Circadian rhythms of insect pheromone titer, calling, emission, and response: a review.

Many insect species have circadian rhythms of pheromone production/titer, calling, emission, and response that are involved in intraspecific communication and impact pest management practices. Rhythms of pheromone biosynthesis, most studied in moths affecting forestry and agriculture, contribute to a periodicity of pheromone concentration or titer within glands or hemolymph. Calling rhythms by the pheromone-emitting sex are physical movements (pumping, vibrating wings) that aid in release and dispersion of the volatile pheromone components attractive to the opposite conspecific sex or both sexes. Circadian rhythms of emission of pheromone also occur as a result of an interaction between calling and the titer of pheromone available for release. Responding individuals usually show a coincidental rhythm of dispersal flight while seeking pheromone plumes in which, by orienting upwind, the insects find mates or food resources. However, some species begin searching an hour or more before the emitting sex initiates calling and emission, which benefits mass trapping control programs because the baited traps do not compete initially with natural pheromone sources. In our review, data of daily rhythms of moths and other insects were extracted from the literature by screen capture software to calculate mean time of activity and standard deviation and fit to normal curves. These methods are illustrated for various insects and as a basis for discussion of interactions of pheromonal circadian rhythms of the well-studied gypsy moth Lymantria dispar, spruce budworm moth Choristoneura fumiferana, turnip moth Agrotis segetum, and cabbage looper moth Trichoplusia ni. The various circadian rhythms are discussed in relation to application of species-specific sex and aggregation pheromones for benign biological control and management of pest insects.

Index of host habitat preference explored by movement-based simulations and trap captures.

Animal species likely have different strengths of host habitat preference (HHP) that might be characterized by a standardized index ranging from 0 (no preference) to 1 (maximum preference). We hypothesized that in some species, HHP may result from individuals dispersing out of the host habitat having a probability of turning back at the boundary, or after entering host habitat by reducing speed or increasing size of turning angles. Computer simulations of individuals moving between various sized patches of host and nonhost habitat were conducted based on these three behaviours hypothesized to affect HHP. In the rebounding model, simulations resulted in equilibria of animal numbers inside and outside of host habitat that depend on sizes of these areas, initial number and the rebounding probability. Curvilinear regression of simulation results suggested an equation that predicted numbers in the host habitat and was solved for rebounding probability. A modified equation that sampled population densities (e.g., insect pheromone trap catches) inside and outside host habitat areas gave the rebounding probability, an index of HHP, without requiring the sizes of the areas. Simulations with traps and moving animals verified that the modified equation could predict the index correctly. The modified equation also estimates an index of HHP from sampled densities due to speed reductions and a combination of this and rebounding. Changes in angular turning size upon entering host habitat, however, did not affect habitat preference. Using pheromone trap captures, we found that the lesser date moth Batrachedra amydraula has a HHP for date Phoenix dactylifera plantations of 0.96. Host habitat preference indexes also were calculated from sampled insect densities reported in the literature. The new index of HHP is useful to characterize habitat patches of many organisms and aid understanding of animal spatial distributions and speciation processes. In addition, the index can be applied in studies of invasive species, trap crops of pest insects and conservation management.

Inhibitory Effects of Semiochemicals on the Attraction of an Ambrosia Beetle Euwallacea nr. fornicatus to Quercivorol.

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. ß-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.

Earwigs (Labidura riparia) mimic rotting-flesh odor to deceive vertebrate predators.

Many insects repel predators with caustic chemicals, while insects mimicking odors of wastes/dead insects to fool predators have not been documented. We found that the shore earwig, Labidura riparia (Dermaptera: Labiduridae) when bitten by anole lizards, Anolis carolinenesus, spits a rotting-flesh odor that deceives these insectivores into rejecting prey. Once a lizard attacked and rejected an earwig, the lizard did not attack another earwig during several weeks despite consuming other prey, indicating associative learning after one trial. The fetid odor was found in the head-prothorax containing salivary glands of both male and female earwigs and was comprised of ∼ 100 ng dimethyl disulfide and ∼ 600 ng dimethyl trisulfide. Nymphs had <5 ng of either compound. Adults also spit odorous sulfides after prolonged attacks by harvester ants, Pogonomyrmex rugosus, who were only deterred by the earwig's forceps. Sulfides released by the earwig are similar to odors of carrion/feces, which may be innately repulsive to some vertebrate predators. The mean initial discharge percentage (IDP) of sulfides from a cohort of earwigs was 62 %; however, IDPs of individuals were highly variable (3-99 %; mean 57 %). The discharge refill time (DRT) to refill 50 % of the earwig's allomone reservoir was estimated at 13 h. A positive relationship in sulfide amounts with body weight was found only in females in 2009, suggesting metabolic cost tradeoffs were revealed when sulfide content was half that in 2010. This is the first report of insects releasing sulfur-containing compounds that may mimic carrion-fecal odors as a deceptive defense against vertebrate predators.

