Peter J. Silk

Reduction in diet breadth results in sequestration of plant chemicals and increases efficacy of chemical defense in a generalist grasshopper

The lubber grasshopper,Romalea guttata, is a generalist feeding on a broad diet of many herbaceous plant species and has a metathoracic defensive secretion normally containing phenolics and quinones synthesized by the insect. When insects were reared on a restricted diet of wild onion, they sequestered sulfur volatiles from the plant into their defensive secretions. These compounds were not detected by gas chromatography-mass spectroscopy in secretions of insects on an artificial diet or a natural, generalist diet of 26 plants that included wild onion as a component, nor were they present in secretions from field-collected insects. Defensive secretions of insects reared on wild onion were significantly more deterrent, by as much as an order of magnitude, to two species of ant predators than secretions from insects on either of the other two diets, despite a reduction in the concentration of autogenous defensive chemicals in secretions of insects on the onion diet. Sequestration of plant chemicals that increased defensive efficacy occurred when diet breadth was reduced. We suggest that this occurs because under conditions of specialization, plant secondary metabolites are more likely to be ingested and bioaccumulated in sufficient concentrations to have biological activity against predators. What we define as casual bioaccumulation of bioactive plant chemicals following dietary specialization may lead to evolution of sequestered defense syndromes in insects, and this process may not necessarily require specific adaptation to or coevolution with a toxic host plant.

Evidence for a Volatile Pheromone in Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) that Increases Attraction to a Host Foliar Volatile

ABSTRACT Analysis by gas chromatography/mass spectrometry (GC/MS) of volatiles from virgin female emerald ash borer, Agrilus planipennis Fairmaire confirmed the emission of (3Z)-lactone [(3Z)-dodecen-12-olide] but not its geometric isomer, (3E)-lactone [(3E)-dodecen-12-olide]. Gas chromatographic/electroantennographic (GC/EAD) analysis of synthetic (3Z)-lactone, which contained 10% (3E) -lactone, showed a strong response of male and female antennae to both isomers. EAG analysis with 0.01 - to100-µg dosages showed a positive dose response, with females giving significantly higher responses than males. In field experiments with sticky purple prism traps, neither lactone isomer affected catches when combined with ash foliar or cortical volatiles (green leaf volatiles or Phoebe oil, respectively). However, on green prism traps, the (3Z)-lactone significantly increased capture of male A. planipennis when traps were deployed in the canopy. Captures of males on traps with both (3E)-lactone and (3Z)-hexenol or with (3Z)-lactone and (3Z)-hexenol were increased by 45–100%, respectively, compared with traps baited with just (3Z)-hexenol. In olfactometer bioassays, males were significantly attracted to (3E) -lactone, but not the (3Z) -lactone or a 60:40 (3E): (3Z) blend. The combination of either (3E)- or (3Z)-lactone with Phoebe oil was not significantly attractive to males. Males were highly attracted to (3Z)-hexenol and the (3Z)-lactone + (3Z)-hexenol combination, providing support for the field trapping results. These data are the first to demonstrate increased attraction with a combination of a pheromone and a green leaf volatile in a Buprestid species.

Attraction of Agrilus planipennis (Coleoptera: Buprestidae) to a Volatile Pheromone: Effects of Release Rate, Host Volatile, and Trap Placement

ABSTRACT Attraction of emerald ash borer, Agrilus planipennis Fairmaire, to a volatile pheromone was demonstrated in three field experiments using baited green sticky traps. A dose-response curve was generated for male A. planipennis to increasing release rates of (3Z)-dodecen-12-olide ((3Z)lactone) in combination with the green leaf volatile, (3Z)-hexenol. Only the lowest release rate (<2.50 µg/d) of (3Z)-lactone significantly increased captures of male A. planipennis, as compared with traps baited with (3Z)-hexenol alone. Effect of trap height, (3Z)-lactone, and (3Z)-hexenol and their interactions on the trap capture of A. planipennis was determined in a factorial experiment. Number of males per trap was significantly and positively affected by (3Z)-lactone, (3Z)-hexenol, and trap height whereas number of females per trap was affected by trap height only; none of the interactions were significant. As predicted, the greatest mean catch of males was in traps baited with (3Z)-lactone and (3Z)-hexenol placed high in the canopy. Electroantennogram tests on the bark volatile, 7-epi-sesquithujene, demonstrated the ability of male and female A. planipennis antennae to detect and respond to this compound, particularly the (+)-7-epi-sesquithujene isomer. Results from an olfactometer bioassay and field testing did not demonstrate attraction of either males or females to (+)-7-epi-sesquithujene. These data increase our understanding of the pheromone ecology of the invasive emerald ash borer, provide further confirmation of the behavioral activity of the female-produced lactone pheromone, and should increase the ability to detect A. planipennis infestations where they are present.

