MCH released in a novel pheromone dispenser prevents spruce beetle,Dendroctonus rufipennis (Coleoptera: Scolytidae), attacks in south-central Alaska

Field tests of 3-methyl-2-cyclohexen-1-one (MCH), the antiaggregation pheromone of the spruce beetle, Dendroctonus rufipennis Kirby, were conducted in south-central Alaska in stands of Lutz spruce, Picea x lutzii Little, and Sitka spruce, P. sitchensis (Bong.) Carr. The deployment of MCH in a novel releaser significantly reduced trap catches and spruce beetle attacks on standing live spruce by 96 and 87%, respectively. The results of this study demonstrate the first successful field test of MCH in Alaska for the prevention of spruce beetle attacks on standing, live spruce.     

Elution rate and spacing of antiaggregation pheromone dispensers for protecting live trees from Dendroctonus pseudotsugae (Coleoptera: Scolytidae)

The antiaggregation pheromone 3-methylcyclohex-2-en-1-one (MCH) is highly effective in preventing the infestation of high-risk trees by Douglas-fir beetle, Dendroctonus pseudotsugae Hopkins. A large portion of the cost of an MCH treatment is related to the time applicators spend walking through an area dispersing the formulated pheromone. Application of fewer MCH dispensers eluting at a higher rate than those currently registered for operational use could potentially reduce treatment costs. Two higher elution rates, 6 and 18 mg/d per dispenser, were compared with the current standard of 2 mg/d per dispenser and an untreated control on 1-ha circular plots. Dispensers were spaced 5,15, and 44 m apart around the plot perimeters eluting 2, 6, and 18 mg/d, respectively. The nominal dose of MCH was 144 mg/ha/d on all plots. Percentages of Douglas-fir trees > or = 20 cm diameter at breast height mass attacked by Douglas-fir beetle were significantly lower on plots treated with dispensers eluting 2 and 6 mg/d and spaced 5 and 15 m apart, respectively, compared with the untreated control. Infestation rate on plots treated with dispensers eluting 18 mg/d and spaced 44 m apart was not significantly different from the control. Douglas-fir beetle abundance and host tree availability were similar on all plots. These results indicate that MCH dispensers eluting 6 mg/d (three times the current standard rate) and spaced 15 m apart (three times existing standard distance) can effectively prevent Douglas-fir beetle infestations.     

Semiochemical emission from individual galleries of the southern pine beetle, (Coleoptera: Curculionidae: Scolytinae), attacking standing trees

We collected, identified, and quantified volatiles arising from individual gallery entrances of the monogamous bark beetle Dendroctonus frontalis Zimmermann. Samples were collected while the insects were mass attacking mature loblolly pines (Pinus taeda L.) in an established infestation in western Mississippi, 1 August through 3 October 2005. Following volatile sample collection, the entrances were dissected and categorized according to those that 1) contained a solitary female (the gallery initiating sex), 2) contained a pair that had not yet produced an egg gallery, 3) led to an egg gallery with niches and/or eggs, or 4) represented failed attacks (either abandoned or containing dead beetles). The greatest mean release rate of the female-produced aggregation pheromone components frontalin (74 ng/h) and trans-verbenol (0.35 microg/h) was detected from entrances of solitary females, whereas the highest mean quantities of the male-produced multifunctional pheromone components endo-brevicomin (18 ng/h) and verbenone (0.15 microg/h) were detected from entrances of preoviposition beetle pairs. Alpha-pinene, a host-produced monoterpene that functions as a synergist for the aggregation attractant for D. frontalis, was detected from entrances of solitary females and preoviposition pairs at a rate of 0.6 mg/h, or 3-4 orders of magnitude greater than the insect-produced components of the attractant. Our results indicate that the release rates of pheromone components used in published field studies of the chemical ecology of D. frontalis (generally > 0.1 mg/h) represent thousands of 'attack equivalents' or production rates on the scale of a beetle mass attack on a single host. Additionally, our data suggest that the loss in attractiveness of host tissue fully colonized by D. frontalis is because of the disappearance of attractants rather than an increase in inhibitors.     

