Vibration‐mediated territoriality in the warty birch caterpillar Drepana bilineata

Abstract The warty birch caterpillar Drepana bilineata produces two distinct types of vibrational signals (mandible drumming and anal scraping) during interactions with conspecifics. Vibrational signalling is characterized using standard and high‐speed videography synchronized with laser‐doppler vibrometry, and behavioural experiments test the hypothesis that signalling functions to advertise occupancy of birch (Betula) leaves. Drumming involves raising the head and striking the leaf with the sharp edges of the open mandibles. Anal scraping involves dragging a pair of specialized oar‐shaped setae against the leaf surface. Staged encounters between leaf residents and conspecific intruders result in the resident signalling, with rates increasing as the intruder moves closer. Intruders signal significantly less often than residents. Conflicts are typically resolved within a few minutes, with the resident winning in 61% of the trials, and the intruder winning in 6%. Contests that last more than 30 min are deemed ‘ties’ and comprise the remaining 33% of trials. The results support the hypothesis that vibrational signals function to advertise leaf occupancy. Vibrational communication is believed to be widespread in Drepanoidea caterpillars, but has only been described in two species to date: D. bilineata (present study) and Drepana arcuata. It is proposed that differences in territorial behaviour and signalling between these species are related to their relative investments in silk leaf mats and shelters. The proximate and ultimate bases for the evolution of vibrational communication in caterpillars are discussed.     

Understanding gregariousness in a larval Lepidopteran: the roles of host plant, predation, and microclimate

Abstract.  1. Many moth and butterfly larvae are gregarious early in development, but become solitary in late instars. This ontogenetic variation in behaviour is probably the result of temporal changes in the costs and benefits associated with gregariousness. This study provides observational and experimental evidence that, in one particular moth species, a series of different ecological factors influence larval behaviour at different times during development.
2. Field observations show that young caterpillars of the limocodid Doratifera casta form large aggregations while foraging, but that mature larvae are largely solitary.
3. A field experiment revealed that individual first to third instar larvae in larger groups develop more rapidly, but that group size had no detectable influence on survival. The developmental advantage associated with gregariousness is affected by host plant species, but not by predator exclusion, suggesting that group living in these cryptic early instar larvae promotes feeding facilitation, but does not provide individuals with protection from natural enemies.
4. Laboratory experiments revealed that aposematic fourth instar caterpillars in large groups were less likely to be attacked by a generalist insect predator than those in small groups.
5. Field observations provided no evidence that group living affects body temperature, suggesting that microclimatic factors do not favour gregariousness in this species.
6. It is concluded that gregariousness in D. casta confers at least two different advantages on larvae at different stages early in development, but that these advantages disappear, or are outweighed by costs associated with intraspecific competition, in final instars.     

Epizootics and behavioral alteration in the arctiid caterpillar Chionarctia nivea (Lepidoptera: Arctiidae) caused by an entomopathogenic fungus, Entomophaga aulicae (Zygomycetes: Entomophthorales)

We observed epizootics and behavioral alteration in the arctiid caterpillar Chionarctia nivea after infection by an entomopathogenic fungus, Entomophaga aulicae, in April at a riverbank in Kyoto Prefecture, central Japan. The density of arctiid cadavers infected with E. aulicae was 1.31 individuals/m². The critically ill caterpillars crawled up the dead stems of grasses and herbs such as the common reed Phragmites communis and the Japanese mugwort Artemisia indica var. maximowiczii to die at and near the highest parts of the stems, while the healthy larvae usually wandered on the ground and fed on the leaves of small herbs. This behavioral difference could be caused by infection with E. aulicae to enhance dispersal of conidia by active discharge in Entomophthorales with the aid of wind and rainfall.     

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.     

