Volatile attractants for three Pteromalid parasitoids attacking concealed spruce bark beetles

Summary. The odour perceptive abilities, and preferences, of three bark beetle parasitoid species; Rhopalicus tutela (Walker), Roptrocerus mirus (Walker), and Roptrocerus xylophagorum (Ratzeburg) (Hymenoptera: Pteromalidae), were investigated to isolate and identify the essential compounds involved in host location. These parasitoids attack several economically important bark beetle species and oviposit preferentially on late larval stages concealed under the bark of conifers. Odours were collected from Norway spruce logs (Picea abies L. Karst.) containing Ips typographus L. (Coleoptera: Scolytidae) larvae. Biologically active compounds were isolated by coupled gas chromatographic-electroantennographic detection (GC-EAD), and identified by GC-mass spectrometry (GC-MS). Based on these analyses, four different synthetic baits were prepared and tested in a Y-tube walking bioassay. In the complex odour samples from spruce logs, only 16 compounds were EAD-active. The tested R. tutela and R. mirus females displayed similar trends in antennal activity to EAD-active compounds, responding mainly to oxygenated monoterpenes that indicate damaged conifers. Consequently, the synthetic baits were exclusively prepared with oxygenated monoterpenes. Parasitoid females (R. tutela and R. mirus) preferred spruce logs containing susceptible hosts over fresh logs, while male parasitoids (R. mirus) did not show any preference. However, when odours from fresh logs were mixed with synthetic baits (mimicking the odour composition of logs containing susceptible hosts), these combinations attracted female parasitoids (R. tutela, R. mirus, and R. xylophagorum). All synthetic baits seemed to be equally attractive to female parasitoids. Received 12 October 2000; accepted 18 January 2001     

Attraction,deterrence or intoxication of bees (Apis mellifera) by plant allelochemicals

The influence of 63 dietary allelochemicals (alkaloids, terpenes, glycosides,etc.) on the feeding behaviour of bees (Apis mellifera) was tested in terms of deterrency and attraction. For 39 compounds a deterrent (mostly alkaloids, coumarins and saponins) and for 3 compounds an attractive response (mostly terpenes) was obtained in choice tests, which allowed the calculation of respective ED₅₀-values. Under no-choice conditions, 17 out of 29 allelochemicals caused mortality at concentrations between 0.003 and 0.6%. Especially toxic were alkaloids, saponins, cardiac glycosides and cyanogenic glycosides. These data show that bees which are confronted with plant allelochemicals in nectar and pollen, are not especially adapted (i.e. insensitive) to the plants' defence chemistry. GLC and GLS-MS data are given on the alkaloid composition of nectar and pollen ofBrugmansia aurea, Atropa belladonna andLupinus polyphyllus.     

Sequestration of plant secondary compounds by butterflies and moths

Summary A number of aposematic butterfly and moth species sequester toxic substances from their host plants. Some of these insects can detect the toxic compounds during food assessment. Some pipevine swallowtails use aristolochic acids among the host finding cues during oviposition and larval feeding and accumulate the toxins in the body tissues throughout all life stages. Likewise, a danaine butterfly,Idea leuconoe, which sequesters high concentrations of pyrrolizidine alkaloids in the body, lays eggs in response to the specific alkaloid components contained in the apocynad host. Insect species sharing the same poisonous host plants may differ in the degree of sequestration of toxins. Two closely ralated aposematic geometrid moth species,Arichanna gaschkevitchii andA. melanaria, sequester a series of highly toxic diterpenoids (grayanotoxins) in different degrees, while a cryptic geometrid species,Biston robstus, does not sequester the toxins, illustrating the diversity in adaptation mechanisms even within the same subfamily. By contrast, a number of lepidopteran species store the same compounds though feeding upon taxonomically diverse plant species. A bitter cyanoglycoside, sarmentosin, was characterised from several moth species in the Geometridae, Zygaenidae and Yponomeutidae, and from the apollo butterflies,Parnassius spp. (Papilionidae), although each species feeds on different groups of plants.Interspecific similarities and differences in life history and ecology are discussed in relation to variable characteristics of sequestration of plant compounds among these lepidopteran insects.     