Response of native and exotic longhorn beetles to common pheromone components provides partial support for the pheromone-free space hypothesis

Longhorn beetles are among the most important groups of invasive forest insects worldwide. In parallel, they represent one of the most well-studied insect groups in terms of chemical ecology. Longhorn beetle aggregation-sex pheromones are commonly used as trap lures for specific and generic surveillance programs at points of entry and may play a key role in determining the success or failure of exotic species establishment. An exotic species might be more likely to establish in a novel habitat if it relies on a pheromone channel that is different to that of native species active at the same time of year and day, allowing for unhindered mate location (i.e., pheromone-free space hypothesis). In this study, we first tested the attractiveness of single pheromone components (i.e., racemic 3-hydroxyhexan-2-one, racemic 3-hydroxyoctan-2-one, and syn-2,3-hexanediol), and their binary and tertiary combinations, to native and exotic longhorn beetle species in Canada and Italy. Second, we exploited trap catches to determine their seasonal flight activity. Third, we used pheromone-baited “timer traps” to determine longhorn beetle daily flight activity. The response to single pheromones and their combinations was mostly species specific but the combination of more than one pheromone component allowed catch of multiple species simultaneously in Italy. The response of the exotic species to pheromone components, coupled with results on seasonal and daily flight activity, provided partial support for the pheromone-free space hypothesis. This study aids in the understanding of longhorn beetle chemical ecology and confirms that pheromones can play a key role in longhorn beetle invasions.     

Synergistic blends of monoterpenes for aggregation pheromones of the mountain pine beetle (Coleoptera: Curculionidae)

The superiority of the host monoterpene myrcene as a synergist for trans-verbenol and exo-brevicomin, aggregation pheromone components of the mountain pine beetle, Dendroctonus ponderosae Hopkins (Coleoptera: Curculionidae), suggests that the ancestral host of the mountain pine beetle is a pine rich in myrcene. A field trapping experiment in British Columbia testing reconstituted bole oleoresin of whitebark pine, Pinus albicaulis Engelmann, composed of mainly myrcene (20.7%), terpinolene (6.8%), and 3-carene (61.9%) showed it to be a better pheromone synergist than reconstituted bole oleoresin of lodgepole pine, Pinus contorta variety latifolia Engelmann, which contained only 2.7, 1.0, and 6.0%, respectively, of the above-mentioned three compounds. In the same experiment myrcene alone was the best synergist. In subsequent experiments, testing myrcene, terpinolene and 3-carene alone and in all possible binary and ternary combinations, a 50:50 blend of myrcene and terpinolene released at the same rate as either compound alone generally resulted in trap catches approximately 3 times higher than with myrcene as a synergist. This result held as long as the terpinolene was free of contaminants, and the traps were in the open, well away from potential interference of semiochemicals emitted by newly attacked trees. 3-Carene seemed to be inert or slightly inhibitory. No single monoterpene tested alone or in binary or ternary combination in the absence of pheromones was attractive. There was no effect of doubling or tripling the release rate of myrcene or terpinolene. In five of nine experiments, adding terpinolene to myrcene caused a significant increase in the percentage of female mountain pine beetles captured. Among host pines, the presence of highly synergistic monoterpenes at various levels in combination with other monoterpenes that are apparently either inert or inhibitory could account for different degrees of pheromone synergism, and thus host preference. The highly synergistic effect of combining myrcene plus terpinolene with the mountain pine beetle aggregation pheromone components opens up the potential for suppression of populations through semiochemical-based mass trapping.     

Determination of Agriotes obscurus (Coleoptera: Elateridae) sex pheromone attraction range using target male behavioural responses

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Long‐term evaluation of field‐wide oriental beetle (Col., Scarabaeidae) mating disruption in blueberries using female‐mimic pheromone lures

