Verbenone reduces landing of the redbay ambrosia beetle,vector of the laurel wilt pathogen,on live standing redbay trees

1. Laurel wilt is a disease that has caused extensive mortality of redbay Persea borbonia in the southeastern U.S.A. The redbay ambrosia beetle Xyleborus glabratus is the vector of the causal agent of laurel wilt, the fungus Raffaelea lauricola.
2. We tested two potential repellents to the redbay ambrosia beetle, verbenone and methyl salicylate (MeSA) in an 8‐month large‐scale experiment conducted in three locations in Florida. In each location, redbay trees were treated with a single or double application of SPLAT (Specialized Pheromone and Lure Application Technology; ISCA Technologies, Riverside, California) verbenone, as well as SPLAT with a 1:2 mix of MeSA and verbenone.
3. The MeSA + verbenone mixes did not reduce beetle captures compared with the control treatment, whereas SPLAT verbenone alone significantly reduced the number of beetles captured on sticky traps placed on redbay trees in the three locations. The reduction of beetle capture was similar regardless of one or two treatments of SPLAT verbenone. The reduction of tree death with the SPLAT verbenone treatment was not statistically significant.
4. The results of the present study suggest that trunk application of verbenone can reduce landing rates of the redbay ambrosia beetle on live redbay trees and shows promise for use in an integrated pest management strategy against laurel wilt.

     

Biology and host associations of redbay ambrosia beetle (Coleoptera: Curculionidae: Scolytinae), exotic vector of laurel wilt killing redbay trees in the southeastern United States

The redbay ambrosia beetle, Xyleborus glabratus Eichhoff (Coleoptera: Curculionidae: Scolytinae), and its fungal symbiont, Raffaelea sp., are new introductions to the southeastern United States responsible for the wilt of mature redbay, Persea borbonia (L.) Spreng., trees. In 2006 and 2007, we investigated the seasonal flight activity of X. glabratus, its host associations, and population levels at eight locations in South Carolina and Georgia where infestations ranged from very recent to at least several years old. Adults were active throughout the year with peak activity in early September. Brood development seems to take 50-60 d. Wood infested with beetles and infected with the Raffaelea sp. was similar in attraction to uninfested redbay wood, whereas both were more attractive than a nonhost species. Sassafras, Sassafras albidium (Nutt.) Nees, another species of Lauraceae, was not attractive to X. glabratus and very few beetle entrance holes were found in sassafras wood compared with redbay. Conversely, avocado, Persea americana Mill., was as attractive to X. glabratus as swampbay, P. palustris (Raf.) Sarg., and both were more attractive than the nonhost red maple, Acer rubrum L. However, avocado had relatively few entrance holes in the wood. In 2007, we compared X. glabratus populations in areas where all mature redbay have died to areas where infestations were very active and more recent. Trap catches of X. glabratus and numbers of entrance holes in trap bolts of redbay were correlated with the number of dead trees with leaves attached. Older infestations where mature host trees had been eliminated by the wilt had low numbers of beetles resulting in trap catches ranging from 0.04 to 0.12 beetles per trap per d compared with 4-7 beetles per trap per d in areas with numerous recently dead trees. Our results indicate beetle populations drop dramatically after suitable host material is gone and provide hope that management strategies can be developed to restore redbay trees. The lack of attraction of X. glabratus to sassafras suggests that spread of X. glabratus may slow once it is outside the range of redbay.     

Evaluation of repellents for the redbay ambrosia beetle,Xyleborus glabratus,vector of the laurel wilt pathogen

The redbay ambrosia beetle, Xyleborus glabratus, is the vector of the laurel wilt disease fungal pathogen, Raffaelea lauricola. Since the vector's initial detection in the USA in the early 2000s, laurel wilt has killed millions of redbay, Persea borbonia, trees and other members of the plant family Lauraceae. To protect host trees from beetle attack and laurel wilt infection, we tested the efficacy of host‐ and non‐host‐derived and commercial compounds as X. glabratus repellents in field experiments. In our first trial, the major constituents of the non‐host tree, longleaf pine, Pinus palustris, and SPLAT Verb (verbenone 10%) were paired with manuka oil attractants and beetle captures were counted. Verbenone and a 1 : 1 blend of myrcene and camphene were intermediate to both the manuka positive and blank negative controls. Subsequently, we tested different blends of methyl salicylate (MeSA), a host defence and signalling compound, and verbenone in SPLAT dispensers using freshly cut redbay bolts as an attractant. All treatments reduced X. glabratus captures and boring holes as compared to the redbay (‐) repellent positive control; however, SPLAT Verb and SPLAT MeSA‐Verb (5% each) achieved the highest repellency, with results comparable to that of the non‐host (laurel oak). These trials establish that host‐derived and commercially available repellent compounds can reduce X. glabratus attacks and therefore have potential as part of an integrated management strategy against laurel wilt and its vector.     

