance of leaf traits as defences against herbivory comes from studies of few model plant species. 2. In a phylogenetically explicit comparison, I explain the differences in preference and performance of tussock moth (Orgyia vetusta Boisduval) larvae on leaves of 27 oak (Quercus) species using nine putative leaf defences. 3. The preference for an oak species correlated positively with the survival of caterpillars. The correlation between preference and performance did not differ between oak species native to the range of tussock moth versus those from outside the herbivore's range. 4. The first principal component of leaf traits predicted survival of caterpillars on oak leaves but only marginally predicted their preference between oak species. A multiple regression model showed that evergreenness, toughness, and condensed tannin content were the best predictors of caterpillar survival, and leaf toughness was the best predictor of host preference. 5. Generalist caterpillars may accurately assess the value of novel food sources. Moreover, many leaf traits that have been found to affect herbivory within a plant species can also be used to predict the fitness of a generalist herbivore between species.     

Effects of trichomes on the behavior and distribution of Platyprepia virginalis caterpillars

Trichomes are an important physical resistance mechanism of plants, as they reduce insect herbivore movement, feeding, and digestion. However, we know little about how trichomes influence herbivore distributions and populations. We conducted laboratory and field experiments to evaluate the preferences of Platyprepia virginalis (Boisduval) (Lepidoptera: Arctiidae) caterpillars to natural and manipulated densities of trichomes on their primary food, Lupinus arboreus Sims (Fabaceae). We then conducted field surveys to determine whether variation in trichome density among lupine bushes affected caterpillar spatial distribution on the landscape. Platyprepia virginalis caterpillars preferred lupine leaves with fewer trichomes in choice and no‐choice experiments. In the field, caterpillar feeding damage was found more often on leaves with fewer trichomes. These preferences scaled up to the level of bushes in the landscape such that more caterpillars were found on bushes with lower trichome densities than on bushes with higher trichome densities. This is one of few studies to show the potential for trichome density to influence herbivore population size and distribution in a natural system at a landscape level. The results are consistent with trichomes functioning as a resistance mechanism with consequences for herbivore choice, performance, and distribution.     

Foraging behavior of Cerulean Warblers during the breeding and non‐breeding seasons: evidence for the breeding currency hypothesis

Birds require additional resources for raising young, and the breeding currency hypothesis predicts that insectivorous species exploit large soft‐bodied prey during the breeding season, but shift to small, likely hard‐bodied, prey during the non‐breeding season. To test this hypothesis, we examined prey use by Cerulean Warblers (Setophaga cerulea), foliage‐gleaning Nearctic‐Neotropical migrants, during the breeding and non‐breeding seasons. We collected data on foraging behavior during the breeding season (including observations of prey items fed to young) in upland mixed‐oak forest in southeastern Ohio in 2009 and 2010 and, during the non‐breeding season, in shade coffee in the Cordillera de Merida, Venezuela, in 2008–2009. Cerulean Warblers captured 7% more large prey (visible prey extending beyond the bill) during the breeding than the non‐breeding season, but foraged at similar rates during both seasons. Large, soft‐bodied prey appeared to be especially important for feeding young. We found that adults delivered large prey on >50% of provisioning visits to nests and 69% of identifiable large prey fed to nestlings were greenish larvae (likely Lepidoptera or caterpillars) that camouflage against leaves where they would tend to be captured by foliage‐gleaning birds. Availability of specific taxa appeared to influence tree species foraging preferences. As reported by other researchers, we found that Cerulean Warblers selected trees in the genus Carya for foraging and our examination of caterpillar counts from the central Appalachian Mountains (Butler and Strazanac 2000 ) showed that caterpillars with greenish coloration, especially Baileya larvae, may be almost twice as abundant on Carya than Quercus. Our results provide evidence for the breeding currency hypothesis, and highlight the importance of caterpillars to a foliage‐gleaning migrant warbler of conservation concern.     

Tracing site‐specific isotopic signatures along a Blue Tit Cyanistes caeruleus food chain