The oriental beetle, Anomala orientalis (Waterhouse) (Col., Scarabaeidae), is the most important root‐feeding pest of blueberries and turfgrass in New Jersey, USA. Previous studies showed that mating disruption is a feasible option for oriental beetle management; however, assessing its efficiency can be challenging, and little is known on its long‐term effects. Accordingly, we conducted studies to investigate low‐dose pheromone lures equivalent to oriental beetle females (i.e. female mimics) as easy‐to‐use indicators of mating disruption success, determine the distance at which oriental beetle males respond to female‐mimic lures and assess the long‐term (3‐year) effects of mating disruption on oriental beetle populations in entire blueberry fields. Our studies showed that rubber septa baited with 0.3 μg of the oriental beetle sex pheromone (Z)‐7‐tetradecen‐2‐one attract similar numbers of males as compared with virgin females and can thus be used as a female mimic. The range of attraction of this lure was found to be also similar to virgin females and <30 m. In blueberries, mating disruption provided 87% inhibition of oriental beetle populations (trap shutdown) over a 3‐year period. Oriental beetle male captures in disrupted fields were threefold higher along the field edges than in the field interiors, indicating movement of males from nearby areas into the pheromone‐treated fields. In addition, mating disruption reduced male attraction to female‐mimic lures by 93% in all 3 years and reduced the number of larvae in sentinel potted plants in 1 of 2 years. These results show for the first time that mating disruption provides consistent long‐term field‐wide control of oriental beetle populations and that female‐mimic pheromone lures can be used as a new tool to assess oriental beetle mating disruption success.     

Field evaluation of effect of temperature on release of disparlure from a pheromone-baited trapping system used to monitor gypsy moth (Lepidoptera: Lymantriidae)

Traps baited with disparlure, the synthetic form of the gypsy moth, Lymantria dispar (L.) (Lepidoptera: Lymantriidae), sex pheromone are used to detect newly founded populations and estimate population density across the United States. The lures used in trapping devices are exposed to field conditions with varying climates, which can affect the rate of disparlure release. We evaluated the release rate of disparlure from delta traps baited with disparlure string dispenser from 1 to 3 yr across a broad geographic gradient, from northern Minnesota to southern North Carolina. Traps were deployed over approximately 12 wk that coincided with the period of male moth flight and the deployment schedule of traps under gypsy moth management programs. We measured a uniform rate of release across all locations when considered over the accumulation of degree-days; however, due to differences in degree-day accumulation across locations, there were significant differences in release rates over time among locations. The initial lure load seemed to be sufficient regardless of climate, although rapid release of the pheromone in warmer climates could affect trap efficacy in late season. Daily rates of release in colder climates, such as Minnesota and northern Wisconsin, may not be optimal in detection efforts. This work highlights the importance of local temperatures when deploying pheromone-baited traps for monitoring a species across a large and climatically diverse landscape.     

The distribution of eggs per host in a herbivorous insect--intersection of oviposition, dispersal and population dynamics

1. The dynamics of parasitic organisms depend critically upon the frequency distribution of parasite individuals per host. However, the processes giving rise to this frequency distribution have rarely been modelled and tested for organisms with complex host selection behaviour. 2. In this study Microrhopala vittata, a chrysomelid beetle, was used to investigate how oviposition behaviour, movement and density of host plants interact in shaping the frequency distribution of egg clusters per host in the field. 3. Enclosures were stocked with two different host species and different beetle densities and various stochastic process models were fitted to egg cluster count data obtained from these enclosures. The different models were derived considering different scenarios, in particular whether or not plant density limits oviposition rate, whether or not ovipositing females actively seek out the most attractive plant within their perception radius and whether a female's oviposition rate is determined by plant intrinsic factors, the plant's egg cluster load or the surrounding beetle density. 4. The model parameters fitted to cage data were used to describe the frequency distribution of egg cluster counts obtained in a release experiment in the field. A total of 220 beetle pairs were released at five locations in a field where this beetle was not observed previously. Each release point was at a border between the two host species. 5. One model predicted for the preferred host species the egg cluster count frequencies in the field from parameters estimated in the cages. This model assumed that egg clusters present on a plant increased subsequent oviposition on this plant. All other models could not describe the distribution of egg cluster counts for either of the two host species. 6. The results suggest that females seek out attractive hosts actively and the attractiveness of a plant increases with its egg cluster load. This behaviour creates a frequency distribution of egg clusters per host that depends only on beetle density but not on plant density. This conclusion has important implications for modelling insect-plant interactions.     

