Manuka Oil Lures Research Articles

Utility of essential oils for development of host-based lures for Xyleborus glabratus (Coleoptera: Curculionidae: Scolytinae), vector of laurel wilt

AbstractRedbay ambrosia beetle, Xyleborus glabratus, is native to Southeast Asia, but subsequent to introduction in Georgia in 2002, it has become a serious invasive pest in the USA, now established in nine southeastern states. Females vector Raffaelea lauricola, the fungus that causes laurel wilt, a lethal vascular disease of trees in the family Lauraceae. Laurel wilt has caused extensive mortality in native Persea species, including redbay (P. borbonia), swampbay (P. palustris), and silkbay (P. humilis). Avocado (P. americana) is now impacted in Florida, and with continued spread, laurel wilt has potential to affect avocado and native Lauraceae in California, Mexico, and throughout the American tropics. Effective lures for detection and control of X. glabratus are critical to slow the spread of laurel wilt. No pheromones are known for this species; primary attractants are volatile terpenoids emitted from host Lauraceae. This report provides a concise summary of the chemical ecology of X. glabratus, highlighting research to identify kairomones used by females for host location. It summarizes development of essential oil lures for pest detection, including discussions of the initial use of phoebe and manuka oil lures, the current cubeb oil lure, and a newly-developed distilled oil lure enriched in (-)-α-copaene.

Attraction of Redbay Ambrosia Beetle, Xyleborus Glabratus, To Leaf Volatiles of its Host Plants in North America.

The redbay ambrosia beetle, Xyleborus glabratus, is an important pest of redbay (Persea borbonia) and swamp bay (P. palustris) trees in forests of the southeastern USA. It is also a threat to commercially grown avocado. The beetle is attracted to host wood volatiles, particularly sesquiterpenes. Contrary to other ambrosia beetles that attack stressed, possibly pathogen-infected, and dying trees, X. glabratus readily attacks healthy trees. To date little is known about the role of leaf volatiles in the host selection behavior and ecology of X. glabratus. To address this question, an olfactometer bioassay was developed to test the behavioral response of X. glabratus to plant leaf volatiles. We found that X. glabratus was attracted to the leaf odors of their hosts, redbay and swamp bay, with no attraction to a non-host tree tested (live oak, Quercus virginiana), which served as a negative control. Gas chromatography-mass spectrometry (GS/MS) analysis of leaves revealed the absence of sesquiterpenes known to be attractive to X. glabratus and present in host wood, suggesting that additional leaf-derived semiochemicals may serve as attractants for this beetle. An artificial blend of chemicals was developed based on GC/MS analyses of leaf volatiles and behavioral assays. This blend was attractive to X. glabratus at a level that rivaled currently used lures for practical monitoring of this pest. This synthetic redbay leaf blend also was tested in the field. Baited traps captured more X. glabratus than unbaited controls and equivalently to manuka oil lures. We hypothesize that leaf volatiles may be used by X. glabratus as an additional cue for host location.

Evaluation of seven essential oils identifies cubeb oil as most effective attractant for detection of Xyleborus glabratus

Redbay ambrosia beetle, Xyleborus glabratus, is an exotic wood-borer that vectors the fungal agent that induces laurel wilt. Since its introduction into Georgia in 2002, X. glabratus has spread throughout the southeastern U.S., and laurel wilt has decimated native Persea trees, particularly redbay (P. borbonia) and swampbay (P. palustris). The lethal disease now threatens avocado (P. americana) in Florida. To control the spread of laurel wilt, effective attractants are needed for early detection of the vector. Phoebe oil lures are the best known attractant, but they are no longer available. Current detection relies on manuka oil lures, but our previous research indicated they have a field life of only 2–3 weeks in Florida. Therefore, we evaluated seven essential oils as attractants for X. glabratus, and ethanol as a potential synergist. Field tests and electroantennography (EAG) were conducted to compare attraction and olfactory response to angelica seed, cubeb, ginger root, manuka, phoebe, tea tree, and orange oils. The highest captures were obtained with cubeb, manuka, and phoebe oils; ginger and angelica oils were intermediate, and tea tree and orange oils were not attractive. Addition of ethanol to oil lures did not increase captures. In subsequent tests with commercial formulations, cubeb lures captured more X. glabratus than manuka lures, were better for early detection at low population levels, and had longevity of at least 8 weeks. The highest EAG responses were elicited with phoebe and cubeb lures. Our results indicate that cubeb lures are the best attractant currently available for detection of X. glabratus.

