Leaf-chewing Insects Research Articles

Application of Entomopathogenic nematodes Suppresses the Root Knot Nematodes, Agrotis ipsilon, Spodoptera litura in Tobacco

Application of Entomopathogenic nematodes (EPNs) is an efficient and novel alternative for management of plant-parasitic nematode and other insects to improve the productivity of tobacco and to protect the tobacco from the disturbing of underground and leaf chewing insects. In the present work, laboratory and field trials were conducted to control root-knot nematode (M. javanica) by using EPNs and the symbiotic bacterial, and the effect of the application was evaluated in tobacco plant. Approximately after inoculation of sixty days, egg masses and galls on tobacco root and agronomic characters of tobacco were calculated. Results showed that both EPNs and the symbiotic bacterium affected the reproduction of M. javanica population and the gall, egg masses and the root-knot nematodes in the tobacco root were significant reduced. EPNs also showed suppressive effect on S. litura larvae, and Agrotis ipsilon larvae as well. It might be the first study about the use of EPNs in the control of M. javanica in tobacco plant.

Root inoculation with soil-borne microorganisms alters gut bacterial communities and performance of the leaf-chewer Spodoptera exigua.

Soil-borne microorganisms can impact leaf-chewing insect fitness by modifying plant nutrition and defence. Whether the altered insect performance is linked to changes in microbial partners of caterpillars remains unclear. We investigated the effects of root inoculation with soil bacteria or fungi on the gut bacterial community and biomass of the folivore Spodoptera exigua. We also explored the potential correlation between both parameters. We performed herbivory bioassay using leaves of tomato plants (Solanum lycopersicum), measured caterpillar weight gain and characterized the gut bacterial communities via 16S rRNA gene metabarcoding. All soil microbes modified the gut bacterial communities, but the extent of these changes depended on the inoculated species. Rhizophagus irregularis and Bacillus amyloliquefaciens had opposite effects on S. exigua weight. While plant inoculation with the fungus influenced gut bacterial diversity, B. amyloliquefaciens also affected the community composition. A reduced abundance of two S. exigua enterococcal symbionts correlated with decreased insect biomass. Our results show that soil microorganisms can induce plant-mediated changes in the gut bacterial community of foliar-feeding caterpillars. We propose that the impact of these alterations on insect performance might rely on specific adaptations within the gut bacteria, rather than solely on the occurrence of changes.

Differential induction of JA/SA determines plant defense against successive leaf-chewing and phloem-feeding insects

Differential induction of JA/SA determines plant defense against successive leaf-chewing and phloem-feeding insects

Biochemical Responses in Populus tremula: Defending against Sucking and Leaf-Chewing Insect Herbivores.

The main biochemical traits were estimated in poplar leaves under biotic attack (aphids and spongy moth infestation). Changes in the abundance of bioactive compounds in genetically uniform individuals of European aspen (Populus tremula), such as proline, polyphenolic compounds, chlorophylls a and b, and volatile compounds, were determined between leaves damaged by sucking insects (aphid-Chaitophorus nassonowi) and chewing insects (spongy moth-Lymantria dispar) compared to uninfected leaves. Among the nine analyzed phenolic compounds, only catechin and procyanidin showed significant differences between the control leaves and leaves affected by spongy moths or aphids. GC-TOF-MS volatile metabolome analysis showed the clear separation of the control versus aphids-infested and moth-infested leaves. In total, the compounds that proved to have the highest explanatory power for aphid-infested leaves were 3-hexenal and 5-methyl-2-furanone, and for moth-infested leaves, trans-α-farnesene and 4-cyanocyclohexane. The aphid-infested leaves contained around half the amount of chlorophylls and twice the amount of proline compared to uninfected leaves, and these results evidenced that aphids influence plant physiology more than chewing insects.

Simultaneous Impact of Rhizobacteria Inoculation and Leaf-Chewing Insect Herbivory on Essential Oil Production and VOC Emissions in Ocimum basilicum.