Effects of diet breadth on autogenous chemical defense of a generalist grasshopper

The lubber grasshopper,Romalea guttata, produces a metathoracic defensive secretion containing primarily phenolics and quinones. This insect feeds on a wide range of plant species. Insects reared on an artificial diet and a diet of onion,Allium canadense, had secretions that contained fewer compounds, lower concentrations of compounds, and markedly altered relative composition of components compared to insects reared on a varied diet of 26 plant species that included onion. The study demonstrates that diet breadth has a major effect on the quality and quantity of the autogenous defensive secretion of this generalist herbivore. The results are compared to diet effects known in chemically defended specialists. Two possible mechanisms explaining the effects of diet breadth are proposed: one involves changes in precursor availability with changing diet breadth; the other suggests that physiological stress due to diet restriction changes allocation of resources to chemical defense.

Idiosyncratic variation in chemical defenses among individual generalist grasshoppers.

The defensive secretion of the lubber grasshopper,Romalea microptera, shows extreme chemical variation among individual adults of the same sex within a single wild population. Certain phenolic compounds were absent in some individuals and present in others. Concentrations of compounds, when present, varied over two to three orders of magnitude. Chemical variation attributable to individuals accounted for 60–88% of the total quantitative variation and was evenly contributed by all individuals in both sexes. Cluster and regression analyses showed no discernible predictable patterns in the defensive secretion variation. The specificity of chemical cues used by predators may explain why these defenses are so idiosyncratic.

Contact Sex Pheromones Identified for Two Species of Longhorned Beetles (Coleoptera: Cerambycidae) Tetropium fuscum and T. cinnamopterum in the Subfamily Spondylidinae

ABSTRACT Male Tetropium fuscum (F.) and T. cinnamopterum Kirby mated with live and dead (freeze-killed) conspecific females upon antennal contact, but did not respond to dead females after cuticular waxes were removed by hexane rinsing. Significantly fewer males of each species attempted to copulate with live or dead heterospecific females than with conspecifics, indicating that mate recognition was mediated by species-specific contact sex pheromones in the females cuticular hydrocarbons. GC/MS analysis of T. fuscum elytra identified n-alkanes and mono-methyl branched alkanes of which 11-methylheptacosane and 3- and 5-methyltricosanes were dominant in females. Full male responses, including copulatory behavior, were restored with application of enantiomerically pure synthetic (S)-11-methyl-heptacosane at 40 µg /female (one female equivalent) but not with racemic or (R)-11-methyl-heptacosane. The cuticular hydrocarbons on T. cinnamopterum elytra included 11-methyl-heptacosane as well as n-alkanes, methyl-branched alkanes, mono-alkenes, and (Z, Z)-6, 9-alkadienes. (Z)-9-pentacosene, (Z)-9-heptacosene, and 11-methyl-heptacosane were female dominant, but only (Z)-9-pentacosene elicited precopulatory behaviors in conspecific males at levels similar to those behaviors elicited by unrinsed females, but elicited copulation in fewer than half of males. At female equivalent dosages (10 µg), neither (Z)-9-heptacosene nor (S)-11-methylheptacosane elicited responses in males that were significantly different from those responses to a rinsed female but when applied together, the proportion of males responding was significantly increased. 11-methyl-heptacosene is thus a contact pheromone component common to both species, which may explain the heterospecific mating attempts by some males.

Efficacy of the pheromone (3Z)-lactone and the host kairomone (3Z)-hexenol at detecting early infestation of the emerald ash borer, Agrilus planipennis

The invasive emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is a major pest of ash trees, Fraxinus spp., in its introduced range in North America. Field studies were conducted to quantify the efficacy of traps baited with kairomone and pheromone lures for early detection of A. planipennis infestation. A trapping experiment demonstrated that green traps baited with the kairomone (3Z)‐hexenol detected at least one adult A. planipennis in 55.3% of plots with ‘nil to low’‐density infestations and in 100% of plots with ‘moderate to high’‐density A. planipennis infestations. Mean trap captures increased significantly with increasing infestation density. In terms of the optimal number of traps per plot, when one (3Z)‐hexenol‐baited trap was placed per plot, the trap detected populations in 62% of the plots with ‘low to moderate’‐density infestations through branch sampling. Detectability was increased to 82% when two traps were placed per plot. Finally, addition of female‐produced (3Z)‐lactone pheromone to traps significantly increased detection rates at both the trap and plot level, as compared with traps baited with the host volatile, (3Z)‐hexenol, alone (88 vs. 60%, respectively). Our results are the first to demonstrate the efficacy of baited green sticky traps for detecting low‐density A. planipennis infestations, particularly when the (3Z)‐lactone pheromone is used. This combination is therefore recommended for development of early‐detection protocols against A. planipennis.

