Behavioral and electrophysiological responses of the banana weevilCosmopolites sordidus to host plant volatiles

Male and femaleCosmopolites sordidus were attracted to freshly cut banana rhizome and pseudostem in a still-air olfactometer. Females responded similarly to odors from a comparatively resistant and from a susceptible cultivar of banana, when presented as either freshly cut tissue or as Porapak-trapped volatiles. Females were also attracted to rotting banana pseudostem and to volatiles collected from it. Males and females gave similar responses to host tissue in both the behavioral bioassay and to collected volatiles in EAG recordings. Weevils did not respond, either behaviorally or electrophysiologically, to a synthetic mixture of mono- and sesqiterpenes, which made up over 9% of the volatiles collected from pseudostem.     

Laboratory and field evidence for maleproduced aggregation pheromone inRhynchophorus cruentatus (F.) (Coleoptera: Curculionidae)

Laboratory and field assays were conducted to determine if palmetto weevil,Rhynchophorus cruentatus (F.), adults produce an aggregation pheromone. Attraction of females in a Y-tube olfactometer to conspecific males was greater than to clean air. Male and female attraction to conspecific male volatiles combined with host-palm,Sabal palmetto (Walter), volatiles was greater than to host-palm volatiles alone. Similarly, more weevils were caught in the field in traps baited with conspecific males plus host-palm tissue than in similar traps baited with only males, or palm tissue, or females, or females plus palm tissue. These results suggest thatR. cruentatus males produce an aggregation pheromone(s) that is highly attractive to conspecific adults of both sexes when combined with host-palm volatiles. This study is an important step towards understanding the chemical ecology ofR. cruentatus.     

Pheromone-based Mating and Aggregation in the Sorghum Chafer, <Emphasis Type="Italic">Pachnoda interrupta</Emphasis>

Adults of the sorghum chafer, Pachnoda interrupta Olivier (Coleoptera: Scarabaeidae: Cetoniinae), form aggregations during the mating period in July, but also in October. The beetles aggregate on food sources, e.g., Acacia spp. trees or sorghum with ripe seeds, to feed and mate. During the mating season, field trapping experiments with live beetles as bait demonstrated attraction of males to unmated females, but not to mated females or males, indicating the presence of a female-emitted sex pheromone. Unmated females combined with banana (food source) attracted significantly more males and females than did unmated females alone. Other combinations of beetles with banana were not more attractive than banana alone. Thus, aggregation behavior appears to be guided by a combination of pheromone and host volatiles. Females and males were extracted with hexane during the mating period, and the extracts were compared by using GC-MS. In a field trapping experiment, 19 compounds found only in females were tested, both singly and in a mixture. Traps baited with one of the female-associated compounds, phenylacetaldehyde, caught significantly more beetles than any other treatment. However, the sex ratio of beetles caught in these traps did not differ from that of control traps, and it is possible that other components may be involved in the sex pheromone signal. Furthermore, traps baited with a mixture of all 19 compounds attracted significantly fewer beetles than did phenylacetaldehyde alone.     

Development of an Efficient Pheromone-Based Trapping Method for the Banana Root Borer <Emphasis Type="Italic">Cosmopolites sordidus</Emphasis>

The banana root borer Cosmopolites sordidus (Germar) (Coleoptera: Curculionidae) is a major pest of bananas throughout the world. Chemical control is both undesirable and expensive, where biological control alternatives are limited, and pheromone-based trapping results in low captures. In this study, several important factors that affect pheromone-based catches, such as trap type, trap dimensions, and color and position of the traps, were optimized. Ground traps were found to be superior to ramp and pitfall traps, and larger traps (40 × 25 cm and above) were more efficient than smaller ones (30 × 15 cm). In a color-choice test, the banana weevil clearly preferred brown traps over yellow, red, gray, blue, black, white, and green, with mahogany being more attractive than other shades of brown. In addition, pheromone baited ground traps positioned in the shade of the canopy caught significantly more adults than those placed in sunlight. Therefore, mahogany-brown ground traps 40 × 25 cm appear to be the most efficient at catching C. sordidus adults and have the greatest potential for use in mass trapping and programs for eradication of this pest.     