Impacts of laurel wilt disease on redbay (Persea borbonia (L.) Spreng.) population structure and forest communities in the coastal plain of Georgia, USA

Laurel wilt disease (LWD), a fungal disease vectored by the non-native redbay ambrosia beetle (Xyleborus glabratus Eichhoff), has caused mortality of redbay (Persea borbonia (L.) Spreng.) in the coastal plain of Georgia since 2003. Despite its rapid spread, little research has evaluated its impacts on redbay population structure and forest communities. Diseased populations of redbay in five sites (2–4 years post infestation) were compared to healthy populations in three uninfested sites in five counties in Georgia. The results showed high redbay mortality, shifts in size structure, and changes in community composition. An average of 90 % of redbay trees ≥3 cm diameter at breast height (DBH) were dead in infested sites, compared to 0–35 % in control sites. Mortality was seen in individuals of the smallest stem diameter category (<1.00 cm diameter at ground height). DBH of live redbay trees in control sites was twice that of those in infested sites. Photosynthetically active radiation was 4.8 times greater at infested sites than control sites due to loss of redbay canopy. Community structure measurements showed redbay trees had the greatest mean importance value (IV) at control sites compared to the 8th mean IV at infested sites for live stems. Two species co-dominant to redbay, sweetbay (Magnolia virginiana L.) and loblolly bay (Gordonia lasianthus (L.) J. Ellis), were of higher importance at infested than control sites, suggesting they are increasing in dominance following the mortality of redbay. This study shows LWD has impacted redbay populations and altered associated forest communities in Georgia.     

Sprouting capacity of <Emphasis Type="Italic">Persea borbonia</Emphasis> and maritime forest community response to simulated laurel wilt disease

There are numerous examples of how exotic insect pests and pathogens have altered the dominance of native tree species. Changes to the structure of associated communities will depend on whether the affected species survives and if so, the degree to which it is diminished. In the southeastern USA, Persea borbonia, a common tree found in many coastal plain habitats, is the primary host of laurel wilt disease (LWD); infection rates and main-stem mortality are catastrophically high (>90%) in invaded populations. We simulated the effects of LWD prior to its arrival in coastal Mississippi by girdling and then removing the main stems of P. borbonia trees. Over a 2-year period, we monitored P. borbonia persistence via basal resprouts, understory light availability, and community structure. Removal of P. borbonia main stems resulted in a 50% increase in light transmission (measured at 1 m above ground level). All treated individuals produced basal resprouts, the size and number of which were positively related to initial tree girth. Post-treatment increases in basal area were greatest for the sub-canopy species, Ilex vomitoria, and were significantly higher in treatment versus control plots. Woody seedlings and herbaceous plants showed no significant trends in composition and abundance over time or between control and treatment plots. Our results suggest that removal of P. borbonia and subsequent resprouting causes shifts in P. borbonia size class frequencies and sub-canopy species dominance but has negligible impacts on understory plant community dynamics.     

Potential for host shifting in Papilio palamedes following invasion of laurel wilt disease

In the southeastern US, laurel wilt disease (LWD) is causing widespread mortality of species in the Lauraceae. The principal target, Persea borbonia, is the primary larval host of Papilio palamedes, which is known to feed on other Lauraceae species. Among these potential hosts, the exotic Cinnamomum camphora is the only species that has shown resistance to LWD. We hypothesized that oviposition preference for C. camphora and P. borbonia would correspond to larval performances on these species and that the relative host suitability of C. camphora would indicate an opportunity for host-switching. We used laboratory experiments and field observations to compare performance and preference of P. palamedes between C. camphora and P. borbonia foliage. Our results indicate moderate survivorship on C. camphora compared to P. borbonia and no differences in first and fourth instar growth rates between treatments. Fourth instars consumed relatively less of C. camphora foliage compared to that of P. borbonia, but metabolic efficiency did not differ between treatments. Rearing on the foliage of P. borbonia stump sprouts from LWD-infected trees resulted in significantly higher growth rates and metabolic efficiency as first and fourth instars, respectively. In the field and laboratory, we found no oviposition preference for C. camphora. While females laid eggs on C. camphora during laboratory trials, the same number of eggs was also laid on inanimate objects. We conclude that C. camphora is suitable for larval development but host-switching to this species by P. palamedes will be primarily constrained by the ecological factors that govern oviposition behaviors.     