Supercooling in the Redbay Ambrosia Beetle (Coleoptera: Curculionidae)

The redbay ambrosia beetle, Xyleborus glabratus Eichoff, (Coleoptera: Curculionidae: Scolytinae) is a severe pest of North American trees and shrubs in the family Lauraceae. Supercooling point (SCP) is an important physiological baseline for cold tolerance studies and could provide useful insights into the invasive potential of X. glabratus in northern latitudes of North America. The supercooling point (SCP) of X. glabratus was experimentally determined on field-collected and artificially cold hardened specimens. Field-collected beetles were captured in Jackson County, Mississippi using Lindgren funnel traps baited with manuka oil lures. Testing was conducted from June through August 2011. The mean SCP for field-collected X. glabratus was -21.7 ± 0.5 °C (± SE). A significant negative trend in the SCPs of field-collected beetles occurred over the summer testing period. Xyleborus glabratus specimens were reared from redbay (Persea borbonia (L.) Sprengel bolts in June 2012 and artificially cold hardened in a low temperature incubator at a thermo-photoperiod of 7 °C:2 °C (10:14 h L:D) for 31 days. Artificially cold hardened X. glabratus supercooled to a mean temperature of -23.9 ± 0.4 °C (± SE), which was significantly lower than that of field-collected beetles. Biometric indices of beetles (size, weight, and size x weight interaction) had no effect on the mean supercooling SCPs of either field-collected or artificially cold hardened beetles. Results from environmentally conditioned beetles suggest that X. glabratus has a high degree of thermal plasticity. Based on the artificially cold hardened mean SCP, X. glabratus and laurel wilt disease have the possibility to impact sassafras and northern spicebush throughout eastern North America. The data, although preliminary, suggests that a previous spatio-temporal model based on climate match data may have substantially underestimated the geographical area that may be affected by X. glabratus. This study will help form the basis of building and validating models to better predict the North American invasion potential of X. glabratus.

Volatiles from the symbiotic fungusRaffaelea lauricolaare synergistic with Manuka lures for increased capture of the Redbay ambrosia beetleXyleborus glabratus

Redbay ambrosia beetle Xyleborus glabratus is an invasive wood boring beetle that has become established in the southeastern U.S.A. and transmits a fungus Raffaelea lauricola that causes lethal laurel wilt. Among susceptible Lauraceae hosts are redbay Persea borbonia and avocado Persea americana. There is a crucial need for detection of this pest as it moves into new areas. Consequently, our goal was to create a better lure for the monitoring and control of redbay ambrosia beetle. We analyzed volatile emissions of R. lauricola, created a synthetic odour blend based on this analysis and tested this odour blend as a potential attractant in a redbay forest infested with X. glabratus. The synthetic Raffaelea odour blend was not attractive to the beetles by itself. However, it synergistically increased attraction to host-mimic volatiles. We tested four commercial release devices for dispensing Raffaelea odour at various release rates. Two prototypes with the highest release rate, when paired with commercial manuka oil lures, captured more beetles than manuka oil lures alone. These results indicate that a synthetic blend of volatiles based on the odour of the symbiotic fungus of X. glabratus may be useful for the development of more sensitive monitoring lures for this invasive pathogen vector.