Evidence for volatile male-produced pheromone in banana weevilCosmopolites sordidus

Females of the banana weevil,Cosmopolites sordidus, were attracted to and made longer visits to live conspecific males, trapped volatiles from males, and dissected male hindguts in a still-air olfactometer. Male weevils were attracted to volatiles trapped from males and made longer visits to live males and volatiles from males. Live females, collected volatiles from females and female hindguts, elicited small or no behavioral responses from either sex. Electroantennogram (EAG) responses from both male and female antennae were elicited by collected volatiles from males and by dichloromethane extracts of male hindguts and bodies but not by surface washes of males. No significant EAG responses were given to equivalent material from females. It is therefore suggested that male banana weevils release an aggregation pheromone via their hindgut.     

Kairomonal Response of Predators to Three Pine Bast Scale Sex Pheromones

The kairomonal activity of the sex pheromones of three pine bast scales, Matsucoccus feytaudi, Matsucoccus josephi, and Matsucoccus matsumurae, as well as a new analog of the M. feytaudi sex pheromone, were investigated in pine forests of France, Portugal, and Italy. The response of the maritime pine bast scale predators, Elatophilus spp. and Hemerobius stigma, was used to test the influence of trapping methods, kairomone composition, and dose. Both predators showed significant attraction to all compounds except to the sex pheromone of M. josephi. Significant increase in captures was observed as a function of dose, and within the studied dose range, up to 2200 μg, no threshold saturation limits were observed for any of the attractive compounds. Trap design and size did not significantly influence predator captures, except for high population levels of Elatophilus crassicornis, when plate traps were more efficient than delta traps. Geographic variations were found in the kairomonal responses patterns of both predators, with the M. matsumurae sex pheromone being more attractive to the oriental populations from Corsica and Italy, whereas the western populations in Aquitaine and Portugal were more attracted to the M. feytaudi sex pheromone.     

Aggregation pheromone of palmetto weevil,Rhynchophorus cruentatus (F.) (Coleoptera: Curculionidae)

5-Methyl-4-octanol is the major aggregation pheromone of the palmetto weevil,Rhynchophorus cruentatus (F.). The pheromone (cruentol) was identified by coupled gas chromatographic-electroantennographic (GC-EAD) analysis of male-produced volatiles, coupled GC-mass spectrometry (MS) in electron impact and chemical ionization mode, and coupled GC-high resolution MS. In laboratory and field assays, a diastereomeric mixture of synthetic cruentol greatly enhanced attraction of weevils to cabbage palmetto,Sabal palmetto (Walter), stem tissue, indicating that cruentol and host volatiles are synergistically attractive. An attractive lure in combination with efficient traps should facilitate development of semiochemical-based management forR. cruentatus.     

Aggregation pheromone of driedfruit beetle,Carpophilus hemipterus Wind-tunnel bioassay and identification of two novel tetraene hydrocarbons

A male-produced aggregation pheromone was demonstrated inCarpophilus hemipterus (L.) (Coleoptera: Nitidulidae) using a wind-tunnel bioassay. Both sexes responded to the pheromone, but the beetles flew in the wind tunnel only after they had been starved for at least several hours. The attractiveness of the pheromone was greatly enhanced by volatiles from a food source, and combinations of pheromone and food volatiles typically attracted 3–10 times more beetles than either source by itself. A variety of food-related sources of volatiles were effective. These included apple juice; a mixture of baker's yeast plus banana; the pinto bean diet used for rearing this beetle; the chemicals propyl acetate, ethanol; and a mixture of acetaldehyde, ethyl acetate, and ethanol. The pheromonal activity resided with a series of 10 male-specific, unsaturated hydrocarbons of 13, 14, and 15 carbon atoms. These were partially separated by HPLC. No single compound was absolutely required for pheromonal activity to be observed, and various subsets of these compounds were active. The most abundant component was (2E,4E,6E,8E)-3,5,7-trimethyl-2,4,6,8-decatetraene. One minor component was (2E,4E,6E,8E)-3,5,7-trimethyl-2,4,6,8-undecatetraene. These structures were proven by synthesis. Together, the synthetic compounds were as active in the wind tunnel as the beetle-derived pheromone.     