Inoculation with rhizobacteria and feeding by herbivores, two types of abiotic stress, have been shown to increase the production of secondary metabolites in plants as part of the defense response. This study explored the simultaneous effects of inoculation with Bacillus amyloliquefaciens GB03 (a PGPR species) and herbivory by third-instar Spodoptera frugiperda larvae on essential oil (EO) yield and volatile organic compound (VOC) emissions in Ocimum basilicum plants. The density of glandular trichomes was also examined, given that they are linked to EO production and VOC emission. Herbivory increased EO content, but inoculation on its own did not. When combined, however, the two treatments led to a 10-fold rise in EO content with respect to non-inoculated plants. VOC emissions did not significantly differ between inoculated and non-inoculated plants, but they doubled in plants chewed by the larvae with respect to their undamaged counterparts. Interestingly, no changes were observed in VOC emissions when the treatments were tested together. In short, the two biotic stressors elicited differing plant defense responses, mainly when EO was concerned. PGPR did not stimulate EO production, while herbivory significantly enhanced it and increased VOC emissions. The combined treatment acted synergistically, and in this case, PGPR inoculation may have had a priming effect that amplified plant response to herbivory. Peltate trichome density was higher in inoculated plants, those damaged by larvae, and those subjected to the combination of both treatments. The findings highlight the intricate nature of plant defense mechanisms against various stressors and hint at a potential strategy to produce essential oil through the combined application of the two stressors tested here.

Rickettsia transmission from whitefly to plants benefits herbivore insects but is detrimental to fungal and viral pathogens.

Most insects are associated with symbiotic bacteria in nature. These symbionts play important roles in the life histories of herbivorous insects by impacting their development, survival, reproduction as well as stress tolerance. Rickettsia is one important symbiont to the agricultural pest whitefly Bemisia tabaci. Here, for the first time, we revealed that the persistence of Rickettsia symbionts in tomato leaves significantly changed the defense pattern of tomato plants. These changes benefit both sap-feeding and leaf-chewing herbivore insects, such as increasing the fecundity of whitefly adults, enhancing the growth and development of the noctuid Spodoptera litura, but reducing the pathogenicity of Verticillium fungi and TYLCV virus to tomato plants distinctively. Our study unraveled a new horizon for the multiple interaction theories among plant-insect-bacterial symbionts.

Herbivore-Dependent Induced Volatiles in Pear Plants Cause Differential Attractive Response by Lacewing Larvae.

Biological control may benefit from the behavioral manipulation of natural enemies using volatile organic compounds (VOCs). Among these, herbivore-induced plant volatiles (HIPVs) provide potential tools for attracting or retaining predators and parasitoids of insect pests. This work aimed to characterize the VOCs emitted by pear plants in response to attack by Cacopsylla bidens (Hemiptera: Psyllidae), a major pest in pear orchards, to compare these with VOCs induced by a leaf chewing insect, Argyrotaenia sphaleropa (Lepidoptera: Tortricidae), and to evaluate the behavioral response of Chrysoperla externa (Neuroptera: Chrysopidae) to HIPVs from pear plants damaged by either herbivore. The results demonstrated that plants damaged by the pear psylla emitted VOC blends with increased amounts of aliphatic aldehydes. Leafroller damage resulted in increased amounts of benzeneacetonitrile, (E)-4,8-dimethylnona-1,3,7-triene, β-ocimene and caryophyllene. In olfactometer bioassays, larvae of C. externa were attracted to herbivore-damaged plants when contrasted with undamaged plants. When plant odors from psylla-damaged were contrasted with those of leafroller-damaged plants, C.externa preferred the former, also showing shorter response lag-times and higher response rates when psylla-damaged plants were present. Our results suggest that pear plants respond to herbivory by modifying their volatile profile, and that psylla-induced volatiles may be used as prey-specific chemical cues by chrysopid larvae. Our study is the first to report HIPVs in pear plants attacked by C. bidens, as well as the attraction of C. externa to psyllid-induced volatiles.

A neotropical mistletoe influences herbivory of its host plant by driving changes in the associated insect community.