Variation in Manuka Oil Lure Efficacy for CapturingXyleborus glabratus(Coleoptera: Curculionidae: Scolytinae), and Cubeb Oil as an Alternative Attractant

Redbay ambrosia beetle, Xyleborus glabratus Eichoff, is an exotic species to North America vectoring a deadly vascular wilt disease of redbay [Persea borbonia (L.) Spreng], swampbay [P. palustris (Raf.) Sarg.], avocado (P. americana Mill.), and sassafras [Sassafras albidum (Nutt.) Nees]. Xyleborus glabratus is attracted to manuka oil lures, which are commercially available, and phoebe oil. Variable efficacy of manuka oil lures and insufficient availability of phoebe oil prompted us to investigate the reasons behind changes in manuka oil lure efficacy and to test cubeb oil, a readily available essential oil from Piper cubeba L. seeds, as an alternative attractant. Attraction, release rates and durations, and volatile composition of manuka oil lures manufactured in 2008 were compared with manuka oil lures manufactured in 2012, and to whole and a distilled fraction of cubeb oil. Manuka oil lures from 2008 were more attractive to X. glabratus than controls for 8 wk, whereas lures from 2012 were attractive for only 2 wk. Cubeb oil and the distilled fraction of it were as attractive as or more attractive than manuka oil in three trials. In gravimetric studies, manuka oil lures from 2008 and cubeb oil lures continued to release volatiles for 57 d, whereas lures from 2012 stopped after 16 d. The chemical composition of volatiles released from new manuka oil lures from 2008 was similar to 2012; however, a preservative (butylated hydroxytoluene) was detected in the 2008 lures. Cubeb oil was an effective attractant for X. glabratus that lasted 8-9 wk when released from bubble lures.

An Uncertain Future for American Lauraceae: A Lethal Threat from Redbay Ambrosia Beetle and Laurel Wilt Disease (A Review)

Laurel wilt is a destructive vascular disease responsible for high mortality of American tree species in the family Lauraceae, particularly redbay (Persea borbonia) and swampbay (P. palustris), two dominant components of Coastal Plain forest communities in the southeastern United States. The disease syndrome emerged as a result of establishment of an exotic wood-boring beetle, Xyleborus glabratus, now known as the redbay ambrosia beetle. During gallery excavation, females of X. glabratus introduce a newly-described, obligatory fungal symbiont, Raffaelea lauricola. This fungus proliferates within the gallery and provides food for the beetles, but it has proven to be pathogenic to American lauraceous hosts, which have had no co-evolved history with R. lauricola. Presence of the foreign fungus elicits secretion of resins and formation of extensive parenchymal tyloses within xylem vessels. The extreme defensive response results in blockage of water transport, systemic wilt, and ultimately tree death. The beetle vector was first detected near Savannah, Georgia in 2002, and since has spread throughout the Southeast to become established in six states. The epidemic spread south through Florida more rapidly than predicted and currently threatens commercial production of avocado (Persea americana). Recent research indicates that California bay laurel (Umbellularia californica) can serve as a reproductive host for X. glabratus and is susceptible to laurel wilt disease. Thus, the US Pacific coastal forest ecosystems (and the California avocado industry) would be negatively impacted should the vector become established along the western coast. This review article summarizes our current understanding of the insect vector, the mycopathogen, and the susceptible host tree species. It also addresses elements of disease management and limitations with our current detection methods for redbay ambrosia beetle, which rely on manuka oil lures. Of the host-based attractants evaluated, cubeb oil shows the most promise as a potential new lure for X. glabratus.

Influence of Trap Color and Host Volatiles on Capture of the Emerald Ash Borer (Coleoptera: Buprestidae)

Field trapping assays were conducted in 2009 and 2010 throughout western Michigan, to evaluate lures for adult emerald ash borer, A. planipennis Fairmaire (Coleoptera: Buprestidae). Several ash tree volatiles were tested on purple prism traps in 2009, and a dark green prism trap in 2010. In 2009, six bark oil distillate lure treatments were tested against manuka oil lures (used in 2008 by USDA APHIS PPQ emerald ash borer cooperative program). Purple traps baited with 80/20 (manuka/phoebe oil) significantly increased beetle catch compared with traps baited with manuka oil alone. In 2010 we monitored emerald ash borer attraction to dark green traps baited with six lure combinations of 80/20 (manuka/phoebe), manuka oil, and (3Z)-hexenol. Traps baited with manuka oil and (3Z)-hexenol caught significantly more male and total count insects than traps baited with manuka oil alone. Traps baited with manuka oil and (3Z)-hexenol did not catch more beetles when compared with traps baited with (3Z)-hexenol alone. When compared with unbaited green traps our results show that (3Z)-hexenol improved male catch significantly in only one of three field experiments using dark green traps. Dark green traps caught a high number of A. planipennis when unbaited while (3Z)-hexenol was seen to have a minimal (nonsignificant) trap catch effect at several different release rates. We hypothesize that the previously reported kairomonal attractancy of (3Z)-hexenol (for males) on light green traps is not as obvious here because of improved male attractancy to the darker green trap.

