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Establishing wildflower strips has been suggested as an effective measure to promote pollination services, pest control or general insect biodiversity, but little is known about the integration of these different objectives when selecting flower seed mixtures. In ten agricultural landscapes in the Netherlands, we established a wildflower strip (0.4 – 4.9 ha) with half of each strip sown with a mixture targeting longer-tongued pollinators and the other half sown with a mixture targeting shorter-tongued pollinators and natural enemies. We determined establishment success of sown wildflowers and evaluated the attractiveness of the established flower communities to multiple functional groups of flower visitors: bumblebees (long-tongued pollinators), hoverflies (short-tongued pollinators and natural enemies), and butterflies and total flower-visitor richness (indicators of wider biodiversity values). Bumblebees clearly preferred the pollinator-targeted seed mixture and were positively associated with cover of Fabaceae and negatively with Apiaceae. Hoverflies consistently preferred the natural enemy mixture and were positively associated with Apiaceae. The other target groups displayed no clear responses to seed mixture type but instead were associated with local flower richness within strips. Across sites, responses of flower-visitors to sown mixture types did not depend on wildflower strip size, proportion of surrounding semi-natural habitat, or flower variables. However, all flower-visitors except butterflies increased with increasing established cover or richness of (sown) flower species across sites. Our results suggest that, although species-rich wildflower strips may benefit several species groups, maximising different objectives involves trade-offs between functional groups that prefer short- or long-corolla flowers. Furthermore, our study suggests that sowing a wildflower mixture does not necessarily result in a vegetation with the same composition as the seed mixture as species may establish poorly or not at all. Selection of flower species for seed mixtures should therefore, in addition to insect target group, take the establishment characteristics of plant species into account.

The ability of an organism to adapt to short-term environmental changes within its lifetime is of fundamental importance. This adaptation may occur through phenotypic plasticity. Insects and mites, in particular, are sensitive to changes in temperature and humidity, especially during the juvenile stages. We studied the role of phenotypic plasticity in the adaptation of eggs to different relative humidity conditions, in the predatory mitePhytoseiulus persimilis, used worldwide as a biological control agent of the spider miteTetranychus urticae. The biocontrol efficacy ofP. persimilisdecreases under dry conditions, partly becauseP. persimiliseggs are sensitive to drought. We exposedP. persimilisadult females from two different strains to constant and variable humidity regimes and evaluated the hatching rate of their eggs in dry conditions, as well as the survival and oviposition rates of these females. Whereas the eggs laid byP. persimilisfemales exposed to constant high humidity did not survive in dry conditions, females exposed to constant low humidity started laying drought-resistant eggs after 24 h of exposure. Survival and oviposition rates of the females were affected by humidity: females laid fewer eggs under constant low humidity and had a shorter lifespan under constant high and constant low humidity. The humidity regimes tested had similar effects across the twoP. persimilisstrains. Our results demonstrate that transgenerational phenotypic plasticity, called maternal effect, allowsP. persimilisfemales to prepare their offspring for dry conditions.

Parasitoid wasps of the genus Trichogramma are used worldwide as biological control agents against lepidopteran pests. Trichogramma wasps develop inside eggs of a wide range of host species, most of them moths. They are generally considered as diurnal insects. Here, we investigated whether Trichogramma wasps can also successfully parasitise host eggs at night under controlled laboratory conditions. Eggs of the moth Ephestia kuehniella were offered under dark conditions (scotophase) to females of Trichogramma brassicae and Trichogramma evanescens either from 9:00 PM to 9:00 AM or from 11:00 AM to 5:00 PM at four different temperatures (5°C, 10°C, 15°C and 20°C). Both species are known to parasitise E. kuehniella eggs in the photophase during daytime. The results show that T. brassicae did not parasitise eggs in the scotophase at night and only very few in the artificially induced scotophase during daytime from 10°C to 20°C. In contrast, T. evanescens parasitised more eggs in the dark both at night and artificially induced scotophase during daytime. Parasitism in the scotophase already started at 5°C, with more eggs being parasitised and more offspring being produced at higher temperatures. T. evanescens displayed higher parasitism activity in the induced scotophase during daytime than in the scotophase at night. The present study suggests that Trichogramma are capable of successfully parasitising host eggs at night, even at low temperatures, but that nocturnal activity with respect to parasitism varies between wasp species.

The predatory mite Phytoseiulus persimilis is widely used for augmentative biological control, as an effective predator of the spider mite Tetranychus urticae. However, the biocontrol efficacy of P. persimilis decreases under dry conditions. One of the reasons for this decline concerns P. persimilis’ eggs, which are sensitive to low humidity. In this study, we investigated the possibility to select for a strain of P. persimilis adapted to dry conditions. To understand the potential sources of phenotypic variation in egg survival under dry conditions, we tested the effects of genetic and environmental factors on variation in this trait. We compared egg hatching of five P. persimilis strains, under constant as well as variable humidity conditions, at 25 °C. The results show no intraspecific genetic variation among the five tested strains in egg hatching under constant and variable humidity conditions. In all five strains, less than 20% of the eggs hatched when they were exposed to constant low (60% RH) humidity conditions. However, when eggs were exposed to successive cycles of low and high humidity, a common pattern observed in the field, significantly higher hatching rates were observed. Under variable humidity conditions, more than 73% of the eggs hatched successfully, even when exposure to high humidity was limited to only 13% of the egg developmental time. Although P. persimilis eggs suffered from a high rate of water loss under constant dry conditions, they were able to compensate for this water loss when exposed to high humidity conditions for a few hours during their development. A decreased biocontrol efficacy of P. persimilis under dry conditions may be explained by a higher egg mortality when relative humidity is constantly low. Yet, when relative humidity exhibits diurnal variation, periods of high humidity may mitigate the effects of periods of low humidity during development of P. persimilis eggs.