Parasitic plants are important sources of stress and can strongly impact their host plants through direct and indirect associations with other herbivores and their associated organisms. In the tropics, mistletoes are frequent parasitic plants, influencing different trophic levels involved with the host plant. Here, we investigated the direct and indirect influences of multiple partners involved in interactions between the mistletoe Psittachantus robustus and its host tree, Vochysia thyrsoidea. More specifically, we assessed if the presence of the mistletoe modified herbivory levels of its host by altering the diversity of associated insects. We found that insect feeding guild modulated mistletoe influence on insect community, and there were fewer species and individuals of leaf-chewing insects in parasitized than non-parasitized trees. Despite this decrease in leaf-chewing insects, there were increased levels of herbivory in parasitized plants. Mistletoes' presence did not influence the hemipteran sap-sucking insects, but this herbivore guild directly responded to the abundance of their associated ants. Overall, our study found empirical support for the crucial role of mistletoes on their host-associated organisms, ultimately shaping the herbivory levels of their tree hosts. By exposing the distinct effects of the different partners involved, our results shed light on the intricated interactions mediated by parasitic plants, opening the path for new investigations.

CRISPR/Cas9-Mediated Targeted Mutagenesis of GmUGT Enhanced Soybean Resistance Against Leaf-Chewing Insects Through Flavonoids Biosynthesis.

Leaf-chewing insects are important pests that cause yield loss and reduce seed quality in soybeans (Glycine max). Breeding soybean varieties that are resistant to leaf-chewing insects can minimize the need for insecticide use and reduce yield loss. The marker gene for QTL-M, Glyma.07g110300 (LOC100775351) that encodes a UDP-glycosyltransferase (UGT) is the major determinant of resistance against leaf-chewing insects in soybean; it exhibits a loss of function in insect-resistant soybean germplasms. In this study, Agrobacterium-mediated transformation introduced the CRISPR/Cas9 expression vector into the soybean cultivar Tianlong No. 1 to generate Glyma.07g110300-gene mutants. We obtained two novel types of mutations, a 33-bp deletion and a single-bp insertion in the GmUGT coding region, which resulted in an enhanced resistance to Helicoverpa armigera and Spodoptera litura. Additionally, overexpressing GmUGT produced soybean varieties that were more sensitive to H. armigera and S. litura. Both mutant and overexpressing lines exhibited no obvious phenotypic changes. The difference in metabolites and gene expression suggested that GmUGT is involved in imparting resistance to leaf-chewing insects by altering the flavonoid content and expression patterns of genes related to flavonoid biosynthesis and defense. Furthermore, ectopic expression of the GmUGT gene in the ugt72b1 mutant of Arabidopsis substantially rescued the phenotype of H. armigera resistance in the atugt72b1 mutant. Our study presents a strategy for increasing resistance against leaf-chewing insects in soybean through CRISPR/Cas9-mediated targeted mutagenesis of the UGT genes.

Spatiotemporal Distribution of Herbivorous Insects Along Always-Green Mountaintop Forest Islands

Insects make up the bulk of terrestrial diversity and about half of insect species are herbivores that have direct relationships with their host plants and are the basis of the entire food chain, on which wildlife and humanity depend. Some herbivorous insect traits, such as their spatio-temporal distribution, are especially relevant in the current scenario of global changes, which are more pronounced in high elevation areas, helping to improve the effectiveness of conservation actions. Here we evaluated the influence that different spatiotemporal scales have on three free-feeding herbivorous insect guilds (fluid-feeding, leaf-chewing, and xylophagous insects) in montane forest islands immersed in a grassland-dominated matrix (campo rupestre). We assessed whether species turnover or nestedness was the main component determining both spatial and temporal species composition variation (β-diversity) of the herbivorous insect community. We also checked the temporal effect on herbivorous insect guilds composition between vertical strata. We sampled herbivorous insects during two summers and two winters in 14 forest islands of different sizes and shapes in a natural mountainous fragment located in southeastern Brazil. A total of 6597 herbivorous insects representing 557 morphospecies were sampled, 290 of which were fluid-feeding, 147 leaf-chewing and 120 xylophagous insects. We found a main contribution of time scale in the organization of the herbivorous insect composition sampled in this study, mainly by turnover, with small differences among guilds. Additionally, we could see that climate determined the local variation of species, corroborating that we have a highly variable always-green system over space and time where the understory community varies less in comparison to the canopy community. Our findings suggest that long-term ecological research on herbivorous community structure in relation to climatic variation is a key element for future investigations, which can be decisive for the conservation of herbivorous insect communities. We also suggest that the effects of anthropogenic pressures must be monitored in this system, since these forest islands may serve as warming refuges in a fragmented landscape holding an invaluable diversity of species that, without these old-growth forest reservoirs, would be doomed to disappear.