Olfactory Responses of the Predatory Mites (N eoseiulus cucumeris) and Insects (O rius strigicollis ) to Two Different Plant Species Infested with Onion Thrips (T hrips tabaci)

Responses of Neoseiulus cucumeris (a predatory mite) and the predatory insect Orius strigicollis to volatiles associated with two different plant species infested with onion thrips, Thrips tabaci, were examined in a Y-tube olfactometer. Both predators species showed a significant preference for volatiles from infested cucumber leaves without T. tabaci over clean air. However, they were not attracted to volatiles from uninfested cucumber leaves, artificially damaged cucumber leaves, or volatiles from T. tabaci plus their visible products collected from cucumber leaves. These results suggest that both predator species are capable of exploiting herbivore-induced volatiles from T. tabaci-infested cucumber leaves as a foraging cue. Neither predator was attracted to volatiles from uninfested spring onion leaves, infested spring onion leaves without T. tabaci, or volatiles from T. tabaci plus their visible products collected from spring onion leaves. Interestingly, they avoided volatiles from artificially damaged spring onion leaves. A possible explanation for the non-significant olfactory responses of the predator species to spring onion plants with infestation damage of T. tabaci is discussed.     

Identification and Field Evaluation of Attractants for the Cranberry Weevil, <Emphasis Type="Italic">Anthonomus musculus</Emphasis> Say

Studies were conducted to develop an attractant for the cranberry weevil, Anthonomus musculus, a pest of blueberry and cranberry flower buds and flowers in the northeastern United States. In previous studies, we showed that cinnamyl alcohol, the most abundant blueberry floral volatile, and the green leaf volatiles (Z)-3-hexenyl acetate and hexyl acetate, emitted from both flowers and flower buds, elicit strong antennal responses from A. musculus. Here, we found that cinnamyl alcohol did not increase capture of A. musculus adults on yellow sticky traps compared with unbaited controls; however, weevils were highly attracted to traps baited with the Anthonomus eugenii Cano aggregation pheromone, indicating that these congeners share common pheromone components. To identify the A. musculus aggregation pheromone, headspace volatiles were collected from adults feeding on blueberry or cranberry flower buds and analyzed by gas chromatography-mass spectrometry. Three male-specific compounds were identified: (Z)-2-(3,3-dimethyl-cyclohexylidene) ethanol (Z grandlure II); (Z)-(3,3-dimethylcyclohexylidene) acetaldehyde (grandlure III); and (E)-(3,3- dimethylcyclohexylidene) acetaldehyde (grandlure IV). A fourth component, (E)-3,7-dimethyl-2,6-octadien-1-ol (geraniol), was emitted in similar quantities by males and females. The emission rates of these volatiles were about 2.8, 1.8, 1.3, and 0.9 ng/adult/d, respectively. Field experiments in highbush blueberry (New Jersey) and cranberry (Massachusetts) examined the attraction of A. musculus to traps baited with the male-produced compounds and geraniol presented alone and combined with (Z)-3-hexenyl acetate and hexyl acetate, and to traps baited with the pheromones of A. eugenii and A. grandis. In both states and crops, traps baited with the A. musculus male-produced compounds attracted the highest number of adults. Addition of the green leaf volatiles did not affect A. musculus attraction to its pheromone but skewed the sex ratio of the captured adults towards females. Although the role of plant volatiles in host-plant location by A. musculus is still unclear, our studies provide the first identification of the primary A. musculus aggregation pheromone components that can be used to monitor this pest in blueberry and cranberry pest management programs.     

Change in Behavioral Response to Herbivore-induced Plant Volatiles in a Predatory Mite Population