The systematics and biology of the Costa Rican species of parasitic wasps in the Thyreodon genus-group (Hymenoptera: Ichneumonidae)

This paper presents an eco‐taxonomic study of the Neotropical representatives of the Thyreodon genus‐group (Hymenoptera: Ichneumonidae) present in Costa Rica, i.e. species of the genera Thyreodon and Rhynchophion. These ichneumonids are koinobiont endoparasitoids of the larvae of Sphingidae and Saturniidae. Intensive sampling in Costa Rica, conducted over 15 years, has revealed the presence of 23 species (12 of which are described here as new –Rhynchophion woodi, Thyreodon woodleyi, T. papei, T. whitfieldi, T. deansi, T. walkerae, T. sharkeyi, T. zitaniae, T. delvarei, T. schauffi, T. darlingi and T. carmeani), whereas earlier studies recognized only nine species in the same area. Several of the additional species we found are simply very scarce and thus unlikely to be encountered except by the type of intensive survey that, in the tropics, has only been conducted in Costa Rica. In some other cases, it has become apparent that more than one species has previously been confused under a single name. Thyreodon rufothorax Cameron is shown to be morphologically and biologically distinct from T. atriventris (Cresson), with which it has long been synonymized, and the relatively well‐known, chromatically distinctive species ‘T. laticinctus Cresson’ and ‘T. morosus Smith’ are both shown to be complexes of sibling species. Reared series have been essential in facilitating the separation of the species in these complexes. An illustrated key is provided to separate all taxa using simple morphological characters. Fourteen of the 23 species have been reared, and all have been found to be restricted to one or a few species of hosts, species of Thyreodon primarily on macroglossine Sphingidae (with one species on Saturniidae) and species of Rhynchophion on sphingine sphingids of the genus Manduca. No two species of these ichneumonids attack the same host caterpillar species, and not all species of macroglossines present in the study area are attacked by species in this genus‐group. Most species are apparently rather rare. Many have only been collected by rearing on a few occasions, other species have been collected at light (six in total) or in Malaise traps (ten in total), or by hand‐netting (20 in total), but no one method collected all of the species present in the principal study area, the Area Conservacion de Guanacaste. The two New World genera Rhynchophion and Thyreodon are found to be related to the Old World genera Dictyonotus and Euryophion, the former of which comprises species that are also sphingid parasitoids, whereas species of the latter attack the caterpillars of other large bombycoid moths, Saturniidae and Eupterotidae. A cladistic analysis shows that Thyreodon is the sister‐lineage to Dictyonotus, and Euryophion is the most basal lineage in the genus‐group. © 2004 The Linnean Society of London, Zoological Journal of the Linnean Society, 2004, 141 , 297–351.     

Temporal and developmental aspects of diet-induced food preferences in larvae of the tobacco hornworm, Manduca sexta

Diet‐induced changes in food preferences in an oligophagous caterpillar were studied in order to characterize the conditions under which this induction occurs. The time course of acquisition and extinction of induced food preferences by larvae of the tobacco hornworm, Manduca sexta (Johan.) (Lepidoptera, Sphingidae), was examined by varying the food and duration of larval food experience. Larvae were given feeding experience with the host plant tomato, Lycopersicon esculentum (Mill.) (Solanaceae), or the acceptable non‐host plant cowpea, Vigna unguiculata (Walp.) (Fabaceae), or switched from one to the other during different instars. Food preferences for V. unguiculata by the fourth or fifth instar larvae were measured individually in two‐choice tests with discs of V. unguiculata and moist filter paper. The acquisition or extinction of an induced food preference for V. unguiculata was indicated by larvae preferring V. unguiculata to filter paper or the reverse, respectively. Results showed that: (1) the period required for the acquisition of induced preference for V. unguiculata can be short (36 h), (2) food experience in either the third or fourth instar period is sufficient, and (3) the most recent feeding experience appears to be important. In contrast, the period required for extinction of induced preference for V. unguiculata appeared to be longer (1–3 instars), and both the most recent feeding experience and total duration of larval experience with the inducing food seem to play a role. Experience is not restricted to a particular instar period to acquire or extinguish such an induced food preference. The induced food preference for V. unguiculata was not very rigid and could be reversed by having one instar of feeding experience on L. esculentum. The findings indicate that diet‐induced food preferences in M. sexta contains elements of habituation and associative learning, but do not support food imprinting and induction of oligophagy.     

