Use of chemical variation and predation as plant defenses byEncelia farinosa against a specialist herbivore

Larvae of the monophagous herbivore,Trirhabda geminata, selectively eat particular plants and plant parts of its natural host,Encelia farinosa. Measurements of leaf damage and larval positions on branches through time support this observation. Time-lapse movie photography revealed that larvae are sufficiently mobile to search most of a plant in a 48-hr period and that aggregations were the result of larval activity and not directly the result of oviposition. Experiments withT. geminata larvae on artificial diets containing a range of natural concentrations of chemical extracts fromE. farinosa leaves showed that the larvae grew significantly slower and had a lower overall survivorship at the high concentration. Combining the results of all choice tests, larvae appeared unable to distinguish between high- and low-concentration agar diets. Considered individually, larval preferences for natural production concentrations changed as the season progressed. Early-season larvae preferred low-concentration leaves, while late-season larvae preferred high-concentrations. Measurements of chemical and nitrogen content of leaves selected by larvae in the field confirmed this pattern. Percent parasitism in field-collected larvae increased with season as the larval population decreased. This combination of slowed growth and increasing parasitism and predation is a putative defense strategy ofEncelia farinosa to prevent adaptation by a specialist herbivore to the total range of compounds elaborated.     

Comparison of volatiles from beetle-transmittedCeratocystis fagacearum and four non-insect-dependent fungi

Ceratocystis fagacearum (Bretz) Hunt is the causative agent of oak wilt disease, which is transmitted primarily by nitidulid beetles. This fungus was compared with four non-insect-dependent fungi for their volatile profiles using gas chromatography and mass spectrometry (GC-MS) and for their attractiveness to nitidulids using a wind-tunnel bioassay. The four additional fungi includedXerula radicata Sing,Pluteus atricapillus Kumm,Tyromyces chioneus Karst, andBotrytis cinerea. Nitidulids have been reported in association with each of these fungi, but unlikeC. fagacearum, they are dispersed primarily by wind or rain. Significant attraction of three nitidulid species,Carpophilus hemipterus (Linne),C. lugubris Murray, andStelidota geminata (Say) was elicited byC. fagacearum and to a lesser extent byX. radicata, but not by the others. A comparison of headspace volatile profiles showed that the odor ofC. fagacearum was the strongest, both with regard to the number of components and in their rates of production. Chemical characterization of the headspace profile ofC. fagacearum revealed 16 components: one aldehyde, one ketone, five alcohols, and nine esters. These components were all common fruit-odor constituents and many of them were previously shown to be attractive to nitidulid beetles. The results of this study suggest that, by mimicking food odors,C. fagacearum odor is an adaptation for attracting nitidulid and possibly other insect vectors.     

Selective Sequestration of Pyrrolizidine Alkaloids from Diverse Host Plants by Longitarsus Flea Beetles

In 11 species of the flea beetle genus Longitarsus we investigated whether the insects sequester the pyrrolizidine alkaloids (PAs) present in their host plants of the families Asteraceae and Boraginaceae. In all cases where PAs could be detected in the leaves of the local host plant, they could also be detected in the corresponding beetles. In one host plant, Pulmonaria officinalis, no PAs could be detected in the leaves, yet were present in the beetles collected from them. We suggest this is due to uptake of PAs during the root-feeding larval stage. By comparing the GC-MS pattern of PAs found in the beetles with those of their hosts, we investigated the specificity of this sequestration. Furthermore, we compared the pattern of sequestered PAs across beetle species that had been feeding on the same plant, and across hosts in Longitarsus species that feed on different plants in the field. This allowed us to analyze to what extent the PA pattern in the insects is specific for the beetle species and depends on the local food plant. Our data indicate that the PAs found in the beetles are largely determined by the host plant, e.g., whether alkaloids typical of the Boraginaceae or Asteraceae are present. However, there are some indications for a selective uptake of PAs and apparently the beetles are able to metabolize them.     

