Spruce budworm growth,development and food utilization on young and old balsam fir trees

Laboratory rearing of spruce budworm, Choristoneura fumiferana, in conjunction with field rearing, gravimetric analyses, a transfer experiment, and foliage chemical analyses at six dates during the period of budworm feeding activity indicated that the age of balsam fir, Abies balsamea, trees (70-year-old mature trees or 30-year-old juvenile trees) affected tree suitability for the spruce budworm via the chemical profile of the foliage. Insects reared on old trees had greater survival and pupal weight, shorter development times, and caused more defoliation than those reared on young trees. Young trees were more suitable for the development of young larvae (instars 2–5), while old trees were more suitable for the development of older, sixth-instar larvae. These results were confirmed by the laboratory transfer experiment. Young larvae fed foliage from young trees had higher relative growth rates (RGR), digestibility (AD), and efficiency of conversion of ingested foliage (ECI) than those fed foliage from old trees. These differences appeared to be related to the high N:tannins ratio, and the high contents of P present in young trees during the development of the young larvae. Old larvae fed foliage from old trees had higher relative growth rates, relative consumption rates (RCR), and digestibility of the foliage than those fed foliage from young trees. The high digestibility of the foliage of old trees was compensated for by a lower efficiency of conversion of digested food (ECD), which in turn resulted in no significant effect of tree age on the efficiency of conversion of ingested foliage by old larvae. The low relative consumption rate of old larvae fed foliage from young trees appeared to be related to the low N:tannins ratio, and the high contents of bornyl acetate, terpinolene, and °-3-carene present in young trees during the budworm sixth instar. Variations in these compounds in relation to tree age may serve as mechanisms of balsam fir resistance to spruce budworm by reducing the feeding rate of sixth instar larvae.     

Nuclear polyhedrosis virus as a biological control agent for Malacosoma americanum (Lepidoptera: Lasiocampidae)

In addition to damaging trees, the eastern tent caterpillar is implicated in early fetal loss and late‐term abortion in horses. In a field study, we evaluated the potential biological control of the caterpillar using eastern tent caterpillar nuclear polyhedrosis virus (ETNPV), a naturally occurring virus that is nearly species‐specific. Egg masses were hatched and second instar larvae were fed virus‐inoculated foliage to propagate the virus in vivo. Then, a viral pesticide was formulated at concentrations of 10⁴, 10⁶ and 10⁸ polyhedral inclusion bodies per ml. The pesticide was applied to foliage on which second, third and fourth instar caterpillars were feeding. When the majority of surviving larvae reached the sixth instar, colonies were collected and the surviving caterpillars counted. Mean numbers of surviving caterpillars per treatment were compared via 95% bootstrap confidence intervals. The data indicate second instar caterpillars were highly susceptible to the virus, but only at the highest concentration tested. Third instar caterpillars were also somewhat susceptible to high virus concentrations, while fourth instar caterpillars were fairly resistant. Our data provide the strongest evidence to date that ETNPV can be propagated, harvested and refined for formulation as a biological control agent for eastern tent caterpillar. Its use on this insect may be merited in circumstances where landowners and managers need to protect trees and horses.     

Preference and performance of Lepidoptera varies with tree age in juniper woodlands

1. As trees age, they undergo significant physiological and morphological changes. Nevertheless, tree ontogeny and its impacts on herbivores are often overlooked as determinants of plant–herbivore population dynamics and the strength of plant–herbivore interactions. 2. Juniperus (Cupressaceae) is a dominant, long‐lived conifer that serves as the sole host to a specialised assemblage of caterpillars. Over the past 150 years, several juniper species in western North America have expanded their geographic occupancy at local and regional scales, which has resulted in an increase in the number of immature trees on the landscape. Using assays in the laboratory, the effects of tree ontogeny on caterpillar performance and oviposition preference for two juniper specialist caterpillars, Callophrys gryneus (Lycaenidae) and Glena quinquelinearia (Geometridae), were examined. The study considered whether responses to tree ontogeny were consistent across caterpillar species and juniper host species. 3. Tree age was found to be a reliable predictor of caterpillar performance, with caterpillars developing more quickly and growing larger when fed foliage from young trees. Differences in the phytochemical diversity between foliage from trees of different ages might help to explain observed differences in caterpillar performance. Interestingly, the specialist butterfly, C. gryneus, displayed an oviposition preference for foliage from old‐growth Juniperus osteosperma trees, despite the fact that larvae of this species performed poorly on older trees. 4. It is concluded that young juniper trees are an important resource for the specialised Lepidopteran community and that tree ontogeny is an important component of intraspecific variation, which contributes to the structure of plant–herbivore communities.     