Food chains culminating with temperate insectivorous passerines are well described, yet whether trophic webs can be site‐specific remains a largely unexplored question. In the case of site‐ or habitat‐specificity of food webs, stable isotope signatures of bird feathers may enable assignment of unmarked individuals to a site or a habitat of origin. We address this question in landscapes that include contrasting forest habitat patches with either deciduous Downy Oak Quercus humilis or evergreen Holm Oak Quercus ilex as dominant tree species. First, we examine the spatial variation across habitats and sites in the stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹⁵N) along the oak leaf–Tortrix moth Tortrix viridana caterpillar–Blue Tit Cyanistes caeruleus food chain. Secondly, we assess whether the isotopic signatures allow for correct assignment of individual birds to their site or habitat of origin. At the scale of the landscape, stable isotope values enabled identification of the different components of the Blue Tit food chain: from oak leaves to Blue Tit nestlings and yearling birds. However, isotopic signatures were site‐specific (i.e. geographical) more than habitat‐specific (i.e. deciduous vs. evergreen oaks). Discriminant analyses correctly assigned 85% of nestlings and 83% of resident yearling birds, indicating a pronounced effect of site on Blue Tit feather isotopic signatures. We thereby demonstrate that isotopes reflect a stronger association of locally born birds to the local features of their habitat than that of un‐ringed yearling birds, whose plumage may have grown while in a wider geographical area. This study provides evidence of site‐specific isotopic signatures from oak leaves to Blue Tit feathers at a fine spatial scale.     

The lethal plant defense paradox remains: inducible host-plant aristolochic acids and the growth and defense of the pipevine swallowtail

Toxic plants with sequestering specialists are presented with a problem because plant derived toxins protect herbivores against natural enemies. It has been suggested that early induction of toxins and later relaxation of these defenses may help the plant resolve this problem because neonate caterpillars incur the physiological cost of dealing with toxins in early life, but are denied toxins when they are able to sequester them efficiently. In California, the pipevine swallowtail, Battus philenor L. (Lepidoptera: Papilionidae), feed exclusively on Aristolochia californica Torrey (Aristolochiaceae), an endemic vine that contains toxic alkaloids called aristolochic acids that caterpillars sequester to provide chemical defense in immature and adult stages. In a field experiment, the concentration of aristolochic acids doubled in the plant following leaf damage and returned to constitutive levels after six days. Neonate pipevine swallowtail caterpillars showed no aversion to high levels of aristolochic acid in a preference test. Caterpillars reared on leaves with supplemented aristolochic acid showed no physiological cost or increased mortality compared to caterpillars reared on un-supplemented leaves. Searching efficiency and capture rate of lacewing larvae (Chrysoperla), a common predator of first instar caterpillars, was compromised significantly after feeding on caterpillars reared on leaves with supplemented concentrations of aristolochic acid compared to caterpillars feeding on control plants. Additionally, mortality of lacewings increased when they were provided with a diet of B. philenor caterpillars reared on supplemented leaves compared to caterpillars reared on control leaves. Thus, the induction of aristolochic acids in the plant following leaf damage does not resolve the problem confronted by the plant and may confer benefits to this sequestering specialist.     

Systemic induction of volatile release in cotton: How specific is the signal to herbivory?

Plants attacked by herbivorous insects release chemical signals that attract natural enemies of the herbivores to the damaged plants. Feeding of Spodoptera exigua larvae on the lower leaves of cotton (Gossypium hirsutum L.) for multiple feeding periods of 9–12 h with a 12 h, interval in between when the caterpillars are removed overnight, will induce a systemic release of volatile compounds that is comparable to the volatiles released in response to continuous feeding damage on the lower leaves for several days. The systemic volatile release in response to herbivory can be mimicked by mechanically damaging the lower leaves and applying caterpillar oral secretion to the injured leaves over 4 days. Cotton plants that are only mechanically damaged systemically release significantly less -pinene, myrcene, (Z)-3-hexenyl acetate, (E)--farnesene and (E,E)--farnesene after 4 days compared to plants damaged mechanically with application of caterpillar regurgitant. However, multiple 9–12 h mechanical damage alone induces a significantly higher systemic release of (Z)-3-hexenyl acetate, myrcene, (E)--ocimene, and (E)--farnesene after 4 days compared to undamaged control plants. This indicates that multiple mechanical damage alone cannot mimic completely the response induced by mechanically injuring the leaves and applying caterpillar regurgitant. A specific elicitor in the regurgitant of the caterpillar enhances the amount of several systemically released volatiles. Thus, the systemic release of volatile compounds by herbivore-damaged cotton plants appears to be regulated by at least two different mechanisms.     

Contribution of insectivorous avifauna to top down control of Lindera benzoin herbivores at forest edge and interior habitats

Predation of herbivorous Lepidoptera larvae by insectivorous avifauna was estimated on Lindera benzoin in edge and interior habitats at two sites in eastern Pennsylvania (USA). Clay baits modeled after Epimecis hortaria (Geometridae) larvae, the primary herbivore of L. benzoin at our study sites, were used to estimate predation by birds. In both habitat types, models were placed on uninjured L. benzoin leaves as well as on leaves that had prior insect herbivore damage. Rates of model attack were greater, and model longevity reduced, in forest edge plots compared to interiors. Naturally occurring herbivore damage on L. benzoin was greater in forest interiors. However, model attack was not significantly greater on leaves with prior herbivory damage, suggesting that birds do not effectively use this type of leaf damage as a cue in their foraging. Our findings are consistent with a contribution of bird predation towards top-down control of herbivory in this system. We further discuss these results in a broader context considering the possible effects of habitat type on leaf quality, leaf defense, and herbivore performance.     