Reduced offspring production in bark beetleTomicus piniperda in pine bolts baited with ethanol and α-pinene,which attract antagonistic insects

Bolts of Scots pine,Pinus sylvestris L., attacked by the bark beetleTomicus piniperda (L.) were baited with ethanol and -pinene to attract antagonistic insects and thereby enhance their detrimental effects on the production of bark beetle progeny. Unbaited and caged bolts were included in the experiments as controls. Attraction of beetles to the bolts and subsequent emergence were estimated using traps. Six phloem-feeding species (potential competitors ofT. piniperda) and four predatory species were caught in significantly higher numbers at the baited bolts than at the unbaited ones. The number of offspring and the productivity ofT. piniperda were four to seven times higher in unbaited bolts than in baited bolts. Exclusion of other insects, by using cages, resulted in a nine-fold increase in the number ofT. piniperda offspring per square meter and productivity (offspring per egg gallery) compared with unbaited, exposed bolts.Hylurgops palliatus (Gyll.) (Scolytidae) andRhagium inquisitor (L.) (Cerambycidae) attacked both the baited and unbaited bolts, whereasAcanthocinus aedilis (L.) (Cerambycidae) andPytho depressus (L.) (Pythidae) reproduced almost exclusively in the baited ones. Large numbers of larvae ofThanasimus (Cleridae) andRhizophagus (Rhizophagidae) emerged from both the baited and unbaited bolts. Adults ofPlegaderus vulneratus (Panzer) andCylister linearis (Er.) (Histeridae) emerged almost exclusively from the baited bolts. The low progeny production ofT. piniperda in the baited bolts was attributed largely to the influence of adults ofRhizophagus andEpuraea (Nitidulidae), and larvae ofThanasimus andA. aedilis.     

Effective attraction radius

The catches of bark beetles (Coleoptera: Scolytidae) were compared between attractive traps releasing semiochemicals and passive traps (cylindrical sticky screens hung, at 10 heights of 0.7–11.5 m, on poles). A central attractive-trap pole was surrounded by three passive-trap poles spaced 50 or 100 m away at the apices of an equilateral triangle. The catches ofTomicus piniperda and other scolytid species on the attractive-trap pole baited with host monoterpenes, or the catches ofIps typographus attracted to synthetic pheromone, were compared to passive trap catches in a Scots pine forest or in a Norway spruce clear-cut, respectively. Information about flight height distributions of the above scolytid species, andHylurgops palliatus, Cryphalus abietis, Pityogenes chalcographus, P. quadridens, P. bidentatus, andTrypodendron domesticum were obtained on the passive and attractive trap poles. A new method is presented for determining the densities of flying insects based on the passive trap's dimensions and catch, duration of test, and speed of insect. Also, a novel concept, the effective attraction radius (EAR), is presented for comparing attractants of species, which is independent of insect density, locality, or duration of test. The EAR is obtained by the ratio of attractive and passive trap catches and the dimensions of the passive trap, and thus should correlate positively with the strength of the attractant and the distance of attraction. EARs are determined from catch data ofT. piniperda andI. typographus as well as from the data of previous investigations on the same or other bark beetles.     

Impact of acidic deposition onEncelia farinosa gray (Compositae: Asteraceae) and feeding preferences ofTrirhabda geminata horn (Coleoptera: Chrysomelidae)

Container grownEncelia farinosa were exposed to three 3-hr episodes of acidic fog (pH 2.5) typical of events in southern California. Adults and larvae of the specialist leaf-feeding herbivore,Trirhabda geminata, preferred to feed on the acidic-treated foliage compared to control fogged (pH 6.3–6.5) foliage. Previous feeding damage on the plants did not affect feeding preference. The acidic-fogged foliage was significantly higher in total nitrogen and soluble protein but not different from control-treated tissue in water content. Stress on native populations of this drought-deciduous shrub caused by atmospheric pollutants may also result in altered feeding ecology of the beetle.     