Metabolomics Differences of Glycine max QTLs Resistant to Soybean Looper.

Quantitative trait loci (QTLs) E and M are major soybean alleles that confer resistance to leaf-chewing insects, and are particularly effective in combination. Flavonoids and/or isoflavonoids are classes of plant secondary metabolites that previous studies agree are the causative agents of resistance of these QTLs. However, all previous studies have compared soybean genotypes that are of dissimilar genetic backgrounds, leaving it questionable what metabolites are a result of the QTL rather than the genetic background. Here, we conducted a non-targeted mass spectrometry approach without liquid chromatography to identify differences in metabolite levels among QTLs E, M, and both (EM) that were introgressed into the background of the susceptible variety Benning. Our results found that E and M mainly confer low-level, global differences in distinct sets of metabolites. The isoflavonoid daidzein was the only metabolite that demonstrated major increases, specifically in insect-treated M and EM. Interestingly, M confers increased daidzein levels in response to insect, whereas E restores M’s depleted daidzein levels in the absence of insect. Since daidzein levels do not parallel levels of resistance, our data suggest a novel mechanism that the QTLs confer resistance to insects by mediating changes in hundreds of metabolites, which would be difficult for the insect to evolve tolerance. Collective global metabolite differences conferred by E and M might explain the increased resistance of EM.

Pith-specific lignification in Nicotiana attenuata as a defense against a stem-boring herbivore.

Plants have developed tissue-specific defense strategies in response to various herbivores with different feeding habits. Although defense responses to leaf-chewing insects have been well studied, little is known about stem-specific responses, particularly in the pith, to stem-boring herbivores. To understand the stem-specific defense, we first conducted a comparative transcriptomic analysis of the wild tobacco Nicotiana attenuata before and after attack by the leaf-chewing herbivore Manduca sexta and the stem borer Trichobaris mucorea. When the stem-boring herbivore attacked, lignin-associated genes were upregulated specifically in the inner parenchymal cells of the stem, the pith; lignin also accumulated highly in the attacked pith. Silencing the lignin biosynthetic gene cinnamyl alcohol dehydrogenase enhanced the performance of the stem-boring herbivore but had no effect on the growth of the leaf-chewing herbivore. Two-dimensional nuclear magnetic resonance results revealed that lignified pith contains feruloyltyramine as an unusual lignin component in the cell wall, as a response against stem-boring herbivore attack. Pith-specific lignification induced by the stem-boring herbivore was modulated by both jasmonate and ethylene signaling. These results suggest that lignin provides a stem-specific inducible barrier, protecting plants against stem-boring insects.

Genotypic Variation in Plant Traits, Chemical Defenses, and Resistance Against Insect Herbivores in Avocado (Persea americana) Across a Domestication Gradient