Host-age discrimination during host location byCotesia glomerata, a larval parasitoid ofPieris brassicae

Some parasitoids are restricted with respect to the host stage that they attack and even to a certain age within a stage. In this paper we investigate whether the parasitoidCotesia glomerata can discriminate between old and young caterpillar instars of its host,Pieris brassicae, before contacting these hosts, since contacts with older instars are very risky with a chance of being killed, due to the aggressive defensive behaviour of the caterpillars. Flight chamber dual choice tests showed that volatile chemicals emitted by Brussels sprouts plants (Brassica oleracea var. gemmifera) after feeding damage by 1st and 5th larval instars are equally attractive to the wasps. Simulated herbivore damage by 2nd and 5th larval instars, obtained by treating mechanically damaged leaves with carterpillar regurgitant, was also equally attractive, even when the wasps were exposed to repeated experience on different larval instars to increase their discriminatory ability. In contrast, single choice contact bioassays showed that the time spent searching on a leaf with feeding damage of 1st instar larvae was significantly longer than the time spent on 5th instar feeding damage or on mechanically damaged leaves. Both flight and contact bioassays did not show any effect of egg-related infochemicals. The results demonstrate thatC. glomerata can discriminate between young and old larval instars ofP. brassicae, without contacting the caterpillars. This is not done through volatile herbivore-induced synomones but through cues that are contacted after arrival at a caterpillar-infested leaf.     

Within-tree variation in density and survival of leafminers on oak Quercus dentata

The density and survival of leafminers were examined on 50 sun leaves from each of 65 Quercus dentata Thunb. individuals in northern Japan in 1997 and 1998. Phyllonorycter (two species), Caloptilia (one species) and Stigmella (three species) were abundant or common on this oak in the study area. These leafminers appeared after mid-June, whereas most externally feeding caterpillars occurred from late May to early June when the water content and nitrogen concentration of leaves were high. The density of these leafminers was about four times higher in 1998 than in 1997. A negative correlation was more often observed between mine density and leaf size, leaf wet weight per area or leaf toughness in the Phyllonorycter species, but the opposite correlation was more frequent for Caloptilia and Stigmella species. Conversely, no clear relation was observed between the survival of Phyllonorycter larvae and leaf traits. In all leafminers except the gregarious Stigmella species, the mine density was more often positively correlated with leaf damage by chewing insects, and also the survival of Phyllonorycter larvae was often positively correlated with leaf damage. In the Phyllonorycter species, the survival of larvae tended to increase with the increase in density at the autumn generation. The correlation in the densities of mines between the summer and autumn generations was more frequently positive in the Phyllonorycter and Caloptilia species. In addition, the densities of different leafminers were often positively correlated. Thus, relations among leafminers, between leafminers and externally feeding caterpillars, and also between herbivores and host plants are complicated.     

Traditional Regulation of Edible Caterpillar Exploitation in the Kopa Area of Mpika District in Northern Zambia

The Traditional Ecological Knowledge (TEK) of the Bisa people of northern Zambia on species, life cycles and host plants of the caterpillars they exploit for food and for household income generation, and the traditional control of caterpillar harvesting, were investigated through a household survey. The accessed indigenous knowledge on caterpillar biology was validated through a forest survey, caterpillar collections and identifications. Eight species of caterpillars were harvested from the surrounding miombo woodlands. Their life systems and host plants were well understood by members of local communities and this knoweldge was communicated amongst them orally. Popular commercial species were Gynanisa maya Strand and Gonimbrasia zambesina Walker. Over 20 miombo tree species hosted the two species but the mean numbers of the caterpillars on Julbernadia paniculata Troupin were significantly larger than those on other host plants (p < 0.05), indicating that J. paniculata was probably the main host plant of the two species. Traditional regulation of caterpillar harvesting involved: (i) monitoring for edible caterpillar development and abundance and for changes in caterpillar habitats, (ii) protection of host plants and moth eggs against late bush fires, through use of traditional fire technology, and (iii) temporal restriction of edible caterpillar harvesting. The possibility of the Zambian Government promoting caterpillar harvesting as an economic incentive to the Bisa people to conserve both this renewable edible caterpillar natural resource and their environment is discussed.     

