Aggregation pheromone ofCarpophilus freemani (Coleoptera: Nitidulidae): A blend of conjugated triene and tetraene hydrocarbons

Summary Males ofCarpophilus freemani Dobson (Coleoptera: Nitidulidae) produce an aggregation pheromone to which both sexes fly in a wind-tunnel bioassay. The major pheromone component (ca. 30 ng per male per day in volatile collections) was identified as (2E,4E,6E)-5-ethyl-3-methyl-2,4,6-nonatriene. A minor component, (2E,4E,6E,8E)-7-ethyl-3,5-dimethyl-2,4,6,8-undecatetraene, was 3–10% as abundant as the major triene and was 5–20% as active when compared at relative doses ranging from natural proportions to 1:1. These compounds act synergistically: a mixture of major and minor components in natural proportions attracted more than twice as many beetles as the major component alone, and the mixture fully accounted for the activity of male-derived volatile collections. Six other male-derived conjugated hydrocarbons, ranging from 2% down to 0.04% as abundant as the major component, were also identified. These are (in order of decreasing bioassay activity when compared on an equal-weight basis): (3E,5E,7E)-6-ethyl-4-methyl-3,5,7-decatriene, (2E,4E,6E)-5-ethyl-3-me-thyl-2,4,6-octatriene, (3E,5E,7E,9E)-8-ethyl-4,6-dimethyl-3, 5,7,9-dodecatetraene, (2E,4E,6E,8E)-3,5,7-trimethyl-2,4,6, 8-undecatetraene, (3E,5E,7E)-5-ethyl-7-methyl-3,5,7-undecatriene, and (2E,4E,6E)-3,5-dimethyl-2,4,6-nonatriene. All structure identifications were confirmed by synthesis. In the wind tunnel, the pheromone acted synergistically with host-type volatiles such as propyl acetate, valeric acid, and ethanol. This concept was verified by fields tests in California, in which there was dramatic synergism between the pheromone and fermenting host materials. Pheromone biosynthesis is discussed.     

加拿大一枝黄花和小飞蓬叶浸提液对大豆幼苗生长的复合化感效应

加拿大一枝黄花(Solidago canadensis)和小飞蓬(Conyza canadensis)常共同入侵至同一农田生态系统。探究了两者对大豆幼苗生长的复合化感作用。加拿大一枝黄花叶浸提液显著抑制大豆幼苗地上生物量。而小飞蓬叶浸提液也明显抑制大豆幼苗地上生物量(未达到显著水平)。与对照处理相比,两者复合叶浸提液处理未显著影响大豆幼苗地上生物量。两者复合叶浸提液处理下大豆幼苗地上生物量显著大于加拿大一枝黄花叶浸提液单一处理,同时也大于小飞蓬叶浸提液单一处理(未达到显著水平)。两者叶浸提液单一处理均明显降低大豆幼苗株高(未达到显著水平)。与对照处理相比,两者复合叶浸提液处理未显著影响大豆幼苗株高。两者复合叶浸提液处理下大豆幼苗株高明显大于两者叶浸提液单一处理(未达到显著水平)。因此,两者叶浸提液对大豆幼苗的生长均具有一定程度的抑制效应,且加拿大一枝黄花叶浸提液对大豆幼苗产生的化感作用(尤其是地上生物量)明显大于小飞蓬。两者复合化感作用明显低于两者单一化感作用,尤其是两者对大豆幼苗地上生物量的复合化感作用显著低于加拿大一枝黄花的单一化感作用。所以,化感效应可能在两者共同入侵(即两者复合作用)进程中所起的贡献低于在两者单一入侵进程中所起的作用。     

New repellent semiochemicals for three species of <Emphasis Type="Italic">Dendroctonus</Emphasis> (Coleoptera: Scolytidae)