Parasitoid learning during interactions with ants: how to deal with an aggressive antagonist

Pauesia picta, P. pinicollis and P. silvestris (Hymenoptera: Aphidiidae) are common parasitoids of the conifer aphid Cinara pinea, which is regularly attended by red wood ants (Formica spp.). In this study, I tested whether females of these parasitoid species learned during interactions with honeydew-collecting Formica polyctena workers that caution is not necessary if searching behaviour is adapted, and whether parasitoids benefit from being able to learn. When searching on Scots pines, naive females of P. picta and P. pinicollis generally retreated to a pine needle when making contact with a honeydew-collecting ant, did not approach ants from the side or from the front and kept a ”safe distance” from ants when sitting on needles. After some non- aggressive ant encounters, experienced female parasitoids changed their behaviour: they reduced their searching speed, approached ants from the side and even from the front, retreated less often in response to an approaching ant and reduced the ”safe distance”. These experienced females had a significantly higher rate of oviposition than naive females or females foraging for an unattended host. Thus, the ability of the parasitoid to learn during interactions with an antagonist led to a prolonged retention time and a higher oviposition rate. By contrast, there was no evidence of learning in P. silvestris. Females of this species showed no behavioural change in response to ant encounters, and there was no difference in the foraging success of naive and experienced female parasitoids. Received: 7 December 1999 / Revised: 23 September 2000 / Accepted: 10 March 2000     

Antennal responses of the western pine beetle, <Emphasis Type="Italic">Dendroctonus brevicomis</Emphasis> (Coleoptera: Curculionidae), to stem volatiles of its primary host, <Emphasis Type="Italic">Pinus ponderosa</Emphasis>, and nine sympatric nonhost angiosperms and conifers

Summary. Stem volatile extracts from ten trees that are sympatric with the western pine beetle, Dendroctonus brevicomis LeConte (Coleoptera: Curculionidae) were assayed by gas chromatographic-electroantennographic detection analysis (GC-EAD). The extracts were from the primary host, ponderosa pine, Pinus ponderosa Dougl. ex Laws. (Pinaceae); two nonhost angiosperms, California black oak, Quercus kelloggii Newb. (Fagaceae), and quaking aspen, Populus tremuloides Michx. (Salicaceae); and seven nonhost conifers, white fir, Abies concolor (Gord. & Glend.) Lindl. ex Hildebr. (Pinaceae), incense cedar, Calocedrus decurrens (Torr.) Florin (Cupressaceae), Sierra lodgepole pine, P. contorta murrayana Grev. & Balf. (Pinaceae), Jeffrey pine, P. jeffreyi Grev. & Balf. (Pinaceae), sugar pine, P. lambertiana Dougl. (Pinaceae), Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco (Pinaceae), and mountain hemlock, Tsuga mertensiana (Bong.) Carr. (Pinaceae). Sixty-four compounds were identified from the ten trees, 42 of which elicited antennal responses in D. brevicomis, usually in both sexes. In addition, several synthetic compounds, including a number of the antennally-active compounds from the extracted trees and some bark beetle pheromone components, elicited antennal responses in a manner similar to that observed with the extracts. Of the antennally-active compounds known to be present in trees sympatric with D. brevicomis, only geraniol was unique to its host. Four antennally-active compounds were found in the host and in other conifers; five compounds were found only in nonhost conifers; eight compounds were found in either or both of the nonhost angiosperms; eight compounds were found in either or both of the angiosperms and in nonhost conifers, but not in the host; and 19 were found in both the host and in angiosperms and/or nonhost conifers. Several bark beetle pheromone components were found in the stem volatile extracts. Conophthorin was identified from both nonhost angiosperms; exo-brevicomin was identified in A. concolor; verbenone was identified from a number of nonhost conifers; and chalcogran was identified from P. tremuloides. The number of nonhost volatile chemicals that D. brevicomis encounters and is capable of detecting, and the diversity of sources from which they emanate, highlight the complexity of the olfactory environment in which D. brevicomis forages. This provides a basis for further work related to chemically-mediated aspects of foraging in this insect and perhaps other coniferophagous bark beetles, and highlights the need to consider foraging context in the design and implementation of semiochemical-based management tactics for tree protection.     