Differences in attraction to semiochemicals present in sympatric pine shoot beetles,Tomicus minor andT. piniperda

The chemical ecology of host- and mate-finding in the pine shoot beetles,Tomicus minor andT. piniperda, was studied in southern Sweden. Beetles were collected in the field from defined attack phases on Scots pine. Using gas chromatography-mass spectroscopy, a number of oxygen-containing monoterpenes, e.g., 3-carene-10-ol, myrtenol,trans-verbenol, and verbenone, were identified from hindgut extracts of both sexes of both species. Compared toT. minor,T. piniperda contained additional compounds and in larger amounts. The amounts were highest in both species at the time when the beetles had bored into contact with the resin-producing xylem-phloem tissue. The synthesis of (1S,6R)-3-carene-10-ol by photooxidatipn of (+)-(1S,6R)-3-carene is described. In comparative electroantennogram (EAG) measurements on males and females of both species, the most active of the tested compounds wastrans-verbenol. Laboratory bioassays of walking beetles showed thatT. piniperda was attracted to uninfestèd pine logs.T. minor was more strongly attracted to pine logs infested with females than to uninfested pine logs, indicating a female-produced aggregation pheromone. Field tests confirmed thatT. piniperda was strongly attracted to pine logs. The attraction ofT. minor to logs was significant only when logs were combined with racemictrans-verbenol and (1S,6R)-3-carene-10-ol.T. minor was also attracted to a combination of these monoterpene alcohols alone. We suggest that host and mate location inT. piniperda is achieved by means of a kairomone composed of host monoterpenes, whileT. minor utilizes a primitive pheromone synergized by host odors. Evolution of host colonization strategies of the two beetles are discussed.Coleoptera: Scolytidae.This study was made within the project Odour signals for control of pest insects.     

Systemic induction of feeding deterrents in cotton plants by feeding ofSpodoptera SPP. Larvae

Cotton,Gossypium hirsutum L., has been shown to exhibit systemic induced resistance to arthropods under certain conditions. We conducted experiments to determine the effects of previous feeding ofSpodoptera exigua Hübner andSpodoptera littoralis (Boisd.) larvae on feeding behavior, growth, and survival of larvae subsequently feeding on cotton. In one feeding choice test,S. exigua larvae preferred young leaves from undamaged control plants to undamaged young leaves from a previously damaged plant. Feeding deterrence was noticeable after only 6 hr of initial feeding damage by larvae, and there was almost complete deterrence after 30 and 54 hr of continuous feeding. In a second feeding choice test,S. littoralis larvae fed more on mature leaves from undamaged control plants than on undamaged mature leaves from previously damaged plants. In no-choice tests, third instars ofS. littoralis fed undamaged young leaves from damaged plants did not gain weight and died by the seventh day, whereas larvae fed young leaves from undamaged control plants gained weight and pupated within 11 days. Sixth instars ofS. littoralis fed either old damaged leaves, old undamaged leaves, or young undamaged leaves all from previously damaged plants gained weight slowly and took more than 12 days to pupate, whereas larvae fed young leaves from undamaged plants gained weight rapidly and pupated within five days of the beginning of the experiment.     

Ancestral semiochemical attraction persists for adjoining populations of siblingIps bark beetles (Coleoptera: Scolytidae)