Summary. Nine compounds identified from captured volatiles of the Douglas-fir beetle, Dendroctonus pseudotsugae, the mountain pine beetle, D. ponderosae, and the spruce beetle, D. rufipennis, that elicited antennal responses in males and females of one or more of these species were tested in the field to determine behavioural activity. 1-Octen-3-ol, found in the volatiles of females of all three species decreased the response of male and female coastal and male interior D. pseudotsugae and both sexes of D. ponderosae to their aggregation pheromones. Acetophenone, identified in the volatiles of females of all three species, significantly decreased the response of interior female D. pseudotsugae. trans-Verbenol, a potent aggregation pheromone of D. ponderosae, decreased the response of both sexes of D. pseudotsugae, while 3-methyl-2-cyclohexen-1- one (MCH), the antiaggregation pheromone of D. pseudotsugae and D. rufipennis decreased the response of both sexes of D. ponderosae. While it has been demonstrated that semiochemical mediated interspecific communication occurs among bark beetles infesting the same host, this study demonstrates that beetles can perceive signals emitted by heterospecifics attacking nonhosts and can potentially use them to avoid attacking the wrong species of conifer.     

Variation in composition of predator-attracting allelochemicals emitted by herbivore-infested plants: Relative influence of plant and herbivore

Summary During foraging, natural enemies of herbivores may employ volatile allelochemicals that originate from an interaction of the herbivore and its host plant. The composition of allelochemical blends emitted by herbivore-infested plants is known to be affected by both the herbivore and the plant. Our chemical data add new evidence to the recent notion that the plants are more important than the herbivore in affecting the composition of the volatile blends. Blends emitted by apple leaves infested with spider mites of 2 different species,T. urticae andP. ulmi, differed less in composition (principally quantitative differences for some compounds) than blends emitted by leaves of two apple cultivars infested by the same spider-mite species,T. urticae (many quantitative and a few qualitative differences). Comparison between three plant species — apple, cucumber and Lima bean — reveals even larger differences between volatile blends emitted upon spider-mite damage (many quantitative differences and several qualitative differences).     

Emission of oilseed rape volatiles after pollen beetle infestation; behavioural and electrophysiological responses in the parasitoid <Emphasis Type="Italic">Phradis morionellus</Emphasis>

Summary. The pollen beetle, Meligethes aeneus, is an important pest of oilseed rape, Brassica napus. Larvae of this species feed only in the buds and flowers of Brassicaceae. One important natural enemy of this beetle is the parasitoid Phradis morionellus that attacks larvae in buds and flowers and also feeds on the flowers. The preferences for odours of non-infested and infested rape were tested for both starved and fed parasitoids in Y-tube olfactometer experiments. The volatile blend released from pollen beetle-infested and non-infested flowering rape and from pollen beetle larvae was identified and quantified. Gas chromatography-electroantennodetection analyses were performed with female P. morionellus. Parasitoids in both treatment groups preferred infested rape, but the proportion of responding female P. morionellus was significantly lower for the group that was starved. Six of the 20 volatiles identified were released at higher rates from infested rape than from non-infested. None of these compounds was found in pollen beetle larvae headspace. P. morionellus antennae detected both major and minor components in the volatile blend. The volatiles released at a significantly higher rate from infested rape and detected by P. morionellus antennae were (Z)-3-hexenylacetate, (Z)-3-hexenol, 3-butenyl isothiocyanate and (E,E)-α-farnesene.     

(S)-2-Methyl-1-hexanol,characteristic mandibular gland substance of ants of theCataglyphis bicolor group

Summary In all the species of theCataglyphis bicolor group examined yet, i.e.C. bicolor, C. diehli, C. isis, C. nodus, andC. viaticus, 2-methyl-1-hexanol is the characteristic substance and almost the only substance found in the mandibular glands. Its chirality has been determined inC. bicolor and shown to be exclusively (S)-2-methyl-1-hexanol.     