Mother’s choice of the oviposition site: balancing risk of egg parasitism and need of food supply for the progeny with an infochemical shelter?

Summary. Oviposition site selection of herbivorous insects depends primarily on host plant presence which is essential for offspring survival. However, parasitoids can exploit host plant cues for host location. In this study, we hypothesised that herbivores can solve this dilemma by ovipositing within high plant diversity. A diverse plant species composition might represent an ‘infochemical shelter’, as a potentially complex volatile blend can negatively affect the host location ability of parasitoids. We examined this exemplarily for the egg-laying response of the generalist leaf beetle, Galeruca tanaceti, in relation to (1) host plant availability and (2) plant species diversity in the field. Further, we investigated the effect of odours from mixed plant species compositions on (3) leaf beetle oviposition site selection and on (4) the orientation of its specialised egg parasitoid, Oomyzus galerucivorus. In the field, egg clutch occurrence was positively related to the presence and quantity of two major host plants, Achillea millefolium (yarrow) and Centaurea jacea, and to the number of herbaceous plant species. In two-choice bioassays, female beetles oviposited more frequently on sites surrounded by an odour blend from a diverse plant species composition (including yarrow) than on sites with a pure grass odour blend. In the presence of yarrow odour and an odour blend from a diverse plant mixture (including yarrow) no difference in the oviposition response was recorded. Experienced parasitoid females were attracted to yarrow odours, but showed no response when yarrow odours were offered simultaneously with odours of a non-host plant. In conclusion, it could be shown in laboratory bioassays that the parasitoid responds only to pure host plant odours but not to complex odour blends. In contrast, the herbivore prefers to oviposit within diverse vegetation in the field and in the laboratory. However, the laboratory results also point to a priority of host plant availability over the selection of a potential ‘infochemical shelter’ for oviposition due to high plant diversity.     

Differential use of conspecific-derived information by sexual and asexual parasitic wasps exploiting partially depleted host patches

When foraging partially depleted patches (i.e., a fraction of hosts are already parasitized), female parasitoids must decide: 1—whether to superparasitize, and 2—whether to stay in their current patch (thus missing the opportunity of finding a better patch elsewhere). To make these decisions, parasitoids may rely on different cues, produced both by the environment and by conspecifics. Animals thriving in different environments may differ in cues they use. In the solitary parasitoid Venturia canescens, thelytokous (asexual) and arrhenotokous (sexual) individuals are found in two contrasting environments. Thelytokous females, from anthropogenic conditions, are known to cope with superparasitism in an adaptive way. On the other hand, little is known about superparasitism by arrhenotokous females. We compared the host exploitation strategies of thelytokous and arrhenotokous females in partially depleted patches. Hosts parasitized by thelytokous females were more frequently avoided than those parasitized by arrhenotokous females, suggesting a stronger chemical marking of the former. Only thelytokous females used information from conspecifics for patch-leaving decisions. The conformity of the differences in the behavior of thelytokous and arrhenotokous females with the environmental conditions they experience in their habitat is discussed.     

