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.     

The search for indigenous dune stabilizers: Germination requirements of selected South African species

The coastline of South Africa is characterized by extensive dune fields which are threatened by development, and thus the demand of stabilization of drift sand will increase. The non-invasive alien grass speciesAmmophila arenaria is at present the dominant sand stabilizer. Due to its foreign origin and invasiveness in North America its use was criticized and indigenous sand-binding species would be preferred. The germination requirements of the nativeArctotheca populifolia, Ipomoea pes-caprae. Myrica cordifolia andScaevola plumieri were investigated. The results showed that for all four species the total germination time and germination phase could be shortened and germination success (number of germinated seeds) improved. The different scarification and stratification treatments broke the dormancy ofI. pes-caprae, M. cordifolia andS. plumieri, whereas the seeds ofA. populifolia were not dormant. Seeds ofI. pes-caprae andM. cordifolia had induced an innate dormancy, respectively, whereas the type of dormancy in seeds ofS. plumieri dormancy processes remained unknown.     

Chemoecological studies of the exocrine glandular larval secretions of two chrysomelid species (Coleoptera):Phaedon cochleariae andChrysomela lapponica

Summary The exocrine glandular secretions of larvae of the subfamily Chrysomelinae are known to repel conspecific adults, other competitive phytophagous insects and natural enemies. InPhaedon cochleariae, the intraspecific activity of tlc fractions of the larval secretion was tested in order to examine the ecological significance of two fractions containing minor components and a fraction containing the major compound, the cyclopentanoid monoterpene (epi)chrysomelidial. InChrysomela lapponica, the defensive activity of the larval secretion against ants is known from specimens feeding upon willow or birch. The feeding preferences of larvae and adults ofC. lapponica from a Finnish and a Czech population were tested. The Finnish individuals significantly preferred feeding uponSalix borealis, whereas they hardly fed upon birch. The Czech specimens clearly preferred birch (Betula pubescens) to willow species. Application of salicin onto leaves of a willow species free of this phenolglycoside revealed that the Finnish individuals preferred feeding upon leaves with salicin. On the other hand, the Czech individuals avoided feeding upon leaves ofB. pubescens treated with salicin. The chemical composition of the glandular secretion of the Finnish larvae differed from the one of the Czech larvae. GC-MS-analyses of the secretions revealed that salicylaldehyde was the only major component of the secretion of Finnish larvae feeding upon the salicin-containing willowS. borealis. The glandular secretion of the Czech larvae feeding upon birch contained numerous esters of isobutyric acid and 2-methylbutyric acid. When Czech larvae had fed upon a salicin-containing willow (S. fragilis), the major compounds of their secretion were benzoic acid, salicylalcohol and benzoic acid esters; salicylaldehyde was only detected in traces. Thus,C. lapponica individuals from the Finland population adapted so closely to a salicincontaining willow that they clearly prefer this plant for food and that they obviously derive their main larval defensive compound (salicylaldehyde) from their host-plant.     

Oviposition stimulants and deterrents control acceptance ofAlliaria petiolata byPieris rapae andP. napi oleracea

Summary Differential acceptance of garlic mustard,Alliaria petiolata byPieris rapae L. andP. napi oleracea is explained by their differential sensitivities to oviposition stimulants and deterrents in the plant. Fractions containing the stimulants and deterrents were isolated by solvent partitioning between water and n-butanol and by open-column chromatography followed by HPLC.P. napi oleracea showed no preference when offered a choice ofA. petiolata or cabbage, but was strongly stimulated to oviposit by post-butanol water extracts ofA. petiolata. The most abundant glucosinolate in this extract was identified as sinigrin, which could explain the high degree of stimulatory activity.P. rapae preferred cabbage plants overA. petiolata, and the relatively low stimulatory activity was also associated with the glucosinolate-containing aqueous extract. However, this species was strongly stimulated by a fraction that contained small amounts of glucotropaeolin along with unknown compounds. Deterrents to both species were found in the butanol extract fromA. petiolata, andP. napi oleracea was more sensitive thanP. rapae to these deterrents. Some HPLC fractions from the BuOH extract were strongly deterrent toP. napi oleracea, but were inactive toP. rapae. The ecological significance of these behavioral differences between the twoPieris species is discussed.     