Domestication promotes divergence between wild and cultivated plants. The “plant domestication-reduced defense” hypothesis proposes that cultivated plants have lower chemical defenses and resistance against herbivores than their wild counterparts. Yet, the effects of domestication on the interactions between perennial crops and insect herbivores have not been well-documented. In this study, we hypothesized that domesticated avocado (Persea americana) has lowered resistance against insect herbivores. To test this hypothesis, we measured variation in plant traits (fruit and seed size, seed germination, and plant growth), chemical defenses (total phenolics), and resistance against two leaf-chewing insect herbivores—a specialist (Copaxa multifenestrata) and a generalist (Spodoptera frugiperda)—among seven avocado genotypes across a domestication gradient: wild (ancestral) genotypes, five (intermediate) landraces (“Blanco,” “Lonjas,” “Vargas,” “Zarcoli,” and “Rodolfo”), and the cultivated (modern) “Hass.” Our results showed that seeds from “Hass” have a lower germination rate and slower growth and have shorter fruits and seeds than the landraces and wild genotypes. “Hass” leaves also had lower amounts of total phenolics than the landraces; however, no differences were found between “Hass” and the wild genotypes. There was no effect of genotype on larval mass gained for both herbivores. However, C. multifenestrata had longer larval longevity on “Hass” and the wild genotypes, whereas S. frugiperda larval longevity showed no differences among genotypes. Moreover, C. multifenestrata inflicted more damage on “Hass,” whereas S. frugiperda inflicted more damage on “Lonjas” than on the other genotypes. In general, bigger fruit and seeds were positively correlated with plant size and phenolic content, and total phenolics were positively correlated with S. frugiperda and negatively correlated with C. multifenestrata larval performance. However, despite the genotypic variation in plant traits, phenolic content, and resistance against two herbivores with different levels of specialization, there was no clear support for the “plant domestication-reduced defense” hypothesis in avocado.

Invasive earthworms reduce chemical defense and increase herbivory and pathogen infection in native trees.

Recent research shows that earthworms can alter defense traits of plants against herbivores and pathogens by affecting soil biochemistry. Yet, the effects of invasive earthworms on defense traits of native plants from previously earthworm‐free ecosystems as well as the consequences for multitrophic interactions are virtually unknown.Here we use a combination of an observational study and a complementary experimental study to investigate the effects of invasive earthworms on leaf defense traits, herbivore damage and pathogen infection in two poplar tree species (Populus balsamifera and Populus tremuloides) native to North American boreal forests.Our observational study showed that earthworm invasion was associated with enhanced leaf herbivory (by leaf‐chewing insects) in saplings of both tree species. However, we only detected significant shifts in the concentration of chemical defense compounds in response to earthworm invasion for P. balsamifera. Specifically, leaf phenolic concentrations, including salicinoids and catechin, were lower in P. balsamifera from earthworm‐invaded sites.Our experimental study confirmed an earthworm‐induced reduction in leaf defense levels in P. balsamifera for one of the defense compounds, tremulacin. The experimental study additionally showed that invasive earthworms reduced leaf dry matter content, potentially increasing leaf palatability, and enhanced susceptibility of trees to infection by a fungal pathogen, but not to aphid infestation, in the same tree species. Synthesis. Our results show that invasive earthworms can decrease the concentrations of some chemical defense compounds in P. balsamifera, which could make them susceptible to leaf‐chewing insects. Such potential impacts of invasive earthworms are likely to have implications for tree survival and competition, native tree biodiversity and ecosystem functioning.

Impact of Soil Rhizobacteria Inoculation and Leaf-Chewing Insect Herbivory on Mentha piperita Leaf Secondary Metabolites.

Secondary metabolites commonly play important physiological roles in plants and can be modified quantitatively and qualitatively by exposure to biotic and abiotic interactions. Plant growth promoting rhizobacteria (PGPR) and herbivory induce systemic resistance. In the present study, we analyzed the induction of secondary metabolites in peppermint plants in response to chewing insect herbivory on PGPR-inoculated Mentha piperita plants. The secondary metabolites of M. piperita plants were increased when plants were inoculated with PGPR and also exposed to caterpillar herbivory. It was found that the total essential oil yield in inoculated plants with insect damage was ~2.6-fold higher than in controls. The yield was similar to that of plants either damaged by insects or inoculated, indicating that there was no synergism. The same trend was observed for phenolic compounds. In contrast, VOC emissions were significantly higher in plants infested by insects, independent of whether they were inoculated. Insect damaged plants had 5.5 times higher monoterpene emissions than control plants, and ~ 2-fold higher emissions than on PGPR-inoculated plants without insects. To gain a better understanding of how herbivory on PGPR-inoculated plants can cause an increase in secondary metabolites of peppermint, we examined changes in plant defense hormones in inoculated plants after herbivory. We found that the combination of both treatments increased the endogenous jasmonic and salicylic acid levels to the same extent as in plants only inoculated or only insect-damaged. Because different interactions can alter the phytochemistry of plants such as M. piperita, this topic is both ecologically and economically relevant.