Three species ofIps pine bark beetles in Hopping's group IX (S.L. Wood'sgrandicollis group),Ips confusus, I. lecontei, andI. paraconfusus, are parapatrically distributed in the American Southwest. They share post-Pleistocene altitudinal ecotones with their host pines. Adjacent to these areas of host overlap, we tested the hypothesis that aggregation pheromones produced by male beetles and/or host volatiles are sufficient to elicit the speciesspecific colonization behaviors typical of these threeIps species in nature. A more distantly related species,I. pini (Hopping's group IV, S.L. Wood'spini group) was used for outgroup comparison. Under the influence of pheromone, males ofI. confusus andI. paraconfusus do not discern among unin-fested log bolts of host and nonhost pine prior to bark contact. Males responding to pheromones emanating from infested bolts are similarly undiscriminating. Females ofI. confusus andI. lecontei olfactorily discern the combination of conspecific males in host pine from other possible beetlepine combinations; females ofI. paraconfusus do not. FemaleI. pini discerned conspecific pheromone from that ofI. lecontei. The bark beetle predator,Enoclerus lecontei, is attracted by, but does not discriminate among, the male-produced volatiles of these Group IXIps species. These results support a hypothesis that divergence in pheromonal responses by these group IXIps species has evolved following their speciation, having been manifested first in the female sex. Evolutionarily, the derived pheromonal messages have preceded their behavioral discrimination by these beetles. Additional speciesspecific cues may operate between the sexes in the field that may preclude heterospecific pairings.     

Stratospheric ozone depletion and plantinsect interactions: Effects of UVB radiation on foliage quality ofCitrus jambhiri forTrichoplusia ni

Projected decreases in stratospheric ozone may result in increases in shortwave ultraviolet (UVB) irradiation at the earth's surface. Furanocoumarins, phototoxic compounds found inCitrus jambhiri foliage, increase in concentration when these plants are grown under enhanced UVB. Survivorship schedules ofTrichoplusia ni (Lepidoptera: Noctuidae) caterpillars reared on plants in the presence and absence of enhanced UVB regimes differ significantly; larvae develop more slowly in early life when reared on plants exposed to increased UVB. This same developmental pattern is observed whenT. ni larvae are reared on artificial diets amended with ecologically appropriate amounts of furanocoumarins. Thus, anthropogenically derived changes in stratospheric ozone and concomitant changes in UV light quality at the earth's surface may influence ecological interactions between insects and their host plants by altering secondary metabolism and hence foliage quality for herbivores.     

Biological amplification for increasing electroantennogram discrimination between two female sex pheromones ofSpodoptera littoralis (Lepidoptera: Noctuidae)

MaleSpodoptera littoralis stimulated by a mixture ofS. littoralis female sex pheromone with a positional isomer and by the pheromone alone, gave greatly amplified electroantennograms (EAGs) when the antennae were connected in series. The technique is described and examples are given of its use in screening extracts for EAG activity. Dose-response curves obtained with this technique are presented and discussed. Through biological amplification, EAG discrimination between the pheromones was increased.Contribution from the Agricultural Research Organization, The Volcani Center, Bet Dagan, Israel, No. 238-E, 1980 series.     

Variation in cardenolide content of the lygaeid bugs,Oncopeltus fasciatus andLygaeus kalmii kalmii and of their milkweed hosts (Asclepias spp.) in central California

A colorimetric assay was used to quantify the amount of cardenolides in the lygaeid bugsOncopeltus fasciatus andLygaeus kalmii kalmii and their milkweed host plants (Asclepias spp.) in central California. The cardenolide content of individual insects, determined in microgram equivalents of digitoxin, varied from zero to over 300 g per insect. Sources of variation of cardenolide content in the insects include interspecific and intraspecific differences in the content of the host plant species and also differences in the content of plant organs on which insects were feeding. This last source of variability may explain temporal variation in the cardenolide content of the insects. Adults ofO. fasciatus, which migrate into California in the late spring and early summer, and adults ofL. k. kalmii, which emerge from winter hibernacula in the early spring, contained small to immeasureable amounts of cardenolides. The colonization pattern ofO. fasciatus on species ofAsclepias in north central California suggests that this species does not maximize its opportunities to sequester large quantities of cardenolides from potential hosts. The emetic potential of lygaeids in California to vertebrate predators is briefly discussed.     

