Utilizing volatile organic compounds for early detection of Fusarium circinatum.

Fusarium circinatum, a fungal pathogen deadly to many Pinus species, can cause significant economic and ecological losses, especially if it were to become more widely established in Europe. Early detection tools with high-throughput capacity can increase our readiness to implement mitigation actions against new incursions. This study sought to develop a disease detection method based on volatile organic compound (VOC) emissions to detect F. circinatum on different Pinus species. The complete pipeline applied here, entailing gas chromatography-mass spectrometry of VOCs, automated data analysis and machine learning, distinguished diseased from healthy seedlings of Pinus sylvestris and Pinus radiata. In P. radiata, this distinction was possible even before the seedlings became visibly symptomatic, suggesting the possibility for this method to identify latently infected, yet healthy looking plants. Pinus pinea, which is known to be relatively resistant to F. circinatum, remained asymptomatic and showed no changes in VOCs over 28 days. In a separate analysis of in vitro VOCs collected from different species of Fusarium, we showed that even closely related Fusarium spp. can be readily distinguished based on their VOC profiles. The results further substantiate the potential for volatilomics to be used for early disease detection and diagnostic recognition.

Ecological fitting: Chemical profiles of plant hosts provide insights on selection cues and preferences for a major buprestid pest.

Specific cues used by emerald ash borer (EAB, Agrilus planipennis) to select hosts are largely unknown. Attractants are likely general and the use of novel host plants provides an opportunity to investigate the commonality of these cues. We examined volatile profiles emitted by five plants that can host EAB and estimated their importance in explaining known oviposition preferences. Foliage volatiles were collected from potted black ash (Fraxinus nigra), Manchurian ash (F. mandshurica), blue ash (F. quadrangulata), white fringetree (Chionanthus virginicus), and olive (Olea europaea) and analyzed using GC-MS. Fifty-nine compounds were detected including eight green leaf volatiles (GLV), 12 monoterpenes, and 21 sesquiterpenes. Ordination plots show separation of species by full foliage profiles, monoterpenes, sesquiterpenes, and known antennally active compounds, but GLVs were similar across hosts. Random Forest (RF) analysis revealed eight compounds that separated plant species with an error rate of ~19%, consisting mostly of sesquiterpenes. Similarity of GLV profiles among known hosts suggests they serve as general cues for host selection. Manchurian ash, a resistant host, produced the highest quantities and variety of sesquiterpenes indicating that some of these chemicals may be antixenotic. All compounds identified by RF have been implicated as deterrents or attractants to woodborers in other studies and should be investigated for adult antennal activity and attraction.

Feeding, Survival, and Fecundity of Adult Emerald Ash Borer (Coleoptera: Buprestidae) on Foliage of Two Novel Hosts and Implications for Host Range Expansion.

Insect herbivores are more likely to successfully use a novel host if the plant is closely related to the ancestral host and the insect is polyphagous. Emerald ash borer (EAB), Agrilus planipennis (Fairmaire) (Coleoptera: Buprestidae), is a specialist wood borer of ash (Fraxinus spp., Lamiales: Oleaceae) trees and one of the most destructive forest pests in North American forests. Recent studies have found that larvae can develop in stems of two ash relatives; white fringetree (Chionanthus virginicus (L.) [Laminales: Oleaceae]) and cultivated olive (Olea europaea (L.) [Laminales: Oleaceae]). For EAB adults, the ability to consume, successfully mate, and lay viable eggs on foliage of these hosts is unknown. Thus, we conducted two no-choice assays with adult EAB on foliage of white fringetree and olive paired with positive controls of susceptible ash. Larval performance was also examined in a reciprocal study with cut stems of white fringetree and green ash (Fraxinus pennsylvanica Marshall) to determine whether adult diet impacted the success of progeny. Longevity, consumption rates, and fecundity of adults were similar on white fringetree and ash foliage. In contrast, adults consuming olive died quickly, consumed more over time, and females laid far fewer eggs compared to those on ash. Adult diet did not impact larval success, but larvae in white fringetree stems grew slower. These results indicate that white fringetree is a suitable host for EAB to complete its lifecycle, although larvae perform more poorly on this host than in susceptible ash species. In contrast, the more distantly related olive appears to be a poor host for adult EAB, although some viable eggs were produced by females.

Larval Performance of a Major Forest Pest on Novel Hosts and the Effect of Stressors.

Novel hosts lacking a coevolutionary history with herbivores can often support improved larval performance over historic hosts; e.g., emerald ash borer [Agrilus planipennis (Fairmaire) Coleoptera: Buprestidae] on North American ash (Fraxinus spp.) trees. Whether trees are novel or ancestral, stress on plants increases emerald ash borer preference and performance. White fringetree [Chionanthus virginicus (L.) Lamiales: Oleaceae] and olive [Olea europaea (L.) Lamiales: Oleaceae] are closely related non-ash hosts that support development of emerald ash borer to adulthood, but their relative suitability as hosts and the impact of plant stress on larval success has not been well studied. In a series of experiments, survival and growth of emerald ash borer larvae on these novel hosts were examined along with the impact of stress. In the first experiment, larvae grew more slowly in cut stems of olive than in green ash [Fraxinus pennsylvanica (Marshall) Lamiales: Oleaceae] and several adults successfully emerged from larger olive stems. In two experiments on young potted olive with photosynthesizing bark, larvae died within a week, but mechanical girdling increased the rate of gallery establishment. The final two experiments on field-grown fringetrees found increased larval survivorship and growth in previously emerald ash borer attacked and mechanically girdled plants than in healthy stems or stems treated with the defense hormone, methyl jasmonate. Our results demonstrate that these non-ash hosts are less suitable for emerald ash borer than preferred ash hosts, but previous emerald ash borer attack or girdling led to better survival and growth demonstrating the importance of stress for larval success. In potted olive, high mortality could be due to higher loads of toxic compounds or the presence of chlorophyllous tissue.

