Alien insect dispersal mediated by the global movement of commodities.

Globalization and economic growth are recognized as key drivers of biological invasions. Alien species have become a feature of almost every biological community worldwide, and rates of new introductions continue to rise as the movement of people and goods accelerates. Insects are among the most numerous and problematic alien organisms, and are mainly introduced unintentionally with imported cargo or arriving passengers. However, the processes occurring prior to insect introductions remain poorly understood. We used a unique dataset of 1,902,392 border interception records from inspections at air, land, and maritime ports in Australia, New Zealand, Europe, Japan, USA, and Canada to identify key commodities associated with insect movement through trade and travel. In total, 8939 species were intercepted, and commodity association data were available for 1242 species recorded between 1960 and 2019. We used rarefaction and extrapolation methods to estimate the total species richness and diversity associated with different commodity types. Plant and wood products were the main commodities associated with insect movement across cargo, passenger baggage, and international mail. Furthermore, certain species were mainly associated with specific commodities within these, and other broad categories. More closely related species tended to share similar commodity associations, but this occurred largely at the genus level rather than within orders or families. These similarities within genera can potentially inform pathway management of new alien species. Combining interception records across regions provides a unique window into the unintentional movement of insects, and provides valuable information on establishment risks associated with different commodity types and pathways.

Improved performance of the eastern spruce budworm on black spruce as warming temperatures disrupt phenological defences.

Phenological shifts, induced by global warming, can lead to mismatch between closely interacting species. The eastern spruce budworm, Choristoneura fumiferana, an important outbreaking insect defoliator in North America, mainly feeds on balsam fir, Abies balsamea, which has historically been well synchronized with the insect. But as climate change pushes the northern range limit of the budworm further north into the boreal forest, the highly valuable black spruce, Picea mariana, historically protected against the budworm by its late budburst phenology, is suffering increased defoliation during the current outbreak. We tested the hypothesis that rising temperatures can lead, not to a mismatch, but to an improved match between the budworm and black spruce through differential phenological advancement. For 3 years, eastern spruce budworm larvae were reared from instar 2 to pupae, on both black spruce and balsam fir, in a temperature free-air controlled enhancement experiment (T-FACE) consisting in 24 field plots, half of which were heated at +2°C from March to October. Our results show that every year, larval development was faster on heated trees and pupation was earlier than on unheated trees. Bud development was also accelerated in heated trees of both species. However, there was no difference in mass between pupae that developed at +2°C and controls at the end of the season. Finally, we found no difference either in development rate or pupal mass between larvae reared on black spruce and those reared on balsam fir. This suggests that under higher temperature regimes, eastern spruce budworm will be as successful on black spruce as on balsam fir, as black spruce budburst becomes better synchronized with the insect's emergence from diapause. This could lead to critical changes in outbreak dynamics and severity with important ecological state shifts at the landscape level.

Worldwide border interceptions provide a window into human-mediated global insect movement.

As part of national biosecurity programs, cargo imports, passenger baggage, and international mail are inspected at ports of entry to verify compliance with phytosanitary regulations and to intercept potentially damaging nonnative species to prevent their introduction. Detection of organisms during inspections may also provide crucial information about the species composition and relative arrival rates in invasion pathways that can inform the implementation of other biosecurity practices such as quarantines and surveillance. In most regions, insects are the main taxonomic group encountered during inspections. We gathered insect interception data from nine world regions collected from 1995 to 2019 to compare the composition of species arriving at ports in these regions. Collectively, 8,716 insect species were intercepted in these regions over the last 25 yr, with the combined international data set comprising 1,899,573 interception events, of which 863,972 were identified to species level. Rarefaction analysis indicated that interceptions comprise only a small fraction of species present in invasion pathways. Despite differences in inspection methodologies, as well as differences in the composition of import source regions and imported commodities, we found strong positive correlations in species interception frequencies between regions, particularly within the Hemiptera and Thysanoptera. There were also significant differences in species frequencies among insects intercepted in different regions. Nevertheless, integrating interception data among multiple regions would be valuable for estimating invasion risks for insect species with high likelihoods of introduction as well as for identifying rare but potentially damaging species.

Evidence for Semiochemical Divergence Between Sibling Bark Beetle Species: Dendroctonus brevicomis and Dendroctonus barberi.