Insect antifeedant activity associated with compounds isolated from species ofLonchocarpus andTephrosia

The antifeedant activity of a series of 21 chalcones, flavanes, and flavanones isolated from the generaLonchocarpus andTephrosia (Leguminosae) was assessed by behavioral and electrophysiological bioassays against larvae ofSpodoptera littoralis andS. exempta. The antifeedant activity is related to the molecular structure of the compounds, and possible modes of interaction with the insect taste receptors are discussed.     

Photooxidation of Nimbin and Salannin, Tetranortriterpenoids from the Neem Tree (Azadirachta indica)

Nimbin and salannin, major triterpenoids accompanying azadirachtin in extracts of neem (Azadirachta indica) seeds, were photooxidized by UV light in the presence of oxygen to more polar, unstable intermediates that rearranged on silica gel to two final products in which the furan ring had been oxidized to isomeric hydroxybutenolides. The isomeric hydroxybutenolides were also readily formed when a crude extract of triterpenoids from neem seeds was irradiated, and both isomers of salannin have been isolated from seeds. Photooxidation of nimbin and salannin proceeded much faster than that of azadirachtin. All photoproducts showed some biological activity againstSpodoptera littoralis, Locusta migratoria, andSchistocerca gregaria. Isonimbinolide was as potent as azadirachtin at inhibiting feeding in all three species, and it also inhibited the growth ofS. littoralis. Isosalanninolide showed potent antifeedant and growth inhibitory activity againstS. littoralis.     

Quantification of cantharidin in canthariphilous ceratopogonidae (Diptera), anthomyiidae (Diptera) and cantharidin-producing oedemeridae (Coleoptera)

Cantharidin contents were determined in several canthariphilous insects by means of quantitative gas chromatography. Usually the ceratopogonidsAtrichopogon oedemerarum andA. trifasciatus caught in the field contained low concentrations of cantharidin, with concentrations in males, in most cases, being lower than in females. When fed in the laboratory with synthetic cantharidin, these species concentrated cantharidin by as much as 100-fold (males) and 40-fold (females). Accumulation in the different body tagmata (head, thorax, abdomen) of these species is similar. Maximal concentrations of cantharidin in tissues ofAtrichopogon are comparable to those known from oedemerid and meloid beetles. InA. trifasciatus about 90% of total cantharidin content is bound in tissues. Investigations using the canthariphilous anthomyiid flyAnthomyia pluvialis and three cantharidin-producing oedemerid species revealed the same pattern of distribution in different body tagmata as inAtrichopogon.     

Defensive secretion of first-instar larvae of rootstalk borer weevil,Diaprepes abbreviatus L. (Coleoptera: Curculionidae), to the fire-antSolenopsis geminata (F.) (Hymenoptera: Formicidae)

Since several species of predatory ants show some kind of repulsion towards the first-instar larvae (FIL) ofDiaprepes abbreviatus L., the predatory behavior ofSolenopsis geminata (F.), a common ant in the citrus groves in Guadeloupe, was studied. Different extracts of larvae were disposed on egg masses ofD. abbreviatus and presented as prey to the ants, both in the field and in the laboratory. The ants are repelled by the FIL extracts. The allelochemicals involved are produced in large amounts, from 5 to 20 ng per larva. Physiochemical analyses have led to the identification of two sesquiterpenes of molecular weight 218 and 234, secreted in the respective proportions of 65 and 35%.     

An estimate of the effectiveness of pheromone-baited traps for the suppression ofScolytus multistriatus (Coleoptera: Scolytidae)

We attempted to suppress a population ofScolytus multistriatus (Marsham), the principal vector of Dutch elm disease, by trapping flying beetles on sticky traps baited with synthetic pheromone. The estimated catch on 421 traps distributed throughout a 1-km² plot in Detroit, Michigan, was nearly 1 million beetles. Because an estimated 5 million beetles emerged in the plot during the study period, we conclude that this preliminary trapping study had no appreciable effect on suppressing the population. Studies that employ improved materials and techniques are continuing.     