Season‐long mating disruption of citrus leafminer,Phyllocnistis citrella Stainton,with an emulsified wax formulation of pheromone

The citrus leafminer, Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae), is a major worldwide pest of citrus. Larval feeding by this insect facilitates proliferation of citrus bacterial canker, Xanthomonas axonopodis pv. citri. Herein, we describe a season‐long disruption trial of P. citrella with a newly developed, emulsified wax dispenser of pheromone (SPLAT‐CLM^TM). A formulation containing a 3 : 1 blend of (Z,Z,E)‐7,11,13‐hexadecatrienal:(Z,Z)‐7,11‐hexadecadienal at a 0.2% loading rate of active ingredient by weight and deployed twice per season (24 weeks total) at 490 g of formulation/ha caused season‐long disruption of male moth catch in pheromone traps as well as reduced leaf infestation. Analysis of pheromone release from dispensers by gas chromatography revealed that effective disruption of P. citrella occurred at a deployment rate of 126 μg of (Z,Z,E)‐7,11,13‐hexadecatrienal/ha/h. Direct observation of moth behaviour in the field suggested that disruption by this formulation occurred by a non‐competitive mechanism. A formulation of the 3 : 1 attractive blend at a 0.02% pheromone loading rate caused only 2–6 weeks of disruption per deployment and did not reduce leaf infestation during mid and end of the season evaluations. A formulation containing 0.2% of (Z,Z)‐7,11‐hexadecadienal alone and deployed at 490 g/ha caused 6–7 weeks of moth disruption to pheromone traps and did not prevent leaf infestation, while an identical formulation loaded with 0.02% (w/w) of (Z,Z)‐7,11‐hexadecadienal alone had no effect on P. citrella orientation to pheromone traps. The SPLAT formulation evaluated herein appears to be an excellent release device for (Z,Z,E)‐7,11,13‐hexadecatrienal given that approximately 100 days of steady release occurred following an initial brief (ca. 7 days) burst of higher release. The advantages of SPLAT as a formulation for P. citrella disruption include low cost of manufacturing, biodegradable and weather resistant characteristics, and flowability allowing machine application. Mating disruption should be an effective alternative to insecticides for management of P. citrella and may reduce the incidence of citrus canker.     

Resistance of fast- and slow-growing subalpine fir to pheromone-induced attack by western balsam bark beetle (Coleoptera: Scolytinae)

Abstract 1 We investigated the resistance of fast‐ and slow‐growing subalpine fir to pheromone‐induced attack by western balsam bark beetle at two sites in the interior of British Columbia, Canada. 2 Attack success by the beetle and subsequent tree mortality were higher in slow‐growing trees than in fast‐growing trees. 3 Fast‐growing trees were more likely to produce secondary resin, and in greater quantities, than slow‐growing trees after attack. 4 Host vigour (indicated by recent radial growth) was positively related to the induced defense response and resistance of subalpine fir to bark beetle attack. These results are discussed in the context of plant defense and plant–herbivore interaction hypotheses. 5 Given the preference of western balsam bark beetle for weakened trees, as well as the reduced defenses and increased mortality rates in less vigorous trees, effective management tactics for this beetle may include strategies that increase the growth and vigour of its subalpine fir host.     

Attractant effect of citral on Platypus koryoensis (Coleoptera: Curculionidae)

An environmentally friendly control strategy for the ambrosia beetle Platypus koryoensis (Murayama), a vector of oak wilt disease, was investigated using the citral as a pheromone component. To develop an effective attractant trap, a semi‐field experiment (Yeoju) and a field experiment (Gunpo) were performed. Different ratios of four different attractant mixtures (ethanol 100 %, ethanol + citral (90:10, 95:5), ethanol + citral + hexane extract of Quercus mongolica (90:5:5)) were investigated to determine their effects on P. koryoensis. The 95:5 mixture of ethanol + citral had the greatest attractant effect on P. koryoensis. However, other ambrosia beetle species did not exhibit a specific response to citral. Our findings indicate that the pheromone citral was effective at attracting P. koryoensis and that certain concentrations of citral could be used to develop an effective and environmentally friendly control agent.     