We investigated geographic variation in the semiochemistry of major disturbance agents of western North American pine forests, Dendroctonus brevicomis Le Conte and Dendroctonus barberi Hopkins (Coleoptera: Curculionidae: Scolytinae), species separated by the Great Basin in the USA that until recently were synonymous. At 15 sites in the western USA and northern Mexico, beetle populations were examined to determine (1) pheromone production by solitary, mining females, (2) male electroantennogram amplitudes in response to known semiochemicals for the genus, or (3) relative attractiveness of two female-produced pheromone components (endo- and exo-brevicomin) and two host odors (alpha-pinene and myrcene) to beetles in the field. Compared to female beetles collected east of the Great Basin (D. barberi), western females (D. brevicomis) produced a consistently higher proportion of, and male antenna were correspondingly more sensitive to, the exo- than the endo-isomer of brevicomin. With the exception of one sampling location (where no preference was observed), beetles west of the Great Basin were more attracted to exo- than endo- brevicomin trap lures, whereas eastern beetles displayed the reverse preference. In contrast, there was not a consistent difference between these populations regarding relative attraction or olfactory response to myrcene or alpha-pinene, although some geographic variability was evident. These data show that the semiochemical systems of D. brevicomis and D. barberi have diverged and corroborate genetic and morphological evidence that they are distinct, allopatric species.

Western Pine Beetle Populations in Arizona and California Differ in the Composition of Their Aggregation Pheromones.

We compared pheromone production and response for populations of western pine beetle, Dendroctonus brevicomis LeConte, from sites in northern Arizona and northern California. Volatiles were collected from individuals of both sexes that had mined as a pair in a Pinus ponderosa log for 1 d, and they were subsequently analyzed by gas chromatography coupled to mass-spectrometry. Principal component analysis of quantities of Dendroctonus pheromone components indicated strong site-associated clustering of blend composition for females but not males. Much of the clustering in females evidently was due to differences in the production of endo- and exo-brevicomin, which occurred in average ratios of 0.1:1 and 19:1 for populations in the California and Arizona sites, respectively. In the California site, exo- was better than endo-brevicomin in enhancing trap catches of both sexes to lures containing the host-tree odor α-pinene and the male-produced aggregation pheromone component frontalin. In an identical test in the Arizona site, endo- was a better adjuvant than exo-brevicomin for male attraction, whereas females did not show a significant preference. At neither location were the isomers antagonistic to one another in activity. Thus, one aggregation pheromone has apparently diverged between these populations, concurrent with published evidence that D. brevicomis on either side of the Great Basin are genetically distinct and are possibly different species. Furthermore, production of and response to the isomers of brevicomin by flying Dendroctonus frontalis Zimmermann in the Arizona site were similar to those of sympatric D. brevicomis. This interspecific signal overlap is likely sustainable since joint species mass-attacks may assist both species in overcoming host defenses, thereby increasing host availability.

An enhanced lure for eastern populations of the North American spruce beetle, Dendroctonus rufipennis Kirby (Coleoptera: Curculionidae).

Regional variation in pheromone production and response has practical implications for the use of semiochemical lures to monitor and control bark beetle populations. We tested 4 lure formulations including 2 new formulations that reflect the pheromone production profiles of western and eastern populations of spruce beetles, Dendroctonus rufipennis Kirby (Coleoptera: Curculionidae), as well as 2 commercially available formulations (current Rocky Mountain lure and current Atlantic lure), in 2 locations in New Brunswick, Canada. In 2 separate years, the new eastern lure containing seudenol, MCOL, and spruce terpenes captured 4 times (2021) and 11 times (2022) more spruce beetles than the current Atlantic lure that consisted of frontalin, seudenol, and spruce terpenes. In 2021, we also captured more eastern larch beetles, Dendroctonus simplex LeConte (Coleoptera: Curculionidae), with the new eastern lure, whereas in 2022, we captured the most D. simplex with the current Atlantic lure, suggesting that more research is needed on D. simplex pheromone production and response across its range. The bark beetle predator, Thanasimus dubius (Fabr.; Coleoptera: Cleridae), did not respond well to the new eastern blend that lacks frontalin, suggesting that response to frontalin is important in finding prey and might be conserved in predator populations. The reduced trap catch of T. dubius to the enhanced lure is beneficial because it does not inhibit natural population control by removing predators from the community. Our study reveals an improved trap lure for eastern populations of spruce beetles and highlights gaps and research needs in bark beetle pheromone ecology.