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.     

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.     

Communication of leaf suitability by gregarious eastern tent caterpillars (Malacosoma americanum)

ABSTRACT. 1. Previous work has shown that leaf age affects recruitment trail marking by eastern tent caterpillars ( Malacosoma americanum Fabr.). Young leaves of host plants elicit trail marking to a greater degree than mature leaves.
2. Experiments were conducted to establish the relationship between the differential behavioural responses of larvae to young and mature leaves and the suitability of foliage for larval growth and survival. Foliage of black cherry ( Prunus serotina Ehrh.), a typical rosaceous host plant, was used for this comparison.
3. Larvae preferred young leaves to mature leaves in choice tests, and marked more to young leaves than to mature leaves in no-choice tests.
4. Mature leaves supported adequate growth through two larval instars of rearing, but thereafter were unsuitable for growth. Larvae fed mature leaves had lower pupal weight, poorer survival, and grew less efficiently than larvae fed young leaves.
5. The results support the hypothesis that the trail communication system of eastern tent caterpillars is an adaptation to efficiently locate leaves which are favourable for larval growth and survival.     

Colorado potato beetle (Coleoptera: Chrysomelidae) feeding, development, and survival to adulthood after continuous exposure to Bacillus thuringiensis subsp. tenebrionis-treated potato foliage from the field

Colorado potato beetle, Leptinotarsa decemlineata (Say), feeding, development, and survival to adulthood were examined after continuously exposing large larvae to Bacillus thuringiensis subsp. tenebrionis-treated potato foliage from the field. In laboratory assays, the overall consumption and the length of period to become prepupae were determined for larvae, which began as 3rd and 4th instars, that were offered potato leaf disks with naturally declining levels of B. thuringiensis residue. In small-cage field experiments, survival to adulthood and the period to adult emergence for beetles confined to potato plants treated with B. thuringiensis beginning as 3rd and 4th instars also were examined. Third instars remaining on plants after a B. thuringiensis application were unlikely to feed and 4th instars consumed only approximately 50% as much foliage as those fed untreated foliage. Many late instars subjected to B. thuringiensis-treated foliage failed to survive to adulthood; 58-83% of these beetles died during the larval stage. Reduced feeding and poor survival of late instars suggest that counts of large larvae after application do not provide a complete picture of the efficacy of the B. thuringiensis treatment. Late instar Colorado potato beetles that were exposed continually to naturally declining levels of B. thuringiensis-treated potato foliage took an average of 1.8-4.5 d longer to become prepupae and 4-8 d longer to emerge as adults compared with those provided with untreated foliage. Delayed emergence of adults that fed on B. thuringiensis-treated potatoes as late instars indicated that development was prolonged in these insects because of ingestion of a sublethal dose of B. thuringiensis.     

Studies on feeding and trophallaxis in the subterranean termite Odontotermes formosanus using rubidium chloride