Attraction of the Redbay Ambrosia Beetle, Xyleborus glabratus, to Avocado, Lychee, and Essential Oil Lures

The redbay ambrosia beetle, Xyleborus glabratus Eichhoff, is an exotic wood-boring insect that vectors the mycopathogen responsible for laurel wilt, a lethal vascular disease of trees in the Lauraceae. High mortality has occurred in native Persea species in the southeastern U.S., and the vector-pathogen complex poses an imminent threat to the production of commercial avocado, P. americana, in south Florida. There is a critical need for effective attractants to detect, monitor, and control this invasive pest. This study combined field tests and laboratory bioassays to evaluate the response of female X. glabratus to host-based volatiles from wood of avocado (cultivars of West Indian, Guatemalan, and Mexican races); from wood of lychee (Litchi chinensis, a presumed non-host that is high in the sesquiterpene α-copaene, a putative attractant); and to commercial lures containing manuka and phoebe oils, two reported attractive baits. Volatile collections and GC-MS analyses were performed to quantify the sesquiterpene content of test substrates. In the field, traps baited with lychee wood captured more beetles than those with wood from avocado cultivars; traps baited with phoebe oil lures captured more beetles than those with manuka oil lures (the current monitoring tool). In field and laboratory tests, X. glabratus did not show a preference among avocado races in either attraction or host acceptance (initiation of boring). In choice tests, lychee was more attractive than avocado initially, but a higher percentage of beetles bored into avocado, suggesting that lychee emits more powerful olfactory/visual cues, but that avocado contains more of the secondary cues necessary for host recognition. Emissions of α-copaene, β-caryophyllene, and α-humulene were correlated with field captures, and lychee wood may be a source of additional semiochemicals for X. glabratus.

Effect of Trap Type, Trap Position, Time of Year, and Beetle Density on Captures of the Redbay Ambrosia Beetle (Coleoptera: Curculionidae: Scolytinae)

The exotic redbay ambrosia beetle, Xyleborus glabratus Eichhoff (Coleoptera: Curculionidae: Scolytinae), and its fungal symbiont Raffaellea lauricola Harrington, Fraedrich, and Aghayeva are responsible for widespread redbay, Persea borbonia (L.) Spreng., mortality in the southern United States. Effective traps and lures are needed to monitor spread of the beetle and for early detection at ports-of-entry, so we conducted a series of experiments to find the best trap design, color, lure, and trap position for detection of X. glabratus. The best trap and lure combination was then tested at seven sites varying in beetle abundance and at one site throughout the year to see how season and beetle population affected performance. Manuka oil proved to be the most effective lure tested, particularly when considering cost and availability. Traps baited with manuka oil lures releasing 5 mg/d caught as many beetles as those baited with lures releasing 200 mg/d. Distributing manuka oil lures from the top to the bottom of eight-unit funnel traps resulted in similar numbers of X. glabratus as a single lure in the middle. Trap color had little effect on captures in sticky traps or cross-vane traps. Funnel traps caught twice as many beetles as cross-vane traps and three times as many as sticky traps but mean catch per trap was not significantly different. When comparing height, traps 1.5 m above the ground captured 85% of the beetles collected but a few were caught at each height up to 15 m. Funnel trap captures exhibited a strong linear relationship (r2 = 0.79) with X. glabratus attack density and they performed well throughout the year. Catching beetles at low densities is important to port of entry monitoring programs where early detection of infestations is essential. Our trials show that multiple funnel traps baited with a single manuka oil lure were effective for capturing X. glabratus even when no infested trees were visible in the area.