Silicon Alters Leaf Surface Morphology and Suppresses Insect Herbivory in a Model Grass Species.

Grasses accumulate large amounts of silicon (Si) which is deposited in trichomes, specialised silica cells and cell walls. This may increase leaf toughness and reduce cell rupture, palatability and digestion. Few studies have measured leaf mechanical traits in response to Si, thus the effect of Si on herbivores can be difficult to disentangle from Si-induced changes in leaf surface morphology. We assessed the effects of Si on Brachypodium distachyon mechanical traits (specific leaf area (SLA), thickness, leaf dry matter content (LDMC), relative electrolyte leakage (REL)) and leaf surface morphology (macrohairs, prickle, silica and epidermal cells) and determined the effects of Si on the growth of two generalist insect herbivores (Helicoverpa armigera and Acheta domesticus). Si had no effect on leaf mechanical traits; however, Si changed leaf surface morphology: silica and prickle cells were on average 127% and 36% larger in Si supplemented plants, respectively. Prickle cell density was significantly reduced by Si, while macrohair density remained unchanged. Caterpillars were more negatively affected by Si compared to crickets, possibly due to the latter having a thicker and thus more protective gut lining. Our data show that Si acts as a direct defence against leaf-chewing insects by changing the morphology of specialised defence structures without altering leaf mechanical traits.

An ERF transcription factor enhances plant resistance to Myzus persicae and Spodoptera litura

Aphids are sap-sucking insects and are pests that cause substantial damage to many crops worldwide. Spodoptera litura is a leaf-chewing insect and poses a severe threat to many species of vegetable and cotton plants. In this paper, we studied an ethylene-responsive factor (ERF) that enhances plant resistance to sap-sucking and leaf-chewing insects. ERF11-like was obtained from Chinese cabbage via reverse transcription-polymerase chain reaction and was termed BrERF11b. The putative protein encoded by the BrERF11b gene includes 168 amino acid residues and contains an EAR motif. The plasmid pBI121-BrERF11b was constructed and transformed into tobacco plants (Nicotiana tabacum cv. K326 and N. benthamiana) by the Agrobacterium-mediated transformation method. The function of the BrERF11b gene in the resistance of the transgenic plants to Myzus persicae and S. litura was assayed. The results showed that the weights of the S. litura larvae fed the transgenic plant line were significantly reduced from 28 to 76% compared to those of the S. litura larvae fed the control plants (wild type), and the transgenic plant line KLine12 significantly reduced the survival rate of S. litura by 62.5%. The colony growth of the apteral aphids fed the transgenic plant line was significantly reduced from 22.5 to 63.4% compared to that of the apteral aphids fed the control plants. Compared to control plants, the transgenic plant line significantly reduced the survival rate of the aphids from 46 to 55%. These results first indicate that the BrERF11b gene significantly enhances plant resistance to S. litura and M. persicae.

Quantitative assessment of plant-arthropod interactions in forest canopies: A plot-based approach.