A comparative study of aggressiveness in eastern North American field cricket species (genus <Emphasis Type="Italic">Gryllus</Emphasis>)

To understand the variation in aggressiveness and factors important for contest outcome, we quantified and compared agonistic interactions of four field cricket species in eastern North America: Gryllus fultoni (Orthoptera; Gryllidae), G. vernalis, G. pennsylvanicus, and G. rubens. The most aggressive behavior that we observed, the grapple, was frequently displayed in agonistic trials of G. pennsylvanicus and G. rubens. By contrast, we never observed this behavior in trials involving G. fultoni and G. vernalis. Consequently, species was the only significant variable affecting the variation in aggressiveness, whereas size difference, age difference, and contest duration had no significant effect. In species with high levels of aggressiveness, G. pennsylvanicus and G. rubens, the factors that might be indicative of resource holding potential such as size or age difference seemed to be significant for contest outcome. In species with low levels of aggressiveness, G. fultoni and G. vernalis, however, there were indications that contest outcomes were determined by aggressiveness itself rather than the size and age differences between opponents. Markov chain analyses revealed that the difference in aggressiveness between species with high and low levels of aggressiveness lay in a sequence of escalating behaviors: antennal fencing, mandible flare, and grapple. The escalated state of aggressiveness characterized by this behavioral sequence in G. pennsylvanicus and G. rubens seems to be the ancestral state in the North American Gryllus phylogeny. We argue that the loss of a tendency to use burrows in G. fultoni and G. vernalis might be related to low levels of aggressiveness in these species.     

GC-EAD responses to semiochemicals by eight beetles in the subcortical community associated with Monterey pine trees in coastal California: similarities and disparities across three trophic levels

Summary.  Antennae of six sympatric bark and ambrosia beetles (Scolytidae), Dendroctonus valens LeConte, Gnathotrichus retusus (LeConte), Hylastes tenuis Eichhoff, Ips mexicanus (Hopkins), Ips plastographus maritimus Lanier, and Pseudohylesinus sericeus (Mannerheim), and two scolytid predators, Enoclerus sphegeus (F.) (Cleridae) and Lascontonus tuberculatus Kraus (Colydiidae), were analyzed by gas chromatographic-electroantennographic detection (GC-EAD) for their responses to synthetic Ips spp. pheromone components, and host and nonhost volatiles. The beetles emerged from cut logs of pitch canker-infected Monterey pine trees, Pinus radiata D. Don. There were significant disparities in EAD response patterns to the hemiterpene and monoterpene alcohol pheromone components that are typically produced by Ips spp. Antennae of I. p. maritimus responded strongly to ( ± )-ipsdienol, ( ± )-ipsenol, amitinol, and lanierone; antennae of I. mexicanus responded strongly to (1S,2S)-(–)-cis-verbenol, with weaker responses to ( ± )-ipsdienol, ( ± )-ipsenol, and amitinol; antennae of H. tenuis responded to (1S, 2R)-(–)-trans-ver-benol, with less pronounced responses to (–)-cis-verbenol and 2-methyl-3-buten-2-ol; and antennae of D. valens, G. retusus, and P. sericeus generally responded to all Ips spp. pheromone components except 2-methyl-3-buten-2-ol (D. valens and G. retusus) and E-myrcenol (G. retusus and P. sericeus). Ips mexicanus responded only to the (–)-enantiomers of ipsenol and ipsdienol, whereas I. p. maritimus responded to (–)-ipsenol, but to both the (+)- and (–)-enantiomers of ipsdienol. The antennae of the two predaceous insects (E. sphegeus and L. tuberculatus) responded to a range of the Ips spp. pheromone components. Host monoterpenes elicited no antennal responses from E. sphegeus, G. retusus, H. tenuis, and I. mexicanus, but several monoterpenes elicited various levels of responses from D. valens and I. p. maritimus antennae. Interestingly, antennae of female D. valens responded to (–), but not (+)-limonene. α- and β-Pinene elicited weak responses from L. tuberculatus antennae. EAD responses to selected nonhost volatiles were almost identical among the six scolytid species, with trans-conophthorin eliciting the strongest response in most cases, followed by three C₆- alcohols and two C₈-alcohols. The antennal responses by most of these species to linalool or geranylacetone were very weak; (E)-2-hexenal, (Z)-3-hexenyl acetate, and benzyl alcohol elicited almost no response. The response pattern of P. sericeus to nonhost volatiles differed slightly from the rest of the scolytids: a strong response to linalool, weaker response to the C₈-alcohols. The two predaceous Coleoptera generally had weak, but detectable, responses to nonhost volatiles, except for a relatively strong response to trans-conophthorin by L. tuberculatus. No notable differences in EAD responses were observed between males and females of the two Ips spp. Our results provide an electrophysiological baseline for future efforts to identify attractive and repellent semiochemicals (aggregation pheromones, host kairomones, or nonhost interruptants) for this guild of scolytids and their key predators that are associated with moribund and pitch canker- infected P. radiata.     