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.

Defoliation-induced changes in foliage quality may trigger broad-scale insect outbreaks.

Top-down effects, like predation, are drivers of insect outbreaks, but bottom-up effects, like host nutritional quality, also influence outbreaks and could in turn be altered by insect-caused defoliation. We evaluated the prediction that herbivory leads to a positive feedback on outbreak severity as nutrient concentration in plant tissues increases through improved soil nutrient availability from frass and litter deposition. Over seven years of a spruce budworm outbreak, we quantified litter nutrient fluxes, soil nitrogen availability, and host tree foliar nutrient status along a forest susceptibility gradient. As the outbreak progressed, both soil nutrient fluxes and availability increased which, in turn, improved foliage quality in surviving host trees. This is consistent with boosted insect fitness and increased population density and defoliation as outbreaks grow. Our results suggest that a positive bottom-up feedback to forest ecosystems from defoliation may result in conditions favorable to self-amplifying population dynamics in insect herbivores that can contribute to driving broad-scale outbreaks.

Host selection and feeding preference of Agrilus planipennis (Coleoptera: Buprestidae) on ash (Fraxinus spp.).

We studied the host selection behavior and feeding preference of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). A. planipennis is an exotic forest insect pest native to Asia that was discovered in North America in 2002 and is causing widespread mortality of ash trees (Fraxinus spp.) in southeast Michigan and surrounding states. We compared host selection and feeding behavior on different species of ash including Manchurian ash (F. mandshurica Rupr.), green ash (F. pennsylvanica Marsh), white ash (F. americana L.), black ash (F. nigra Marsh), blue ash (F. quadrangulata Michx.), and European ash (F. excelsior L). Manchurian ash is native to Asia, whereas the other species (native to North America or Europe) represent novel hosts for A. planipennis. Beetles distributed themselves more frequently and fed to a greater extent on green, black, and white ash compared with blue, European, and Manchurian ash. Although beetles consumed every ash species offered to them, Manchurian ash and blue ash were least preferred in feeding bioassays. When we analyzed the volatile content of intact and girdled ash for quantitative variation in 11 compounds that elicited antennal activity, we found that the overall volatile profiles of the six ash species differed significantly in their relative amounts of antennally active compounds. Green ash has lower relative amounts of volatiles compared with Manchurian ash, which might render it more attractive and less resistant to A. planipennis. Lower tolerance and resistance of green ash might make it more susceptible to mortality compared with Manchurian ash, which coevolved with the beetle in its native range. Repellent odors, potential antifeedants, and genes for resistance in Manchurian ash could be explored for methods to manage A. planipennis populations.

Phenological synchrony between eastern spruce budworm and its host trees increases with warmer temperatures in the boreal forest.

Climate change is predicted to alter relationships between trophic levels by changing the phenology of interacting species. We tested whether synchrony between two critical phenological events, budburst of host species and larval emergence from diapause of eastern spruce budworm, increased at warmer temperatures in the boreal forest in northeastern Canada. Budburst was up to 4.6 ± 0.7 days earlier in balsam fir and up to 2.8 ± 0.8 days earlier in black spruce per degree increase in temperature, in naturally occurring microclimates. Larval emergence from diapause did not exhibit a similar response. Instead, larvae emerged once average ambient temperatures reached 10°C, regardless of differences in microclimate. Phenological synchrony increased with warmer microclimates, tightening the relationship between spruce budworm and its host species. Synchrony increased by up to 4.5 ± 0.7 days for balsam fir and up to 2.8 ± 0.8 days for black spruce per degree increase in temperature. Under a warmer climate, defoliation could potentially begin earlier in the season, in which case, damage on the primary host, balsam fir may increase. Black spruce, which escapes severe herbivory because of a 2-week delay in budburst, would become more suitable as a resource for the spruce budworm. The northern boreal forest could become more vulnerable to outbreaks in the future.

High individual variation in pheromone production by tree-killing bark beetles (Coleoptera: Curculionidae: Scolytinae).