Damage by herbivorous spider mites induces plants to produce volatiles that attract predatory mites that consume the spider mites. A clear attraction to volatiles from Lima bean plants infested with the spider mite Tetranychus urticae has been consistently reported during more than 15 years for the predatory mite Phytoseiulus persimilis. We have monitored the response to volatiles from spider-mite infested Lima bean plants for a laboratory population of the predatory mite from 1991 to 1995 on a regular basis. A reduction in the level of attraction in the laboratory population of P. persimilis was recorded in mid-1992. The attraction of the laboratory population was weaker than that of a commercial population in the latter part of 1992, but the responses of these two populations were similarly weak in 1994 and 1995. Therefore, a behavioral change has also occurred in this commercial population. Experiments were carried out to address the potential causes of this change in attraction. The attraction of predators from a commercial population with a strong response decreased after being reared in our laboratory. Within a predator population with a low degree of attraction, strongly responding predators were present and they could be isolated on the basis of their behavior: predators that stayed on spider-mite infested plants in the rearing set-up had a strong attraction, while predators that had dispersed from the rearing set-up were not attracted to prey-infested bean plants. From our laboratory population with a low degree of attraction, isofemale lines were initiated and maintained for more than 20 generations. All isofemale lines exhibited a consistently strong attraction to spider mite-induced plant volatiles, similar to the attraction recorded for several populations in the past 15 years. Neither in a population with a strong attraction nor in two with a weak attraction was the response of the predators affected by a starvation period of 1–3 hr. Based on these results, possible causes for the observed reduction in predator attraction to spider mite-induced bean volatiles are discussed. The predatory mite P. persimilis is a cornerstone of biological control in many crops worldwide. Therefore, the change in foraging behavior recorded in this predator may have serious consequences for biological control of spider mites.     

The Involvement of Volatile Infochemicals from Spider Mites and from Food-Plants in Prey Location of the Generalist Predatory Mite <Emphasis Type="Italic">Neoseiulus californicus</Emphasis>

We investigated volatile infochemicals possibly involved in location of the generalist predatory mite Neoseiulus californicus to plants infested with spider mites in a Y-tube olfactometer. The predators significantly preferred volatiles from lima bean leaves infested with Tetranychus urticae to uninfested lima bean leaves. Likewise, they were attracted to volatiles from artificially damaged lima bean leaves and those from T. urticae plus their visible products. Significantly more predators chose infested lima bean leaves from which T. urticae plus their visible products had been removed than artificially damaged leaves, T. urticae, and their visible products. These results suggest that N. californicus is capable of exploiting a variety of volatile infochemicals originating from their prey, from the prey-foodplants themselves, and from the complex of the prey and the host plants (e.g., herbivore-induced volatiles). We also investigated predator response to some of the synthetic samples identified as volatile components emitted from T. urticae-infested lima bean leaves and/or artificially damaged lima bean leaves. The predators were attracted to each of the five synthetic volatile components: linalool, methyl salicylate, (Z)-3-hexen-1-ol, (E)-2-hexenal, and (Z)-3-hexenyl acetate. The role of each volatile compound in prey-searching behavior is discussed.     

Responses of Flea Beetle <Emphasis Type="Italic">Phyllotreta cruciferae</Emphasis> to Synthetic Aggregation Pheromone Components and Host Plant Volatiles in Field Trials

Male-specific compounds, previously identified from Phyllotreta cruciferae and synthesized or isolated from natural sources, attracted both sexes of the beetle in field trials and therefore function as components of a male-produced aggregation pheromone. Six field experiments of 7 to 10 d duration each were conducted over 2 yr using modified boll weevil traps and two doses of pheromone. Treatments containing two doses of allyl isothiocyanate (AITC), a breakdown product of glucosinolates in Brassica napus L., a host plant of the beetles, were included in the study. A dose response was observed for both the pheromone components and AITC, and combinations of the pheromone and AITC generally attracted greater numbers of flea beetles than did either component itself. This increased attraction to a combination of beetle-produced compounds and host odors has not been previously demonstrated in halticine beetles and could help explain patterns of movement by P. cruciferae into field crops.     

Aphid and Plant Volatiles Induce Oviposition in an Aphidophagous Hoverfly

Episyrphus balteatus DeGeer (Diptera, Syrphidae) is an abundant and efficient aphid-specific predator. We tested the electroantennographic (EAG) response of this syrphid fly to the common aphid alarm pheromone, (E)-β-farnesene (EβF), and to several plant volatiles, including terpenoids (mono- and sesquiterpenes) and green leaf volatiles (C6 and C9 alcohols and aldehydes). Monoterpenes evoked significant EAG responses, whereas sesquiterpenes were inactive, except for the aphid alarm pheromone (EβF). The most pronounced antennal responses were elicited by six and nine carbon green leaf alcohols and aldehydes [i.e., (Z)-3-hexenol, (E)-2-hexenol, (E)-2-hexenal, and hexanal]. To investigate the behavioral activity of some of these EAG-active compounds, E. balteatus females were exposed to R-(+)-limonene (monoterpene), (Z)-3-hexenol (green leaf alcohol), and EβF (sesquiterpene, common aphid alarm pheromone). A single E. balteatus gravid female was exposed for 10 min to an aphid-free Vicia faba plant that was co-located with a semiochemical dispenser. Without additional semiochemical, hoverfly females were not attracted to this plant, and no oviposition was observed. The monoterpene R-(+)-limonene did not affect the females’ foraging behavior, whereas (Z)-3-hexenol and EβF increased the time of flight and acceptance of the host plant. Moreover, these two chemicals induced oviposition on aphid-free plants, suggesting that selection of the oviposition site by predatory hoverflies relies on the perception of a volatile blend composed of prey pheromone and typical plant green leaf volatiles.     