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.     

Host-plant leaf surface compounds influencing oviposition in Delia antiqua

Summary. Delia antiqua (Diptera: Anthomyiidae) females lay eggs between the leaves of onion plants or in the soil around the base of the plants, then the maggots feed on the onion bulb and roots causing rapid secondary infection by fungi and bacteria. It is well known that the first sensory modality used by the onion fly is vision, therefore the shape (vertical narrow cylinders) and colour (yellow) of the plant play a crucial role in the recognition of a potential host plant. In the past it has been shown that n-dipropyl disulfide (Pr₂S₂), a typical component of onion volatiles, is an important chemical host plant cue. We extracted host leaf surface to verify if Pr₂S₂ is the major chemical oviposition stimulant and to determine if other as yet unknown substances may play a role in host-plant selection. We confirmed that the females laid more eggs around onion plants with leaves than when only the onion bulb was present and that the odour of chopped onion stimulates oviposition. Extraction of the surface of onion leaves revealed that only the apolar fraction contained substances that stimulate egg-laying in D. antiqua. GC-EAD analysis indicated that a minor constituent, Pr₂S₂, is perceived by the olfactory receptor on the antennae of the onion fly females. This confirmed the importance of Pr₂S₂ as oviposition stimulant. Contact with the polar fraction did not stimulate egg-laying behaviour in this Delia species. We discuss the oviposition strategy of D. antiqua in comparison with its closely related species, D. radicum, in which the oviposition behaviour is stimulated mainly through contact with the cabbage leaf surface and only partially by the host volatiles.     

Responses of nymphs of desert locust,Schistocerca gregaria to volatiles of plants used as rearing diet

Summary Gregarious nymphs of the desert locust,Schistocerca gregaria (Forsk.) (Orthoptera: Acrididae) were more attracted to volatiles from mechanically damaged food plants used for rearing than to either the undamaged or damaged food plants not used as diet in Y-tube olfactometer assays. Comparative analysis of the volatile emissions from plants used for rearing and food plants not used for rearing,e.g. Sorghum bicolor, Pennisetum clandestinum, Schouwia thebaica, wheat (Triticum sp., var. Nyangumi),Zygophyllum simplex, Heliotropium undulatum andTribulus terrestris was carried out by GC, GC-EAD and GC-MS. Significant quantitative and qualitative differences were found in the volatile emissions and olfactory responses of nymphs in GC-EAD assays. Up to 33 compounds were identified in volatiles of the plants of which 9 evoked EAGs. EAG-active components included common green leaf compounds (E)-2-pentenal, (E)-2-hexenal, 4-methyl-3-pentenal, (E)-3-hexenyl acetate, (Z)-3-hexenyl acetate, (Z)-2-hexenyl acetate, (Z)-3-hexen-1-ol and (Z)-2-hexen-1-ol. (Z)-3-Hexenyl butyrate and (Z)-3-hexenyl isovalerate were detected in stimulatory amounts only in the volatiles ofS. thebaica. (E, Z)-2,6-Nonadienal was detected as a component in the volatiles ofT. terrestris and was highly stimulatory. In EAG assays with seven common green leaf volatiles, (Z)-3-hexenyl acetate was most stimulatory while hexanal was the least. No significant differences were recorded between antennal responses of males and females to the tested compounds. These results are discussed with regard to current hypotheses on host plant recognition through detection of their airborne volatiles and the learning behaviour by nymphs ofS. gregaria.     