Aggregation via pheromone signalling is essential for tree-killing bark beetles to overcome tree defenses and reproduce within hosts. Pheromone production is a trait that is linked to fitness, so high individual variation is paradoxical. One explanation is that the technique of measuring static pheromone pools overestimates true variation among individuals. An alternative hypothesis is that aggregation behaviour dilutes the contribution of individuals to the trait under selection and reduces the efficacy of natural selection on pheromone production by individuals. We compared pheromone measurements from traditional hindgut extractions of female southern pine beetles with those obtained by aerating individuals till they died. Aerations showed greater total pheromone production than hindgut extractions, but coefficients of variation (CV) remained high (60-182%) regardless of collection technique. This leaves the puzzle of high variation unresolved. A novel but simple explanation emerges from considering bark beetle aggregation behaviour. The phenotype visible to natural selection is the collective pheromone plume from hundreds of colonisers. The influence of a single beetle on this plume is enhanced by high variation among individuals but constrained by large group sizes. We estimated the average contribution of an individual to the pheromone plume across a range of aggregation sizes and showed that large aggregation sizes typical in mass attacks limit the potential of natural selection because each individual has so little effect on the overall plume. Genetic variation in pheromone production could accumulate via mutation and recombination, despite strong effects of the pheromone plume on the fitness of individuals within the aggregation. Thus, aggregation behaviour, by limiting the efficacy of natural selection, can allow the persistence of extreme phenotypes in nature.

Attraction of the southern pine beetle, Dendroctonus frontalis, to pheromone components of the western pine beetle, Dendroctonus brevicomis (Coleoptera: Curculionidae: Scolytinae), in an allopatric zone.

Subtle differences in pheromone components of sympatric species should be attractive only to the producing species and unattractive or repellent to the nonproducing species, and thereby maintain reproductive isolation and reduce competition between species. Bark beetles Dendroctonus brevicomis and D. frontalis (Coleoptera: Curculionidae) are known to have common pheromone components, except for exo-brevicomin, which is produced by D. brevicomis. We predicted that D. frontalis would not respond to exo-brevicomin outside of the zone of sympatry with D. brevicomis. We conducted a field experiment to determine the effect of exo-brevicomin on attraction of D. frontalis and associated species in Mississippi. We determined whether D. frontalis pheromone production differed inside and outside the sympatric zone and compared the pheromone profiles with D. brevicomis within the sympatric zone. Trapping studies revealed that D. frontalis can perceive and respond positively to exo-brevicomin, an aggregation pheromone of a sympatric congener (D. brevicomis), at locations hundreds of kilometers outside the sympatric zone. Qualitative pheromone profiles showed that both species emit similar pheromone components: frontalin, endo-brevicomin, exo-brevicomin, trans-verbenol, verbenone, and myrtenol. Although not previously reported, D. frontalis males from Arizona produced exo-brevicomin. The predator Thanasimus dubius did not discriminate traps baited with exo-brevicomin and was most attracted to traps with frontalin. Hylastes beetles were significantly attracted to traps baited with exo-brevicomin in combination with other compounds. Our results raise new practical and evolutionary questions on the role of exo-brevicomin in the behavioral ecology of D. frontalis. The addition of exo-brevicomin to the current lure might increase the efficiency of trapping programs in the southeastern United States.

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.

Impact of an Invasive Longhorned Beetle, Tetropium fuscum (Coleoptera: Cerambycidae), on Community Structure of Subcortical and Wood-Associated Insects in Eastern Canada.

Tetropium fuscum (Fabricius) (Coleoptera: Cerambycidae), a phloem-feeding and wood-boring beetle introduced from Eurasia, attacks spruce in eastern Canada alongside its native congener Tetropium cinnamopterum Kirby. We reared phloem- and wood-feeding insects (and their predators) from bolts of red and Norway spruce (Picea rubens and Picea abies) in Nova Scotia, comparing insect communities between bolts with added eggs of T. fuscum or T. cinnamopterum and bolts without added Tetropium (controls). We tested for impacts of each Tetropium on insect community structure (Simpson's diversity, richness, and evenness). We also asked whether, consistent with Darwin's Naturalization Hypothesis, Tetropium spp. would have greater impacts on emergence of its closer relatives (which might be most likely to compete and/or share natural enemies). Addition of Tetropium eggs (either species) to bolts lowered insect diversity in both host trees. Both richness and evenness components of diversity were always lower in +Tetropium treatments, although different components reached statistical significance in different Tetropium species × host combinations. Addition of Tetropium spp. significantly reduced emergence of some species: Evodinus monticola (Randall) (Coleoptera: Cerambycidae) was reduced by T. fuscum on both hosts and by T. cinnamopterum on Norway spruce; Hylobius congener Dalla Torre, Schenkling, and Marshall was reduced by T. fuscum on red spruce; and Xylophagus sp. (Diptera: Xylophagidae) was reduced by T. cinnamopterum on Norway spruce. However, there was no relationship between Tetropium's impact on a community member and their phylogenetic relatedness, and the overall impacts of Tetropium presence were not very different between T. fuscum and T. cinnamopterum.