Primer effects of a brood pheromone on honeybee behavioural development

Primer pheromones are thought to act in a variety of vertebrates and invertebrates but only a few have been chemically identified. We report that a blend of ten fatty-acid esters found on the cuticles of honeybee larvae, already known as a kairomone, releaser pheromone and primer pheromone, also act as a primer pheromone in the regulation of division of labour among adult workers. Bees in colonies receiving brood pheromone initiated foraging at significantly older ages than did bees in control colonies in five out of five trials. Laboratory and additional field tests also showed that exposure to brood pheromone significantly depressed blood titres of juvenile hormone. Brood pheromone exerted more consistent effects on age at first foraging than on juvenile hormone, suggesting that the primer effects of this pheromone may occur via other, unknown, mechanisms besides juvenile hormone. These results bring the number of social factors known to influence honeybee division of labour to three: worker-worker interactions, queen mandibular pheromone and brood pheromone.     

A novel pheromone dispenser for mating disruption of the leafminer Phyllocnistis citrella (Lepidoptera: Gracillariidae)

The sex pheromone of the leafminer Phyllocnistis citrella Stainton (Lepidoptera: Gracillariidae) was deployed in a Florida citrus (Citrus spp.) grove by using a novel deployment device (IFM-413) containing SPLAT, a flowable formulation of an emulsified wax compound designed to provide slow release of semiochemicals. The device consisted of two disks connected by string. Each disk was loaded with 1 g of SPLAT containing either 0.15% (Z,Z,E)-7,11,13-hexadecatrienal (triene) or 2% (Z,Z)-7,11-hexadecadienal (diene). The devices were deployed using a two-dimensional multivariate design to determine the optimal rate of pheromone per unit area and degree of aggregation of the deployment devices (number of trees treated per unit area). The IFM-413 device proved effective at becoming securely entangled in tree branches. Furthermore, the devices effectively delivered pheromone-loaded SPLAT that resulted in disruption of trap catch of male P. citrella. Response surfaces showed a quadratic response of trap catch disruption to both total amount of pheromone per unit area and the degree of aggregation of the deployed devices (number of treated trees per unit area). The response surfaces for 0.15% triene or 2.0% diene were similar. The diene produced an effect similar to that of the triene at approximately 13 times the rate of the triene. The greatest disruption of trap catch occurred when the number of treated trees per unit area was greatest (no aggregation of deployment devices). Manufacturing, packaging, and mechanical deployment of the devices remain to be investigated.     

Mass attack of trees by Ips typographus induced by sex-specific pheromone: a model of attack dynamics

A model of attack dynamics on a single tree under mass attack by the spruce bark beetle Ips typographus is developed from earlier studies of pheromone production and response of the flying population to the pheromone signal. The model is explored for different numbers and sex-ratios of pioneer beetles and responding beetles. Both numbers of pioneers and responders were important for the dynamics, while the sex-ratio of pioneers was less important. A decreased proportion of males among the responding beetles was important for the result of the model. In the early stage of the attack the model predictions of both numbers and sex-ratio of beetles landing were similar to examples from naturally mass-attacked trees, although the attack proceeded more slowly than in the field. Several aspects of the attack dynamics, such as density-regulation of the number of attacks on the tree, as observed in the field, were not predicted by the model. Gaps in the knowledge of the chemical ecology of the tree-insect relation, such as qualitative change in odours from the tree, spacing mechanisms on the bark surface, and mortality due to host resistence, are pointed out with help of the model.     

Differing behavioral and endocrinological effects of two female sex pheromones on male goldfish

Ovulatory female goldfish sequentially release at least two sex pheromones: 17 alpha, 20 beta-dihydroxy-4-pregnen-3-one (17,20 beta P) and a mixture of F prostaglandins (PGFs). This study sought to determine whether these pheromones have different endocrinological and behavioral actions and whether the PGF pheromone, which is released by spawning females, is responsible for increasing the gonadotropin (GtH) and milt (sperm and seminal fluid) levels of spawning males. Grouped and isolated males were exposed to combinations of these pheromones, food odor, and spawning and nonspawning females. 17,20 beta P stimulated GtH increases in both grouped and isolated males but had only minor effects on behavior; because its principal function appears to be physiological it may be considered a "primer" pheromone. In contrast, exposue to the PGFs elicited large increases in sexual behavior but increased GtH only when fish were exposed as groups; this pheromone's principal action appears to be behavioral and it should be considered a "releaser" pheromone. Although males had increased GtH and milt levels after 1 hr of spawning, males allowed to interact with nonspawning females also had elevated GtH; thus, behavioral interactions appear capable of elevating GtH in the absence of either pheromone. The existence of an independent behavioral mechanism which stimulates GtH was supported by the fact that males exposed to 17,20 beta P while spawning had GtH levels much greater than males exposed to only one of these stimuli.     