Evolutionary strategies of chemical defense in aposematic butterflies: Cyanogenesis in Asteraceae-feeding American Acraeinae

Summary American Acraeinae butterflies often ingest large amounts of dehydropyrrolizidine alkaloids (PAs) from their Asteraceae hostplants in both larval and adult stages, but do not normally store these compounds for defence, instead biosynthesizing large amounts of the cyanogenic glucoside linamarin in all stages. This defence syndrome (rejection of plant toxins andde novo synthesis of protective chemicals) is considered to be the most evolved among aposematic (unpalatable mimicry-model) butterflies, as are the Acraeinae and Heliconiini which also synthesize cyanogens. Storage or minimal processing of larval hostplant-derived defensive chemicals is widespread and characterizes the most primitive model groups; an intermediate series (Danainae/Ithomiinae) also obtains the principal defensive chemicals (PAs) from plants, but mostly in the adult stage. These syndromes are discussed and contrasted with the pattern seen in Chrysomelidae beetles, wherede novo synthesis is widespread and considered primitive.Originally presented at the 9th Meeting on Micromolecular Evolution, Systematics and Ecology (Ribeirão Preto, SP, April 1987); see also Brownet al. (1990)     

Specificity of odour-mediated avoidance of competition in Drosophila parasitoids

Although there are many examples of the role of volatile infochemicals in interactions between trophic levels of insect communities, surprisingly little is known of volatile interactions between species within the third trophic level. Recently it was found that Leptopilina heterotoma, an endoparasitoid that attacks Drosophila larvae, avoids one type of patches (decaying stinkhorn mushrooms) when parasitoids of another species (L. clavipes) are present on these patches. L. heterotoma is able to smell the presence of L. clavipes from a distance (Fig. 1). In this paper we investigate the source of the odour that induces avoidance behaviour, by varying the host species and parasitoid species present on stinkhorn mushrooms, and by using another type of patch (sap-fluxes of wounded trees). L. heterotoma was found to avoid stinkhorn patches with conspecific as well as heterospecific parasitoids (Fig. 2). Hosts had to be present in the patch to elicit avoidance, but avoidance behaviour was also found with another host species present in the patch (Fig. 3). No avoidance behaviour was found with sap-flux patches with hosts and parasitoids on them (Fig. 4). Avoidance of stinkhorn patches only occurred when the parasitoids present on the patch were able to contact hosts (Figs. 5 and 6). The exact source of the odour that elicits avoidance is still unclear, so that one can only speculate on the function of the signal. However, there is a clear benefit to the receiver, because it is able to avoid superior competitors. Avoidance can lead to non-aggregated parasitoid distributions. The importance of avoidance behaviour for population dynamics and stability of parasitoid-host systems is discussed.     

农药类化合物对大型溞的毒性作用模式:与基线毒性化合物比较研究

农药在控制有害生物的同时,对水生生态系统产生较大的毒性效应。本文通过实验获得了25种基线化合物对大型溞的急性毒性数据,并与57种农药类化合物对大型溞的急性毒性数据进行比较研究,同时根据体外浓度LC_(50)、生物富集因子BCF和体内临界浓度CBR的关系,计算了这些化合物在大型溞体内的临界浓度,研究了农药类化合物对大型溞的毒性作用机理。结果表明,基线化合物在大型溞体内的临界浓度log1/CBR值在一个很窄的范围波动,而农药类化合物在大型溞体内的临界浓度log1/CBR值范围广且较高。这说明多数农药类化合物对大型溞为反应性毒性作用模式。其中,除草剂对大型溞的毒性显著低于杀菌剂和杀虫剂对大型溞的毒性。这可能是因为除草剂主要通过干扰植物光合作用、植物激素或植物分子合成发挥毒性效应,从而导致其对大型溞的生理系统难以发生反应性毒性效应。而杀虫剂和杀菌剂主要通过干扰生物神经系统、生殖系统、呼吸作用或大分子合成发挥毒性效应,因此易与大型溞生理系统发生生物化学反应,从而具有较高的毒性效应。本文分别建立了除草剂、杀菌剂和杀虫剂对大型溞的急性毒性QSAR模型。除草剂对大型溞的急性毒性机理较简单,其毒性与化合物疏水性程度和离子化程度有关;而杀菌剂对大型溞的急性毒性主要与化合物的标准生成热和极性表面积有关;杀虫剂对大型溞的急性毒性作用机理较复杂,它们对大型溞的毒性效应与其和生物分子之间的氢键和范德华力有关。     