1. Declining pollinator populations have caused concern about consequences for food production, and have initiated an increasing number of initiatives that aim to mitigate pollinator loss through enhancement of floral resources. Studies evaluating effects of mitigation measures generally demonstrate positive responses of pollinators to floral resource enhancement. However, it remains unclear whether this represents landscape-level population effects or results from a spatial redistribution of individuals from otherwise unaffected populations. 2. Here we present a method for estimating landscape-level population effects using data from commonly used standardized pollinator transect surveys. The approach links local density responses of pollinators in both mitigation sites and surrounding landscape elements to the area these habitats occupy in mitigation landscapes as well as control landscapes to obtain landscape-level population estimates. 3. We demonstrate the method using data from a two-year study examining the effects of experimental wildflower enhancements on wild bumble bees and solitary bees in Dutch agricultural landscapes. The results show that conclusions based on local responses may differ significantly from those based on landscape-level responses. 4. Wildflower enhancements significantly enhanced landscape-level abundance of both bumble bees and solitary bees. Bumble bees showed a pronounced positive local density response in mitigation sites and the surrounding landscape that was in line with significant landscape-level increases in abundance. However, solitary bees showed no local response to mitigation sites, and the landscape-level increases in abundance only became apparent when the area of bee habitat was taken into account. 5. Incorporating the area of both newly created and pre-existing pollinator habitats into effect estimates accounts for density-dependent processes such as dilution, spill-over and local concentration of individuals. It therefore results in more reliable estimates of the response to mitigation measures of pollinators, as well as other mobile arthropod groups that are often being surveyed using transect surveys.

Neighboring plants can influence arthropods on a focal plant and this can result in associational resistance or associational susceptibility. These effects can be mediated by above- and belowground interactions between the neighbor and focal plant, but determining the relative contribution of the above- and belowground effects remains an open challenge. We performed a common garden experiment with a design that enabled us to disentangle the above- and belowground effects of five different plant species on the growth and chemistry of the focal plant ragwort (Jacobaea vulgaris), and the arthropod community associated to this plant. Aboveground effects of different neighboring plant species were more important for the growth and quality of J. vulgaris and for the arthropod abundance on this plant than belowground effects of neighbors. This remained true when only indirect neighbor effects (via affecting the biomass or quality of the focal plant) were considered. The aboveground neighbor effects on arthropod abundance on the focal plant were strongly negative. However, the magnitude of the effect depended on the identity of the neighboring species, and herbivore abundance on the focal plant was higher when surrounded by conspecific than by heterospecific plants. We also observed interactions between above- and belowground neighbor effects, indicating that these effects may be non-additive. We conclude that above- and belowground associational effects are not equally strong, and that neighbor effects on plant-arthropod interactions occur predominantly aboveground. Read More: http://www.esajournals.org/doi/abs/10.1890/14-0563.1?af=R

The effects of floral species composition on offspring performance of solitary bees are rarely studied under conditions where foraging behaviour of mothers is allowed to play a role. In a semi-field experiment, we restricted foraging choices of the polylectic mason bee . Osmia bicornis L. to flower species belonging to plant families presumably used to different extent: . Borago officinalis L. (Boraginaceae), . Centaurea cyanus L. (Asteraceae) and . Brassica napus L. (Brassicaceae). We quantified the foraging behaviour and brood cell production by mother bees, and compared the quality of offspring in pure and mixed flower species stands. Offspring survival in pure stands was expected to reflect the mothers' foraging preferences in the mixed stand. Pure stands of . B. napus supported highest offspring survival, body mass and fraction of females produced. Offspring survival on . C. cyanus and . B. officinalis was very low. Larval mortality occurred earlier in brood cells provided with . B. officinalis pollen than in brood cells provided with . C. cyanus pollen suggesting different effects of pollen quality on early larval and later development. The time spent on different foraging activities correlated with lifetime reproductive output. However, in mixed stands, the proportion of time the bees were foraging on the different flower species did not differ significantly. Foraging behaviour may therefore not generally be a good proxy for the quality of floral resources for offspring production. Our results suggest that resources collected from one plant species may influence the usefulness of resources from another plant species. Bees may therefore overcome potentially deleterious effects of the suboptimal resources by mixing low- and high-quality resources. This may help generalist bees, such as . O. bicornis, to cope with an unpredictable environment. Die Effekte der Artenzusammensetzung von blühenden Pflanzen auf das Gedeihen der Nachkommen von solitären Bienen werden selten unter Bedingungen untersucht, die zulassen, dass das Sammelverhalten der Mütter eine Rolle spielt. In einem Semi-Freilandexperiment, beschränkten wir die Sammeloptionen der polylektischen Mauerbiene . Osmia bicornis L. auf Blütenarten aus vermutlich unterschiedlich stark genutzten Familien: . Borago officinalis L. (Boraginaceae), . Centaurea cyanus L. (Asteraceae) und . Brassica napus L. (Brassicaceae). Wir bestimmten das Sammelverhalten und die Brutzellenproduktion von Bienenweibchen und verglichen die Qualität der Nachkommen bei reinem und gemischtem Blütenangebot. Die Überlebensrate der Nachkommen bei Angebot von nur einer Blütenart sollte die Präferenz der Mütter in gemischten Blütenbeständen wiederspiegeln. Reine Bestände von . B. napus ergaben die höchste Überlebensrate, Körpermasse und den höchsten Weibchenanteil bei den Nachkommen. Bei . C. cyanus und . B. officinalis war die Überlebensrate sehr gering. Die Larvensterblichkeit setzte in mit . B. officinalis verproviantierten Brutzellen früher ein als in Brutzellen mit . C. cyanus als Proviant. Somit sollte es unterschiedliche Effekte der Pollenqualität auf die frühe larvale und die spätere Entwicklung geben. Die für die verschiedenen Sammelaktivitäten aufgewendete Zeit korrelierte mit der Gesamtproduktion an Nachkommen einer Mutterbiene. Indessen unterschieden sich die Zeitanteile, die bei gemischtem Angebot mit dem Sammeln an unterschiedlichen Blütenarten verbracht wurden, nicht signifikant voneinander. Das Sammelverhalten könnte deshalb kein guter Indikator für die Qualität von Blütenressourcen für die Produktion von Nachkommen sein. Unsere Ergebnisse legen nahe, dass Ressourcen, die von einer Pflanzenart gesammelt wurden, den Nutzen der Ressourcen von einer anderen Art beeinflussen könnten. Die Bienen könnten somit mögliche abträgliche Effekte einer suboptimalen Ressource durch das Mischen von qualitativ hoch- und minderwertigen Ressourcen kompensieren. Dies könnte Generalisten wie der Mauerbiene . O. bicornis helfen, mit einer nicht vorhersagbaren Umwelt zurechtzukommen.