The potential for host switching via ecological fitting in the emerald ash borer-host plant system.

The traits used by phytophagous insects to find and utilize their ancestral hosts can lead to host range expansions, generally to closely related hosts that share visual and chemical features with ancestral hosts. Host range expansions often result from ecological fitting, which is the process whereby organisms colonize and persist in novel environments, use novel resources, or form novel associations with other species because of the suites of traits that they carry at the time they encounter the novel environment. Our objective in this review is to discuss the potential and constraints on host switching via ecological fitting in emerald ash borer, Agrilus planipennis, an ecologically and economically important invasive wood boring beetle. Once thought of as an ash (Fraxinus spp.) tree specialist, recent studies have revealed a broader potential host range than was expected for this insect. We discuss the demonstrated host-use capabilities of this beetle, as well as the potential for and barriers to the adoption of additional hosts by this beetle. We place our observations in the context of biochemical mechanisms that mediate the interaction of these beetles with their host plants and discuss whether evolutionary host shifts are a possible outcome of the interaction of this insect with novel hosts.

Feeding and Development of Emerald Ash Borer (Coleoptera: Buprestidae) on Cultivated Olive, Olea europaea.

We examined the suitability of cultivated olive, Olea europaea L., as a host for emerald ash borer, Agrilus planipennis Fairmaire. In a bioassay using cut stems from a field-grown olive tree (cv. 'Manzanilla') we found that 45% of larvae that had emerged from eggs used to inoculate stems, were recovered alive, many as larvae or prepupae, during periodic debarking of a subset of stems. Three intact stems that 19 larvae successfully entered were exposed to a simulated overwintering treatment. Four live adults emerged afterwards, and an additional pupa and several prepupae were discovered after debarking these stems. Cultivated olive joins white fringetree as one of the two species outside of the genus Fraxinus capable of supporting the development of emerald ash borer from neonate to adult.

Distribution, Predictors, and Impacts of Emerald Ash Borer (Agrilus planipennis) (Coleoptera: Buprestidae) Infestation of White Fringetree (Chionanthus virginicus).

Emerald ash borer, Agrilus planipennis (Fairmaire), is an invasive pest of ash trees (Fraxinus spp.) in North America that was recently found infesting white fringetree (Chionanthus virginicus L.). Initial reports of the infestation of white fringetree by emerald ash borer occurred in southwestern Ohio and Chicago, IL. We examined white fringetrees at additional sites in Illinois, Indiana, Ohio, and Pennsylvania in Summer and Fall 2015 and Winter 2016 for emerald ash borer infestation. Our aim was to examine white fringetrees at a limited number of sites with emerald ash borer infestation and to relate tree size, crown dieback, epicormic sprouting, tree sex, and adjacency to ash or white fringetrees with the likelihood of beetle infestation. A higher proportion of infested trees exhibited epicormic sprouting and the likelihood that a tree was infested increased with increasing crown dieback, variables that may be both predictors and responses to attack. The proportion of trees infested with emerald ash borer increased with increasing tree size. Signs consistent with emerald ash borer infestation were found in 26% of 178 white fringetrees, with at least one host infested at each site in all states. Infestation rates of white fringetrees increased with the density of white fringetrees at each site. The Chicago Botanic Garden site had a significantly lower infestation (3.7%) than other sites, which may be due to proactive management of ash. Overall, these data indicate white fringetree has been utilized by emerald ash borer throughout their overlapping ranges in the United States in ornamental settings likely due to ecological fitting.

Growth of Larval Agrilus planipennis (Coleoptera: Buprestidae) and Fitness of Tetrastichus planipennisi (Hymenoptera: Eulophidae) in Blue Ash (Fraxinus quadrangulata) and Green Ash (F. pennsylvanica).

Emerald ash borer (Agrilus planipennis Fairmaire) is an invasive primary pest of North American ash (Fraxinus spp.) trees. Blue ash (F. quadrangulata) is less susceptible to emerald ash borer infestations in the forest than other species of North American ash. Whereas other studies have examined adult host preferences, we compared the capacity of emerald ash borer larvae reared from emerald ash borer eggs in the field and in the laboratory to survive and grow in blue ash and the more susceptible green ash (F. pennsylvanica). Emerald ash borer larval survivorship was the same on both ash species. Mortality due to wound periderm formation was only observed in living field grown trees, but was low (<4%) in both green and blue ash. No difference in larval mortality in the absence of natural enemies suggests that both green and blue ash can support the development of emerald ash borer. Larvae reared from eggs on blue ash were smaller than on green ash growing in the field and also in bolts that were infested under laboratory conditions. In a laboratory study, parasitism rates of confined Tetrastichus planipennisi were similar on emerald ash borer larvae reared in blue and green ash bolts, as were fitness measures of the parasitoid including brood size, sex ratio, and adult female size. Thus, we postulate that emerald ash borer larvae infesting blue ash could support populations of T. planipennisi and serve as a potential reservoir for this introduced natural enemy after most of the other native ash trees have been killed.