Sex pheromone component ratios and mating isolation among three Lygus plant bug species of North America.

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.

Modeling and regression analysis of semiochemical dose-response curves of insect antennal reception and behavior.

Dose-response curves of the effects of semiochemicals on neurophysiology and behavior are reported in many articles in insect chemical ecology. Most curves are shown in figures representing points connected by straight lines, in which the x-axis has order of magnitude increases in dosage vs. responses on the y-axis. The lack of regression curves indicates that the nature of the dose-response relationship is not well understood. Thus, a computer model was developed to simulate a flux of various numbers of pheromone molecules (10(3) to 5 × 10(6)) passing by 10(4) receptors distributed among 10(6) positions along an insect antenna. Each receptor was depolarized by at least one strike by a molecule, and subsequent strikes had no additional effect. The simulations showed that with an increase in pheromone release rate, the antennal response would increase in a convex fashion and not in a logarithmic relation as suggested previously. Non-linear regression showed that a family of kinetic formation functions fit the simulated data nearly perfectly (R(2) >0.999). This is reasonable because olfactory receptors have proteins that bind to the pheromone molecule and are expected to exhibit enzyme kinetics. Over 90 dose-response relationships reported in the literature of electroantennographic and behavioral bioassays in the laboratory and field were analyzed by the logarithmic and kinetic formation functions. This analysis showed that in 95% of the cases, the kinetic functions explained the relationships better than the logarithmic (mean of about 20% better). The kinetic curves become sigmoid when graphed on a log scale on the x-axis. Dose-catch relationships in the field are similar to dose-EAR (effective attraction radius, in which a spherical radius indicates the trapping effect of a lure) and the circular EARc in two dimensions used in mass trapping models. The use of kinetic formation functions for dose-response curves of attractants, and kinetic decay curves for inhibitors, will allow more accurate predictions of insect catch in monitoring and control programs.

Genetic versus census estimators of the opportunity for sexual selection in the wild.

Abstract The existence of a direct link between intensity of sexual selection and mating-system type is widely accepted. However, the quantification of sexual selection has proven problematic. Several measures of sexual selection have been proposed, including the operational sex ratio (OSR), the breeding sex ratio (BSR), and the opportunity for sexual selection (I(mates)). For a wild population of pronghorn (Antilocapra americana), we calculated OSR and BSR. We estimated I(mates) from census data on the spatial and temporal distribution of receptive females in rut and from a multigenerational genetic pedigree. OSR and BSR indicated weak sexual selection on males, but census and pedigree I(mates) suggested stronger sexual selection on males than on females. OSR and BSR correlated with census but not pedigree estimates of I(mates), and census I(mates) did not correlate with pedigree estimates. This suggests that the behavioral mating system, as deduced from the spatial and temporal distribution of females, does not predict the genetic mating system of pronghorn. The differences we observed between estimators were primarily due to female mate sampling and choice and to the sex ratio. For most species, behavioral data are not perfectly accurate and therefore will be an insufficient alternative to using multigenerational pedigrees to quantify sexual selection.

Estimating insect flight densities from attractive trap catches and flight height distributions.

Methods and equations have not been developed previously to estimate insect flight densities, a key factor in decisions regarding trap and lure deployment in programs of monitoring, mass trapping, and mating disruption with semiochemicals. An equation to estimate densities of flying insects per hectare is presented that uses the standard deviation (SD) of the vertical flight distribution, trapping time, the trap's spherical effective radius (ER), catch at the mean flight height (as estimated from a best-fitting normal distribution with SD), and an estimated average flight speed. Data from previous reports were used to estimate flight densities with the equations. The same equations can use traps with pheromone lures or attractive colors with a measured effective attraction radius (EAR) instead of the ER. In practice, EAR is more useful than ER for flight density calculations since attractive traps catch higher numbers of insects and thus can measure lower populations more readily. Computer simulations in three dimensions with varying numbers of insects (density) and varying EAR were used to validate the equations for density estimates of insects in the field. Few studies have provided data to obtain EAR, SD, speed, and trapping time to estimate flight densities per hectare. However, the necessary parameters can be measured more precisely in future studies.

