Mating Disruption of a Flighted Spongy Moth, Lymantria Dispar Japonica (Motchulsky) in Japan.

Mating disruption of a flighted spongy moth, Lymantria dispar japonica (Motchulsky)(Lepidoptera: Lymantridae), with a synthetic version of its sex pheromone, (+)-disparlure ([7R,8S] -cis-7,8-epoxy-2- methyloctadecane), was tested in the forests in Japan. Pheromone trap catches and the percentage mating of tethered females were measured in the pheromone-treated and untreated control forests. The attraction of male moths to pheromone traps placed at a height of 1.5 m was significantly disrupted when the pheromone dispensers were placed at 1.5 m height, but many moths were captured in control plots. Mating of tethered females placed at 1.5 m was inhibited entirely, while 44% of females were mated in an untreated control forest. We report the first trial of mating disruption against a flighted spongy moth, and these results suggest that mating disruption with the synthetic sex pheromone appears promising for reducing damage caused by L. dispar japonica.

Perception of Heterospecific Sex Pheromone Causes Less Effective Mating Disruption in the Beet Semilooper, Autographa nigrisigna (Lepidoptera: Noctuidae).

Confuser® V is a pheromone-based mating disruptant designed to reduce damage caused by seven species of moth pests, including the beet semilooper, Autographa nigrisigna (Lepidoptera: Noctuidae). Eggs and larvae of A. nigrisigna are often found in fields treated with Confuser® V, suggesting that some components in the Confuser® V blend may have adverse effects on the efficacy of mating disruption of this species. Therefore, we examined whether A. nigrisigna perceives heterospecific pheromone components in the Confuser® V blend and delineated the roles of these components with respect to attraction and communication disruption. We found that several heterospecific pheromone components in the Confuser® V blend were perceived by male A. nigrisigna, and the addition of these components to the pheromone blend of A. nigrisigna reduced the attraction of males in the field, and tended to reduce the efficacy of mating disruption in cage bioassays.

Sex Attractant Pheromone of the Japanese Slave-Making Ant, Polyergus samurai.

The objective of our project was to identify the sex attractant pheromone of queens of the slave-making ant Polyergus samurai, which was characterized as a blend of methyl 6-methylsalicylate and methyl 3-ethyl-4-methylpentanoate. Both compounds were identified in volatiles collected from squashed heads of queens, and in field trials, a blend of the two compounds was highly attractive to males. In contrast, males were not attracted to a blend of methyl 6-methylsalicylate with 3-ethyl-4-methylpentanol. Thus, the pheromone blend of this species is analogous to that of species in the Polyergus lucidus complex from eastern North America, rather than that of the Polyergus breviceps complex from western North America and the European species P. rufescens. These results are discussed in the context of speciation within the genus.

Plant odor and sex pheromone are integral elements of specific mate recognition in an insect herbivore.

Specific mate recognition relies on the chemical senses in most animals, and especially in nocturnal insects. Two signal types mediate premating olfactory communication in terrestrial habitats: sex pheromones, which blend into an atmosphere of plant odorants. We show that host plant volatiles affect the perception of sex pheromone in males of the African cotton leafworm Spodoptera littoralis and that pheromone and plant volatiles are not perceived as independent messages. In clean air, S. littoralis males are attracted to single synthetic pheromone components or even the pheromone of a sibling species, oriental cotton leafworm S. litura. Presence of host plant volatiles, however, reduces the male response to deficient or heterospecific pheromone signals. That plant cues enhance discrimination of sex pheromone quality confirms the idea that specific mate recognition in noctuid moths has evolved in concert with adaptation to host plants. Shifts in either female host preference or sex pheromone biosynthesis give rise to new communication channels that have the potential to initiate or contribute to reproductive isolation.

Tetramorium tsushimae Ants Use Methyl Branched Hydrocarbons of Aphids for Partner Recognition.