Potential parasitoid attractants,volatile composition throughout a bark beetle attack

Several hymenopteran parasitoids prey on mature bark beetle larvae (Scolytidae), concealed under the bark of conifers. According to previous results, certain oxygenated monoterpenes are crucial in host location. However, synthetic baits tested in laboratory bioassays are not quite as attractive to the parasitoids as the natural volatile bouquet from conifers containing susceptible bark beetle larvae. The aim of the current study was to identify the "missing components" in previously tested synthetic baits. Volatiles were sampled throughout bark beetle attacks (Ips typographus) on Norway spruce logs (Picea abies), and analyzed (qualitatively and quantitatively) by coupled gas chromatography-mass spectrometry (GC-MS). In addition, a pilot test was done employing the relatively new technique, fast GC-surface acoustic wave detector (SAW), zNose^TM, described in the methods. Quantitative differences in volatile composition were found between infested and uninfested logs, and between different stages of the bark beetle attack. In addition, the volatile composition of logs with or without mature bark beetle larvae (susceptible to parasitism) was compared, isolating odor differences at this most suitable host stage. In general, the quantity of monoterpenes (the predominant phytochemical volatiles in conifers) was found to decrease with time after initial infestation. In contrast, certain oxygenated monoterpenes and benzenoid compounds increased as susceptible bark beetle larvae developed, which therefore could be candidates for improving parasitoid baits. To the best of our knowledge, this is the first report on long-term dynamics of the volatile composition of bark beetle infested P. abies. The results should be interpreted in the light of previous electrophysiological and behavioral investigations of larval parasitoids, and present a number of compounds that could improve synthetic parasitoid baits.     

Antennal responses of four species of tree-killing bark beetles (Coleoptera: Scolytidae) to volatiles collected from beetles,and their host and nonhost conifers

Summary. Host selection in tree-killing bark beetles (Coleoptera: Scolytidae) is mediated by a complex of semiochemical cues. Using gas chromatographic-electroantennographic detection (GC-EAD) and GC-mass spectrometric analyses, we conducted a comparative study of the electrophysiological responses of four species of tree-killing bark beetles, the Douglas-fir beetle, Dendroctonus pseudotsugae, Hopkins, the mountain pine beetle, D. ponderosae Hopkins, the spruce beetle, D. rufipennis Kirby, and the western balsam bark beetle, Dryocoetes confusus Swaine, to volatiles captured by aeration of 1) bole and foliage of four sympatric species of conifers, Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco, lodgepole pine, Pinus contorta var. latifolia Engelm., interior spruce, Picea engelmannii x glauca, and interior fir, Abies lasiocarpa x bifolia, and 2) con- and heterospecific beetles at three stages of attack. We identified 13 monoterpenes in the conifers and nine compounds in the volatiles of beetles that elicited antennal responses. There was no qualitative difference in the terpene constitution of the four species of conifers and very little difference across beetle species in their antennal response to compounds from conifers or beetles. The lack of species-specific major or minor components in conifers suggests that beetles would need to detect differences in the ratios of different compounds in conifers to discriminate among them. Attraction to hosts and avoidance of nonhost conifers may be accentuated by perception of compounds emitted by con- and heterospecific beetles, respectively. The 22 compounds identified are candidate semiochemicals with potential behavioural roles in host location and discrimination.     

