The plant bugs Lygus hesperus, Lygus lineolaris, and Lygus elisus (Hemiptera: Miridae) are major pests of many agricultural crops in North America. Previous studies suggested that females release a sex pheromone attractive to males. Other studies showed that males and females contain microgram amounts of (E)-4-oxo-2-hexenal, hexyl butyrate, and (E)-2-hexenyl butyrate that are emitted as a defense against predators. Using gas chromatography-mass spectrometry, we found that female L. lineolaris and L. elisus have a 4:10 ratio of hexyl butyrate to (E)-2-hexenyl butyrate that is reversed from the 10:1 ratio in female L. hesperus (males of the three species have ~10:1 ratio). These reversed ratios among females of the species suggest a behavioral role. Because both sexes have nearly equal amounts of the major volatiles, females should release more to attract males. This expectation was supported because L. hesperus females released more hexyl butyrate (mean of 86 ng/h) during the night (1800-0700 hours) than did males (<1 ng/h). We used slow-rotating pairs of traps to test the attraction of species to blends of the volatiles with a subtractive method to detect synergism. Each species' major butyrate ester was released at 3 μg/h, the minor butyrate according to its ratio, and (E)-4-oxo-2-hexenal at 2 μg/h. The resulting catches of only Lygus males suggest that (E)-4-oxo-2-hexenal is an essential sex pheromone component for all three species, (E)-2-hexenyl butyrate is essential for L. elisus and L. lineolaris, and hexyl butyrate is essential for L. hesperus. However, all three components are recognized by each species since ratios of the butyrate esters are critical for conspecific attraction and heterospecific avoidance by males and thus play a role in reproductive isolation among the three species. Because L. hesperus males and females are known to emit these major volatiles for repelling ant predators, our study links defensive allomones in Lygus bugs with an additional use as sex pheromones.
The Euwallacea fornicatus (Eichhoff) species complex includes the polyphagous shot hole borer (PSHB), an ambrosia beetle infesting avocado limbs, Persea americana Mill. Synthetic quercivorol, a monoterpene alcohol, is known to attract females (males are flightless) over a range of release rates spanning three orders of magnitude. The upper release dose was extended 10-fold using sticky traps baited with quercivorol released at 1× (0.126 mg/day), 10×, and 108× relative rates to obtain a dose-response curve fitting a kinetic formation function. Naturally infested limbs of living avocado trees were wrapped with netting to exclude the possibility of catching emerging beetles on the encircling sticky traps. The results indicate PSHB are significantly attracted to infested limbs. Ethanol released over a 64-fold range (lowest rate of 7.5 mg/day) was moderately inhibitory of PSHB attraction to 1× quercivorol. β-caryophyllene and eucalyptol did not appear to affect attraction at the rates tested. A field test of potential inhibitors of 1× quercivorol was done using ~1 mg/day releases of monoterpene ketones: (-)-(S)-verbenone, (+)-(R)-verbenone, 3-methyl-2-cyclo-hexen-1-one (MCH or seudenone), piperitone, (+)-(S)-carvone, and racemic cryptone. Only piperitone and the two enantiomers of verbenone were strongly inhibitory. A blend of piperitone and verbenone tested together at different distances (0, 0.5, 1, 2, and 4 m) from a 1× quercivorol baited sticky trap became increasingly ineffective in inhibiting the attractant as separation distance increased. Due to the relatively short-range repellency (<1 m), the inhibitors would need to be released from several places on each tree to effectively repel PSHB from avocado trees. Effective attraction radii, EAR, and circular EARc are estimated for the quercivorol baits released at 1×, 10× and 108× rates. Push-pull simulations of moving beetles were performed in 1 ha plots with 2, 4, or 16 traps of 10× EARc and 400 trees (0, 1, or 3 inhibitors per tree) of which ten had an infested limb (EARc = 0.5 m). The simulations indicate that push-pull methods would be more effective in reducing PSHB mating than simply using mass-trapping alone.
