Interactions between Ecological Traits and Host Plant Type Explain Distribution Change in Noctuid Moths

Abstract:  The ecological traits of species determine how well a species can withstand threats to which it is exposed. If these predisposing traits can be identified, species that are most at risk of decline can be identified and an understanding of the processes behind the declines can be gained. We sought to determine how body size, specificity of larval host plant, overwintering stage, type of host plant, and the interactions of these traits are related to the distribution change in noctuid moths. We used data derived from the literature and analyzed the effects of traits both separately and simultaneously in the same model. When we analyzed the traits separately, it seemed the most important determinants of distribution change were overwintering stage and type of host plant. Nevertheless, ecological traits are often correlated and the independent effect of each trait may not be seen in analyses in which traits are analyzed separately. When we accounted for other correlated traits, the results were substantially different. Only one trait (body size), but 3 interactions, explained distribution change. This finding suggests that distribution change is not determined by 1 or 2 traits; rather, the effect of the traits depends on other interacting traits. Such complexity makes it difficult to understand the processes behind distribution changes and emphasizes the need for basic ecological knowledge of species. With such basic knowledge, a more accurate picture of the factors causing distribution changes and increasing risk of extinction might be attainable.     

Leaf surface chemicals stimulating oviposition byPieris rapae (Lepidoptera: Pieridae) on cabbage

Summary The chemical stimulation of oviposition byPieris rapae on cabbage was investigated by leaf washing and extraction. Isolation of the stimulant by various chromatographic techniques was monitored by a bioassay using Sieva bean as a surrogate host plant. Cold water, chloroform, or chloroform followed by cold water washes failed to release the stimulant from leaf surfaces. Boiling water or chloroform followed by methanol was required. The most active stimulatory compound was identified as 3-indolylmethyl glucosinolate (glucobrassicin). Other glucosinolates were identified as sinigrin, which was only slightly active, and glucoiberin, which was completely inactive as a stimulant. The significance of the selective response ofP. rapae andP. brassicae to different glucosinolates and the implications of the binding of polar allelochemicals to leaf surfaces is discussed with respect to host utilization and perception mechanisms of pierids.     

Chemical basis for differential acceptance of two sympatric rutaceous plants by ovipositing females of a swallowtail butterfly, Papilio polytes (Lepidoptera, Papilionidae)

Summary. In the natural habitat of Papilio polytes, a Rutaceae feeder utilizing Toddalia asiatica as a major host plant, some other rutaceous plants such as Murraya paniculata (abundant) and Glycosmis citrifolia (relatively rare) occur sympatrically as potential hosts. Whereas G. citrifolia is occasionally infested in the field, M. paniculata remains entirely unexploited by the butterfly. We thus examined the phytochemical mechanisms that can explain the differential acceptance of the two plants by ovipositing females of P. polytes. The foliage of G. citrifolia was found to readily induce oviposition and females deposited eggs in response to a methanolic extract of the plant. Stimulatory activity-directed fractionation of the extract revealed the presence of two characteristic compounds, trans-4-hydroxy-N-methylproline and 2-C-methylerythronic acid, known to serve as oviposition stimulants for the butterfly. In addition, larvae performed as well or better onG. citrifolia as on T. asiatica. Similar examination of the inhibitory chemical constituents of M. paniculata led to the isolation of an oviposition deterrent. The compound, identified as trigonelline (N-methylpyridine- 3-carboxylic acid), exhibited moderate oviposition deterrency to females. These results combined with our previous data are in agreement with the observed differential utilization of the two plants by P. polytes in the field.     

Sex pheromones and their potential role in the evolution of reproductive isolation in small ermine moths (Yponomeutidae)