Evaluation of Double-Decker Traps for Emerald Ash Borer (Coleoptera: Buprestidae)

Improved detection tools are needed for the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive forest insect from Asia that has killed millions of ash (Fraxinus spp.) trees in North America since its discovery in Michigan in 2002. We evaluated attraction of adult A. planipennis to artificial traps incorporating visual (e.g., height, color, silhouette) and olfactory cues (e.g., host volatiles) at field sites in Michigan. We developed a double-decker trap consisting of a 3-m-tall polyvinyl pipe with two purple prisms attached near the top. In 2006, we compared A. planipennis attraction to double-decker traps baited with various combinations of manuka oil (containing sesquiterpenes present in ash bark), a blend of four ash leaf volatiles (leaf blend), and a rough texture to simulate bark. Significantly more A. planipennis were captured per trap when traps without the rough texture were baited with the leaf blend and manuka oil lures than on traps with texture and manuka oil but no leaf blend. In 2007, we also tested single prism traps set 1.5 m above ground and tower traps, similar to double-decker traps but 6 m tall. Double-decker traps baited with the leaf blend and manuka oil, with or without the addition of ash leaf and bark extracts, captured significantly more A. planipennis than similarly baited single prism traps, tower traps, or unbaited double-decker traps. A baited double-decker trap captured A. planipennis at a field site that was not previously known to be infested, representing the first detection event using artificial traps and lures. In 2008, we compared purple or green double-decker traps, single prisms suspended 3-5 m above ground in the ash canopy (canopy traps), and large flat purple traps (billboard traps). Significantly more A. planipennis were captured in purple versus green traps, baited traps versus unbaited traps, and double-decker versus canopy traps, whereas billboard traps were intermediate. At sites with very low A. planipennis densities, more A. planipennis were captured on baited double-decker traps than on other traps and a higher percentage of the baited double-decker traps captured beetles than any other trap design. In all 3 yr, peak A. planipennis activity occurred during late June to mid-July, corresponding to 800-1200 growing degree-days base 10 degrees C (DD10). Nearly all (95%) beetles were captured by the end of July at approximately 1400 DD10.

First Occurrence of Laurel Wilt Disease Caused by Raffaelea lauricola on Redbay Trees in Mississippi.

Laurel wilt is a lethal, nonnative vascular wilt disease of redbay (Persea borbonia), sassafras (Sassafras albidum), and other trees in the Lauraceae (1,4). It is caused by a fungus (Raffaelea lauricola) and transmitted by the redbay ambrosia beetle (Xyleborus glabratus), a nonnative insect first detected in Georgia in 2002 (1,2). Since introduction of the pathogen and vector (presumably from Asia), laurel wilt has caused extensive mortality to redbays in Georgia, Florida, and South Carolina (1). In June 2009, a landowner in Gautier, MS reported dead redbay trees. Signs and symptoms were identical to those reported for laurel wilt along the Atlantic Coast (wilted, bronze red foliage, and dark gray-to-black vascular discoloration) (1). Infected trees have subsequently been confirmed in and near the Pascagoula River Basin. Size of infected redbays ranged from 5 to 20 cm (diameter at breast height). No heavily decomposed or fallen redbays were noted. Many individual specimens exhibited extensive drying of stem wood and dry, wilted, light brown foliage. This indicates that introduction to the area may have occurred within the last 3 years. X. glabratus adults were collected (30°26'44.45″N, 88°39'41.83″W) in a Lindgren funnel trap baited with phoebe and manuka oil lures. Beetle identification was confirmed by USDA-APHIS, and voucher specimens were submitted to the Smithsonian National Museum of Natural History and the Mississippi Entomological Museum. Symptomatic redbay wood chips from the same location were surface sterilized and plated on cycloheximide-streptomycin malt agar and R. lauricola was isolated. A 1,026-bp portion of 18S rDNA (GenBank No. GQ996063) was amplified by PCR and sequenced using primers NS1 and NS4. BLASTn searches revealed perfect homology to R. lauricola isolate PL 697 (GQ329704). Two isolates of R. lauricola were recovered and prepared into separate spore suspensions (1 × 108 CFU/ml). Each isolate was inoculated into two healthy redbays. The inoculated redbays were placed in a growth chamber with two water-only controls. All inoculated plants, and none of the controls, exhibited wilt symptoms and died within 20 days. R. lauricola was recovered from the discolored sapwood of the inoculated plants, completing Koch's postulates. A model prediction for the natural dispersion of X. glabratus and R. lauricola estimated that these organisms may not reach Mississippi for 10 to 15 years (3). The current detection of laurel wilt in Mississippi is substantially ahead of this estimate. Currently, no records of laurel wilt have been reported from western Georgia, all of Alabama, or the panhandle of Florida. Confirmed locations in Mississippi are in Jackson County, along the Interstate 10 corridor and the Pascagoula River drainage. Due to the relatively large extent of the infestation (~64 km2, including hundreds of infected trees) eradication is not being attempted. Surveys, remote sensing, and phylogeographic analysis are underway to delineate the extent of infestation and discover the mode of introduction. The current outbreak of laurel wilt in Mississippi is likely the result of human transport of infested wood, either from Asia as a separate, new introduction or from previously infested areas in the southeastern United States.