Ipsenol: an aggregation pheromone forIps latidens (Leconte) (Coleoptera: Scolytidae)

Ipsenol was identified from the frass of male, but not female,Ips latidens from British Columbia, feeding in phloem tissue of lodgepole pine,Pinus contorta var.latifolia. The responses ofJ. latidens to sources of ips-enol andcis-verbenol were determined with multiple-funnel traps in stands of lodgepole pine in British Columbia. Ipsenol attracted both male and femaleI. latidens, verifying that it is a pheromone for this species. MaleI. latidens showed a slight preference for (S)-(–)-ipsenol.cis-Verbenol was not produced by beetles of either sex and, in contrast to an earlier report, both enantiomers inhibited attraction to ipsenol-baited traps. The predators,Enoclerus sphegeus andThanasimus undatulus (Cleridae), were attracted to traps baited withcis-verbenol and ipsenol.     

Olfactory Responses of Plutella xylostella Natural Enemies to Host Pheromone, Larval Frass, and Green Leaf Cabbage Volatiles

The parasitoids Trichogramma chilonis (Hymenoptera: Trichogrammatidae) and Cotesia plutellae (Hymenoptera: Braconidae), and the predator Chrysoperla carnea (Neuroptera: Chrysopidae), are potential biological control agents for the diamondback moth, Plutella xylostella (Lepidoptera: Yponomeutidae). We present studies on the interactions between these bioagents and various host-associated volatiles using a Y olfactometer. T. chilonis was attracted to a synthetic pheromone blend (Z11–16:Ald, Z11–16:Ac, and Z11–16:OH in a 1:1:0.01 ratio), to Z11–16:Ac alone, and to a 1:1 blend of Z11–16:Ac and Z11–16:Ald. C. plutellae responded to the blend and to Z11–16:Ac and Z11–16:Ald. Male and female C. carnea responded to the blend and to a 1:1 blend of the major components of the pheromone, although no response was elicited by single compounds. Among the four host larval frass volatiles tested (dipropyl disulfide, dimethyl disulfide, allyl isothiocyanate, and dimethyl trisulfide), only allyl isothiocyanate elicited significant responses in the parasitoids and predator, but C. plutellae and both sexes of C. carnea did respond to all four volatiles. Among the green leaf volatiles of cabbage (Brassica oleracea subsp. capitata), only Z3–6:Ac elicited significant responses from T. chilonis, C. plutellae, and C. carnea, but C. plutellae also responded to E2–6:Ald and Z3–6:OH. When these volatiles were blended with the pheromone, the responses were similar to those elicited by the pheromone alone, except for C. carnea males, which had an increased response. The effect of temperature on the response of the biological agents to a mixture of the pheromone blend and Z3–6:Ac was also studied. T. chilonis was attracted at temperatures of 25–35°C, while C. plutellae and C. carnea responded optimally at 30–35°C and 20–25°C, respectively. These results indicate that the sex pheromone and larval frass volatiles from the diamondback moth, as well as volatile compounds from cabbage, may be used by these natural enemies to locate their diamondback moth host.     

