Evaluation ofHadena perplexa [Lepidoptera: Phalaenidae] as a biocontrol agent of bladder campionSilene vulgaris [Caryophyllaceae] in Canada: Rearing and host specificity

Rearing techniques and results of preliminary host range tests are reported forHadena perplexa (Denis & Schiffermuller) (Lep.: Phalaenidae) a candidate biocontrol agent against the weed bladder campion,Silene vulgaris (Moench) Garcke, in Manitoba, Canada. In the laboratory, it was necessary to pipette a 15% honey solution in water into the flowers as food for the adult moths. When reared singly to avoid cannibalism, 56% of the 1^st instar larvae developed to pupae. Larvae fed on a natural diet for 10 days can then be reared on either one of 2 artificial diets. Choice oviposition tests and no-choice larval feeding tests were conducted with plant species closely related toS. vulgaris in the generaSilene, Dianthus, Gypsophila, Lychnis, Saponaria. Species in 4 of 5 of these genera were accepted for oviposition, and species in all 5 genera supported the development of 1^st instar larvae to the pupal stage.H. perplexa should not be introduced into Canada.      

Distribution and colonisation ability of three parasitoids and their herbivorous host in a fragmented landscape

Habitat fragmentation can disrupt communities of interacting species even if only some of the species are directly affected by fragmentation. For instance, if parasitoids disperse less well than their herbivorous hosts, habitat fragmentation may lead to higher herbivory in isolated plant patches due to the absence of the third trophic level. Community-level studies suggest that parasitoids tend to have limited dispersal abilities, on the order of tens of metres, much smaller than that of their hosts, while species-oriented studies document dispersal by parasitoids on the scale of kilometres. In this study the distribution patterns of three parasitoid species with different life histories and their moth host, Hadena bicruris, a specialist herbivore of Silene latifolia, were compared in a large-scale network of natural fragmented plant patches along the rivers Rhine and Waal in the Netherlands. We examined how patch size and isolation affect the presence of each species. Additionally, experimental plots were used to study the colonisation abilities of the species at different distances from source populations.In the natural plant patches the presence of the herbivore and two of the parasitoids, the gregarious specialist Microplitis tristis and the gregarious generalist Bracon variator were not affected by patch isolation at the scale of the study, while the solitary specialist Eurylabus tristis was. In contrast to the herbivore, the presence of all parasitoid species declined with plant patch size. The colonisation experiment confirmed that the herbivore and M. tristis are good dispersers, able to travel at least 2 km within a season. B. variator showed intermediate colonisation ability and E. tristis showed very limited colonisation ability at this spatial scale. Characteristics of parasitoid species that may contribute to differences in their dispersal abilities are discussed.     

Pollination and seed predation by moths on Silene and allied Caryophyllaceae: evaluating a model system to study the evolution of mutualisms

Nursery pollinators, and the plants they use as hosts for offspring development, function as exemplary models of coevolutionary mutualism. The two pre-eminent examples--fig wasps and yucca moths--show little variation in the interaction: the primary pollinator is an obligate mutualist. By contrast, nursery pollination of certain Caryophyllaceae, including Silene spp., by two nocturnal moth genera, Hadena and Perizoma, ranges from antagonistic to potentially mutualistic, offering an opportunity to test hypotheses about the factors that promote or discourage the evolution of mutualism. Here, we review nursery pollination and host-plant interactions in over 30 caryophyllaceous plants, based on published studies and a survey of researchers investigating pollination, seed predation, and moth morphology and behavior. We detected little direct evidence of mutualism in these moth-plant interactions, but found traits and patterns in both that are nonetheless consistent with the evolution of mutualism and merit further attention.     

Sex‐biased oviposition by a nursery pollinator on a gynodioecious host plant: Implications for breeding system evolution and evolution of mutualism

Dioecy, a breeding system where individual plants are exclusively male or female, has evolved repeatedly. Extensive theory describes when dioecy should arise from hermaphroditism, frequently through gynodioecy, where females and hermaphrodites coexist, and when gynodioecy should be stable. Both pollinators and herbivores often prefer the pollen‐bearing sex, with sex‐specific fitness effects that can affect breeding system evolution. Nursery pollination, where adult insects pollinate flowers but their larvae feed on plant reproductive tissues, is a model for understanding mutualism evolution but could also yield insights into plant breeding system evolution. We studied a recently established nursery pollination interaction between native Hadena ectypa moths and introduced gynodioecious Silene vulgaris plants in North America to assess whether oviposition was biased toward females or hermaphrodites, which traits were associated with oviposition, and the effect of oviposition on host plant fitness. Oviposition was hermaphrodite‐biased and associated with deeper flowers and more stems. Sexual dimorphism in flower depth, a trait also associated with oviposition on the native host plant (Silene stellata), explained the hermaphrodite bias. Egg‐receiving plants experienced more fruit predation than plants that received no eggs, but relatively few fruits were lost, and egg receipt did not significantly alter total fruit production at the plant level. Oviposition did not enhance pollination; egg‐receiving flowers usually failed to expand and produce seeds. Together, our results suggest that H. ectypa oviposition does not exert a large fitness cost on host plants, sex‐biased interactions can emerge from preferences developed on a hermaphroditic host species, and new nursery pollination interactions can arise as negative or neutral rather than as mutualistic for the plant.     