Differences in feeding and trophallaxis among castes of the fungus‐cultivating higher subterranean termite Odontotermes formosanus Shiraki (Isoptera: Termitidae) were determined using rubidium (Rb) as a tracer. In the feeding study, workers and third instars fed directly on Rb‐treated filter paper and acquired the highest levels of Rb content. Although fifth and sixth instars obtained the Rb content over the marked level, the feeding ability of the caste was very poor. However, soldiers did not feed on Rb‐treated filter paper. In the trophallaxis study, worker, soldier, and fifth‐ and sixth‐instar recipients paired with Rb‐fed worker donors had a significantly higher Rb content than those paired with control donors. Results indicate that soldiers are completely dependent on workers for nutrition. Workers and fifth and sixth instars may obtain nutritional material from foraging workers. When third‐instar recipients were paired with Rb‐fed worker donors, they did not have a significantly higher Rb content than when they were paired with control donors, suggesting that the frequency of worker/third instar trophallaxis was very low. Transfer efficiency from the four trophallactic combinations ranged from 1.2 (worker donors to third‐instar recipients) to 12.9% (worker donors to soldier recipients). Accordingly, nutritional material from foraging workers could be transferred to non‐foraging workers, soldiers, and fifth and sixth instars by trophallaxis; however, third instars rarely obtain nutritional material from foraging workers, because they are capable of feeding within the nest.     

Herbivory meets fungivory: insect herbivores feed on plant pathogenic fungi for their own benefit

Plants are regularly colonised by fungi and bacteria, but plant‐inhabiting microbes are rarely considered in studies on plant–herbivore interactions. Here we show that young gypsy moth (Lymantria dispar) caterpillars prefer to feed on black poplar (Populus nigra) foliage infected by the rust fungus Melampsora larici‐populina instead of uninfected control foliage, and selectively consume fungal spores. This consumption, also observed in a related lepidopteran species, is stimulated by the sugar alcohol mannitol, found in much higher concentration in fungal tissue and infected leaves than uninfected plant foliage. Gypsy moth larvae developed more rapidly on rust‐infected leaves, which cannot be attributed to mannitol but rather to greater levels of total nitrogen, essential amino acids and B vitamins in fungal tissue and fungus‐infected leaves. Herbivore consumption of fungi and other microbes may be much more widespread than commonly believed with important consequences for the ecology and evolution of plant–herbivore interactions.     

Induced preference for host plant chemicals in the tobacco hornworm: contribution of olfaction and taste

Many herbivorous insects induce preferences for host plants. Recent work in Manduca sexta indicates that induced preferences are mediated by a “tuning” of the peripheral taste system to chemicals within host plant foliage. We tested this hypothesis by rearing caterpillars on artificial diet or potato foliage, and then examining olfactory- and taste-mediated responses to potato foliage extract. First, we confirmed earlier reports that consumption of potato foliage tunes the peripheral taste system by reducing responsiveness to glucose and increasing responsiveness to foliage extract. Second, we offered caterpillars a choice between disks treated with foliage extract (experimental) or solvent alone (control). The foliage-reared caterpillars approached and consumed the experimental disks disproportionately, whereas the diet-reared caterpillars approached and consumed both disks indiscriminately. This indicated that induced preferences involve olfaction and taste. Third, we ran choice tests with foliage-reared caterpillars deprived of either olfactory or gustatory input. Caterpillars lacking olfactory input approached both disks indiscriminately, but fed selectively on experimental disks. In contrast, caterpillars lacking gustatory input approached experimental disks selectively, but fed indiscriminately on both types of disk. We conclude that even though olfaction helps caterpillars locate potato foliage, it is the “tuned” gustatory response that ultimately mediates the induced preference.     

Inhibition of herbivory on young holly leaves: evidence for the defensive role of saponins

Summary Response of the southern red mite, Oligonychus ilicis McGregor, to young and one year old leaves of Ilex opaca Aiton was studied on three dates during the period of leaf expansion in the spring. Young foliage, which is rich in nutrients but also contains high levels of saponins, was found to be unsuitable for colonization by this oligophagous herbivore until the leaves had matured and levels of saponins had declined. Mites preferentially colonized and had higher survival and reproduction on sclerophyllous, one-year old leaves than on young leaves in early spring. High levels of saponins in young, second-flush leaves of shoots that had earlier been damaged by frost were again correlated with low mite survival in June. Laboratory preference tests with the fall webworm, Hyphantria cunea Drury, and the eastern tent caterpillar, Malacosoma americanum (F.), indicated that young, saponin-rich holly foliage is especially unpalatable to these highly polyphagous caterpillars. Fifth instar fall webworms required significantly longer to complete their development, attained lower weights, and had greater mortality on artificial diet amended with low concentrations of purified holly saponins than on control diet. These results support the hypothesis that the high levels of saponins in young holly leaves provide protection from herbovores until the leaves have matured and their structural defenses are developed.The investigation reported in this paper (No. 88-7-8-196) is in connection with a project of the Kentucky Agricultural Experiment Station and is published with the approval of the Director     

What physiological processes permit insects to eat Eucalyptus leaves?