Modelling push-pull management of pest insects using repellents and attractive traps in fruit tree orchards.

BACKGROUND: Push-pull with semiochemicals in pest management uses repellents to reduce response of pests to food-mate resources (push) and attractive traps to reduce populations (pull). Simulation models of push-pull can aid understanding of plant-insect interactions in nature and suggest hypotheses for field tests that improve management. A previous model indicated advantages of push-pull for controlling ambrosia beetle, Euwallacea fornicatus, pest of avocado trees. However, the simulated behavior of repellency was inconsistent with field observations. RESULTS: We simulated individual-based movement of insects in push-pull to reveal relationships between parameters of radii (strength) of attractive traps, pest aggregations, and repellents with densities of each in an avocado orchard to visualize and understand the interactions and significance. Simulations indicated placement of traps along a 1-ha area periphery as a barrier resulted in similar trapping and mating as when traps were in a grid, either when insects originated randomly inside the plot or came from outside the plot. However, when insects originated from outside, both arrangements caught slightly more than when insects originated within the plot. CONCLUSION: There were no differences in capture rates whether traps were spaced in a barrier along the plot's periphery or in a grid covering the plot. Push-pull was more effective than pull (mass trapping) alone. Repellent behavior of individuals when encountering a repellent radius was modelled by approximate 90° turns (right or left at random) when about to enter an infestation, consistent with earlier observations of effects of repellents on bark beetles orienting to aggregation pheromone. © 2022 Society of Chemical Industry.

Inbreeding depression in pronghorn (Antilocapra americana) fawns.

Although inbreeding depression affects survival, fitness and population viability, the extent of inbreeding depression in wild populations remains unclear. We examined inbreeding depression in the small, isolated National Bison Range (NBR), MT, pronghorn (Antilocapra americana) population following a bottleneck. We have studied the National Bison Range pronghorn extensively since 1981, and we have detailed birth, survival and mate choice data. We genotyped all animals in the population between 1999 and 2010 at 19 microsatellite loci, assigned paternities to all fawns based on genotype and constructed a genetic-based pedigree to calculate inbreeding coefficients (f). We found an increase in the frequency of inbreeding following the bottleneck. We detected evidence of inbreeding depression on fawn survival to weaning, birth mass, foot length and condition. We estimated the number of diploid lethal equivalents on survival to weaning as 24.17-28.72. Standardized heterozygosity (H) had a relatively small influence on survival, mass, length and condition compared with f, and H was not correlated with f. We conclude that for pronghorn, H was not a good predictor of pedigree-estimates of f.

Methyl Salicylate Fails to Enhance Arthropod Predator Abundance or Predator to Pest Ratios in Cotton.

Conservation biological control is a fundamental tactic in integrated pest management (IPM). Greater biological control services can be achieved by enhancing agroecosystems to be more favorable to the presence, survival, and growth of natural enemy populations. One approach that has been tested in numerous agricultural systems is the deployment of synthetic chemicals that mimic those produced by the plant when under attack by pests. These signals may attract arthropod natural enemies to crop habitats and thus potentially improve biological control activity locally. A 2-yr field study was conducted in the cotton agroecosystem to evaluate the potential of synthetic methyl salicylate (MeSA) to attract native arthropod natural enemies and to enhance biological control services on two key pests. Slow-release packets of MeSA were deployed in replicated cotton plots season long. The abundance of multiple taxa of natural enemies and two major pests were monitored weekly by several sampling methods. The deployment of MeSA failed to increase natural enemy abundance and pest densities did not decline. Predator to prey ratios, used as a proxy to estimate biological control function, also largely failed to increase with MeSA deployment. One exception was a season-long increase in the ratio of Orius tristicolor (White) (Hemiptera: Anthocoridae) to Bemisia argentifolii Bellows and Perring (= Bemisia tabaci MEAM1) (Hemiptera: Aleyrodidae) adults within the context of biological control informed action thresholds. Overall results suggest that MeSA would not likely enhance conservation biological control by the natural enemy community typical of U.S. western cotton production systems.