Pollinator communities exhibit variable responses to changing landscape composition. A general expectation is that a decreasing cover of semi-natural habitats negatively affects pollinator reproduction, population size and pollination services, but few studies have investigated the simultaneous effects of landscape complexity on different aspects of pollinator communities and functioning. In 20 agricultural landscape plots the size of an average Dutch farm, we studied how changing landscape complexity affected wild bee abundance, species richness and reproduction. To measure pollination, we placed potted strawberry plants as phytometers in landscapes. Landscape complexity was characterized as the area of semi-natural habitats. In addition, we estimated floral resource abundance in each landscape plot. We expected that i) bee species richness, reproduction and pollination would be positively related to area of semi-natural habitats and flower abundance, and that ii) species richness and reproduction would be positively related to pollination. An increase in semi-natural habitats in landscapes increased both the abundance of cavity-nesting bees colonizing trap nests, and the growth rates of experimental Bombus terrestris L. colonies, but not the species richness of wild bees measured by pan traps. There was only a tendency for higher pollination levels of strawberry plants with higher cover of semi-natural habitats. There was no relationship between species richness and bee reproduction in a landscape and the pollination services. Estimated flower abundance in landscape had a positive effect on bumblebee colony growth only and not on the other variables. Our results suggest that, by improving habitat quality on their farms through establishing more semi-natural habitats or enhancing the flower availability in semi-natural habitats, farmers can promote reproduction of a number of functionally important bee species and the pollination services they provide. Bee species richness, however, seems to be more difficult to enhance and requires more than just creating more of the same type of habitats or flowers.

Behaviour of nocturnal insects is routinely observed under red light, but it is unclear how the behaviour under red light compares to behaviour in complete darkness, or under a source of white light. Here, we measure movement behaviour of the nocturnal carabid beetle Pterostichus melanarius Illiger (Coleoptera: Carabidae) using camera recording under a near-infrared (nir), red or white radiation source. Red light significantly reduced movement speed in females similar to the effect of white light and different from nir. Also movement activity and pause length were affected by radiation source, with a significant difference between nir and white light, and with intermediate values in red light. The results presented here indicate that P. melanarius has different movement behaviour under the three radiation sources and suggest that nir rather than red radiation is most appropriate for measuring behaviour in total darkness. However, in the field total darkness is rare both because of natural light sources such as the moon and stars but increasingly also because of ecological light pollution, and therefore red light may still be of use for observing ecologically and practically relevant natural night-time behaviour.

Co-evolutionary theory underpins our understanding of interactions in nature involving plant–herbivore and host–parasite interactions. However, many studies that are published in the empirical literature that have explored life history and development strategies between endoparasitoid wasps and their hosts are based on species that have no evolutionary history with one another. Here, we investigated novel associations involving two closely related solitary endoparasitoids that originate from Europe and North America and several of their natural and factitious hosts from both continents. The natural hosts of both species are also closely related, all being members of the same family. We compared development and survival of both parasitoids on the four host species and predicted that parasitoid performance is better on their own natural hosts. In contrast with this expectation, survival, adult size and development time of both parasitoids were similar on all (with one exception) hosts, irrespective as to their geographic origin. Our results show that phylogenetic affinity among the natural and factitious hosts plays an important role in their nutritional suitability for related parasitoids. Evolved traits in parasitoids, such as immune suppression and development, thus enable them to successfully develop in novel host species with which they have no evolutionary history. Our results show that host suitability for specialized organisms like endoparasitoids is closely linked with phylogenetic history and macro-evolution as well as local adaptation and micro-evolution. We argue that the importance of novel interactions and ‘ecological fitting’ based on phylogeny is a greatly underappreciated concept in many resource–consumer studies.

Resource patch size and patch nutritional quality are both important factors influencing local densities of herbivores. The responses of herbivores to resource patch size have been mostly studied in aboveground plant–insect interactions, whereas belowground organisms have received little attention. We studied responses of different root-feeding nematode species associated with marram grass (Ammophila arenaria (L.) Link) to resource patch size and quality. Different nematode species were released in experimental mesocosms filled with dune sand in which we established marram grass patches of varying sizes. Half of the patches of small, medium and large size were fertilized to test if immigration probabilities of nematodes depended on patch quality. We tested the hypotheses that (1) nematodes should aggregate on larger patches and (2) colonization of patches would also depend on patch nutritional quality, with higher nematode recapture rates expected in fertilized patches. Two species (Helicotylenchus pseudorobustus, Hemicycliophora thornei) of the five released species were recaptured in the experiment. The fraction of nematodes immigrating into the rhizosphere of a plant patch increased with patch size (i.e. root biomass), which was in line with predictions of the Resource Concentration Hypothesis. When fractions were recalculated to represent recapture rates per liter of soil, recapture rates of nematodes did not differ among patch sizes, indicating that the increase in recapture rates was directly proportional to the increase in patch size. This suggests that the process through which nematodes located patches was not distinguishable from a random process where entering patches is based on random encounters with patch boundaries. In contrast to our expectation, fertilization had a strong negative effect on patch responses of both nematode species. Our study represents an approach that may be used to explore whether belowground biota behave in similar ways as aboveground biota, in order to determine how perceived differences in environments affect ecological interactions.