Many species of insects eat Eucalyptus foliage despite its relatively low nutritional value and the many plant secondary metabolites (PSMs) present, for example, terpenes, phenols and formylated phloroglucinols (FPGs). Formylated phloroglucinols are a new class of PSMs that act as antifeedants for possums and koalas. What physiological processes are present that permit insects to eat eucalypt foliage and how do PSMs influence insect feeding or digestion? Some trees seem to be repeatedly infested with eucalypt‐feeding insects, possibly as a result of previous chemosensory cues remaining from parental selection of a plant. Avoidance or storage of PSMs permit jarrah leafminers (Perthida glyphopa) and sawflies (Perga sp.) to consume eucalypt foliage without dealing with the majority of these compounds. Some PSMs can be metabolized by polysubstrate membrane oxidases as found in caterpillars or sawflies that feed on eucalypts. High midgut pH may be advantageous for nutrient extraction and PSM metabolism, and midgut pH ranges between 8.5 and 8.9 for caterpillars of Hyalarcta huebneri. Plant secondary metabolites may not be absorbed as a result of the combined presence of the peritrophic matrix and endogenous surfactants. Excretion of PSMs can be as metabolites or intact compounds. Both putative metabolites and sideroxylonal‐A, an FPG, are present in the faeces of larvae of the case moth, H. huebneri. The presence of sideroxylonal‐A in the food had an effect on the presence of 5‐hydroxytryptamine (5HT) in the central nervous system of caterpillars, as larvae fed leaves with a high concentration of sideroxylonal‐A had relatively more 5HT in the brain and central nervous system ganglia than larvae fed leaves containing a low concentration. Further work is necessary to clarify how PSMs are handled by eucalypt‐feeding insects and what effect FPGs have on feeding and digestion.     

Adoption of lycaenid Niphanda fusca (Lepidoptera: Lycaenidae) caterpillars by the host ant Camponotus japonicus (Hymenoptera: Formicidae)

Caterpillars of the parasitic lycaenid butterfly are often adopted by host ants. It has been proposed that this adoption occurs because the caterpillars mimic the cuticular hydrocarbons of the host ant. This study aimed to examine whether caterpillars of the Japanese lycaenid butterfly Niphanda fusca induce adoption by mimicking their host ant Camponotus japonicus. Behavioral observations conducted in the laboratory showed that most second‐instar caterpillars were not adopted, whereas most third‐instar caterpillars were successfully adopted by host workers. A chemical comparison detected no characteristic differences in the cuticular hydrocarbon profiles between second‐ and third‐instar caterpillars. However, morphological features of the caterpillars differed between the second and third instars; third‐instar caterpillars developed exocrine glands (ant organs) such as tentacle organs and a dorsal nectary organ. These results suggest that multiple chemical signatures, not only cuticular hydrocarbons, may be important for invasion of the host ant nest.     