In mutualisms, partner discrimination is often the most important challenge for interacting organisms. The interaction between ants and aphids is a model system for studying mutualisms; ants are provided with honeydew by aphids and, in turn, the ants offer beneficial services to the aphids. To establish and maintain this system, ants must discriminate mutualistic aphid species correctly. Although recent studies have shown that ants recognize aphids as mutualistic partners based on their cuticular hydrocarbons (CHCs), it was unclear which CHCs are involved in recognition. Here, we tested whether the n-alkane or methylalkane fraction, or both, of aphid CHCs were utilized as partner recognition cues by measuring ant aggressiveness toward these fractions. When workers of Tetramorium tsushimae ants were presented with dummies coated with n-alkanes of their mutualistic aphid Aphis craccivora, ants displayed higher levels of aggression than to dummies treated with total CHCs or methyl alkanes of A. craccivora; responses to dummies treated with n-alkanes of A. craccivora were similar to those to control dummies or dummies treated with the CHCs of the non-mutualistic aphid Acyrthosiphon pisum. By contrast, ants exhibited lower aggression to dummies treated with either total CHCs or the methylalkane fraction of the mutualistic aphid than to control dummies or dummies treated with CHCs of the non-mutualistic aphid. These results suggest that T. tsushimae ants use methylalkanes of the mutualistic aphid's CHCs to recognize partners, and that these ants do not recognize aphids as partners on the basis of n-alkanes.

Substrate vibrations mediate behavioral responses via femoral chordotonal organs in a cerambycid beetle.

BACKGROUND: Vibrational senses are vital for plant-dwelling animals because vibrations transmitted through plants allow them to detect approaching predators or conspecifics. Little is known, however, about how coleopteran insects detect vibrations. RESULTS: We investigated vibrational responses of the Japanese pine sawyer beetle, Monochamus alternatus, and its putative sense organs. This beetle showed startle responses, stridulation, freezing, and walking in response to vibrations below 1 kHz, indicating that they are able to detect low-frequency vibrations. For the first time in a coleopteran species, we have identified the sense organ involved in the freezing behavior. The femoral chordotonal organ (FCO), located in the mid-femur, contained 60-70 sensory neurons and was distally attached to the proximal tibia via a cuticular apodeme. Beetles with operated FCOs did not freeze in response to low-frequency vibrations during walking, whereas intact beetles did. These results indicate that the FCO is responsible for detecting low-frequency vibrations and mediating the behavioral responses. We discuss the behavioral significance of vibrational responses and physiological functions of FCOs in M. alternatus. CONCLUSIONS: Our findings revealed that substrate vibrations mediate behavioral responses via femoral chordotonal organs in M. alternatus.

Mating Disruption of a Carpenter Moth, Cossus insularis (Lepidoptera: Cossidae) in Apple Orchards with Synthetic Sex Pheromone, and Registration of the Pheromone as an Agrochemical.

Mating disruption of the carpenter moth, Cossus insularis (Staudinger) (Lepidoptera: Cossidae), with a synthetic version of its sex pheromone, a mixture of (E)-3-tetradecenyl acetate and (Z)-3-tetradecenyl acetate, was tested for three successive years in apple (Malus domestica Borkh.) orchards. Pheromone trap catches, percentage mating of tethered females and females enclosed with males in a mating cage, and tree damage were measured in both the pheromone-treated and untreated control orchards. The attraction of male moths to pheromone traps at heights of 1.5, 3, and 5 m was strongly disrupted when the pheromone dispensers were placed at 1.5 m height. Mating of tethered females placed at 1 m was completely inhibited, and the mating of tethered females at a height of 3 m was significantly reduced by the treatment in comparison to matings in an untreated control orchard. Similarly, mating of pairs of moths enclosed in mating cages was significantly reduced by the synthetic pheromone treatment in comparison to controls. The percentage of damaged trees in the pheromone-treated orchard also decreased significantly over the course of the experiment. These results suggest that mating disruption with the synthetic sex pheromone appears promising for reducing damage caused by C. insularis in apple orchards in Japan, and a commercial mating disruption product has been developed and registered.