Nursery pollination by a moth in Silene latifolia: the role of odours in eliciting antennal and behavioural responses

Since the 1970s it has been known that the nursery pollinator Hadena bicruris is attracted to the flowers of its most important host plant, Silene latifolia, by their scent. Here we identified important compounds for attraction of this noctuid moth. Gas chromatographic and electroantennographic methods were used to detect compounds eliciting signals in the antennae of the moth. Electrophysiologically active compounds were tested in wind-tunnel bioassays to foraging na?ve moths, and the attractivity of these compounds was compared with that to the natural scent of whole S. latifolia flowers. The antennae of moths detected substances of several classes. Phenylacetaldehyde elicited the strongest signals in the antennae, but lilac aldehydes were the most attractive compounds in wind-tunnel bioassays and attracted 90% of the moths tested, as did the scent of single flowers. Our results show that the most common and abundant floral scent compounds in S. latifolia, lilac aldehydes, attracted most of the moths tested, indicating a specific adaptation of H. bicruris to its host plant.     

Age-dependent clutch size in a koinobiont parasitoid

Abstract.  1. The Lack clutch size theory predicts how many eggs a female should lay to maximise her fitness gain per clutch. However, for parasitoids that lay multiple clutches it can overestimate optimal clutch size because it does not take into account the future reproductive success of the parasitoid.
2. From egg-limitation and time-limitation models, it is theoretically expected that (i) clutch size decreases with age if host encounter rate is constant, and (ii) clutch size should increase with host deprivation and hence with age in host-deprived individuals.
3. Clutch sizes produced by ageing females of the koinobiont gregarious parasitoid Microplitis tristis Nees (Hymenoptera: Braconidae) that were provided daily with hosts, and of females ageing with different periods of host deprivation were measured.
4. Contrary to expectations, during the first 2 weeks, clutch size did not change with the age of the female parasitoid, neither with nor without increasing host-deprivation time.
5. After the age of 2 weeks, clutch size decreased for parasitoids that parasitised hosts daily. The decrease was accompanied by a strong decrease in available eggs. However, a similar decrease occurred in host-deprived parasitoids that did not experience egg depletion, suggesting that egg limitation was not the only factor causing the decrease in clutch size.
6. For koinobiont parasitoids like M. tristis that have low natural host encounter rates and short oviposition times, the costs of reproduction due to egg limitation, time limitation, or other factors are relatively small, if the natural lifespan is relatively short.
7. Koinobiont parasitoid species that in natural situations experience little variation in host density and host quality might not have strongly evolved the ability to adjust clutch size.     

Stereoisomeric pattern of lilac aldehyde in Silene latifolia, a plant involved in a nursery pollination system

The monoterpene lilac aldehyde is found in floral scent of several plants species, among them Silene latifolia. This plant is involved in a nursery pollination system, because a noctuid moth, Hadena bicruris, is not only pollinator but also seed predator. Lilac aldehyde is the key floral scent compound of S. latifolia for attracting Hadena. This monoterpene has three stereogenic centers, and eight different isomers are possible. Here, we analysed the ratio of lilac aldehyde isomers from plants originating from 18 different populations of S. latifolia using enantioselective multidimensional GC-MS (enantio-MDGC-MS), and compared resulting variability with variability found in total scent emitted by specimen under study. Though variability in total emitted scent was high, ratio of lilac aldehyde isomers was a more conservative trait. There was no correlation between the ratio of lilac aldehyde isomers and the total emitted floral scent pattern. Both, ratio of stereoisomers and total emitted scent were independent from the geographic origin of the plants. In conclusion, the ratio of lilac aldehyde stereoisomers in S. latifolia is a reliable trait, and may used by the nursery pollinator H. bicruris for host-plant detection.     

The contribution of a pollinating seed predator to selection on Silene latifolia females

Interactions, antagonistic or mutualistic, can exert selection on plant traits. We explored the role of Hadena bicruris, a pollinating seed predator, as a selective agent on its host, the dioecious plant Silene latifolia. We exposed females from artificial-selection lines (many, small flowers (SF) vs. few, large flowers (LF)) to this moth. Infestation did not differ significantly between lines, but the odds of attacked fruits aborting were higher in SF females. We partitioned selection between that caused by moth attack and that resulting from all other factors. In both lines, selection via moth attack for fewer, smaller flowers contrasted with selection via other factors for more flowers. In LF females, selection via the two components was strongest and selection via moth attack also favoured increased fruit abortion. This suggests that the moths act as more of a selective force on flower size and number via their predating than their pollinating role.