Responses of Metaphycussp. nr. flavus to semiochemicals released from a scale host, Coccus hesperidum

Summary. Metaphycus sp. nr. flavus (Encyrtidae: Hymenoptera) is a parasitoid species collected from the Mediterranean region which lays its eggs in the immature stages of several economically important soft scale insects (Hemiptera: Coccidae), including brown soft scale, Coccus hesperidum L. (= host insect). Preliminary tests suggested that the parasitoid is most successful in producing offspring when it oviposits in the younger stages of brown soft scale. In Y-olfactometer bioassays measuring wasp choices and residence times, naïve parasitoids were significantly more attracted to yucca leaves infested with 26, 27, or 28 d-old scale than to uninfested leaves, whereas leaves with older (29-30 d-old) scale were no more attractive than uninfested leaves. Parasitoids also spent significantly more time in the arm with yucca leaves infested with 26 d-old scale than in the arm with uninfested leaves. These results are consistent with observations of the parasitoids reproductive success on scale of different ages, whereby older scale are more likely to encapsulate the developing eggs of M. sp. nr. flavusfemales than are younger scale. Further bioassays determined that yucca leaves that had been infested with 26 d-old scale but from which the scale had been removed were as attractive as infested leaves. In contrast, infested yucca leaves from which scale had been removed and the leaves subsequently washed with distilled water were less attractive than infested leaves. Furthermore, the wash water containing scale residues was attractive to female wasps. In total, these results suggest that Metaphycussp. nr. flavus females utilize volatile, water soluble compounds produced by brown soft scale as cues to locate suitable hosts.     

Hydroquinones from defensive secretion of a giant Pacific millipede, <Emphasis Type="Italic">Acladocricus setigerus</Emphasis> (Diplopoda: Spirobolida)

Summary. The presence of considerable quantities of hydroquinones including hydroquinone, 2-methylhydroquinone, 2,3- dimethoxyhydroquinone, 2-methoxy-3-methylhydroquinone, and 2,3-dimethoxy-5-methylhydroquinone renders the defensive secretion of Acladocricus setigerus (Silvestri, 1897) significantly different from that of other quinone-producing millipedes. In addition, two hitherto undescribed compounds, namely, 2,3-dimethoxy-5-methylhydroquinone and 2-methyl-3,4-methylenedioxyphenol, were characterized from the defensive secretion. However, it is uncertain if the latter compound is formed after the release of the secretion. The two major compounds, 2-methyl-1,4-benzoquinone and 2-methoxy-3-methyl-1,4-benzoquinone, constitute about 75% of the defensive fluid. Furthermore, hydrocarbons, which are typically present in the secretions of most other arthropods that use benzoquinones as repellents, are notably absent in the secretion of A. setigerus.     

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.     

Green leaf volatiles enhance sex attractant pheromone of the tobacco budworm,Heliothis virescens (Lep.: Noctuidae)

Summary Components of the green leaf volatile complex (Z-3-hexenyl acetate andE-2-hexenyl acetate) were shown to enhance responses of tobacco budworm,Heliothis virescens, males to the sex attractant pheromone of conspecific females in the field. The results are discussed with regard to green leaf volatiles which enhance the attractant pheromone of a cohabiting species, and serve as attractants of a parasitoid of conspecific larvae.     

The antifungal activity of the leaf gland volatile oil of sweet gale (Myrica gale) (Myricaceae)

Summary The antifungal activity of volatile oil obtained from the leaves of sweet gale (Myrica gale) has been investigated. Fungi, isolated from the leaves ofM. gale, were grown in liquid medium containing the oil and its effect on biomass production was gauged. The oil inhibited growth of all species of fungi to greater or lesser extent. Sesquiterpenes were more active than monoterpenes, and among the sesquiterpenes germacrone was marginally more active than -elemenone. These results support the proposition that the volatile oil ofM. gale has a role in resistance to fungal pathogens.     