Shifts in developmental diet breadth of Lymantria xylina (Lepidoptera: Lymantriidae)

The moth Lymantria xylina Swinhoe (1903) (Lepidoptera: Lymantriidae) is a major defoliator of hardwood and fruit trees in Taiwan. Although the plants identified as host plants of L. xylina usually refer to plants used as food or as shelter, most of the host plant identifications have not considered the role of these hosts on larval development. This study investigated various instars feeding on different plants to assess the developmental diet breath of L. xylina. Forty-seven plant species, belonging to 25 families were used in feeding trials. Various bioassays, including first instar survival and long-term feeding trials, indicated the most suitable host plants for the different developmental stages. Results of the first instar survival trial indicated that first instars could survive only on 13 of the tested plant species. In addition, first instars could only successfully grow to pupa on seven of these 13 test plants species. To assess the developmental diet breath shifts of this moth, 38 plant species (excluding those nine plant species that the first instars did not feed upon) were fed to third and fifth instars in long-term feeding trials. Survival to pupa was noted on 12 and 13 test plant species for the third and fifth instars, respectively. In short, we found that the larvae performed differently when fed on various host plants and that the host plant range increased with the larval stage. Therefore, it is necessary to adjust the host plant range of this moth and to consider host plant breadth together with the developmental stages of caterpillars.     

Group size effects on survivorship and adult development in the gregarious larvae of Euselasia chrysippe (Lepidoptera, Riodinidae)

Caterpillars living in aggregations may derive several benefits that outweigh the costs, including better survivorship and improved growth rates. I tested whether larval group size had an effect on these two vital rates in Euselasia chrysippe. These caterpillars feed gregariously during all instars and move in processionary form over the host plant and even pupate together. There was a positive relationship between group size and larval survivorship in the field, although genetic variability was not taken into account in this experiment. Under laboratory conditions, there was also a positive relationship between group size, and larval growth rate and adult weight. This supports the hypothesis that aggregations facilitate feeding and larval growth. Single sixth instar larvae in the laboratory also had a lower survivorship than larvae in groups. These results provide further evidence of the benefits of group living for gregarious caterpillars.     

Diet mixing enhances the performance of a generalist caterpillar,Platyprepia virginalis

1. Platyprepia virginalis caterpillars are dietary generalists and feed on multiple host species within a single day. We conducted field experiments to evaluate their performance on diets consisting of only their primary food, Lupinus arboreus, or diets consisting of L. arboreus plus other acceptable host species. 2. We found that relative growth rates and rates of survival were higher when they fed on mixed diets compared to lupine only. These results were consistent with hypotheses that mixed diets provided balanced nutrition, diluted toxins, and/or allowed recovery from parasitoids, although our data did not allow us to separate these non‐exclusive explanations. 3. We assayed alkaloids in their host foliage, in the caterpillars themselves, in parasitoids within caterpillars, in food boluses passing through their guts, and in frass that they excreted. We consistently found positive assays for alkaloids in foliage and in frass but negative assays in caterpillars, parasitoids, and food boluses. This suggests that the alkaloids that they ingest are metabolised or rendered non‐reactive by unknown means during passage through the gut. We found no support for the hypothesis that mixed diets prevented caterpillars from exhausting food supplies or allowed them to sequester chemicals from their alkaloid‐containing hosts. 4. Behavioural observations revealed that previous experience influenced a caterpillar's likelihood of moving to a different host. Caterpillars that had previously fed on other hosts were more likely to move to lupine while caterpillars previously collected on lupine were equally likely to choose more lupine or a different host. 5. These results are unusual in providing a clear and consistent benefit of diet mixing in a natural field setting where multiple ecological factors act upon the caterpillars.     

Cascading effects of host size and host plant species on parasitoid resource allocation

1. The bottom‐up factors that determine parasitoid host use are an important area of research in insect ecology. Host size is likely to be a primary cue for foraging parasitoids due to its potential influence on offspring development time, the risk of multiparasitism, and host immunocompetence. Host size is mediated in part by host‐plant traits that influence herbivore growth and potentially affect a herbivore's quality as a host for parasitoids. 2. Here, we tested how caterpillar host size and host plant species influence adult fly parasitoid size and whether host size influences wasp parasitoid sex allocation. We measured the hind tibia lengths and determined the sex of wasp and fly parasitoids reared from 11 common host species of polyphagous caterpillars (Limacodidae) that were in turn reared on foliage of seven different host plant species. 3. We also tested how host caterpillar species, host caterpillar size, and host and parasitoid phenology affect how the parasitoid community partitions host resources. We found evidence that parasitoids primarily partition their shared hosts based on size, but not by host species or phenology. One index of specialisation (d′) supports our observation that these parasitoids are quite generalised within the Limacodidae. In general, wasps were reared from caterpillars collected in early instars, while flies were reared from caterpillars collected in late instars. Furthermore, for at least one species of solitary wasp, host size influenced sex allocation of offspring by ovipositing females. 4. Host‐plant quality indirectly affected the size attained by a tachinid fly parasitoid through its direct effects on the size and performance of the caterpillar host. The host plants that resulted in the highest caterpillar host performance in the absence of enemies also yielded the largest parasitoid flies, which suggests that host plant quality can cascade up to influence the third trophic level.     