Efficacy of Chemical Mimicry by Aphid Predators Depends on Aphid-Learning by Ants.

Chemical mimicry is an effective strategy when signal receivers recognize and discriminate models by relying on chemical cues. Some aphid enemies mimic the cuticular chemicals of aphids through various means thus avoiding detection and attack by aphid-tending ants. However, because ants have been reported to learn the chemical signatures of aphids in order to distinguish the aphids, the efficacy of chemical mimicry is predicted to depend on the experience of the ants that had tended aphids. The present study tested this hypothesis using two predator species: larvae of the green lacewing Mallada desjardinsi, and larvae of the ladybeetle Scymnus posticalis. Lacewing larvae carry the carcasses of aphids on which they have preyed upon their backs, and these function via chemical camouflage to reduce the aggressiveness of aphid-tending ants toward the larvae. Ladybeetle larvae reportedly produce a covering of wax structures, and their chemicals appear to attenuate ant aggression. We examined whether the behavior of the ant Tetramorium tsushimae toward these predators changed depending on their aphid-tending experience. Ants moderated their aggressiveness toward both predators when they had previously tended aphids, indicating that chemical mimicry by both aphid predators is dependent on previous experience of the ants in tending aphids. Chemical mimicry by the predators of ant-tended aphids is therefore considered to exploit learning-dependent aphid recognition systems of ants.

Identification of a Pheromone Component and a Critical Synergist for the Invasive Beetle Callidiellum rufipenne (Coleoptera: Cerambycidae).

The invasive Asian cerambycid beetle Callidiellum rufipenne (Motschulsky), informally known as the Japanese cedar longhorned beetle, was first detected in North America in North Carolina in 1997. The beetle has since been detected in neighboring states and is expected to further expand its range. However, delineating the current distribution of C. rufipenne has been hindered by the lack of efficient sampling methods. Here, we present the results of research on the chemistry of volatile pheromones of C. rufipenne. Analyses of headspace odors revealed that males produce (R)-3-hydroxyhexan-2-one, with lesser amounts of (S)-3-hydroxyhexan-2-one, and (R)- and (S)-2-hydroxyhexan-3-one. In field bioassays conducted over several years in Connecticut, where populations of the beetle were well established, no reconstructed blend of these compounds was significantly attractive to beetles of either sex. However, during field trials in Japan that targeted another species, we discovered that adult male and female C. rufipenne were attracted to a blend of racemic 3-hydroxyhexan-2-one and a novel natural product, 1-(1H-pyrrol-2-yl)-1,2-propanedione. Attraction to (R)-3-hydroxyhexan-2-one and the pyrrole subsequently was confirmed in field trials in Connecticut. Although it is unclear why the pyrrole acts as a synergist for a species that apparently does not produce it, the serendipitous discovery that adult C. rufipenne are attracted by the blend of ketone and pyrrole provides a badly needed method for monitoring its ongoing range expansion within North America, and for detecting new introductions in other parts of the world.

Ants Learn Aphid Species as Mutualistic Partners: Is the Learning Behavior Species-Specific?

In ant-aphid associations, many aphid species provide ants with honeydew and are tended by ants, whereas others are never tended and are frequently preyed upon by ants. In these relationships, ants must have the ability to discriminate among aphid species, with mutualistic aphids being accepted as partners rather than prey. Although ants reportedly use cuticular hydrocarbons (CHCs) of aphids to differentiate between mutualistic and non-mutualistic species, it is unclear whether the ability to recognize mutualistic aphid species as partners is innate or involves learning. Therefore, we tested whether aphid recognition by ants depends on learning, and whether the learning behavior is species-specific. When workers of the ant Tetramorium tsushimae had previously tended the cowpea aphid, Aphis craccivora, they were less aggressive toward this species. In addition, ants also reduced their aggressiveness toward another mutualistic aphid species, Aphis fabae, after tending A. craccivora, whereas ants remained aggressive toward the non-mutualistic aphid, Acyrthosiphon pisum, regardless of whether or not they had previous experience in tending A. craccivora. When ants were offered glass dummies treated with CHCs of these aphid species, ants that had tended A. craccivora displayed reduced aggression toward CHCs of A. craccivora and A. fabae. Chemical analyses showed the similarity of the CHC profiles between A. craccivora and A. fabae but not with A. pisum. These results suggest that aphid recognition of ants involves learning, and that the learning behavior may not be species-specific because of the similarity of CHCs between different aphid species with which they form mutualisms.