Invasive species represent one of the most important threats to biodiversity worldwide, with consequences for ecosystem functioning and the delivery of important ecological services to society. Several hypotheses have been generated to explain the success of exotic plants in their new ranges, with escape from their old natural enemies, such as pathogens and herbivores (the ‘enemy release hypothesis’) and novel defensive chemistry (the ‘novel weapons hypothesis’) receiving considerable attention. Thus far, virtually all studies of exotic plants and insects have been conducted in a strictly bi-trophic framework involving plants and herbivores. On the other hand, it has been argued that a better understanding of the forces regulating community structure and function should include natural enemies of the herbivores. Furthermore, indirect interactions between organisms in the plant roots (below-ground) and shoots (above-ground) are known to strongly effect the behaviour and performance of consumers in the opposite ‘compartment’. Here, we discuss a range of physiological, evolutionary and ecological aspects of plant–herbivore-natural enemy interactions involving exotic plants. Further, interactions between soil and above-ground organisms are explored with respect to studies with exotic plants. We argue that it is important to link population and community ecology to individual-level variation in the physiology and behaviour of insects across several trophic levels in studies with invasive plants. Future research with invaders should also aim to integrate physically separated compartments (e.g. plant roots and shoots). This will facilitate a more complete understanding of the factors underlying the success (or failure) of exotic plants to spread and become dominant in their new ranges. Moreover, these data will also help to unravel the relative importance of top-down and bottom-up processes in regulating communities in which exotic plants have become established. Lastly, we discuss consequences for conservation.

Plant–insect interactions occur in spatially heterogeneous habitats. Understanding how such interactions shape density distributions of herbivores requires knowledge on how variation in plant traits (e.g. nutritional quality) affects herbivore abundance through, for example, affecting movement rates and aggregation behaviour. We studied the effects of plant patch size and herbivore-induced differences in plant nutritional quality on local densities of insect herbivores for two Brassica oleracea cultivars, i.e. white cabbage and Brussels sprouts. Early season herbivory as a treatment resulted in measurable differences in glucosinolate concentrations in both cultivars throughout the season. Herbivore induction and patch size both influenced community composition of herbivores in both cultivars, but the effects differed between species. Flea beetles (Phyllotreta spp.) were more abundant in large than in small patches, and this patch response was more pronounced on white cabbage than on Brussels sprouts. Herbivore-induction increased densities in all patches. Thrips tabaci was also more abundant in large patches and densities of this species were higher on Brussels sprouts than on white cabbage. Thrips densities were lower on induced than on control plants of both cultivars and this negative effect of induction tended to be more pronounced in large than in small patches. Densities of the cabbage moth (Mamestra brassicae) were lower on Brussels sprouts than on white cabbage and lower on herbivore-induced than on uninduced plants, with no effect of patch size. No clear effects of patch size and induction were found for aphids. This study shows that constitutive and herbivore-induced differences in plant traits interact with patch responses of insect herbivores.

Parasitic wasps employ a wide range of chemical cues to find their hosts. Very recently, we discovered how 2 closely related egg parasitoids, Trichogramma brassicae and Trichogramma evanescens, exploit the anti-aphrodisiac pheromone benzyl cyanide of one of their hosts, the gregarious large cabbage white butterfly Pieris brassicae that deposits a clutch of more than 20 eggs per oviposition bout. The pheromone is transferred by male butterflies to females during mating to enforce female monogamy. On detecting the anti-aphrodisiac, the tiny parasitic wasps ride on a mated female butterfly to a host plant and then parasitize her freshly laid eggs. The present study demonstrates that both wasp species similarly exploit the anti-aphrodisiac mixture of methyl salicylate and indole of another host, the more common solitary small cabbage white butterfly Pieris rapae that deposits only one egg at a time. Interestingly, this behavior is innate in T. brassicae, whereas T. evanescens learns it after one successful ride on a mated female butterfly. Moreover, we show that the wasps only respond to the anti-aphrodisiacs of the 2 cabbage white butterflies when the ubiquitous compounds are part of a complete mated female odor blend. Obviously, parasitic wasps use the sophisticated espionage-and-ride strategy to find eggs of different gregarious and solitary host species. From the wasps' perspective there seems to be a trade-off between the abundance and egg-laying behavior of the butterflies. Our findings suggest that Pieris butterflies are under strong selective pressure to minimize the use of an anti-aphrodisiac. Copyright 2010, Oxford University Press.

The mechanisms through which trophic interactions between species are indirectly mediated by distant members in a food web have received increasing attention in the field of ecology of multitrophic interactions. Scarcely studied aspects include the effects of varying plant chemistry on herbivore immune defences against parasitoids. We investigated the effects of constitutive and herbivore-induced variation in the nutritional quality of wild and cultivated populations of cabbage (Brassica oleracea) on the ability of small cabbage white Pieris rapae (Lepidoptera, Pieridae) larvae to encapsulate eggs of the parasitoid Cotesia glomerata (Hymenoptera, Braconidae). Average encapsulation rates in caterpillars parasitised as first instars were low and did not differ among plant populations, with caterpillar weight positively correlating with the rates of encapsulation. When caterpillars were parasitised as second instar larvae, encapsulation of eggs increased. Caterpillars were larger on the cultivated Brussels sprouts plants and exhibited higher levels of encapsulation compared with caterpillars on plants of either of the wild cabbage populations. Observed differences in encapsulation rates between plant populations could not be explained exclusively by differences in host growth on the different Brassica populations. Previous herbivore damage resulted in a reduction in the larval weight of subsequent herbivores with a concomitant reduction in encapsulation responses on both Brussels sprouts and wild cabbage plants. To our knowledge this is the first study demonstrating that constitutive and herbivore-induced changes in plant chemistry act in concert, affecting the immune response of herbivores to parasitism. We argue that plant-mediated immune responses of herbivores may be important in the evaluation of fitness costs and benefits of herbivore diet on the third trophic level.