Research on canopy arthropods has progressed from species inventories to the study of their interactions and networks, enhancing our understanding of how hyper-diverse communities are maintained. Previous studies often focused on sampling individual tree species, individual trees or their parts. We argue that such selective sampling is not ideal when analyzing interaction network structure, and may lead to erroneous conclusions. We developed practical and reproducible sampling guidelines for the plot-based analysis of arthropod interaction networks in forest canopies. Our sampling protocol focused on insect herbivores (leaf-chewing insect larvae, miners and gallers) and non-flying invertebrate predators (spiders and ants). We quantitatively sampled the focal arthropods from felled trees, or from trees accessed by canopy cranes or cherry pickers in 53 0.1 ha forest plots in five biogeographic regions, comprising 6,280 trees in total. All three methods required a similar sampling effort and provided good foliage accessibility. Furthermore, we compared interaction networks derived from plot-based data to interaction networks derived from simulated non-plot-based data focusing either on common tree species or a representative selection of tree families. All types of non-plot-based data showed highly biased network structure towards higher connectance, higher web asymmetry, and higher nestedness temperature when compared with plot-based data. Furthermore, some types of non-plot-based data showed biased diversity of the associated herbivore species and specificity of their interactions. Plot-based sampling thus appears to be the most rigorous approach for reconstructing realistic, quantitative plant-arthropod interaction networks that are comparable across sites and regions. Studies of plant interactions have greatly benefited from a plot-based approach and we argue that studies of arthropod interactions would benefit in the same way. We conclude that plot-based studies on canopy arthropods would yield important insights into the processes of interaction network assembly and dynamics, which could be maximised via a coordinated network of plot-based study sites.

Declines and Resilience of Communities of Leaf Chewing Insects on Missouri Oaks Following Spring Frost and Summer Drought

Extreme weather events dramatically impact populations of individual insect species but the consequences of such events for entire insect communities are not well documented. We present evidence that late spring frosts and summer drought negatively affect the community of insects found on Missouri oaks (Quercus alba and Q. velutina), amounting to a 23-186 fold decrease depending on the oak species, insect size and feeding guild, and the specific weather event. Spring faunas required three years or more following spring frosts to reach pre-event levels, whereas summer faunas were more resilient, rebounding within 1-2 years. The impact of summer drought on leaf tying caterpillars, however, lasted longer than that of the spring frost; it was five years before numbers of leaf ties reached pre-drought levels. Smaller species of leaf tying Lepidoptera took longer to recover than larger species following the drought. Overall, we found no evidence for a general decline in abundance, even a modest one, during the 20 years of study of faunas on oak trees in southeastern Missouri. However, the effects of drought reported here are likely to increase with time as the climate in the Midwest U.S. is only expected to become warmer and drier during the summer months. Similarly, the risk of late season frost damage to trees is expected to increase with predicted earlier onset of spring. Understanding the impact of such weather events on insect communities influences our ability to predict how habitat and landscape management, or lack thereof, will influence future patterns of insect abundance and diversity.

Caterpillars Cooperate to Overcome Plant Glandular Trichome Defenses

One common advantage proposed for group-living in animals is social facilitation of feeding, by which food acquisition by an individual is made easier by feeding neighbours. The present paper provides an explanation of social facilitation of feeding in a gregarious leaf-chewing insect, demonstrating how caterpillars cooperate to overwhelm plant trichome defenses and pierce a hole in the leaf on which they feed collectively. Specifically, it tests the hypotheses that the caterpillars feed collectively, that the glandular trichomes of Solanum acerifolium are effective defenses against the specialist caterpillar Mechanitis menapis, and that feeding by early-instar M. menapis is socially facilitated in the presence of glandular trichomes. A field survey showed that Mechanitis menapis on trichome–bearing plants feed collectively on the abaxial surface of leaves during the first larval instars. In a lab experiment comparing feeding on control and ethanol-washed leaves, caterpillars on the glandular-trichome-free washed sides of leaves initiated feeding sooner and had higher survival rates, suggesting that glandular trichomes are effective defenses. Behavioural observations showed that feeding is socially facilitated in response to glandular trichomes: caterpillars on the washed sides of leaves were more likely to begin feeding alone and initiated more separate feeding sites, suggesting that caterpillars are less able to initiate feeding independently in the presence of glandular trichomes. These results demonstrate a novel mechanism of cooperation among folivores, showing how they can benefit from grouping to tackle food sources that would be unavailable to isolated individuals. This study thus demonstrates that social facilitation of feeding extends to insect folivores, a hyper-diverse and abundant feeding guild.