D-Pulegone: A feeding deterrent to deer mice (Peromyscus maniculatus) and prairie voles (Microtus ochrogaster)

Summary D-Pulegone, a monoterpene present in pennyroyålMentha sp. is repellent to a number of vertebrates, decreasing consumption of feed and granular particles. In the present study, several tests were used to determine the reliability of this substance as a feeding deterrent. D-Pulegone (10mM—1M) applied as a coating to apple pieces significantly (P < 0.001) decreased ingestion by deer mice,Peromyscus maniculatus, and prairie voles,Microtus ochrogaster. The deterrent effect of d-pulegone was not diminished following multiple exposures suggesting that this compound may be a promising candidate for use a a general vertebrate repellent. The inhibitory effect of 1M d-pulegone on apple ingestion in voles was abolished when contact was prevented by encasing the stimulus solution in plastic mesh capsules. We conclude that while d-pulegone has repellent properties following direct contact, volatile exposure alone may not be sufficient to elicit avoidance behavior.     

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

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

Differential predation by males on moulted females may explain the competitive displacement of Gammarus duebeni by G. pulex (Amphipoda)

Summary Several studies have reported the ability of Gammarus pulex to successfully compete with and ultimately displace G. duebeni in freshwater. These studies have been unable, however, to provide adequate explanations for this displacement based on mechanisms of exploitative competition and/or differences in the environmental tolerances of the species. The present study investigates the role of predation as an extreme form of interference competition between G. pulex and G. duebeni. The data presented show that: (1) given the opportunity, males of both species prey on females of the other species, but only on females that have recently moulted; (2) mate guarding in both species significantly reduces the incidence of predation on moulted females; (3) mate guarding by male G. pulex is more successful in reducing predation on moulted females than is guarding by male G. duebeni. We propose that differential predation rate on moulted females may constitute the primary mechanism by which G. pulex displaces G. duebeni. Offprint requests to: J.T.A. Dick     

Resistance of the generalist moth Trichoplusia ni (Noctuidae) to a novel chemical defense in the invasive plant Conium maculatum

Summary. Conium maculatum is an apiaceous species native to Eurasia that is highly toxic to vertebrates due to the presence of piperidine alkaloids, including coniine and γ-coniceine. More than 200 years after invading the United States this species remains mostly free from generalist insect herbivores. The presence of novel chemical defenses in the introduced range could provide invasive species with a competitive advantage relative to native plants. The cabbage looper (Trichoplusia ni) is a generalist lepidopteran found throughout the US that occasionally feeds on C. maculatum. We evaluated the toxicity of piperidine alkaloids to T. ni and determined putative resistance mechanisms, both behavioral and physiological, that allows this insect to develop successfully on C. maculatum foliage. T. ni larvae raised on diets enriched with coniine and γ-coniceine showed a decrease in consumption and longer development time, but no effects on growth were found at any alkaloid concentration. In a diet choice experiment T. ni larvae showed no avoidance of alkaloid-enriched diets, suggesting that the deterrence produced by alkaloids was related to a post-ingestive metabolic response. The ability of T. ni to consume diets high in alkaloid content could be due to at least three different mechanisms: 1) a decreased consumption rate, 2) efficient excretion of at least 1/3 of ingested alkaloids unmetabolized in frass, and 3) partial detoxification of alkaloids by cytochrome P450 s, as shown by the decreased larval growth in the presence of piperonyl butoxide, a P450 inhibitor. Even though T. ni tolerates C. maculatum alkaloids, the use of this species as a host plant could be ecologically disadvantageous due to prolonged larval growth and thus increased exposure to predators. Novel plant secondary compounds do not guarantee increased resistance to generalist herbivores.