Through artificial selection, domesticated plants often contain modified levels of primary and secondary metabolites compared to their wild progenitors. It is hypothesized that the changed chemistry of cultivated plants will affect the performance of insects associated with these plants. In this paper, the development of several specialist and generalist herbivores and their endoparasitoids were compared when reared on a wild and cultivated population of cabbage, Brassica oleracea, and a recently established feral Brassica species. Irrespective of insect species or the degree of dietary specialization, herbivores and parasitoids developed most poorly on the wild population. For the specialists, plant population influenced only development time and adult body mass, whereas for the generalists, plant populations also affected egg-to-adult survival. Two parasitoid species, a generalist (Diadegma fenestrale) and a specialist (D. semiclausum), were reared from the same host (Plutella xylostella). Performance of D. semiclausum was closely linked to that of its host, whereas the correlation between survival of D. fenestrale and host performance was less clear. Plants in the Brassicaceae characteristically produce defense-related glucosinolates (GS). Levels of GS in leaves of undamaged plants were significantly higher in plants from the wild population than from the domesticated populations. Moreover, total GS concentrations increased significantly in wild plants after herbivory, but not in domesticated or feral plants. The results of this study reveal that a cabbage cultivar and plants from a wild cabbage population exhibit significant differences in quality in terms of their effects on the growth and development of insect herbivores and their natural enemies. Although cultivated plants have proved to be model systems in agroecology, we argue that some caution should be applied to evolutionary explanations derived from studies on domesticated plants, unless some knowledge exists on the history of the system under investigation.

In nature, individuals of short-lived plant species (e.g. annuals, biennials) may grow at different times during the growing season. These plants are therefore exposed to different season-related conditions such as light and temperature, which in turn may have consequences for primary and secondary chemistry of the plant. Despite this, many studies examining plant–consumer interactions are performed in single replicates, which may thus ignore temporal variation in the expression of phenotypic plant traits that affect multitrophic interactions. In the present study, we demonstrated that even under strictly controlled conditions in a greenhouse, secondary plant chemistry changes dramatically in plants growing at different times in a single year, i.e. July, August and November. Glucosinolate (GS) contents in herbivore-damaged leaves of two different crucifer species, Brassica oleracea and Sinapis alba were higher in the August and November replicates than in the July replicate and GS concentrations were 10–25 times higher in S. alba than in B. oleracea. The development of a specialist herbivore, Plutella xylostella, also varied significantly over the three replicates. Larvae of P. xylostella that had fed upon either S. alba or B. oleracea, attained the largest biomass and had the fastest development rate in the November replicate. Female P. xylostella moths grew larger on S. alba than on B. oleracea, whereas male biomass was not significantly affected by host-plant species. Plant species, but not season also affected performance of the endoparasitoid, Diadegma semiclausum. Similar to the performance of host females, parasitoid males developed faster and attained the highest biomass when attacking P. xylostella larvae feeding on S. alba. Most importantly, the performance of the herbivore and its parasitoid only appeared to partially conform to levels of GS in damaged leaves, indicating that there is a complex of factors involved in determining the effects of plant quality on higher trophic levels.

Long-term memory (LTM) formation usually requires repeated, spaced learning events and is achieved by the synthesis of specific proteins. Other memory forms require a single learning experience and are independent of protein synthesis. We investigated in two closely related parasitic wasp species,Cotesia glomerataandCotesia rubecula, whether natural differences in foraging behaviour are correlated with differences in LTM acquisition and formation. These parasitic wasp species lay their eggs in young caterpillars of pierid butterflies and can learn to associate plant odours with a successful egg laying experience on caterpillars on the odour-producing plant. We used a classical conditioning set-up, while interfering with LTM formation through translation or transcription inhibitors. We show here thatC. rubeculaformed LTM after three spaced learning trials, whereasC. glomeratarequired only a single trial for LTM formation. After three spaced learning trials, LTM formation was complete within 4 h inC. glomerata, whereas inC. rubecula, LTM formation took 3 days. Linking neurobiology with ecology, we argue that this species-specific difference in LTM acquisition and formation is adaptive given the extreme differences in both the number of foraging decisions of the two wasp species and in the spatial distributions of their respective hosts in nature.

Differences in allelochemistry of plants may influence their ability to attract parasitoids.We studied responses of Diadegma semiclausum (Hellén), a parasitoid of the diamondback moth (Plutella xylostella L.), to inter- and intraspecific variation in odor blends of crucifers and a non-crucifer species. Uninfested Brussels sprout (Brassica oleracea L. gemmifera), white mustard (Sinapis alba L.), a feral Brassica oleracea, and malting barley (Hordeum vulgare L.) were compared for their attractivity to D. semiclausum in a Y-tube bioassay. Odors from all plants were more attractive to the parasitoid than clean air. However, tested against each other, parasitoids preferred the volatile blend from the three cruciferous species over that of malting barley.Wasps also discriminated between uninfested crucifers: mustard was as attractive as feral B. oleracea, and both were more attractive than Brussels sprout. Attractivity of uninfested plants was compared with that of plants infested by larvae of the host P. xylostella. Host-infested mustard and Brussels sprout were more attractive than uninfested conspecifics. Interestingly, the volatile blends of uninfested white mustard and infested Brussels sprout were equally attractive.We also compared the volatile composition of different plant sources by collecting headspace samples and analysing them with GC-MS. Similarities of volatile profiles were determined by hierarchic clustering and non-metric scaling based on the Horn-index. Due to the absence of several compounds in its blend, the volatile profile of barley showed dissimilarities from blends of crucifers. The odor profile of white mustard was distinctly different from the two Brassicaceae.Feral Brassica oleracea odor profile was different from infested Brussels sprout, but showed overlap with uninfested Brussels sprout. Odor blends from infested and uninfested Brussels sprout were similar, and mainly quantitative differences were found. D. semiclausum appears to discriminate based on subtle differences in volatile composition of odor blends from infested and uninfested plants. [KEYWORDS: Diadegma semiclausum ; Plutella xylostella ; Hordeum vulgare ; Sinapis alba ; Brassica oleracea ; olfactometer ; headspace volatiles ; GC-MS]