Characterization of the benthic vegetation in the Farwà Lagoon (Libya)

This study forms part of the Action Plan for the Conservation of Marine Vegetation in the Mediterranean SEa (United Nation Environmental Program). It was carried out in June 2000 in the Farwà Lagoon, Libya. The mapping of the main benthic vegetation was achieved by compiling the field observations (transect method), and remote sensing of SPOT satellite images. The phytobenthos in the Farwà lagoon covers an area of 1820 ha (65%). Three benthic macrophyte species dominate, namely the marine phanerogamsCymodocea nodosa andPosidonia oceanica, and the algaCaulerpa prolifera. DeadPosidonia oceanica leaves (litter) form veritable mounds in the vicinity of the openings leading to open sea. These leaves, which come from the coastal sea, are brought into the lagoon by currents and tides; their decomposition will lead to high oxygen consumption and the release of hydrogen sulphide. The phenological data ofPosidonia oceanica shoots sampled in the lagoon are similar to those from other stations in the Mediterranean. Conversely, the lepidochronological parameters of shoots sampled in the central part of the lagoon exhibit values that are substantially higher than those generally recorded in the Mediterranean. The mean number of leaves produced annually is 9.9 (mean value for the Mediterranean: 7.5) and the rhizome growth rate is of 35.7 mm.yr⁻¹ (mean value for the Mediterranean: 7.5 mm.yr⁻¹). This hypersaline environment would seem to provide optimum growth conditions for the speciesPosidonia oceanica.     

Nest-size variation reflecting anti-predator strategies in social spider mites of<Emphasis Type="Italic"> Stigmaeopsis</Emphasis> (Acari: Tetranychidae)

The social spider mites (Acari: Tetranychidae) of Stigmaeopsis weave dense nests on the underside of host leaves. Four species occur on the leaves of bamboo in Japan: Stigmaeopsis longus, S. celarius, S. takahashii and S. saharai. We initially reconfirmed the occurrence of distinct variation in nest size among the species. Based on the hypothesis that this variation plays a role in protecting the spider mites from predators, we looked at the behavior of the natural enemies that occur on the host plants along with members of Stigmaeopsis. We found considerable variation in the ability of nests to protect the spider-mite eggs. The smallest nests protected the eggs against three predators, whereas the largest nests protected the eggs against only one predator species. So, decreases in nest size increased egg defense. Thus we concluded that nest-size variation reflects a strategy for reducing predation.Communicated by D. Gwynne     

Leaf volatiles and polyphenols in pear trees infested byPsylla pyricola. Evidence of simultaneously induced responses

Summary Feeding by the homopteranPsylla pyricola on leaves of pear trees induces the production of volatile compounds, such as (E,E)--farnesene and methyl-salicylate, as well as the production of polyphenols. The inference on induction is based on GC-MS and HPLC chromatograms from the same samples ofPsylla infested leaves, leaves from the same pear tree beforePsylla infestation and uninfested leaves from other pear trees.Psylla infestation greatly enhanced the production of volatiles ((E,E)--farnesene, methyl-salicylate and others) and triggered the production of new polyphenols, characterized by much longer retention times.However, the responses to infestation depend critically on leaf age (defined by leaf distance to apex). With respect to the leaf volatiles it appears that infested, old leaves produce fewer compounds and lower amounts of the volatiles than infested, young leaves. Moreover, there seem to be differences in pattern. Relative to (E,E)--farnesene, methyl-salicylate was found in much lower amounts in heavily infested, old leaves. With respect to polyphenols it was found that infested old leaves collected in August have polyphenols with the same retention times, but more or less equal amounts as uninfested young leaves collected in May. This shows thatPsylla infestation causes the induced response mostly in young leaves.The induced leaf volatiles may act as synomones to heteropteran bugs. As shown elsewhere,Anthocoris nemoralis responds significantly to (E,E)--farnesene and methyl-salicylate when offered in pure form against clean air in a Y-tube olfactometer. The effect of polyphenols on the performance ofP. pyricola is not yet known. Hence, a role in direct defence is still to be investigated.     

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

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

Protein and amino acid composition of the tergal gland secretions ofBlatta orientalis andEurycotis floridana (Dictyoptera: Blattidae)

Summary Nymphs and adult females ofBlatta orientalis and nymphs ofEurycotis floridana produce a proteinaceous sticky secretion which accumulates on the last abdominal tergites. The proteic patterns do not differ between individuals of the same species. HPLC analyses show that all the common amino acids are found in both species, aspartic and glutamic acids representing 24 to 37% of the total amount of amino acids. InB. orientalis, glutamic acid is the more abundant amino acid whereas inE. floridana it is the aspartic acid. The secretion appears and accumulates rapidly on isolated insects. Behavioural assays revealed that these secretions have a defensive role.     