Sex pheromones and attractants in the Eucosmini and Grapholitini (Lepidoptera,Tortricidae)

Summary The geometric isomers (E,E)-, (E,Z)-, (Z,E)-, and (Z,Z)-8,10-dodecadien-1-yl acetate were identified as sex pheromone components or sex attractants in the tribes Eucosmini and Grapholitini of the tortricid subfamily Olethreutinae. Species belonging to the more ancestral Tortricinae were not attracted. Each one isomer was behaviourally active in males ofCydia andGrapholita (Grapholitini), either as main pheromone compound, attraction synergist or attraction inhibitor. Their reciprocal attractive/antagonistic activity in a number of species enables specific communication with these four compounds.Pammene, as well as otherGrapholita andCydia responded to the monoenic 8- or 10-dodecen-1-yl acetates. Of the tribes Olethreutini and Eucosmini,Hedya, Epiblema, Eucosma, andNotocelia trimaculana were also attracted to 8,10-dodecadien-1-yl acetates, but several otherNotocelia to 10,12-tetradecadien-1-yl acetates. The female sex pheromones ofC. fagiglandana, C. pyrivora, C. splendana, Epiblema foenella andNotocelia roborana were identified. (E,E)- and (E,Z)-8,10-dodecadien-1-yl acetate are producedvia a commonE9 desaturation pathway inC. splendana. CallingC. nigricana andC. fagiglandana females are attracted to wingfanning males.     

Calanoid copepod,cladoceran, and rotifer eggs in sea-bottom sediments of northern Californian coastal waters: Identification,occurrence and hatching

In Ireland, mussels on exposed rocky shores constitute an interbreeding mixture of two forms of mussels,Mytilus edulis L. and the Mediterranean musselM. galloprovincialis Lmk. This paper presents an in-depth analysis, carried out between October 1984 and December 1986, of genetic variability at two partially diagnostic loci,Odh andEst-D, in two exposed-shore populations ofMytilus spp. in the west of Ireland. Significant differences at theOdh locus were observed in the genetic composition of adult mussels from different tidal levels. These differences were repeatable whether one was analysing replicate samples at a single point in time, samples collected at different points in time, i.e., in different years, or samples collected from different shores. Mussels recruiting to artificial substrates set out for a period of one month at different tidal levels at one of these sites were also observed to be genetically different; mussels higher up the shore exhibited higher frequencies of those alleles characteristically at high frequency inM. galloprovincialis for both theOdh andEst-D loci. Hence, the genetic differences observed in adult mussels are much more exaggerated in juveniles and are already apparent within the first month of benthic life. Possible reasons for the observed microgeographic differentiation are discussed. It is concluded that the observed genetic differences between mussels at different tidal levels arise either in the pelagic/attachment stage or very shortly after settlement.     

Faecal shield chemical defence is not important in larvae of the tortoise beetle Chelymorpha reimoseri (Chrysomelidae: Cassidinae: Stolaini)

The most frequently investigated defence mechanism among larvae of tortoise beetles is protection by faecal shields, which generally present chemicals that are directly sequestered and/or modified from larval host-plants. In the present work we investigate the tortoise beetle Chelymorpha reimoseri that feeds on the leaves of Ipomoea carnea fistulosa (Convolvulaceae), seeking for the importance of this mechanism to their larvae. We show that 4th instar larvae suffer low predation regardless of the presence of shields in field and laboratory bioassays with ants and chicks. Chloroform extract from larvae without shields provided high protection against predation as shown in bioassays in the field, as well as against chicks, suggesting that C. reimoseri does not rely on the shield for protection. The aqueous extract of the shields did not show activity in such bioassays. The compounds responsible for this protection have yet to be identified, and it remains an open question as to whether they are sequestered from the host plant or de novo biosynthesized. This is the first record of chemical defence in cassidine beetles without the need for faecal shields. These findings indicate that more attention should be paid to chemicals present in the tissues of larvae and/or adults of tortoise beetles; the protective compounds sequestered from host plants or de novo biosynthesized can provide an alternative or complementary strategy against predation in these insects.     