Summary Sex pheromone communication in the nine European species of small ermine moths (Yponomeuta) is reviewed in regard to the potential role of pheromones in the speciation process. Six of the nine species studied (viz.,Y. evonymellus, Y. cagnagellus, Y. padellus, Y. irrorellus, Y. plumbellus, andY. vigintipunctatus) use a mixture of (E)-11-and (Z)-11-tetradecenyl acetate in different ratios as primary pheromone components, with combinations of tetradecyl acetate, (Z)-9-tetradecenyl acetate, (Z)-11-hexadecenyl acetate and the corresponding alcohols of the acetates as additional pheromone components. Analysis of (Z)- to (E)-11-tetradecenyl acetate ratios produced by individual females of these species demonstrated significant variation among females of all species. However, the ranges of ratios produced byY. cagnagellus, Y. irrorellus, andY. plumbellus, sharing the same host-plant species, spindle tree, did not overlap. Niche separation of all six species mentioned required consideration of at least one additional pheromone component or of temporal aspects. The remaining three species,i.e. Y. malinellus, Y. mahalebellus andY. rorellus, have pheromones that differ qualitatively.Biosynthetic routes to the pheromone components identified are proposed on the basis of fatty acid pheromone precursors found in the pheromone glands. A phylogenetic tree for the genus is constructed based on allozyme frequency data and changes in pheromone composition are superimposed on this tree. We suggest that the ancestral ermine moth pheromone is a mixture of (Z)-11- and (E)-11-tetradecenyl acetate and the corresponding alcohols, and a scenario of how present-day patterns evolved is outlined. The pheromone differences among the three species using spindle tree as their host-plant might have evolved throughreproductive character displacement upon secondary contact between populations that had already diverged genetically in allopatry. Pheromone differences within the so-calledpadellus-complex (includingY. cagnagellus, Y. mahalebellus, Y. malinellus, Y. padellus, andY. rorellus) in which species might have originated sympatrically, may have evolved byreinforcing selection as these species still hybridise and produce viable offspring when confined in cages. The role of pheromones in reproductive isolation amongYponomeuta species is emphasised by (1) the function of pheromone components of some of the species as behavioural antagonists to other species, (2) the cross-attraction under experimental conditions between allochronic species with similar pheromones, and (3) the formation of hybrids in the laboratory between species that are isolated in nature by pheromone differences.     

Ontogenetic changes in social behaviour in the forest tent caterpillar,<Emphasis Type="Italic"> Malacosoma disstria</Emphasis>

Many animals, including gregarious caterpillars, begin life in groups and become increasingly solitary as they grow larger. These ontogenetic changes in social behaviour suggest that the costs and benefits of grouping change with increasing individual size. Forest tent caterpillar (Malacosoma disstria) colonies exhibit complex social behaviour during the early larval instars, using pheromone trails to move together between temporary bivouacs and feeding sites, and break up as the caterpillars grow. We demonstrate changes in individual responses to cues from conspecifics that explain changes in aggregation during caterpillar development. We used Markov chain analysis to test the influence of pheromone trails and colony-mates on an individual caterpillars tendency to switch between quiescence, searching, walking and spinning. Pheromone-laden silk trails increased the tendency to begin locomotion, whereas colony-mates increased switching from activity to quiescence. Trails also influenced the form taken by locomotor behaviour, and promoted directed walking over searching. Social cues thus increase the efficiency of individual locomotion. Younger larvae were more quiescent and more reluctant to walk in the absence of trails than were older insects. An increase in independent locomotion as the larvae grow provides a mechanism to explain colony break-up and points to an ontogenetic shift in the internal processes driving behaviour. Scaling relationships suggest that many of the benefits associated with group-living in caterpillars decrease as individuals grow larger, providing an adaptive explanation for observed ontogenetic changes in social behaviour.Communicated by N. Wedell     

(<Emphasis Type="Italic">Z,Z</Emphasis>)-11,13-Hexadecadienyl acetate and (<Emphasis Type="Italic">Z,E</Emphasis>)-11,13,15-hexadecatrienyl acetate: synergistic sex pheromone components of oak processionary moth,Thaumetopoea processionea (Lepidoptera: Thaumetopoeidae)

Summary. Our objective was to identify sex pheromone components of the oak processionary moth, Thaumetopoea processionea (Lepidoptera: Thaumetopoeidae), whose larvae defoliate oak, Quercus spp., forests in Eurasia and impact human health. Coupled gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectrometric (MS) analyses of pheromone gland extract of female T. processionea revealed two consistently EAD-active compounds. They were identified as (Z,Z)-11,13-hexadecadienyl acetate (Z11,Z13-16:OAc) and (Z,E)-11,13,15-hexadecatrienyl acetate (Z11,E13,15-16:OAc) by comparative GC, GC-MS and GC-EAD analyses of insect-produced compounds and authentic standards. In replicated field experiments (2000, 2001) in Nordbaden, Südbaden and Sachsen-Anhalt (Germany), Z11,Z13-16:OAc and Z11,E13,15-16:OAc in combination, but not singly, attracted significant numbers of male moths. It will now be intriguing to investigate whether Z11,E13,15-16:OAc, or its corresponding alcohol or aldehyde, serves as a pheromone component also in other species of the Thaumetopoeidae.     