Sequestration of plant pyrrolizidine alkaloids by chrysomelid beetles and selective transfer into the defensive secretions

Summary Oreina cacaliae andO. speciosissima (Coleoptera, Chrysomelidae) sequester in their elytral and pronotal defensive secretions pyrrolizidine alkaloids (PAs) as Noxides (PA N-oxides). The PA N-oxide patterns found in the beetles and their host plants were evaluated qualitatively and quantitatively by capillary gas chromatography/mass spectrometry (GC-MS). Of the three host plantsAdenostyles alliariae (Asteraceae) is the exclusive source for PA N-oxide sequestration in the defensive secretions of the beetles. With the exception of O-acetylseneciphylline the N-oxides of all PAs ofA. alliariae, i.e. senecionine, seneciphylline, spartioidine, integerrimine, platyphylline and neoplatyphylline were identified in the secretion. PA N-oxides typical ofSenecio fuchsii (Asteraceae) were detected in the bodies of the beetles but not in their secretion. No PAs were found in the leaves of the third host plant,Petasites paradoxus (Asteraceae). The results suggest the existence of two distinctive storage compartments for PA N-oxides in the beetle: (1) the defensive secretion, containing specifically PA N-oxides acquired fromA. alliariae; (2) the body of the beetle, sequestering additionally but less selectively PA N-oxides from other sources,e.g. S. fuchsii or monocrotaline N-oxide fed in the laboratory. The concentration of PA N-oxides in the defensive secretion is in the range of 0.1 to 0.3 mol/1, which is more than 2.5 orders of magnitude higher than that found in the body of the beetle. No significant differences exist in the ability of the two species of beetles to sequester PA N-oxides fromA. alliariae, althoughO. speciosissima, but notO. cacaliae, produces autogenous cardenolides. A negative correlation seems to exist between the concentrations of plant-derived PA N-oxides andde novo synthesized cardenolides in the defensive secretion ofO. speciosissima.     

Some polyphagous Homoptera gain protection from their natural enemies by feeding on the toxic plantsSpartium junceum andErythrina corallodendrum (Leguminosae)

Summary Interactions were studied among alkaloid-containing legumes (Erythrina corallodendrum andSpartium junceum) and non-toxic plants (Citrus sinensis, Cucurbita moschata andEuphorbia tirucalli), several polyphagous homopterans,Aphis craccivora (Aphididae),Icerya purchasi, I. aegyptiaca (Margarodidae),Lepidosaphes ulmi (Diaspididae) andPlanococcus citri (Pseudococcidae), and some major natural enemies of these homopterans. Significant reductions in survival due to negative effects of alkaloid containing as compared with non-alkaloidal plants were recorded for the predatorsRodolia cardinalis andChilocorus bipustulatus, but not forCryptolaemus montrouzieri (Coleoptera: Coccinellidae),Chrysoperla carnea (Neuroptera: Chrysopidae) andSympherobius sanctus (Neuroptera: Sympherobiidae). The development time of the larvae or pupae ofR. cardinalis, C. carnea andS. sanctus was longer on the toxic plants than on the non-toxic ones. The percentage of parasitism ofA. craccivora collected from the non-alkaloidal plantsVicia palaestina andMelilotus albus was much higher than that onS. junceum. The parasitoid complexes ofA. craccivora differed between both plant groups. The nutritive value of honeydew ofI. purchasi andA. craccivora, as expressed by the life span ofEncyrtus infelix (Hymenoptera: Encyrtidae) adults, was also investigated. Life spans were significantly longer when the wasps fed on honeydew produced on non-alkaloidal plants (C. sinensis andPittosporum tobira) than on alkaloid containing plants whenI. purchasi — but notA. craccivora — was the producer. It is suggested that the chemical defense ofE. corallodendrum andS. junceum is exploited by polyphagous phytophages to reduce predation. In nature, population growth and density of four of the investigated homopterans are conspicuously high when they developed on the alkaloid containing plant species, and very low on non-alkaloid plants. The efficiency of their natural enemies may be reduced by sequestration of alkaloids (or other toxic plant compounds) or their transfer into excreted honeydew. Therefore it is assumed that a generalist phytophagous homopteran may be protected from its natural enemies, although at different rates of efficiency, if it can safely sequester the host allelochemical when it develops on toxic species within its host range.     