Learning and natal host influence host preference,handling time and sex allocation behaviour in a pupal parasitoid

The host choice and sex allocation decisions of a foraging female parasitoid will have an enormous influence on the life-history characteristics of her offspring. The pteromalid Pachycrepoideus vindemiae is a generalist idiobiont pupal parasitoid of many species of cyclorrhaphous Diptera. Wasps reared in Musca domestica were larger, had higher attack rates and greater male mating success than those reared in Drosophila melanogaster. In no-choice situations, na?ve female P. vindemiae took significantly less time to accept hosts conspecific with their natal host. Parasitoids that emerged from M. domestica pupae spent similar amounts of time ovipositing in both D. melanogaster and M. domestica. Those parasitoids that had emerged from D. melanogaster spent significantly longer attacking M. domestica pupae. The host choice behaviour of female P. vindemiae was influenced by an interaction between natal host and experience. Female P. vindemiae reared in M. domestica only showed a preference among hosts when allowed to gain experience attacking M. domestica, preferentially attacking that species. Similarly, female parasitoids reared on D. melanogaster only showed a preference among hosts when allowed to gain experience attacking D. melanogaster, again preferentially attacking that species. Wasp natal host also influenced sex allocation behaviour. While wasps from both hosts oviposited more females in the larger host, M. domestica, wasps that emerged from M. domestica had significantly more male-biased offspring sex ratios. These results indicate the importance of learning and natal host size in determining P. vindemiae attack rates, mating success, host preference and sex allocation behaviour, all critical components of parasitoid fitness. Electronic Publication     

Vegetation succession and lichen diversity on dry coastal calcium-poor dunes and the impact of management experiments

The negative impact of grass and moss encroachment on the botanical diversity of West European coastal dunes attracted increasing attention in the 1990s. This paper focuses on moss encroachment during primary succession in the xeroseries. Until the mid-1970s, vegetation types rich in species of the lichen generaCladonia andCladina were found on the fixed,Corynephorus canescens-dominated, so-called grey dunes all over the island of Terschelling, The Netherlands. In addition, species ofHypogymnia, Parmelia andUsnea, which usually grow on trees, occurred here terrestrially on moss carpets or bare sand. These vegetation types are still present on the Noordsvaarder, a nature reserve in the western part of the island. They occur on parts of seven dune ridges parallel to the coast and form a chronosequence in which age increases with distance from the sea. Our study found the highest lichen diversity on the second and third dune ridges in a stage of primary succession that can be assigned to theViolo-Corynephoretum. The changes from lichen-rich to moss-dominated stadia were significantly related to soil development and acidification in connection with the ageing of the dune soil. The superficial cutting of sods in moss-encroached vegetation appeared to be unsuccessful as a management technique for restoring the biodiversity of cryptogams. Our findings suggest that the best option for maintaining lichen vegetation in theViolo-Corynephoretum is the blow-in of sand with a subneutral or neutral pH from reactivated and natural blowouts or from foredunes, with increasing lime content respectively.     

Aphid alarm pheromone (E)-β-farnesene: A host finding kairomone for the aphid primary parasitoidAphidius uzbekistanicus (Hymenoptera: Aphidiinae)

Summary The present study aimed to test the possible function of the aphid alarm pheromone (E)--farnesene (EBF) as a host finding kairomone for aphid primary parasitoids. Extracts of volatile emissions of undisturbed aphids and of aphids under parasitoid attack were obtained by air entrainment. Extracts of cornicle secretions were gained by disturbing aphids and taking their secretions into solution. Extracts were compared by gas chromatography. Only air entrainments of aphids under attack and solvent extracts of cornicle secretions contained the alarm pheromone. In Y-tube olfactometer bioassays, femaleA. uzbekistanicus were attracted to aphid groups under attack of parasitoids, presumably by released EBF. High concentrations of synthetic EBF (1.4 µg to 5.7 µg) also attractedA. uzbekistanicus females. Females with oviposition experience reacted more readily to lower concentrations of EBF than females without experience. In experiments designed as Petri-dish bioassays, the test animals could contact filter paper discs that were treated with solutions containing EBF. Behavioural effects like antennation or stinging attack were not observed. With computer video analysis of parasitoid movements, some effects onA. uzbekistanicus behaviour were detected, again indicating attraction to EBF.As the volatile aphid alarm pheromone attractedA. uzbekistanicus females, it can be concluded that this chemical stimulus acts as a host finding kairomone for this parasitoid species. However, its effect over long distances seems to be limited due to the relatively high concentrations required for reactions. Of two other parasitoid species examined (P. volucre andL. testaceipes) onlyP. volucre was also significantly attracted to the volatile EBF in the Y-tube olfactometer.     