In order to explore the behavioral mechanisms underlying aggregation of foragers on local resource patches, it is necessary to manipulate the location, quality and quantity of food patches. This requires careful control over the conditions in the foraging arena, which may be a challenging task in the case of aquatic resource-consumer systems, like that of freshwater zooplankton feeding on suspended algal cells. We present an experimental tool designed to aid behavioral ecologists in exploring the consequences of resource characteristics for zooplankton aggregation behavior and movement decisions under conditions where the boundaries and characteristics (quantity and quality) of food patches can be standardized. The aggregation behavior of Daphnia magna and D. galeata x hyalina was tested in relation to i) the presence or absence of food or ii) food quality, where algae of high or low nutrient (phosphorus) content were offered in distinct patches. Individuals of both Daphnia species chose tubes containing food patches and D. galeata x hyalina also showed a preference towards food patches of high nutrient content. We discuss how the described equipment complements other behavioral approaches providing a useful tool to understand animal foraging decisions in environments with heterogeneous resource distributions.

Trait-mediated effects modify patch-size density relationships in insect herbivores and parasitoids

Constitutive and induced changes in plant quality impact higher trophic levels, such as the development 27 of parasitoids, in different ways. An efficient way to study how plant quality affects parasitoids is to 28 examine how the parasitoid larva is integrated within the host during the growth process. In two exper- 29 iments, we investigated the effects of varying nutritional quality of Brassica oleracea on parasitoid larval 30 development inside the host, the diamondback moth (Plutella xylostella). First, we compared larval 31 growth of the specialist Diadegma semiclausum and the generalist Diadegma fenestrale, when the host 32 was feeding on Brussels sprout plants that were either undamaged or were previously induced by cater- 33 pillar damage. Larvae of the generalist D. fenestrale showed lower growth rates than larvae of the special- 34 ist D. semiclausum, and this difference was more pronounced on herbivore-induced plants, suggesting 35 differences in host-use efficiency between parasitoid species. The growth of D. semiclausum larvae was 36 also analyzed in relation to herbivore induction on Brussels sprouts and on a wild B. oleracea strain. Par- 37 asitoid growth wasmore depressed on induced than on undamaged control plants, andmore on wild cab- 38 bage than on Brussels sprouts, which was largely explained by differences in host mass. The effects of 39 induction of wild Brassica on parasitoid development were pronounced early on, but as P. xylostella feed- 40 ing began inducing the previously undamaged control plants, the effect of induction disappeared, reveal- 41 ing a temporal component of plant–parasitoid interactions. This study demonstrates how insights into 42 the physiological aspects of host–parasitoid interactions can improve our understanding of the effects 43 of plant-related traits on parasitoid wasps.

In the field, plants are attacked by several herbivore species both simultaneously and in isolation. Spatial variation in damage to plants by different herbivores may affect the search behaviour of parasitoid wasps, but the consequences of this variation for hosteparasitoid interactions are still little understood. We examined the effects of multiple herbivory on the search behaviour of the parasitoid Cotesia glomerata. Reduced parasitism was found in a field tent experiment, when both the host small cabbage white, Pieris rapae, and the nonhost cabbage moth, Mamestra brassicae, were present on Brassica oleracea plants. When hosts and nonhosts occurred on the same or neighbouring plants, this reduction in parasitism was similar, but the underlying behavioural mechanisms were different. In wind tunnel bioassays, parasitoids were equally attracted towards plants infested by the two herbivore species but were more attracted to plants with both herbivore species than to plants with only one. Differences in arrival tendencies therefore could not explain the reduced parasitism in the tent experiment with mixed infestations. Experiments showed that parasitoids readily left nonhost patches, whereas leaving tendencies from mixed patches of hosts and nonhosts were the same as from pure host patches. Therefore, reduced leaving tendencies and reduced host encounters explained the lower parasitism rate in mixed infestations in the tent experiment. Our results show that the spatial context in which hosts and nonhosts attack plants determines the foraging efficiency of parasitoids, with consequences for hosteparasitoid interactions.

Herbivore-induced plant volatiles (HIPV) provide important information that influences host location behaviour for insect natural enemies, such as parasitoid wasps, that develop in the bodies of herbivorous insects. The dietary breadth of both the parasitoid and its host may affect the extent to which a searching parasitoid relies on HIPV. Specialist species are expected to rely on specific volatile cues to which they respond innately, whereas generalists are expected to show a higher degree of phenotypic plasticity that depends on foraging experience in the parasitoid. We compared the response to HIPV emitted by different plant species damaged by host and nonhost caterpillars for two congeneric parasitoid species, the specialist Diadegma semiclausum and the generalist Diadegma fenestrale, attacking caterpillars of the diamondback moth, Plutella xylostella. For the three tested plant species, Brassica oleracea, a feral Brassica population and Sinapis alba, both parasitoid species preferred volatiles from host-infested plants over those produced by undamaged plants. However, both parasitoid species only distinguished between volatiles induced by host and nonhosts when the caterpillars had been feeding on B. oleracea, the plant on which they had been reared. Chemical analysis of the volatile blends could not explain volatile preferences of the parasitoids. Despite the difference in their dietary breadth, the two parasitoids responded similarly to HIPV and experience treatments. A flexible response to a wide array of volatile blends by parasitoids is probably important in nature, given that different generations of the host and the parasitoid probably develop on different food plants.