Isolations from the redbay ambrosia beetle, Xyleborus glabratus, confirm that the laurel wilt pathogen, Raffaelea lauricola, originated in Asia

The laurel wilt pathogen Raffaelea lauricola was hypothesized to have been introduced to the southeastern USA in the mycangium of the redbay ambrosia beetle, Xyleborus glabratus, which is native to Asia. To test this hypothesis adult X. glabratus were trapped in Taiwan and on Kyushu Island, Japan, in 2009, and dead beetles were sent to USA for isolation of fungal symbionts. Individual X. glabratus were macerated in glass tissue grinders, and the slurry was serially diluted and plated onto malt agar medium amended with cycloheximide, a medium semiselective for Ophiostoma species and their anamorphs, including members of Raffaelea. R. lauricola was isolated from 56 of 85 beetles in Taiwan and 10 of 16 beetles in Japan at up to an estimated 10 000 CFUs per beetle. The next most commonly isolated species was R. ellipticospora, which also has been recovered from X. glabratus trapped in the USA, as were two other fungi isolated from beetles in Taiwan, R. fusca and R. subfusca. Three unidentified Raffaelea spp. and three unidentified Ophiostoma spp. were isolated rarely from X. glabratus collected in Taiwan. Isolations from beetles similarly trapped in Georgia, USA, yielded R. lauricola and R. ellipticospora in numbers similar to those from beetles trapped in Taiwan and Japan. The results support the hypothesis that R. lauricola was introduced into the USA in mycangia of X. glabratus shipped to USA in solid wood packing material from Asia. However differences in the mycangial mycoflora of X. glabratus in Taiwan, Japan and USA suggest that the X. glabratus population established in USA originated in another part of Asia.     

Entomopathogenic fungi as biological control agents for the vector of the laurel wilt disease,the redbay ambrosia beetle,Xyleborus glabratus (Coleoptera: Curculionidae)

The redbay ambrosia beetle (RAB), Xyleborus glabratus, is a wood-boring insect that vectors the fungal pathogen, Raffaelea lauricola, which causes laurel wilt, a lethal disease of avocado. The objective of this study was to determine the susceptibility of RAB to infection and subsequent death by exposure to three commercial strains of entomopathogenic fungi [two strains of Isaria fumosorosea (Ifr 3581 and PFR), and strain GHA of Beauveria bassiana]. RAB females were dipped in fungal spore solutions and their median survivorship times (MST) determined. Contact with any of the biopesticides resulted in death of all RAB females. MSTs of RAB females ranged from 3 days (B. bassiana) to 5 days (I. fumosorosea PFR). B. bassiana killed RAB females faster, followed by Ifr 3581 and PFR. RAB females dipped in B. bassiana suspensions had the highest number of viable spores attached to their bodies, followed by Ifr 3581. Beetles dipped in PFR suspension had significantly less viable spores attached to their bodies. No significant differences were observed in the mortality of beetles exposed to entomopathogenic fungi by dipping in a fungal suspension or walking on treated avocado bolts. Beetles bored into the logs and constructed galleries, but they were found dead inside the galleries a few days after exposure to the entomopathogens. Entomopathogenic fungal infection in dead beetles was confirmed through molecular techniques. This is the first study to demonstrate that entomopathogenic fungi are potential biological control agents against RAB.     

Establishment of laurel forest vegetation in adjacent secondary forest in central Japan