Photosynthesis and assimilate partitioning between carbohydrates and isoprenoid products in vegetatively active and dormant guayule: physiological and environmental constraints on rubber accumulation in a semiarid shrub.

The stems and roots of the semiarid shrub guayule, Parthenium argentatum, contain a significant amount of natural rubber. Rubber accumulates in guayule when plants are vegetatively and reproductively dormant, complicating the relationship between growth/reproduction and product synthesis. To evaluate the factors regulating the partitioning of carbon to rubber, carbon assimilation and partitioning were measured in guayule plants that were grown under simulated summer- and winter-like conditions and under winter-like conditions with CO(2) enrichment. These conditions were used to induce vegetatively active and dormant states and to increase the source strength of vegetatively dormant plants, respectively. Rates of CO(2) assimilation, measured under growth temperatures and CO(2) , were similar for plants grown under summer- and winter-like conditions, but were higher with elevated CO(2) . After 5 months, plants grown under summer-like conditions had the greatest aboveground biomass, but the lowest levels of non-structural carbohydrates and rubber. In contrast, the amount of resin in the stems was similar under all growth conditions. Emission of biogenic volatile compounds was more than three-fold higher in plants grown under summer- compared with winter-like conditions. Taken together, the results show that guayule plants maintain a high rate of photosynthesis and accumulate non-structural carbohydrates and rubber in the vegetatively dormant state, but emit volatile compounds at a lower rate when compared with more vegetatively active plants. Enrichment with CO(2) in the vegetatively dormant state increased carbohydrate content but not the amount of rubber, suggesting that partitioning of assimilate to rubber is limited by sink strength in guayule.

Semiochemicals Affecting Attraction of Ambrosia Beetle Euwallacea fornicatus (Coleoptera: Curculionidae: Scolytinae) to Quercivorol: Developing Push-Pull Control.

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.

Modeling distributions of flying insects: effective attraction radius of pheromone in two and three dimensions.

The effective attraction radius (EAR) of an attractive pheromone-baited trap was defined as the radius of a passive "sticky" sphere that would intercept the same number of flying insects as the attractant. The EAR for a particular attractant and insect species in nature is easily determined by a catch ratio on attractive and passive (unbaited) traps, and the interception area of the passive trap. The spherical EAR can be transformed into a circular EAR(c) that is convenient to use in two-dimensional encounter rate models of mass trapping and mating disruption with semiochemicals to control insects. The EAR(c) equation requires an estimate of the effective thickness of the layer where the insect flies in search of mates and food/habitat. The standard deviation (SD) of flight height of several insect species was determined from their catches on traps of increasing heights reported in the literature. The thickness of the effective flight layer (F(L)) was assumed to be SD x square root of 2pi, because the probability area equal to the height of the normal distribution,1/(SD x square root of 2pi), times the F(L) is equal to the area under the normal curve. To test this assumption, 2000 simulated insects were allowed to fly in a three-dimensional correlated random walk in a 10-m thick layer where an algorithm caused them to redistribute according to a normal distribution with specified SD and mean at the midpoint of this layer. Under the same conditions, a spherical EAR was placed at the center of the 10-m layer and intercepted flying insects distributed normally for a set period. The number caught was equivalent to that caught in another simulation with a uniform flight density in a narrower layer equal to F(L), thus verifying the equation to calculate F(L). The EAR and F(L) were used to obtain a smaller EAR(c) for use in a two-dimensional model that caught an equivalent number of insects as that with EAR in three dimensions. This verifies that the F(L) estimation equation and EAR to EAR(c) conversion methods are appropriate.

Lygus spp. (Heteroptera: Miridae) host-plant interactions with Lesquerella fendleri (Brassicaceae), a new crop in the arid southwest.