Candidate Attractant Pheromones of Two Potentially Invasive Asian Cerambycid Species in the Genus Xylotrechus.

Research during the 1980s showed that male Xylotrechus pyrrhoderus Bates (Coleoptera: Cerambycidae), a vineyard pest, produce (2S,3S)-2,3-octanediol and (S)-2-hydroxyoctan-3-one as possible pheromone components, but to our knowledge, field tests were not carried out. We confirmed that at least female X. pyrrhoderus were attracted by a 1:1 blend of these two compounds in field trials in Japan. Furthermore, more than 200 males and females of the congener Xylotrechus rufilius Bates were attracted by racemic 2-hydroxyoctan-3-one, and inhibited by syn-2,3-octanediol. Adult X. rufilius recently were intercepted in a shipment from China entering Baltimore, Maryland, raising concerns that this polyphagous species could establish in North America. Our results suggest that traps baited with 2-hydroxyoctan-3-one would be a valuable tool to assess whether X. rufilius has indeed become established, and to monitor for future introductions of X. rufilius.

Do aphid carcasses on the backs of larvae of green lacewing work as chemical mimicry against aphid-tending ants?

Ants attack and exclude natural enemies of aphids in ant-aphid mutualisms. However, larvae of the green lacewing, Mallada desjardinsi, prey on the cowpea aphid, Aphis craccivora, without exclusion by aphid-tending ants. Lacewing larvae are protected from ants by carrying aphid carcasses on their backs. Here, we tested whether cuticular hydrocarbons (CHCs) of aphid carcasses affected the aggressiveness of aphid-tending ants. Aphid carcasses were washed with n-hexane to remove lipids. Lacewing larvae with washed aphid carcasses were attacked by aphid-tending ants more frequently than those with untreated aphid carcasses. We measured the aggressiveness of aphid-tending ants to lacewing larvae that were either carrying a piece of cotton wool (a dummy aphid carcass) treated with CHCs from aphids or lacewing larvae, or carrying aphid carcasses. The rates of attack by ants on lacewing larvae carrying CHCs of aphids or aphid carcasses were lower than that of attack on lacewing larvae with conspecific CHCs. Chemical analysis by gas chromatography/mass spectrometry showed similarity of CHCs between aphids and aphid carcasses. These results suggest that aphid carcasses on the backs of lacewing larvae function via chemical camouflage to limit attacks by aphid-tending ants.

Female sex pheromone of a carpenter moth, Cossus insularis (Lepidoptera: Cossidae).

This study describes the identification of a sex pheromone component of a cossid moth, Cossus insularis. Coupled gas chromatographic-electroantennographic detection (GC-EAD) analysis of solid-phase microextraction (SPME) collections of volatiles released by live female moths showed that two compounds elicited EAG responses from the antennae of male moths. These compounds were identified as (E)-3-tetradecenyl acetate (E3-14:Ac) and (Z)-3-tetradecenyl acetate (Z3-14:Ac) by mass spectral analysis and retention index comparisons with synthetic standards. The ratio of E3-14:Ac and Z3-14:Ac was 95:5 in the effluvia of a female. In field bioassays, sticky traps baited with blends of E3-14:Ac and Z3-14:Ac showed that E3-14:Ac is an essential component of the pheromone. However, the role of Z3-14:Ac is unclear, because E3-14:Ac as a single component was as attractive to male moths as blends of E3-14:Ac and Z3-14:Ac, including the 95:5 blend released by live female moths.