Paradigms in Eastern Spruce Budworm (Lepidoptera: Tortricidae) Population Ecology: A Century of Debate.

Three main hypotheses have been postulated over the past century to explain the outbreaking population dynamics of eastern spruce budworm, Choristoneura fumiferana (Clemens). The Silviculture Hypothesis first arose in the 1920s, with the idea that outbreaks were driven by forestry practices favoring susceptible softwood species. In the 1960s, it was proposed that populations were governed by Multiple Equilibria, with warm weather conditions releasing low-density populations from the regulatory control of natural enemies. Dispersal from outbreak foci, or "epicenters," was seen as causing widespread outbreaks that eventually collapsed following resource depletion. However, in the 1980s, following the re-analysis of data from the 1940s outbreak in New Brunswick, this interpretation was challenged. The alternative Oscillatory Hypothesis proposed that budworm population dynamics were governed by a second-order density-dependent process, with oscillations being driven by natural enemy-victim interactions. Under this hypothesis, weather and resource availability contribute to secondary fluctuations around the main oscillation, and weather and moth dispersal serve to synchronize population cycles regionally. Intensive, independent population studies during the peak and declining phases of the 1980s outbreak supported the principal tenet of the Oscillatory Hypothesis, but concluded that host plant quality played a more important role than this hypothesis proposed. More recent research on the early phase of spruce budworm cycles suggests that mate-finding and natural-enemy-driven Allee effects in low-density populations might be overcome by immigration of moths, which can facilitate the onset of outbreaks. Even more recent research has supported components of all three hypotheses attempting to explain spruce budworm dynamics. In the midst of a new rising outbreak (2006-present), we discuss the evolution of debates surrounding these hypotheses from a historic perspective, examine gaps in current knowledge, and suggest avenues for future research (e.g., intensive studies on low-density populations) to better understand and manage spruce budworm populations.

Displacement of Tetropium cinnamopterum (Coleoptera: Cerambycidae) by Its Invasive Congener Tetropium fuscum.

We examined the native community of insects interacting with an invasive species, Tetropium fuscum (F.) (Coleoptera: Cerambycidae), in its new range to explore reasons for the invader's relatively slow spread. Tetropium fuscum is a European spruce borer established in Nova Scotia since at least 1990, but it has spread only about 125 km from its site of introduction. We compared the densities of Tetropium spp., their known parasitoids, and the community of wood-boring insects at sites located within the invasion zone in Nova Scotia versus well outside this zone, in New Brunswick, Canada. Using red spruce trees stressed by girdling or felling, we tested whether: 1) T. fuscum had altered the native wood-boring community; 2) T. fuscum displaced a native congener, Tetropium cinnamopterum (Kirby); and 3) parasitism rates of Tetropium spp. differed between the invaded and noninvaded zones. Both Tetropium spp. and their parasitoid wasps emerged exclusively from felled trees as opposed to girdled trees. We found no difference in community diversity inside versus outside the invasion zone. The combined densities of both Tetropium spp. and their overall parasitism rates also did not differ between zones, but T. cinnamopterum density was significantly greater outside the invasion zone, suggesting T. fuscum may displace the native congener where they are sympatric. Our results suggest that the native and invasive Tetropium spp. act as a single functional species in the invasion zone. We speculate that natural control agents (predators, parasitoids, and competitors) might be limiting the rate of spread of T. fuscum.

The Role of Multimodal Signals in Species Recognition Between Tree-Killing Bark Beetles in a Narrow Sympatric Zone.