Animals form memory types that differ in duration and stability. The initial anaesthesia-sensitive memory (ASM) can be replaced by anaesthesia-resistant memory (ARM), and/or by protein synthesis-dependent, long-term memory (LTM). We previously showed that two closely related parasitic wasp species differ in learning rate and memory consolidation. In Cotesia glomerata, LTM lasting at least 24 h was formed after single-trial conditioning, whereas single-trial conditioning led to ARM that waned before 24 h in Cotesia rubecula. This species formed LTM only after repeated conditioning trials spaced in time. Here, we used artificial selection on learning rate to investigate whether selection for a low learning rate in C. glomerata would result in C. rubecula-like memory dynamics. Memory consolidation was tested by using cold-shock anaesthesia and protein synthesis inhibitors. After single-trial conditioning, ARM was consolidated within hours in unselected C. rubecula, but directly, without an intermediate ARM phase, into LTM in unselected C. glomerata. We obtained low learning rate selection lines of C. glomerata wasps that, like C. rubecula, did not form LTM after single-trial conditioning, to see whether such wasps would then consolidate ARM instead of LTM. We showed that this was not the case. The selected wasps formed LTM after repeated, spaced conditioning trials, but formed only ASM without consolidation of ARM or LTM after single-trial learning. Ecological consequences of this type of memory formation are discussed.

Despite a large number of studies on herbivore-induced plant volatiles (HIPVs), little is known about which specific compounds are used by natural enemies to locate prey- or host- infested plants. In addition, the role of HIPVs in attracting natural enemies has been restricted largely to agricultural systems. Differences in volatile blends emitted by cultivars and plants that originate from wild populations may be attributed to potentially contrasting selection regimes: natural selection among the wild types and artificial selection among cultivars. A more realistic understanding of these interactions in a broader ecological and evolutionary framework should include studies that involve insect herbivores, parasitoids, and wild plants on which they naturally interact in the field. We compared the attractiveness of HIPVs emitted by wild and cultivated cabbage to the parasitoid Cotesia rubecula, and determined the chemical composition of the HIPV blends to elucidate which compounds are involved in parasitoid attraction. Wild and cultivated cabbage differed significantly in their volatile emissions. Cotesia rubecula was differentially attracted to the wild cabbage populations and preferred wild over cultivated cabbage. Isothiocyanates, which were only emitted by the wild cabbages, may be the key components that explain the preference for wild over cultivated cabbage, whereas terpenes may be important for the differential attraction among the wild populations. Volatile analysis revealed that parasitoid attraction cannot be explained by simple linear relationships. Our results suggest that unraveling which compound(s) are innately attractive to parasitoids of cabbage pests should include wild Brassicaceae.

Many insects possess a sexual communication system that is vulnerable to chemical espionage by parasitic wasps. We recently discovered that a hitch-hiking (H) egg parasitoid exploits the antiaphrodisiac pheromone benzyl cyanide (BC) of the Large Cabbage White butterfly Pieris brassicae . This pheromone is passed from male butterflies to females during mating to render them less attractive to conspecific males. When the tiny parasitic wasp Trichogramma brassicae detects the antiaphrodisiac, it rides on a mated female butterfly to a host plant and then parasitizes her freshly laid eggs. The present study demonstrates that a closely related generalist wasp, Trichogramma evanescens , exploits BC in a similar way, but only after learning. Interestingly, the wasp learns to associate an H response to the odors of a mated female P. brassicae butterfly with reinforcement by parasitizing freshly laid butterfly eggs. Behavioral assays, before which we specifically inhibited long-term memory (LTM) formation with a translation inhibitor, reveal that the wasp has formed protein synthesis-dependent LTM at 24 h after learning. To our knowledge, the combination of associatively learning to exploit the sexual communication system of a host and the formation of protein synthesis-dependent LTM after a single learning event has not been documented before. We expect it to be widespread in nature, because it is highly adaptive in many species of egg parasitoids. Our finding of the exploitation of an antiaphrodisiac by multiple species of parasitic wasps suggests its use by Pieris butterflies to be under strong selective pressure.

The diversity and complexity of food webs (the networks of feeding relationships within an ecological community) are considered to be important factors determining ecosystem function and stability. However, the biological processes driving these factors are poorly understood. Resource quality affects species interactions by limiting energy transfer to consumers and their predators, affecting life history and morphological traits. We show that differences in plant traits affect the structure of an entire food web through a series of direct and indirect effects. Three trophic levels of consumers were influenced by plant quality, as shown by quantitative herbivore–parasitoid–secondary parasitoid food webs. We conclude, on the basis of our data, that changes in the food web are dependent on both trait- and density-mediated interactions among species.

1. Changing plant composition in a community can have profound consequences for herbivore and parasitoid population dynamics. To understand such effects, studies are needed that unravel the underlying behavioural decisions determining the responses of parasitoids to complex habitats. 2. The searching behaviour of the parasitoid Diadegma semiclausum was followed in environments with different plant species composition. In the middle of these environments, two Brassica oleracea plants infested by the host Plutella xylostella were placed. The control set-up contained B. oleracea plants only. In the more complex set-ups, B. oleracea plants were interspersed by either Sinapis alba or Hordeum vulgare. 3. Parasitoids did not find the first host-infested plant with the same speed in the different environments. Sinapis alba plants were preferentially searched by parasitoids, resulting in fewer initial host encounters, possibly creating a dynamic enemy-free space for the host on adjacent B. oleracea plants. In set-ups with H. vulgare, also, fewer initial host encounters were found, but in this case plant structure was more likely than infochemicals to interfere with the searching behaviour of parasitoids. 4. On discovering a host-infested plant, parasitoids located the second host-infested plant with equal speed, demonstrating the effect of experience on time allocation. Further encounters with host-infested plants that had already been visited decreased residence times and increased the tendency to leave the environment. 5. Due to the intensive search of S. alba plants, hosts were encountered at lower rates here than in the other set-ups. However, because parasitoids left the set-up with S. alba last, the same number of hosts were encountered as in the other treatments. 6. Plant composition of a community influences the distribution of parasitoid attacks via its effects on arrival and leaving tendencies. Foraging experiences can reduce or increase the importance of enemy-free space for hosts on less attractive plants.