Efficacy of trap and lure types for detection of Agrilus planipennis (Col., Buprestidae) at low density

AbstractDevelopment of effective trapping tools for forest pests and evaluating the key components of these tools is necessary to locate early‐stage infestations and develop management responses to them. Agrilus planipennis Fairmaire (emerald ash borer) is an introduced pest of ash (Fraxinus spp. L.) in North America. The effectiveness of different trap and lure combinations were tested in areas with low and high density populations of A. planipennis. At low density sites, purple prism traps outperformed green traps and girdled ash trap trees in capture rates (adults per day) and rates of detection of A. planipennis. Also, manuka oil lures, used as a standard lure in a national survey programme, captured higher rates of A. planipennis than did previous standards of girdled ash trap trees. There was no logistic relationship between the detection of A. planipennis on a trap and the diameter of the ash tree from which the trap was suspended, possibly because of the use of artificial lures with these traps. There was also no difference in the mean number of A. lanipennis captured per day between ash species and between vigour rating of ash associated with the traps. However, traps placed in open grown and dominant trees captured more beetles than traps placed in lower canopy class trees. At sites defined as low and high density, there was no difference in the larval density per cm3 of phloem. This suggests that exposure time to A. planipennis has been shorter at those low density sites. By exploiting the trap and tree characteristics that improve A. planipennis capture rates and detection efficacy, there can be future improvement in management of this pest. If detection can occur before infested ash trees exhibit signs and symptoms, there may be a potential for reducing the mortality of ash within stands.

Emerald Ash Borer (Coleoptera: Buprestidae) Attraction to Stressed or Baited Ash Trees

Emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), has killed millions of ash (Fraxinus sp.) trees in North America since its discovery in Michigan in 2002. Efficient methods to detect low-density A. planipennis populations remain a critical priority for regulatory and resource management agencies. We compared the density of adult A. planipennis captured on sticky bands and larval density among ash trees that were girdled for 1 or 2 yr, wounded, exposed to the stress-elicitor methyl jasmonate, baited with Manuka oil lures, or left untreated. Studies were conducted at four sites in 2006 and 2007, where A. planipennis densities on untreated trees ranged from very low to moderate. In 2006, 1-yr girdled trees captured significantly more adult A. planipennis and had higher larval densities than untreated control trees or trees treated with methyl jasmonate or Manuka oil. Open-grown trees captured significantly more A. planipennis beetles than partially or fully shaded trees. In 2007, A. planipennis population levels and captures of adult A. planipennis were substantially higher than in 2006. The density of adults captured on sticky bands did not differ significantly among canopy exposure classes or treatments in 2007. Larval density was significantly higher in untreated, wounded, and 1-yr girdled trees (girdled in 2007) than in 2-yr girdled trees (girdled in 2006), where most phloem was consumed by A. planipennis larvae the previous year. A total of 36 trees (32 in 2006, 4 in 2007) caught no beetles, but 16 of those trees (13 in 2006, 3 in 2007) had A. planipennis larvae. In 2006, there was a positive linear relationship between the density of adults captured on sticky bands and larval density in trees. Our results show that freshly girdled and open grown trees were most attractive to A. planipennis, especially at low-density sites. If girdled trees are used for A. planipennis detection or survey, debarking trees to locate larval galleries is crucial.