Allocation of nitrogen and carbon in an estuarine salt marsh in Portugal

Above and below-ground biomass and nitrogen and carbon composition ofSpartina maritima, Halimione portulacoides andArthrocnemum perenne, dominating species in plant communities of the lower, middle and higher salt marsh, respectively, were compared in an estuarine salt marsh in Portugal. Plant and soil nitrogen and carbon pools were estimated. For all three species root biomass was significantly higher (70–92% of total biomass) than above-ground biomass. The percentage of root biomass was related to the location of the plants in the marsh: higher values were found in plants growing in the lower salt marsh where the sediment was more unstable and subject to tidal action, which stresses the role of the roots as an anchor. For all three species nitrogen concentrations were highest in leaves, reflecting the photosynthetic role of the tissue. For carbon higher concentrations were found in the stems, with the exception ofS. maritima. In general, lower nitrogen concentrations were found in summer, which can be explained by dilution processes due to plant growth. For both nitrogen and carbon, higher concentrations were found in the soil surface layers. Higher soil nitrogen and carbon levels were associated with higher organic matter contents. Most of the nitrogen in the salt marsh occurred in the sediments (0–40 cm) and only ca. 5.7–13.3% of the total was found in the plants. The greater portion (76.5%–86%) of carbon was found in the sediment.     

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.     

Oily substances from gastral intersegmental glands of the antPachycondyla tridenta (Ponerinae): Lack of pheromone function in tandem running and antibiotic effects but further evidence for lubricative function

Summary Gas chromatographic analyses of volatizable material from gaster intersegmental complex glands ofP. tridentata revealed the presence of linoleic acid, palmitic acid, methyl oleate, and several long-chain hydrocarbons as major constituents, which form an oily fluid mixture. The nest relocation communication ofP. tridentata is tandem running. Tandem following is mediated by pheromones as demonstrated by dummy experiments with isolated gasters and thoraces and with glass balls. The secretions of all gaster complex glands, as well as hind gut contents and metapleural gland secretions, were unable to evoke tandem following. Morphology and position of the glands, lack of pheromonal function, oily properties, low volatility, and lack of antibiotic effects of these secretions strongly suggest a function as lubricants for the ants' gastral segments.     

Leaf surface compound fromBrassica oleracea (Cruciferae) induces oviposition byPieris brassicae (Lepidoptera: Pieridae)

Summary Chemicals present on the surface of cabbage (Brassica oleracea L.) leaves were extracted by dipping these leaves for 3 s in dichloromethane followed by a 3 s dip in methanol. When offered in dual choice bioassays using green paper cards as a substrate, the methanol extract stimulated oviposition activity byPieris brassicae L. (Lepidoptera: Pieridae) females. The oviposition stimulant was isolated using medium pressure liquid chromatography, reversed-phase HPLC, ion-pair HPLC and ion exchange chromatography. Using¹H-NMR spectroscopy, the stimulant could be identified as glucobrassicin (3-indolyl-methyl-glucosinolate). When pure glucobrassicin was offered at a dose identical to that in the crude methanol extract, butterflies did not discriminate between these two substrates in a dual choice test. It is argued that a high sensitivity for indole glucosinolates as host recognition factors may confer an adaptive value for these specialist crucifer feeders. The nutritional significance of their precursor tryptophan and the non-volatile nature of the aglycones formed upon enzymic hydrolysis in damaged tissues are proposed as properties of indole glucosinolates that contribute to this possible adaptive advantage.     

Behavior and ecology of Old World Luperini beetles of the genusAulacophora (Coleoptera: Chrysomelidae)

Summary The attraction of Old World leaf beetles in the genusAulacophora to kairomones and parakairomones which are effective lures for New WorldDiabrotica andAcalymma were investigated. Beetles captured on sticky traps baited with single and multicomponent lures were no different from the control traps for two species ofAulacophora. Yellow colored traps and squash blossoms are attractive toAulacophora beetles which detect sub-microgram quantities of cucurbitacins on silica gel. Leaf feeding behavior and flight activity data are correlated with varietal preference of threeAulacophora species. The common response byDiabrotica andAulacophora to cucurbitacins reinforces the two groups' coevolutionary association with the Cucurbitaceae. The apparent lack of a common response toCucurbita blossom volatiles suggests recent evolutionary pathways are substantially different for these two groups of beetles.     

Oviposition by butterflies on young leaves: Investigation of leaf volatiles

Summary Various butterflies select young foliage on which to lay their eggs; volatiles emitted by young and old leaves have been compared (by sorption enrichment, followed by GC-MS) to gauge possible qualitative and quantitative differences between the two age groups. The plants investigated are cabbage (Brassica oleracea), two milkweeds (Asclepias syriaca andA. curassavica), the bitter orange (Citrus aurantium) and the lime (C. aurantiifolia). The chemical compounds identified belong to three classes, isoprenoids, fatty acid derivatives and benzenoids. Quantitative differences were found between young and old leaves, of which a few may be characteristic of young leaves only. Thirty-four single trials withDanaus plexippus exposed to volatiles from young and old leaves are recorded.     