Distributions of lucidophyllous species are limited due to the fragmentation of laurel forest. On Komayama Hill in central Japan, we evaluated the colonization of typical lucidophyllous vascular plants from a 350-year-old laurel forest into adjacent abandoned secondary forest for conservation and restoration purposes. A total of 14 consecutive subplots were established along the vegetation border between the two forests (length, 30 m; width, 5 m), extending 70 m into the secondary forest; 18 quadrats of old-growth forest were surveyed. Edge effects of old-growth forest were found to play an important role in re-establishing lucidophyllous saplings and seedlings in the secondary forest. In particular, the abundances of the four dominant canopy species of the old-growth forest significantly decreased with increasing distance. Hence, they are expected to colonize further into the secondary forest and, ultimately, to dominate the canopy. However, the number of lucidophyllous species did not change with distance. Species such as Ficus nipponica, Damnacanthus indicus, Ilex integra, and Lemmaphyllum microphyllum were near-completely or completely limited to the old-growth forest. They are known to be negatively affected by forest fragmentation and were observed to be struggling to colonize the exterior of the old-growth forest even after 60 years of abandonment. Their absence highlighted the limited colonization capacities of some old-growth forest species and underlined the time required for habitat restoration following human disturbance. We conclude that it is important to consider the population dynamics of dominant canopy species and the colonization of these interior species when assessing the habitat expansion of lucidophyllous species and hence the restoration of degraded lands.     

Treefall gap characteristics and regeneration in the laurel forest of Tenerife

Abstract. We conducted a study in the laurel forest of Tenerife (Canary Islands, Spain) to describe the characteristics of natural gaps and to assess the role of treefall gaps in forest dynamics. Very little is left of the natural laurel forest with i.a. Laurus azorica, Ilex canariensis and Prunus lusitanica. We looked for treefall gaps in 80 randomly located 2500 m² plots. These plots represented ca. 1% of the remaining and protected laurel forest of Tenerife. We recorded the characteristics of the species causing the gaps, gap architecture and gap age in all observed gaps larger than 10 m². We inventoried the regeneration in each gap and in a neighbouring control plot with the same topography. Large gaps (>75 m²) were not common in the laurel forest. The absence of large gaps could be due to the physiognomy of the vegetation, the mild weather or the rarity of disturbances. Instead of forming gaps, many trees decompose in place and branches from neighbouring trees and suckers from the decomposed trees occupy the free space. Also, the high rate of asexual regeneration could contribute to the fast closing of the gap. The number of gaps created by Prunus lusitanica was higher than expected (based on canopy composition) while Ilex canariensis and Laurus azorica created fewer gaps. In this evergreen forest, differences between gap and non-gap conditions are not as distinct as in other forest types. Only 0.4% of the canopy is in the gap phase (0.6% including gaps smaller than 10m²). No differences were found in patterns of regeneration between gap and non-gap phases in the forest. Gaps do not explain the persistence of pioneer species in the laurel forest.     

Spatial patterns of trees and juveniles in a laurel forest of Tenerife, Canary Islands

Spatial patterns are important characteristics of the forest and theycan reveal such things as successional status and ecological characteristics ofthe species. We tested the hypothesis that spatial distribution will bedifferent, depending on whether the species is intolerant or tolerant to shade.We assessed the spatial distribution of trees (> 4 cm dbh) andjuveniles in eight laurel forest plots. A univariate spatial analysis(performed with Ripley's K₁) showed that all tree species havesignificantaggregation at short distances (2 m). Nevertheless, two groups ofspecies could be differentiated: Erica scoparia,Myrica faya and Ilex canariensisshowed a tendency for aggregation at large distances (larger than 6m)while L. azorica and Prunuslusitanicashowed aggregation only at shorter distances. Ripley's BivariateK_1,2 analyses showed no significant differences in the spatialdistribution ofanalyzed species pairs from a null model. Only Laurusazoricahad a sufficient sample size for analysis of juvenile distribution. Aunivariateanalysis revealed thatL. azorica seedlings (stems < 50 cm high)were clumped in some plots up to 5 m, but this was not consistent.Saplings (stems > 50 cm high and < 4 cm dbh)didnot show strong clumping even at short distances. L.azoricasaplings had no significant aggregation with, nor repulsionfrom, adults of the same or different species. Spatial patterns of the speciesshould be considered in the development of restoration plans of the laurelforest 90%of which has disappeared or been intensively disturbed on Tenerife Island.     