The Euwallacea sp. near fornicatus (Euwallacea sp. 1 hereafter) feeds on many woody shrubs and trees and is a pest of avocado, Persea americana Mill., in several countries including Israel and the United States. Quercivorol baits are commercially available for Euwallacea sp. 1 females (males do not fly), but their attractive strength compared to other pheromones and potential for mass trapping are unknown. We used sticky traps baited with quercivorol released at 0.126 mg/d (1×) and at 0.01×, 0.1×, and 10× relative rates to obtain a dose-response curve of Euwallacea sp. 1 attraction. The curve fitted well a kinetic formation function of first order. Naturally infested limbs of living avocado trees had attraction rates equivalent to 1× quercivorol. An effective attraction radius (EAR) was calculated according to previous equations for each of the various baits (1× EAR = 1.18 m; 10× EAR = 2.00 m). A pole with six sticky traps spaced from 0.25-5.75 m in height had captures of Euwallacea sp. 1 yielding a mean flight height of 1.24 m with vertical flight distribution SD of 0.88 m (0.82-0.96 m, 95% CI). The SD with specific EAR was used to calculate EARc, two-dimensional EAR (1× EARc = 0.99 m; 10× EARc = 2.86 m), for comparison with other insect pheromone traps and for use in simulations. The simulation methods described previously were performed with combinations of 1-16 traps with 1-50 aggregations per 9-ha plot. The simulations indicate mass trapping with quercivorol could be effective if begun in spring before Euwallacea sp. 1 establishes competing sources of attraction.
The lesser date moth (LDM) Batrachedra amydraula is a significant pest of date palm fruits. Previously, detection and monitoring of the pest was inaccurate due to high costs of sampling with lifting machines. We report a practical system for detection and monitoring of LDM based on pheromone traps and relevant models. Dose-response experiments with LDM pheromone traps indicated a 1 mg lure is optimal for monitoring. Delta traps with adhesive covering their entire inner surface gave the highest captures while trap colour was unimportant. Sampling pheromone traps throughout the night indicated male flight began at 1:00-2:00 and reached a peak 2 h before sunrise. Monitoring traps exposed all year long in Israel revealed three generations with different abundance. Trapping transects in a date plantation indicated interference from a monitoring trap became minimal at distances >27 m away. Inter-trap distances closer than this may lower efficiency of monitoring and mass trapping in control programs. Our estimate of the circular effective attraction radius (EARc) of a 1 mg delta trap for LDM (3.43 m) shows this bait is among the most attractive compared with baits for other insects. We developed encounter-rate equations with the pheromone trap EARc to model the interplay between population levels, trap density and captures that are useful for detection of invasive LDM and its control by mass trapping. The integrated methodologies are applicable to many pest species.
Euwallacea fornicatus (Eichhoff), the polyphagous shot hole borer (PSHB), is an ambrosia beetle infesting avocado Persea americana Mill. limbs in North America and Israel. We conducted field experiments with sticky traps in avocado orchards to develop push–pull semiochemical methods of managing PSHB. Traps baited with 10-fold increasing doses (0.01 to 100× or 1.26 µg to 12.6 mg/d) of attractant quercivorol were previously shown to increasingly capture female PSHB (males flightless). We converted trap catch of this relationship to a standardized effective attraction radius (EAR) that predicts capture power of baited-traps regardless of insect flight density. Earlier, piperitone and verbenone were shown to strongly inhibit attraction of PSHB to quercivorol-traps. We tested increasing numbers of 1× piperitone dispensers at 0.75-m distance surrounding a quercivorol-trap and found PSHB catch to decline exponentially. Increasing decadic doses (0.01 to 10×) of either verbenone or piperitone released at 1× quercivorol-traps caused a sigmoidal first-order kinetic-decay in catch. Verbenone (1×) placed at increasing distances (0, 0.25, 0.5, 1, and 2 m) from a 1× quercivorol-trap became increasingly ineffective in reducing catch of PSHB. We found no evidence that ethanol released from 7.5 to 480 mg/d affected attraction of PSHB, but Scobicia chevrieri (Villa and Villa) (Coleoptera: Bostrichidae) was increasingly attracted. Due to their relatively short-range (<0.5 m) inhibition of attractive sources, piperitone, and verbenone dispensers should be placed on avocado trunks where PSHB aggregations occur before the flight season.
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