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.     

Influence of intra‐tree variation in phenology and oviposition site on the distribution and performance of Ennomos subsignaria on mature sycamore maple

Abstract 1. Field surveys and a manipulative experiment were conducted to examine the hypotheses that intra‐tree heterogeneity in natural enemy activity, foliar quality (independent of phenology), or phenology influence the intra‐tree distribution and performance of Ennomos subsignaria on mature sycamore maple, Acer pseudoplatanus. 2. Ennomos subsignaria intra‐tree distribution was distinctly clumped. Egg mass density was 85% higher on the lower bole than in the crown. Most early instars were found on lower crown proximal branches while most late instars and pupae were found on lower crown distal branches. This resulted in high levels of defoliation in the lower crown, especially on proximal branches. 3. No parasitoids were reared from eggs or late‐instar larvae and only one pupa was parasitised, suggesting that preference for the bole and lower crown was not a response to parasitism. Similarly, E. subsignaria performance was not influenced by variable foliage quality (independent of phenology) within the crown. However, sycamore maple phenology had a large influence on E. subsignaria survival. More than 90% of newly emerged larvae survived to the adult stage when they fed on foliage with three pairs of leaves expanded per bud, whereas survival on younger foliage was reduced by >45%. 4. The peak period of E. subsignaria egg hatch was approximately 2 weeks after the peak period of sycamore maple budburst, which occurred acropetally. Egg hatch was closely synchronised with the availability of most suitable leaves for insect development on proximal branches of the lower crown, the location where most larvae initiated feeding. 5. The results support the phenology hypothesis and suggest that intra‐tree variation in oviposition site and host phenological development influence the intra‐tree distribution and performance of this generalist herbivore.     

Development and feeding efficiency of Malacosoma neustrium larvae reared with Quercus spp. leaves

Observations on larval development of Malacosoma neustrium were conducted both in a cork oak stand and in the laboratory by using leaves of different host trees (cork oak, holm oak and downy oak) as food source. Instars were determined using head capsule and frass measurements. In the field the larvae progressed up to the fifth instar before pupating, and the increase in head capsule width followed Dyars Rule with a rate of increase (R.I.) value of 1.74. The same number of instars was determined for the larvae reared with cork oak (R.I. = 1.73) and holm oak (R.I. = 1.70) leaves. The caterpillars reared with downy oak foliage completed larval development in five, six and seven instars and the R.I. values obtained were 1.60, 1.52 and 1.44 respectively. A lower mortality was recorded for the larvae reared on holm oak. Growth and feeding indices were determined for the larvae from the third up to the last instar. The highest leaf consumption was detected for the fifth instar larvae reared on holm oak. For the caterpillars which completed five instars before pupating, the relative consumption rate (RCR) decreased from the third up to the fifth instar: from 4.8 to 1.7 (cork oak), from 7.4 to 3.3 (holm oak) and from 14.3 to 2.1 (downy oak). The relative growth rate (RGR) was highest during the fourth stadium (0.24, 0.27 and 0.33 for larvae reared with cork oak, holm oak and downy oak leaves respectively) and decreased in the fifth instar (0.09, 0.14 and 0.14 for larvae reared with cork oak, holm oak and downy oak leaves respectively), probably because of greater expense of energy due to the approach of maturity. Feeding and growth indices could be useful to define a defoliation prediction model.