Lesquerella fendleri is a new crop being cultivated in the arid Southwest for the hydroxy fatty acids found in its seed oils; however, little is know about the agricultural pests that will affect the seed production of this crop or the role the crop may have as a possible source or sink for current agricultural pests in the Arizona landscape. In the early spring, lygus bugs, some of our most important agricultural pests, are normally found in relatively small numbers on weeds; however, as lesquerella production increases, it may serve as an important early-season host for lygus. Here we present results from olfactometer bioassays that showed a significant attraction by Lygus hesperus females to volatiles associated with flowering lesquerella. Headspace volatiles of flowering lesquerella were collected and identified by gas chromatography-mass spectrometry, and phenylacetaldehyde (PAA) was found to be the major component, followed by benzaldehyde and Z-3- hexenyl acetate. In the field, we examined the Lygus spp. complex, as well as the prevalence of other herbivores and select predators, and monitored their responses to unbaited sticky traps of various colors and PAA-baited green and blue sticky traps. Green, blue, and clear sticky traps captured significantly more Lygus spp. (L. elisus in particular) than opaque yellow and red traps, but PAA-baited blue and green traps did not capture more lygus than unbaited traps. Collops spp., however, were collected in higher numbers on PAA-baited traps, suggesting that this compound might provide a means of recruiting and/or retaining this particular natural enemy.

Simulation of mating disruption and mass trapping with competitive attraction and camouflage.

Simulation models of mass trapping and mating disruption were developed based on correlated random walks (CRW) of flying male moths searching for females. Males encountered pheromone plumes, transformed into a circular probability surface represented as an effective attraction radius (EAR), from females and from dispensers with or without traps. In simulations, parameters of dispenser EAR and density, female EAR and density, female stationary periods, male density, and male orienting times in EAR of dispensers or females were varied, whereas the male CRW parameters (speed, turning angle, and step size) remained constant to evaluate effects on the percentages of females mating. When male orienting time was constant regardless of EAR, the models indicated no difference in mating disruption efficacy between either a higher density of dispensers with smaller EAR or a lower density of dispensers with a compensating larger EAR. However, when the orienting time was increased in proportion to dispenser EAR, fewer dispensers with larger EAR were more effective in reducing female mating than were more numerous ones with smaller EAR. When costs of pheromone are substantial, however, more numerous dispensers of smaller EAR would be more economical because dose-response curves in previous studies indicate release rate must increase exponentially to achieve a linear increase in EAR. The models are useful in understanding the variables affecting the success of insect control programs. More precise measurements of the above parameters in the field are needed before the models can precisely predict outcomes of mating disruption and mass trapping.

Attraction of the Euwallacea sp. near fornicatus (Coleoptera: Curculionidae) to Quercivorol and to Infestations in Avocado.

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.

An insect anti-antiaphrodisiac.

Passive mechanisms of mate guarding are used by males to promote sperm precedence with little cost, but these tactics can be disadvantageous for their mates and other males. Mated females of the plant bug Lygus hesperus are rendered temporarily unattractive by seminal fluids containing myristyl acetate and geranylgeranyl acetate. These antiaphrodisiac pheromones are gradually released from the female's gonopore, declining until they no longer suppress male courtship. Because starting quantities of these compounds can vary widely, the repellant signal becomes less reliable over time. Evidence was found of a complimentary mechanism that more accurately conveys female mating status. Once inside the female, geranylgeranyl acetate is progressively converted to geranylgeraniol then externalized. Geranylgeraniol counteracts the antiaphrodisiac effect despite having no inherent attractant properties of its own. This is the first evidence for such an anti-antiaphrodisiac pheromone, adding a new element to the communication mechanisms regulating reproductive behaviors.

Pollinator-prey conflicts in carnivorous plants: When flower and trap properties mean life or death.

Insect-pollinated carnivorous plants are expected to have higher fitness if they resolve pollinator-prey conflicts by sparing insects pollinating their flowers while trapping prey insects. We examined whether separation between flowers and traps of the carnivorous sundew species or pollinator preferences for colours of flowers enable these plants to spare pollinators. In addition, we collected odours from flowers and traps of each carnivorous species in order to identify volatile chemicals that are attractive or repellent to pollinators and prey insects. In Drosera spatulata and D. arcturi, no volatiles were detected from either their flowers or traps that could serve as kairomone attractants for insects. However, behavioural experiments indicated white colour and spatial separation between flowers and traps aid in reducing pollinator entrapment while capturing prey. In contrast, D. auriculata have flowers that are adjacent to their traps. In this species we identified chemical signals emanating from flowers that comprised an eight-component blend, while the plant's traps emitted a unique four-component blend. The floral odour attracted both pollinator and prey insects, while trap odour only attracted prey. This is the first scientific report to demonstrate that carnivorous plants utilize visual, spatial, and chemical signals to spare flower visitors while trapping prey insects.