Primary Attraction of the Fir Engraver, Scolytus ventralis

In laboratory bioassays, Porapak Q-captured and steam-distilled volatiles from the bark of host trees, Abies grandis, particularly from root-rot-infected trees, attracted 50–70% of male and female fir engravers, Scolytus ventralis. Gas chromatographic–electroantennographic detection (GC-EAD) analyses of Porapak Q-captured bark volatiles revealed 19 EAD-active compounds of which 13 (mostly monoterpenes) were identified by GC–mass spectrometry (GC-MS). In separate field experiments, multiple-funnel traps baited with two blends of these 13 synthetic volatiles released at 280 and 340 mg/ 24 hr attracted 66 and 93% of the total S. ventralis captured, respectively. The clerid predator, Thanasimus undulatus, also responded strongly to the kairomonal volatiles. Additional experiments produced no evidence for aggregation pheromones in S. ventralis. These included laboratory bioassays and GC and GC-EAD analyses of Porapak Q-captured volatiles from male- and female-infested logs or trees undergoing mass attack in the field, GC analyses and/or bioassays of extracts from female accessory glands, extracted volatiles from emerged, attacking and juvenile hormone-treated beetles of both sexes, and videotape analysis of the behavior of attacking beetles on the bark surface. We argue against the hypothesis of pheromone-mediated secondary attraction in S. ventralis and conclude that the attack dynamics of this species can be explained solely by its sensitive primary attraction response to host volatiles.     

Behavioral Responses of Adult <Emphasis Type="Italic">Sitophilus granarius</Emphasis> to Individual Cereal Volatiles

The antennae of Sitophilus granarius (L.) (Coleoptera: Curculionidae) adults detect a wide variety of compounds in the odor blend of various cereal grains (Germinara et al., Tec. Molit., 53:27–34, 2002). In the present study, we looked at the behavioral responses of the granary weevil to 20 of these individual volatiles (aliphatic alcohols, aldehydes, ketones, and aromatics) in a two-choice pitfall olfactometer, using the aggregation pheromone and propionic acid as the attractant and repellent controls, respectively. Five doses, ranging from 1 μg to 1 mg, of each compound were tested. At least one concentration of eight compounds attracted beetles but required doses 1,000- to 5,000-fold higher than the concentration of aggregation pheromone to elicit a response. Three compounds, while attractive at lower concentrations, acted as repellents at higher doses. Twelve compounds were repellent at concentrations similar to the quantity of propionic acid that significantly repelled beetles. The data show that granary weevil adults have the ability to respond behaviorally to a wide range of cereal volatiles and that responses may change as a function of concentration. The results suggest that host finding behavior of weevils will depend on the balance of positive and negative volatile stimuli from grain as the relative concentrations of volatiles may change during storage. An understanding of how the weevils respond to such changes could be useful for the development of effective integrated pest management strategies.     

Field evaluation ofAnomala schonfeldti ohaus (Coleoptera: Scarabaeidae) synthetic sex pheromone

Synthetic 2-(E)-nonenol, previously identified as the sex pheromone ofAnomala schonfeldti (Coleoptera: Scarabaeidae), is demonstrated to be very attractive to males in the field. Nevertheless, no significant differences were found between treatments with 1, 5, 10, and 20 mg dosages. Males ofA. schonfeldti were more significantly attracted to traps at 30 cm high than at 90 cm. Although the observed behavior seemed to indicate a trend of more attraction to buried traps than those placed at 30 cm, there was no statistical difference between the two treatments. Pheromone-baited traps caught significantly more beetles than traps containing three virgin females. Over 70% of released beetles were recaptured in six traps surrounding the point of release and separated from each other by 50 m, suggesting a possible use of the pheromone (in combination with floral compounds) in mass trapping.     

Mechanism of aggregation behavior inMaladera matrida Argaman (Coleoptera: Scarabaeidae)

AdultMaladera matrida Argaman (Coleoptera: Scarabaeidae: Melolonthinae) males emerge from soil for an active period at dusk, a few minutes before the females. Adults are found during most of the active hours on the foliage in aggregations composed of an equal sex ratio. The mechanism of aggregation behavior ofM. matrida beetles was studied in a Y-shaped olfactometer. No evidence was found for the existence of an aggregation pheromone released either by males or by females, but behavior tests indicate that adultM. matrida beetles, males as well as females, are attracted to volatiles of an injured host plant. The following scenario is suggested: Males emerge daily from soil at dusk, a few minutes before the females, and immediately start feeding. Additional males are attracted to the injured host's volatiles and form aggregations. When females emerge from soil, the attractant volatiles are concentrated in spots, and the females join the aggregations, forming an equal sex ratio.