When related species coexist, selection pressure should favor evolution of species recognition mechanisms to prevent interspecific pairing and wasteful reproductive encounters. We investigated the potential role of pheromone and acoustic signals in species recognition between two species of tree-killing bark beetles, the southern pine beetle, Dendroctonus frontalis Zimmermann, and the western pine beetle, Dendroctonus brevicomis LeConte, in a narrow zone of sympatry, using reciprocal pairing experiments. Given the choice of adjacent con- or heterospecific female gallery entrance in a log, at least 85% of walking males chose the entrance of the conspecific, and half the males that initially entered heterospecific galleries re-emerged and entered the conspecific gallery within 15 min. Waveform analysis of female acoustic "chirps" indicated interspecific differences in chirp timing. Males may use information from female acoustic signals to decide whether to enter or remain in the gallery. Individuals in forced heterospecific pairings (produced by confinement of a heterospecific male within the female entrance) did not differ in pheromone production from individuals of conspecific pairs. However, due to the absence of the right species of male, galleries with heterospecific pairs released an abnormal pheromone blend that lacked at least one key component of the aggregation pheromone of either species. The complete aggregation pheromone (i.e., the pheromone blend from entrances with pairs) does not appear to deter interspecific encounters or confer premating reproductive isolation per se; however, it may confer selective pressure for the maintenance of other reproductive isolation mechanisms.

Evidence that (+)-endo-brevicomin is a male-produced component of the Southern pine beetle aggregation pheromone.

Previous research indicated that the aggregation pheromone of the southern pine beetle, Dendroctonus frontalis, is produced only by females, the sex that initiates attacks. We provide evidence indicating that secondarily arriving males augment mass aggregation by releasing the attractive synergist (+)-endo-brevicomin. Healthy pines artificially infested with both sexes of D. frontalis were significantly more attractive to conspecifics than trees infested solely with females. Coupled gas chromatography-electroantennographic detection (GC-EAD) analyses of volatiles isolated from male beetles revealed substantially greater olfactory sensitivity by D. frontalis to endo-brevicomin than to any other component. The threshold of detection of both sexes for (+)-endo-brevicomin was four orders of magnitude lower than for its antipode and at least one order of magnitude lower than for either enantiomer of frontalin, the major female-produced aggregation pheromone component. Pairing with a female in a gallery stimulated individual male beetles to produce hundreds of nanograms of (+)-endo-brevicomin. (+)-endo-Brevicomin was detected in a small percentage of female D. frontalis, whereas (-)-endo-brevicomin was never detected in either sex. In field trapping bioassays, we confirmed that (+)-endo-brevicomin is a potent synergist for attractive combinations of frontalin and pine turpentine. However, (+)-endo-brevicomin failed to attract D. frontalis either when presented alone or in combination with turpentine. We postulate that mass colonization of host trees by D. frontalis is mediated by distinct semiochemicals from both sexes rather than females alone. Our discovery of a key aggregation pheromone component in such an apparently well-studied species implies that the pheromone models of other bark beetles could benefit from systematic reexamination using newer technologies. Additionally, baits fortified with (+)-endo-brevicomin may enhance pest management strategies that exploit attractants for D. frontalis.

Fitness consequences of pheromone production and host selection strategies in a tree-killing bark beetle (Coleoptera: Curculionidae: Scolytinae).

Timing of arrival at a resource often determines an individual's reproductive success. Tree-killing bark beetles can reproduce in healthy trees by attacking in adequate numbers to overcome host defences that could otherwise be lethal. This process is mediated by aggregation and antiaggregation pheromones. Beetles that arrive early in such a "mass attack" must contend with undiminished tree defences, and produce enough pheromones to attract more beetles, but have a head start on gallery construction and egg-laying. Beetles that arrive late may be impeded by competition and diminishing availability of phloem, but should experience fewer costs associated with pheromone production and battling tree defences. We investigated relationships between timing of arrival, body size, pheromone production and fitness in the southern pine beetle, Dendroctonus frontalis. In field experiments, we captured beetles that arrived early (pioneers) and late on slash pine trees, Pinus elliottii, and measured pheromone amounts in their hindguts. We marked gallery entrances of beetles as they landed on a tree and measured their reproductive success after the attack terminated. We found no difference in body size or pheromone amounts between early and late arrivers. Most beetles arrived at the middle of the attack sequence, and excavated longer galleries per day than early arrivers. The number of offspring produced per day by beetles that established galleries midway through mass attack was higher than those that arrived early or very late in the sequence. Our results suggest that beetles do not exhibit adaptive phenotypic plasticity in pre-landing pheromone production, depending on the extent of previous colonisation of a host. Rather, it appears that stabilising selection favours beetles that attack in the middle of the sequence, and contributes to attack synchrony. Synchronous attack on trees is essential before population booms characteristic of tree-killing bark beetles can occur in nature.