Sequestration of pyrrolizidine alkaloids by larvae ofTellervo zoilus (Lepidoptera: Ithomiinae) and their role in the chemical protection of adults against the spiderNephila maculata (Araneidae)

Summary Life stages of the primitive Australian ithomiine butterflyTellervo zoilus and its larval hostplant, the apocynaceous vineParsonsia straminea, were quantitatively assayed for pyrrolizidine alkaloids (PAs). PAs were found in all stages, mainly as N-oxides, being most concentrated in larvae and freshly-emerged adults. Although adults feed at various confirmed PA sources this probably does not compensate for losses, as wild-caught adults had considerably lower concentrations of PAs. The main alkaloid present in both freshly-emerged adults and in leaves of the host-plant was lycopsamine (1b), stored by butterflies in the N-oxide form. Its presence in higher proportion, in relation to intermedine (1a), in larvae, pupae and adults ofTellervo in relation to the host-plants suggests the inversion of intermedine to lycopsamine by the insects. No 14-member ring macrocyclic PAs were detected in either food-plant or butterflies. Several other PAs were found in wild-caught adults reflecting visits to other PA sources. PAs were also found in high concentrations in freshly-emerged individuals of the danaineEuploea core bred onParsonsia straminea. Wild-caughtDanaus affinis had high PA levels acquired from adult feeding. Freshly emergedEuploea raised onIschnocarpus frutescens andDanaus raised onIschnostemma carnosum (both PA-free) were preyed on by the orb weaving spiderNephila maculata, and showed no PAs. In all cases where PAs were present, most butterflies were liberated, usually cut out of the web unharmed, byNephila. The spider's response was not closely linked to PA concentration, however, and may also depend on hunger levels and previous experience with PA-containing butterflies. All control and other non-PA containing butterflies were consumed although rejection of some body parts of freshly-emergedDanaus affinis suggests that compounds other than PAs may be involved.     

Extension of twoCaulerpa species along the Tunisian coast

Two species of the green alga genusCaulerpa are considered as invasive in Tunisia:Caulerpa racemosa andC. taxifolia. C. racemosa is found in a many new localities at the central and southern Tunisian coast, and even in the North of the country in colder water. These new observations lead to the hypothesis of the multiplicity of the origins of the introduction of the species and suggest possible invasive potentialities. The general thallus, and more particularly the number, the shape as well as the arrangement of ramelli on fronds allowed us to identify four morphologically different groups. These groups could be either different systematic entities or polymorphic variations of the same taxon due to different ecological conditions.Caulerpa taxifolia was observed for the first time in Tunisia at the roadstead of Sousse in the beginning of the year 2000. It is recognized as identical to that found in the other Mediterranean localities (GenBank number AJ228960). Up to the end of March 2000, a submarine prospecting campaign revealed an affected area of about 350 ha in front of 2 km of coast. Colonies ofCaulerpa taxifolia disseminating in this area appear in isolated spots. form, In the growing stage, they form surfaces varying between 0.5 to 6 m² and located 20 to 100 m from each other. The total covered surface was estimated to be between 0.5 and 1 ha.     

Daphne gnidium, a possible native host plant of the European grapevine moth Lobesia botrana, stimulates its oviposition. Is a host shift relevant?

Summary. We tested the hypothesis that Daphne gnidium is an ancestral host plant of Lobesia botrana. To this end, we studied the oviposition response of this moth to various aerial organs of this plant. Our results show that females prefer to lay eggs on daphne berries rather than on grapes (cv. chasselas) and that polar surface extracts of daphne berries and leaves strongly stimulate oviposition in a dosedependant way, whereas flower extracts are weakly stimulant and branch extracts are inactive or deterrent for oviposition. Furthermore, a total extract of daphne berries stimulates oviposition in semi-natural conditions when applied onto fresh grapes. Oviposition stimulants from berries were isolated by HPLC coupled with a dual-choice oviposition bioassay. The ecological value of D. gnidium for L. botrana is discussed and the possible use of oviposition stimulants from daphne, contrasting with the oviposition signal of the cultivated host plant, is proposed in the perspective of developing behavioural manipulation methods such as stimulo-deterrent diversionary strategies compatible with IPM programs.