In vitro and in vivo host range of Anticarsia Gemmatalis multiple nuclear polyhedrosis virus

Summary A clone of the wild type (wt) Anticarsia gemmatalis multiple nuclear polyhedrosis virus AgMNPV, derived from a geographical isolate (Hondrina, Brazil) and designated AgMNPV-CL4-3A1, was used to determine the host range of this virus in six established lepidopteran cell lines: Anticarsia gemmatalis (BCIRL-AG-AM1), Helicoverpa zea (BCIRL-HZ-AM1), Heliothis virescens (BCIRL-HV-AM1), Helicoverpa armigera (BCIRL-HA-AM1), Trichoplusia ni (TN-CL1), Bombyx mori (BMN), and a coleopteran cell line Anthonomus grandis (BRL-AG-1). In addition, the in vivo host range of this clone was also assayed in larvae of Helicoverpa zea, Heliothis virescens, Trichoplusia ni, and the homologous species Anticarsia gemmatalis by probit analysis. On the basis of temporal studies of TCID₅₀ values, BCIRL-HV-AM1 cells gave the highest extracellular virus (ECV) titer (9.7×10⁶ TCID₅₀/ml) followed by BCIRL-HA-AM1 cells (8.3×10⁵ TCID₅₀/ml) and BCIRL-AG-AM1 cells (3.2×10⁵ TCID₅₀/ml). In addition, a low ECV titer of 1.37×10³ TCID₅₀/ml was detected from TN-CL1 cells 96 h postinoculation, while BRL-AG-1, BMN, and BCIRL-HZ-AM1 cells were nonpermissive to AgMNPV-CL4-3A1 on the basis of TCID₅₀ results. AgMNPV-CL4-3A1 and the wild type AgMNPV had similar restriction profiles that were different from wild type AcMNPV. The LC₅₀ values were 96.9, 564.6, 733.3, and 1.1×10⁴ occlusion bodies/cm² of diet for A. gemmatalis, Helicoverpa zea, Heliothis virescens, and T. ni, respectively. This article presents the results of research only. Mention of proprietary products in this article does not indicate endorsement or a recommendation for use by USDA-ARS. All programs and services of the USDA are offered on a nondiscriminatory basis without regard to race, color, national origin, religion, sex, marital status or handicap.     

Field observations of mating,oviposition, and feeding behavior ofAnticarsia gemmatalis (Lepidoptera: Noctuidae) in a soybean field

The temporal occurrence and frequency of mating, oviposition, and feeding of wild velvetbean caterpillar moths, Anticarsia gemmatalis,in a soybean field (Glycine max)were documented. Mating occurred predominately during the first 4 h after sunset and primarily on leaflet bottoms. Oviposition was concentrated within the first 6 hr after sunset. Eggs were laid singly, and in areas of high trichome density, on leaflets, pulvini, and petioles. Feeding occurred primarily at night, with most males feeding during the first half of the night and females during the second half. Males and females shared some food sites but others were visited strictly by males.     

Phenology of Predation on Insects in a Tropical Forest: Temporal Variation in Attack Rate on Dummy Caterpillars

In communities of tropical insects, adult abundance tends to fluctuate widely, perhaps in part owing to predator–prey dynamics. Yet, temporal patterns of attack rates in tropical forest habitats have not been studied systematically; the identity of predators of insects in tropical forests is poorly known; and their responses to temporal variation in prey abundance have rarely been explored. We recorded incidence and shape of marks of attacks on dummy caterpillars (proxy of predation rate) in a sub‐montane tropical forest in Uganda during a yearlong experiment, and explored correlations with inferred caterpillar abundance. Applying the highest and lowest observed daily attack rates on clay dummies over a realistic duration of the larval stage of butterflies, indicates that the temporal variation in attack rate could cause more than 10‐fold temporal variation in caterpillar survival. Inferred predators were almost exclusively invertebrates, and beak marks of birds were very scarce. Attack rates by wasps varied more over time than those of ants. Attack rates on dummies peaked during the two wet seasons, and appeared congruent with inferred peaks in caterpillar density. This suggests (1) a functional response (predators shifting to more abundant resource) or adaptive timed phenology (predators timing activity or breeding to coincide with seasonal peaks in prey abundance) of predators, rather than a numerical response (predator populations increasing following peaks in prey abundance); and (2) that predation would dampen abundance fluctuations of tropical Lepidoptera communities.