Responses of Cerambycidae and other insects to traps baited with ethanol, 2,3-hexanediol, and 3,2-hydroxyketone lures in north-central Georgia

ABSTRACT In north-central Georgia, 13 species of woodboring beetles (Coleoptera: Cerambycidae: Cerambycinae) were attracted to multiple-funnel traps baited with ethanol and one of the following pheromones: (1) racemic 3-hydroxyhexan-2-one; (2) racemic 3-hydroxyoctan-2-one; and (3) syn-2,3-hexanediol. The following species were attracted to traps baited with ethanol and 3-hydroxyhexan-2-one: Anelaphus pumilus (Newman), Eburia quadrigeminata (Say), Euderces pini (Olivier), Knulliana cincta (Drury), Neoclytus mucronatus (F.), Neoclytus scutellaris (Olivier), and Xylotrechus colonus (F.). Clytus marginicollis Castelnau & Gory, and Anelaphus parallelus (Newman) were attracted to traps baited with ethanol and 3-hydroxyoctan-2-one, whereas traps baited with ethanol and syn-2,3-hexanediol were attractive to Anelaphus villosus (F.), A. parallelus, Neoclytus acuminatus (F.), Neoclytus jouteli jouteli Davis, and Megacyllene caryae (Gahan). Ethanol enhanced catches of seven cerambycid species in traps baited with syn-2,3-hexanediol and 3,2-hydroxyketones. Catches of bark and ambrosia beetles (Curculionidae: Scolytinae) in ethanol-baited traps were largely unaffected by the addition of syn-2,3-hexanediol and 3,2-hydroxyketone lures, except for two species. The mean catches of Hypothenemus rotundicollis Wood & Bright and Dryoxylon onoharaensum (Murayama) in ethanol-baited traps increased and decreased, respectively, with the addition of racemic 3-hydroxyoctan-2-one. Traps baited with ethanol and syn-2,3-hexanediol were attractive to Xylobiops basilaris (Say) (Bostrichidae) and Chariessa pilosa (Forster) (Cleridae), whereas Temnoscheila virescens (F.) (Trogossitidae) were attracted to traps baited with ethanol and 3-hydroxyhexan-2-one. The assassin bug, Apiomerus crassipes (F.) (Hemiptera: Reduviidae), was attracted to traps baited with ethanol and 3,2-hydroxyketones.

Phenology and Spatial Distribution of Native and Exotic Tetropium Longhorned Beetles (Coleoptera: Cerambycidae)

ABSTRACT The co-existence of two closely related Tetropium species in eastern Canada, invasive T. fuscum and native T. cinnamopterum (TF and TC, respectively), provides a model system to investigate seasonal and spatial demographic parameters of biological invasions at the interspecific level. In this study, we take advantage of the similar semiochemical communication of TF and TC to evaluate the abundance of adults of the two species concurrently using grids of traps baited with pheromone and host volatiles in stands of spruce. Adult TF emerged on average 2 wk before TC both in the field and under controlled laboratory conditions. This observation, combined with the early reproduction of emergent females, implies that the smaller (younger) larvae of native TC may be at increased risk of intra-guild predation by TF. The high spatial association between male and female TF in dense, aggregated populations suggests that the rate of mate encounter is depressed in sparse populations toward the edge of the invasive range. The higher level of spatial aggregation for TF than TC, particularly at high population density, suggests a higher propensity of adult TF to congregate at “landmarks.” Considering the broader range of host conditions, earlier seasonal emergence, and presumably more effective mate encounter for TF than TC, the exotic TF may be a superior competitor with the potential to displace or reduce the abundance of TC.

Behavioral and electrophysiological responses of Tetropium fuscum (Coleoptera: Cerambycidae) to pheromone and spruce volatiles.

ABSTRACT The brown spruce longhorn beetle, Tetropium fuscum (F.), is an invasive wood-boring species in eastern Canada. Gas chromatographic/electroantennographic (GC/EAD) analyses of Norway and red spruce volatiles detected a number of consistent EAD-active responses to compounds that are known to be stress-induced in spruce. The effects of these EAD-active compounds on various aspects of adult behavior were tested. In two-choice olfactometer assays, a monoterpene spruce blend, (R) - (-) -linalool, (3Z,6E) -&agr;-farnesene, (E) -&bgr;-farnesene and spruce essential oil were attractive to both sexes. However, when they were combined with the male-produced pheromone (fuscumol), they elicited a sex-specific response: females were significantly attracted to combinations of fuscumol plus either (3Z,6E)-&agr;-farnesene, (E)-&bgr;-farnesene and spruce essential oil but males were not. Fuscumol alone was unattractive to either sex in the olfactometer. Males exposed to fuscumol, (3Z,6E)-&agr;farnesene, or a combination of both, but not (E)-&bgr;-farnesene, were more likely to engage in the pheromone calling posture relative to controls. Both the monoterpene spruce blend and spruce essential oil elicited significantly greater trap capture of both sexes of T. fuscum in the presence of fuscumol and ethanol than (3Z,6E)-&agr;-farnesene or (R)-(-)-linalool, which did not elicit trap capture alone or in combination with fuscumol. The data support the hypothesis that stress-induced sesquiterpene components, such as (3Z,6E)-&agr;-farnesene, are important for mediating close-range attraction and behavior in T. fuscum while the monoterpene components are important for long-range processes (trap capture).