Foliar quality influences tree-herbivore-parasitoid interactions: effects of elevated CO<Subscript>2</Subscript>, O<Subscript>3</Subscript>, and plant genotype

This study examined the effects of carbon dioxide (CO₂)-, ozone (O₃)-, and genotype-mediated changes in quaking aspen (Populus tremuloides) chemistry on performance of the forest tent caterpillar (Malacosoma disstria) and its dipteran parasitoid (Compsilura concinnata) at the Aspen Free-Air CO₂ Enrichment (FACE) site. Parasitized and non-parasitized forest tent caterpillars were reared on two aspen genotypes under elevated levels of CO₂ and O₃, alone and in combination. Foliage was collected for determination of the chemical composition of leaves fed upon by forest tent caterpillars during the period of endoparasitoid larval development. Elevated CO₂ decreased nitrogen levels but had no effect on concentrations of carbon-based compounds. In contrast, elevated O₃ decreased nitrogen and phenolic glycoside levels, but increased concentrations of starch and condensed tannins. Foliar chemistry also differed between aspen genotypes. CO₂, O₃, genotype, and their interactions altered forest tent caterpillar performance, and differentially so between sexes. In general, enriched CO₂ had little effect on forest tent caterpillar performance under ambient O₃, but reduced performance (for insects on one aspen genotype) under elevated O₃. Conversely, elevated O₃ improved forest tent caterpillar performance under ambient, but not elevated, CO₂. Parasitoid larval survivorship decreased under elevated O₃, depending upon levels of CO₂ and aspen genotype. Additionally, larval performance and masses of mature female parasitoids differed between aspen genotypes. These results suggest that host-parasitoid interactions in forest systems may be altered by atmospheric conditions anticipated for the future, and that the degree of change may be influenced by plant genotype.     

Foliage of oaks grown under elevated CO2 reduces performance of Antheraea polyphemus (Lepidoptera: Saturniidae)

To understand how the increase in atmospheric CO2 from human activity may affect leaf damage by forest insects, we examined host plant preference and larval performance of a generalist herbivore, Antheraea polyphemus Cram., that consumed foliage developed under ambient or elevated CO2. Larvae were fed leaves from Quercus alba L. and Quercus velutina Lam. grown under ambient or plus 200 microl/liter CO2 using free air carbon dioxide enrichment (FACE). Lower digestibility of foliage, greater protein precipitation capacity in frass, and lower nitrogen concentration of larvae indicate that growth under elevated CO2 reduced the food quality of oak leaves for caterpillars. Consuming leaves of either oak species grown under elevated CO2 slowed the rate of development of A. polyphemus larvae. When given a choice, A. polyphemus larvae preferred Q. velutina leaves grown under ambient CO2; feeding on foliage of this species grown under elevated CO2 led to reduced consumption, slower growth, and greater mortality. Larvae compensated for the lower digestibility of Q. alba leaves grown under elevated CO2 by increasing the efficiency of conversion of ingested food into larval mass. Despite equivalent consumption rates, larvae grew larger when they consumed Q. alba leaves grown under elevated compared with ambient CO2. Reduced consumption, slower growth rates, and increased mortality of insect larvae may explain lower total leaf damage observed previously in plots in this forest exposed to elevated CO2. By subtly altering aspects of leaf chemistry, the ever-increasing concentration of CO2 in the atmosphere will change the trophic dynamics in forest ecosystems.     

Forest Fragmentation Drives the Loss of Insectivorous Birds and an Associated Increase in Herbivory

Insectivorous birds are known to play a decisive role for the natural control of herbivorous insects. Thus, they enhance the growth, reproduction, and survival of plant individuals and in the long‐term benefit plant regeneration. However, particularly in the tropics, forest fragmentation has been suggested to cause a loss of insectivorous birds. Yet, it is unclear whether this hampers the trophic control of herbivorous insects with potential consequences for plants. Therefore, we investigated the effect of increasing forest fragmentation on tritrophic interactions between insectivorous birds, herbivorous insects, and plants in a subtropical forest landscape, South Africa. We monitored the community composition of birds and estimated insectivorous bird abundances along a gradient of forest fragmentation. In the same sites, we installed bird exclosures on a common plant species (Englerophytum natalense) to assess effects of the trophic control of insectivorous birds on herbivorous insects and leaf area loss (LAL). Forest fragmentation strongly shaped the functional composition of bird communities, particularly through a loss of forest‐dependent insectivorous birds. Moreover, LAL was higher within bird exclosures than on control branches and increased with increasing forest fragmentation on the control branches. Altogether, forest fragmentation seems to hamper the trophic control of herbivorous insects by insectivorous birds through changes in the community composition. This, in turn, may interfere with tritrophic interactions and ecological processes. Thus, conservation efforts aiming at enhancing the natural control of herbivorous insects should focus on the maintenance of continuous indigenous forests that are well‐connected to smaller forest fragments on the landscape scale.