Phloeophagous and predaceous insects responding to synthetic pheromones of bark beetles inhabiting white spruce stands in the Great Lakes region

Tree killing and saprophytic bark beetles exert important ecological and economic roles in North American spruce forests. Chemical signaling among bark beetles, and responses by associate insects such as predators and competitors, have significant effects on the population dynamics and ecology of this community. Synthetic pheromones of primary (tree killing) and secondary (saprophytic) bark beetle species and blank controls were tested using multiple funnel and lower stem flight traps in white spruce forests in Wisconsin, Michigan, and Minnesota. Six phloeophagous and four predaceous species were collected with significant attraction by the bark beetles Dryocoetes affaber, Dryocoetes autographus, and Polygraphus rufipennis, and the predatory checkered beetles (Coleoptera: Cleridae) Thanasimus dubius and Enoclerus nigrifrons. In general, trap catches to synthetic lures resembled the species composition obtained by felling trees and collecting emerging beetles in a companion study, although several species showed differing trends. Some cross attraction occurred among bark beetles and between bark beetles and predatory beetles. For example, P. rufipennis was abundant in traps baited with Dryocoetes spp. pheromones. Thanasimus dubius and E. nigrifrons were collected in significant numbers in traps baited with the pheromone of the spruce beetle (Dendroctonus rufipennis), frontalin plus -pinene. This is a new observation for E. nigrifrons. Attraction of T. dubius to the pheromones of at least three bark beetle species in the Great Lakes region, as well as to several southern and western species, reflects its role as a habitat specialist and feeding generalist. Several other important predators and competitors commonly obtained in pine forests in this region were not obtained in these spruce stands, either in response to synthetic pheromones of spruce colonizing beetles, or in host material colonized by these beetles. Potential differences in predator prey dynamics between spruce and pine ecosystems in the Great Lakes region are discussed.     

Evaluation of two attempts to trap defined populations ofScolytus multistriatus

Two experiments were conducted to investigate the use of a pheromone-based trap-out technique for suppressing populations ofScolytus multistriatus. In the first experiment, elm bolts containing the immature stages ofS. multistriatus were placed in an isolated community that contained elm trees, but which was essentially devoid of a resident beetle population. The infested bolts produced a total of 46,485 adult beetles of which 20% were recovered on traps baited with synthetic pheromone. In the second experiment beetles were released in a desert valley containing no resident elms or beetles. Only 1% of 20,000 released beetles were recaptured on traps erected on vertical cardboard cylinders and on elm logs. These rates of recapture are related to the attraction of beetles to naturally occurring brood sources versus pheromone-baited traps.Coleoptera: Scolytidae.     

Foreign-language Skills in Rove-Beetles? Evidence for Chemical Mimicry of Ant Alarm Pheromones in Myrmecophilous <Emphasis Type="Italic">Pella</Emphasis> Beetles (Coleoptera: Staphylinidae)

By using chemical analyses, as well as laboratory and field behavioral tests, we tested the hypothesis that rove beetles of the myrmecophilous genus Pella use alarm pheromone compounds to avert attacks by their host ant Lasius fuliginosus. The secretions of Pella funestus and P. humeralis contain quinones and different aliphatic compounds, mainly undecane and 6-methyl-5-hepten-2-one (sulcatone). The latter two chemicals are also found in L. fuliginosus pheromone glands. Behavioral tests confirmed that undecane serves as an “aggressive alarm”-inducing pheromone in L. fuliginosus, whereas sulcatone most likely is a “panic-alarm”-inducing pheromone. The main tergal-secretion compounds, various quinones and undecane, individually and in mixtures induced aggression in L. fuliginosus workers. When sulcatone was added to these compounds, the space around the odor source was avoided and a reduced number of aggressive acts observed, suggesting that sulcatone blocks the aggression-inducing effect of undecane and the quinones. These results support the hypothesis that Pella beetles mimic alarm pheromones of their hosts. This is a rare example of chemical mimicry in myrmecophilous insects in which chemicals other than cuticular hydrocarbons are used.