Several methods are available for analyzing different aspects of behavioral transition matrices, but a comprehensive framework for their use is lacking. We analyzed parasitoid foraging behavior in environments with different plant species compositions. The resulting complex data sets were analyzed using the following stepwise procedure. We detected abrupt changes in the event log files of parasitoids, using a maximum likelihood method. This served as a criterion for splitting the event log files into two parts. For both parts, Mantel’s test was used to detect differences between first-order transition matrices, whereas an iterative proportional fitting method was used to find behavioral flows that deviated from random transitions. In addition, hidden repetitive sequences were detected in the transition matrices on the basis of their relative timing, using Theme. We discuss the results for the example from a biological context and the comprehensive use of the different methods. We stress the importance of such a combined stepwise analysis for detecting differences in some parts of event log files.

1. Habitat complexity may stabilize interactions among species of different trophic levels by providing refuges to organisms of lower trophic levels. 2. Searching behaviour of the parasitoid, Diadegma semiclausum, was followed in different semifield set-ups, a low and high-density monoculture of Brassica oleracea and two intercrops, B. oleracea with Sinapis alba (also a member of the Brassicaceae) and B. oleracea with Hordeum vulgare (Poaceae). 3. When a low-density monocrop of B. oleracea was compared with a high-density monocrop, no differences were found in the ability of the female wasps to locate a host-infested plant, B. oleracea, infested with Plutella xylostella that was placed in the centre of the set-up. 4. The efficiency of the parasitoid to locate the host-infested plant was differentially affected by the species composition of the vegetation. Wasps entered the Sinapis-Brassica set-up faster, but took more time to find the host-infested plant than in the Hordeum-Brassica set-up. 5. The horizontal arrangement, i.e. by mixing S. alba or H. vulgare with, or placing them as rows between B. oleracea, did not affect host-finding efficiency. 6. Plant height did influence host finding. Wasps found the host-infested plants earlier in the set-up with short Sinapis plants compared with tall Sinapis plants. 7. Once the wasps had landed on the host-infested plant, the surrounding vegetation did not affect time needed to parasitize five consecutive hosts on the same infested plant, regardless of the composition or horizontal/vertical arrangement of the set-up. 8. Chemical and structural refuges in complex landscapes may play an important role in the persistence of this system through dampening oscillations of parasitoid and host populations. [KEYWORDS: crucifers ; Diadegma semiclausum ; Plutella xylostella ; proportional hazards model ; refuges]

Experimental data on the relationship between plant patch size and population density of herbivores within fields often deviates from predictions of the theory of island biogeography and the resource concentration hypothesis. Here we argue that basic features of foraging behaviour can explain different responses of specialist herbivores to habitat heterogeneity. In a combination of field and simulation studies, we applied basic knowledge on the foraging strategies of three specialist herbivores: the cabbage aphid (Brevicoryne brassicae), the cabbage butterfly (Pieris rapae L.) and the diamondback moth (Plutella xylostella L.), to explain differences in their responses to small scale fragmentation of their habitat. In our field study, populations of the three species responded to different sizes of host plant patches (9 plants and 100 plants) in different ways. Densities of winged cabbage aphids were independent of patch size. Egg-densities of the cabbage butterfly were higher in small than in large patches. Densities of diamondback moth adults were higher in large patches than in small patches. When patches in a background of barley were compared with those in grass, densities of the cabbage aphid and the diamondback moth were reduced, but not cabbage butterfly densities. To explore the role of foraging behaviour of herbivores on their response to patch size, a spatially explicit individual-based simulation framework was used. The sensory abilities of the insects to detect and respond to contact, olfactory or visual cues were varied. Species with a post-alighting host recognition behaviour (cabbage aphid) could only use contact cues from host plants encountered after landing. In contrast, species capable with a pre-alighting recognition behaviour, based on visual (cabbage butterfly) or olfactory (diamondback moth) cues, were able to recognise a preferred host plant whilst in flight. These three searching modalities were studied by varying the in flight detection abilities, the displacement speed and the arrestment response to host plants by individuals. Simulated patch size - density relationships were similar to those observed in the field. The importance of pre- and post- alighting detection in the responses of herbivores to spatial heterogeneity of the habitat is discussed

In the light of current theories on the effects of intercropping on pest reduction, population responses of the diamondback moth (Plutella xylostella), the cabbage aphid (Brevicoryne brassicae) and the life history traits of the large white butterfly (Pieris brassicae) were studied in a Brussels sprout (Brassica oleracea gemmifera)/malting barley (Hordeum vulgare) additive row intercrop and a Brussels sprout monoculture. More P. xylostella adults were caught in the monoculture than in the intercrop. Numbers of P. xylostella larvae and pupae per sprout plant were lower in intercropped plots than in monocultures. However, more larvae and pupae were found per m2 leaf area in the inter- than in the monocrop. Both the densities per plant and per m2 leaf area of B. brassicae populations were lower in the inter- than in the monocrop. After the barley withered and competition with Brussels sprout abated, aphid densities became higher in the inter- than in the monocrop. These findings may be explained by interspecific plant competition resulting in stressed sprout plants with a smaller size and delayed phenology relative to monocropped plants. Effects of differences in plant nutritional quality on herbivore performance were studied by offering leaves of inter- and monocropped sprout plants to larval P. brassicae. Performance and food utilisation were significantly better on leaves from the intercrop (lower dry weight consumption, higher growth rates) than from the monocrop. Defoliation rate was also higher on leaves of intercropped plants than on monocropped ones. The studies indicate that plant stress and consequent changes in developmental rate and nutritional quality of plants are playing a role in herbivore population responses to intercropping. It is argued that such confounding effects of plant competition in intercropping designs can hamper the evaluation of herbivore responses in pest-suppressive agro-ecosystems. [KEYWORDS: Intercropping; Interspecific competition; "Host plant quality hypothesis"; Brevicoryne brassicae; Plutella xylostella; Pieris brassicae]

Populations of wild Brassica oleracea L. grow naturally along the Atlantic coastlines of the United Kingdom and France. Over a very small spatial scale (i.e.