Foraging responses in the butterflies <Emphasis Type="Italic">Inachis io</Emphasis>, <Emphasis Type="Italic">Aglais urticae</Emphasis> (Nymphalidae), and <Emphasis Type="Italic">Gonepteryx rhamni</Emphasis> (Pieridae) to floral scents

Summary. For butterflies to be efficient foragers, they need to be able to recognize rewarding flowers. Flower signals such as colours and scents assist this recognition process. For plant species to attract and keep butterflies as pollinators, species-specific floral signals are crucial. The aim of this study is to investigate foraging responses to floral scents in three temperate butterfly species, Inachis io L. (Nymphalidae), Aglais urticae L. (Nymphalidae), and Gonepteryx rhamni L. (Pieridae), in behavioural choice bioassays. The butterflies were allowed to choose bet-ween flower models varying in scent and colour (mauve or green). Flowers or vegetative parts from the plants Centaurea scabiosa L. (Asteraceae), Cirsium arvense (L.) (Asteraceae), Knautia arvensis (L.) (Dipsacaceae), Buddleja davidii Franchet (Loganicaeae), Origanum vulgareL. (Lamiaceae), Achillea millefolium L. (Asteraceae), and Philadelphus coronarius L. (Hydrangiaceae) were used as scent sources. All visits to the models — those that included probing and those that did not — were counted, as was the duration of these behaviours. Both flower-naive and flower-experienced (conditioned to sugar-water rewards, the colour mauve, and specific floral scents) butterflies were tested for their preference for floral versus vegetative scents, and to floral scent versus colour. The butterflies were also tested for their ability to switch floral scent preferences in response to rewards. Flower-naive butterflies demonstrated a preference for the floral scent of the butterfly-favourable plants C. arvense and K. arvensis over the floral scent of the non-favourable plants Achillea millefolium (Asteraceae), and Philadelphus coronarius cv. (Hydrangiaceae). Most of the butterflies that were conditioned to floral scents of either C. arvense, K. arvensis, or B. davidii readily switched theirfloral scent preferences to the one most recently associated with reward, thus demonstrating that floral scent constancy is a result from learning. These findings suggest that these butterflies use floral scent as an important cue signal to initially identify and subsequently recognize and distinguish among rewarding plants. Received 2 September 2001; accepted 9 September 2002.     

The roles of nicotine and natural enemies in determining larval feeding site distributions of Manduca sexta L. and Manduca quinquemaculata (Haworth) on tobacco

Summary. Field observations indicated that hornworms select feeding sites non-randomly on tobacco. We tested the hypotheses that differences in feeding site locations of larvae of Manduca sexta L. and Manduca quinquemaculata (Haworth) (Lepidoptera: Sphingidae) on tobacco could be explained by differential nicotine concentrations within plants and leaves, species-specific responses to nicotine, or pressure exerted by natural enemies. Results showed that third-instar larvae of M. sexta fed more proximally and centrally on the leaf, whereas M. quinquemaculata fed more distally. Within-plant selection of leaves did not differ; both species selected leaves in the middle region of the plant. Nicotine concentrations in a high nicotine genotype, NC95, varied within each leaf, increasing 2—3 fold from the basal to apical portion of the leaf, and within each plant, increasing 7—10 fold from the first fully expanded leaf to the twelfth (lowest) leaf. In laboratory bioassays, both Manduca species responded to nicotine as a feeding deterrent. Electrophysiological studies demonstrated that gustatory organs of both species responded to nicotine at concentrations found in tobacco leaves and that M. quinquemaculata generally showed a less vigorous response to nicotine than M. sexta. Field mortality of M. sexta due to parasitism by Cotesia congregata (Say) and to parasitism and predation combined differed among feeding sites; predation alone did not. Results suggest that although nicotine concentration and species specific responses to nicotine play a role in determining feeding site locations, pressure exerted by natural enemies, especially parasitism by C. congregata, is more important. Received 22 February 2000; accepted 20 July 2001.     

Host-plant green-leaf volatiles synergize the synthetic sex pheromones of the corn earworm and codling moth (Lepidoptera)

Summary The capture of adult male moths in female sex pheromone traps of two key agricultural pests, the corn earworm (Helicoverpa zea) and the codling moth (Cydia pomonella), is enhanced or synergized by a certain group of host-plant volatiles, the green-leaf volatiles (GLVs). Since female adults of both species call and release their sex pheromones while perched upon the leaves of their host-plants, the volatile constituents from the leaves of a number of host-plants were compared. Sex pheromone traps containing one of the prominent leaf volatiles of certainH. zea hosts, (Z)-3-hexenyl acetate, not only significantly increased the capture ofH. zea males but were preferred over traps baited only with sex pheromone. Similarly, traps baited with synthetic sex pheromome ofC. pomonella plus a blend of GLVs captured significantly more males than traps baited only with sex pheromone. Since male moths are not captured in traps baited only with these GLVs, it appears that these GLVs act as pheromone synergists which increase or enhance the attraction or arrestment of male moths in pheromone traps.