Suitability of California bay laurel and other species as hosts for the non‐native redbay ambrosia beetle and granulate ambrosia beetle

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Changes in arbuscular mycorrhizal fungus community along an exotic plant Eupatorium adenophorum invasion in a chinese secondary forest

Knowledge of the changes in arbuscular mycorrhizal (AM) fungi is fundamental for understanding the success of exotic plant invasions in natural ecosystems. In this study, AM fungal colonization and spore community were examined along an invasive gradient of the exotic plant Eupatorium adenophorum in a secondary forest in southwestern China. With increasing E. adenophorum invasion, the density of arbuscules in the roots of E. adenophorum significantly increased, but the AM root colonization rate and the densities of vesicles and hyphal coils in roots of E. adenophorum were not significantly different. A total of 29 AM fungi belonging to nine genera were identified based on spore morphology. Claroideoglomus etunicatum, Funneliformis geosporus, and Glomus aggregatum were the most common AM fungal species. The E. adenophorum invasion significantly decreased the AM fungal spore density in the soil. Furthermore, with increasing of E. adenophorum invasion the spore densities of C. etunicatum, G. aggregatum, and G. arenarium significantly decreased, whereas F. geosporus significantly increased. Nonmetric multidimensional scaling demonstrated that the AM fungus community composition was significantly different (P=0.003) in the different invasive levels of E. adenophorum, and significantly correlated with plant species richness, soil total P, and soil NO₃ ^?-N. The results suggest that the alteration in AM fungus community might be caused by E. adenophorum invasion via changing the local plant community and soil properties in a Chinese secondary forest ecosystem.     

Effect of tree host species on fungal community composition in a tropical rain forest in Panama

Abstract.  To determine whether host species influence the composition of fungal communities, the ascomycetes and basidiomycetes present on three tree species ( Prioria copaifera (Fabaceae), Quararibea asterolepis (Bombacaceae), and Trichilia tuberculata (Meliaceae)) were sampled on the 50-ha Forest Dynamics Project plot in lowland moist tropical forest on Barro Colorado Island, Panama. The most abundant fungal morphotypes of both ascomycetes and basidiomycetes were generalists found on all three hosts, but detrended correspondence analysis revealed distinct differences in fungal community composition among host trees. These differences among hosts were constant across census years. Randomization tests revealed that there were significantly fewer host-generalist fungi than expected for ascomycetes but not for basidiomycetes. These results indicate that host composition plays a role in structuring both ascomycete and basidiomycete fungal communities, but that the most successful fungal morphotypes are capable of colonizing multiple host species.     

Effects of pine wilt disease invasion on soil properties and Masson pine forest communities in the Three Gorges reservoir region,China

Pine wilt disease (PWD) has caused significant Masson pine mortality in the Three Gorges reservoir region in central China. In this study, five uniform Masson pine stand types infected by PWD were selected and surveyed on slopes and aspects with similar environmental conditions. In sites that had been infected, soil bulk density was reduced, and the difference among the groups was statistically significant (P < 0.05) at the 0–10 cm and 10–20 cm soil layers, but not at 20–40 cm. Other soil water‐related physical properties, excluding noncapillary porosity, significantly differed among the groups in all soil layers. Additionally, the values of available phosphorus, sodium, potassium, calcium, and magnesium were higher in the invaded stands, but the total nitrogen and organic matter contents were lower. Masson pine does not become reestablished following PWD‐induced mortality but is instead replaced by broad‐leaved tree species. Among the 19 examined environmental variables, five were found to be significantly related with the ordination of plant community structure: Masson pine stumps (MPS), K⁺, capillary water holding capacity (CWHC), capillary porosity (CP), and soil water content (SWC). Among these factors, the plant community structure was principally related to MPS and K⁺. The findings of this study show that the outbreak of PWD has impacted Masson pine forest soil properties and altered forest community composition. The disease is negatively related with the presence of Masson pine and positively associated with that of broad‐leaved tree species.     

A preliminary study on collembolan populations in a pine forest

1. A study was made of the populations of Collembola in the pine forest. Among the eight species commonly found, Folsomia octoculataHandschin was numerically dominant.
2. Seasonal population changes of these eight species were described over a year. It was found that many species show two peaks in number. As for F. octoculata, the dominant species, the changes in number was discussed in connection with the variation in age structure.
3. The census data for each species were examined for the distribution pattern by using the regression method. None of these eight species was distributed at random in the soil and they could be classified into three groups based on the differences in the degree of aggregation in terms of coefficient β of the regression. It was suggested that such differences in the distribution pattern are largely due to differences in the response to heterogeneity of the habitat. In the case of F. octoculata, the changes of distribution pattern in the course of post-embryonic development were examined and it was found that (a) the component of distribution is a single individual rather than a colony, and (b) the degree of aggregation is decreased with the development of individuals.
4. Using the relationship, the relation of the necessary sample size to the population density was derived for a fixed level of presision (D=S.E./m=0.2) and some suggestions were given concerning sampling plans for these insects.

     

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