Furanocoumarins and their detoxification in a tri-trophic interaction

The parsnip webworm, Depressaria pastinacella, specializes on wild parsnip, Pastinaca sativa, and several species of Heracleum, hostplants rich in toxic furanocoumarins. Rates of furanocoumarin metabolism in this species are among the highest known for any insect. Within its native range in Europe, webworms are heavily parasitized by the polyembryonic parasitoid wasp Copidosoma sosares. In this study, we determined whether these parasitoids are exposed to furanocoumarins in host hemolymph, whether they can metabolize furanocoumarins, and whether parasitism influences the ability of webworms to detoxify furanocoumarins. Hemolymph of webworms fed artificial diet containing 0.3 % fresh weight xanthotoxin, a furanocoumarin prevalent in wild parsnip hosts, contained trace amounts of this toxin; as well, hemolymph of webworms consuming P. sativa flowers and fruits contained trace amounts of six of seven furanocoumarins present in the hostplant. Thus, parasitoids likely encounter furanocoumarins in host hemolymph. Assays of xanthotoxin metabolism in C. sosares failed to show any ability to metabolize this compound. Parasitized webworms, collected from populations of Heracleum sphondylium in the Netherlands in 2004, were on average 55 % larger by weight than unparasitized individuals. This weight is inclusive of host and parasitoid masses. Absolute rates of detoxification (nmoles min⁻¹) of five different furanocoumarins were indistinguishable between parasitized and unparasitized ultimate instars, suggesting that the intrinsic rates of metabolism are fixed. Thus, although parasitized larvae are larger, detoxification rates are not commensurate with size; rates in parasitized larvae expressed per gram of larval mass were 25 % lower than in unparasitized larvae.     

Small but smart: the interaction between environmental cues and internal state modulates host-patch exploitation in a parasitic wasp

The reproductive success of insect parasitoids depends on two activities, searching for hosts to obtain immediate fitness rewards (offspring) and searching for food to extend life span and enhance future reproductive opportunities. Models suggest that to deal with this trade-off and to cope with the variability of the resources they depend on, parasitoids should simultaneously integrate information originating from three distinct sources: host and food availabilities in the environment (environmental information) and the metabolic reserves of the parasitoid (internal information). We tested whether, in the parasitic wasp Venturia canescens, these three types of information are perceived simultaneously and interact during host-patch exploitation. Experiments were performed with strains originating from two different geographical locations and with individuals of the two reproductive modes, arrhenotoky and thelytoky. We hypothesised that there would be differences between modes of reproduction as they thrive in different ecological conditions. Our results show that metabolic state, food availability and the number of ovipositions performed simultaneously modulate patch residence time in V. canescens of both populations and reproductive modes. Unfed wasps left their host patch earlier than fed ones. The tendency to leave the patch increased with an increasing probability of finding food. Our study confirms that globally, each oviposition increases the patch-leaving tendency (decremental mechanism). This effect was modulated by both the metabolic reserves and food availability, and the relationship depended on the geographic origin of populations. Individuals of one of the populations switched from a decremental to an incremental patch-leaving mechanism depending on the presence or absence of food in the vicinity. Differences between reproductive modes in the responses to environmental cues can be explained by the different ecological conditions they live in.