SPLAT-OrB reveals competitive attraction as a mechanism of mating disruption in oriental beetle (Coleoptera: Scarabaeidae)

This study compared the efficacy of SPLAT-OrB, a new pheromone formulation for oriental beetle mating disruption that can be mechanically applied, with hand-applied plastic dispensers in commercial blueberry fields. Both formulations were tested at 2.5 and 5 g of the major sex pheromone component (Z)-7-tetradecen-2-one per hectare, and evaluated by measuring trap shut-down, mating success of caged females, and the number of grubs in sentinel blueberry pots baited with tethered females. All pheromone-treated plots had fewer male captures in traps and lower mating success of caged females compared with untreated plots. SPLAT-OrB, and plastic dispensers at the higher rate, also reduced the number of grubs in sentinel pots. To understand the mechanism of mating disruption in oriental beetle, males were observed approaching the pheromone sources in disrupted plots. In addition, male oriental beetle captures were quantified in plots treated with varying SPLAT-OrB dollop densities per ha. Consistent with predictions for competitive attraction, field observations revealed males approaching the pheromone source and male captures decreasing concavely with increasing dollop density. In a mark-release-recapture study, male oriental beetles responded to SPLAT-OrB dollops and plastic dispensers at least 60 m from the source. Additionally, SPLAT-OrB emitted pheromone that was attractive to male oriental beetles for >5 wk; however, emission rates and attraction dropped rapidly during the first 3-4 wk. This study demonstrates the feasibility of using SPLAT-OrB for oriental beetle mating disruption as an alternative to hand-applied plastic dispensers, and conclusively reveals that a principal mechanism is the competitive attraction of males.     

A releaser pheromone that attracts males in the urine of mature female masu salmon

A releaser pheromone of masu salmon Oncorhynchus masou was investigated using Y-maze behavioural experiments. During the reproductive season, urine of mature females contains a releaser pheromone which acts as a sex attractant for spermiating mature male parr. The releaser pheromone in mature female urine is one or more low molecular weight substances (less than 10 000) soluble in ether under basic conditions. The attractant was not present in either the coelomic fluid of ovulated females nor in neutral or acidic extracts of female urine which contain free steroids and F-type prostaglandins, respectively.     

Comparative analyses of the pigment-aggregating and -dispersing actions of MCH on fish chromatophores

In melanophores of the peppered catfish and the Nile tilapia, melanin-concentrating hormone (MCH) at low doses (<1 μM) induced pigment aggregation, and the aggregated state was maintained in the presence of MCH. However, at higher MCH concentrations (such as 1 and 10 μM), pigment aggregation was immediately followed by some re-dispersion, even in the continued presence of MCH, which led to an apparent decrease in aggregation. This pigment-dispersing activity at higher concentrations of MCH required extracellular Ca²⁺ ions. By contrast, medaka melanophores responded to MCH only by pigment aggregation, even at the highest concentration employed (10 μM). Since it is known that medaka melanophores possess specific receptors for α-melanophore-stimulating hormone (α-MSH), the possibility that interaction between MSH receptors and MCH at high doses in the presence of Ca²⁺ might cause pigment dispersion is ruled out. Cyclic MCH analogs, MCH (1–14) and MCH (5–17), failed to induce pigment dispersion, whereas they induced aggregation of melanin granules. These results suggest that another type of MCH receptor that mediates pigment dispersion is present in catfish and tilapia melanophores, and that intact MCH may be the only molecule that can bind to these receptors. Determinations of cAMP content in melanophores, which were isolated from the skin of three fish species and treated with 10 nM or 10 μM MCH, indicate that MCH receptors mediating aggregation may be coupled with Gi protein, whereas MCH receptors that mediate dispersion may be linked to Gs. The response of erythrophores, xanthophores and leucophores to MCH at various concentrations was also examined, and the results suggest that the distribution patterns of the two types of MCH receptors may differ among fish species and among types of chromatophore in the same fish.