Rail transport as a vector of emerald ash borer

1. The emerald ash borer (EAB) Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) is an invasive woodboring beetle native to northeastern Asia that continues to expand its range within North America and European Russia. The insect reproduces within and kills most North American species of ash Fraxinus spp.
2. Because both the adult and larval life stages of EAB are difficult to detect prior to development of tree symptoms, much work has focused on quantifying spread and clarifying the potential movement pathways to improve early detection and monitoring strategies.
3. In the present study, we retrospectively analyzed county‐level infestations of emerald ash borer in the state of Iowa, U.S.A., subsequent to the initial detection of EAB in 2010. Visual data analysis had suggested that new infestations were not in accordance with the expected patterns of establishment along roads, near campgrounds or by large population centres.
4. We found a positive correlation between the establishment and detection of EAB in Iowa counties and the length of railroads in each county. To our knowledge, this is the first statistically significant association between rail pathways and the spread of EAB on the North American continent.

     

Girdling increases survival and growth of emerald ash borer larvae on Manchurian ash

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Drought stress increased survival and development of emerald ash borer larvae on coevolved Manchurian ash and implicates phloem‐based traits in resistance

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Differential utilization of ash phloem by emerald ash borer larvae: ash species and larval stage effects

• 1 Two experiments were performed to determine the extent to which ash species (black, green and white) and larval developmental stage (second, third and fourth instar) affect the efficiency of phloem amino acid utilization by emerald ash borer (EAB) Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) larvae.
• 2 EAB larvae generally utilized green ash amino acids more efficiently than those of the other two species. For example, the concentrations of only six (two essential) and seven (two essential) amino acids were lower in frass from EAB that fed upon black and white ash than in the corresponding phloem, respectively. By contrast, concentrations of 16 (eight essential) amino acids were lower in the frass from EAB that fed upon green ash than in the phloem. In addition, in green ash, the frass : phloem ratios of 13 amino acids were lower than their counterparts in black and white ash.
• 3 The concentrations of non‐essential amino acids glycine and hydroxylproline were greater in frass than in phloem when EAB fed on black ash, although not when EAB fed on green or white ash.
• 4 The concentration of total phenolics (a group of putative defensive compounds to EAB, expressed as antioxidant activity of acetone extraction) was high in EAB frass but even higher in the phloem samples when the data were pooled across ash species and EAB larval stages. This suggests EAB larvae may eliminate phenolics through a combination of direct excretion and enzymatic conversion of phenolics to nonphenolics before excretion. Because the ratio of frass total phenolics to phloem total phenolics in white ash was lower than the ratios in black and green ash, the ability to destroy phenolics or convert them to nonphenolics was greater when EAB larvae fed on white ash.
• 5 Fourth‐instar EAB extracted phloem amino acids, including threonine, more efficiently than third‐instar EAB. The different larval developmental stages of EAB did not differ in their apparent ability to destroy phenolics or convert them to nonphenolics.

     

Optimizing early detection strategies: defining the effective attraction radius of attractants for emerald ash borer Agrilus planipennis Fairmaire

1. Adult emerald ash borers are attracted to green prism traps baited with the ash host volatile (3Z)-hexenol and the sex pheromone of emerald ash borer (3Z)-dodecen-12-olide [(3Z)-lactone]. Quantifying the heretofore unknown range of attraction of such traps would help optimize deployment strategies for early detection.
2. Examining trap captures of traps deployed in pairs at variable distances offers insight into the range of attraction. Recent work has shown the range of attraction can be estimated as half the intertrap distance at which trap catch begins to decrease, which should occur when proximate traps overlap their respective attractive ranges.
3. We estimated these traps' attractive range for emerald ash borer using 98 baited dark green prism traps deployed in pairs, one trap per tree, in an urban park in Saint Paul, Minnesota, USA in summer 2020.
4. We estimate attractive range by fitting a logistic model to trap catch data using Bayesian inferential methods and describe advantages thereof.
5. The attractive range of these baited traps was estimated to be between 16 and 73 m, with a median of 28 m. We recommend that dark green prism traps baited with these semiochemicals be placed 25–35 m apart near high-risk entry points.

     

Effects of grafting on host plant resistance in ash (Fraxinus spp.) to emerald ash borer (Agrilus planipennis Fairmaire)

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Interspecific variation in resistance to emerald ash borer (Coleoptera: Buprestidae) among North American and Asian ash (Fraxinus spp.)

We conducted a 3-yr study to compare the susceptibility of selected North American ash and an Asian ash species to emerald ash borer, Agrilus planipennis Fairmaire, an invasive wood-boring beetle introduced to North America from Asia. Because of a coevolutionary relationship between Asian ashes and emerald ash borer, we hypothesized an Asian ash species, Manchurian ash, is more resistant to the beetle than its North American congeners. Consistent with our hypothesis, Manchurian ash experienced far less mortality and yielded far fewer adult beetles than several cultivars of North American green and white ash. Surprisingly, a black ash (North American) x Manchurian ash hybrid was highly susceptible to emerald ash borer, indicating this cultivar did not inherit emerald ash borer resistance from its Asian parent. A corollary study investigated the efficacy of soil-applied imidacloprid, a systemic, neonicotinoid insecticide, for controlling emerald ash borer in each of the five cultivars. Imidacloprid had no effect on emerald ash borer colonization of Manchurian ash, which was low in untreated and treated trees. In contrast, imidacloprid did enhance survival of the North American and hybrid cultivars and significantly reduced the number of emerald ash borer adults emerging from green and white ash cultivars. We identify a possible mechanism of resistance of Manchurian ash to emerald ash borer, which may prove useful for screening, selecting, and breeding emerald ash borer-resistant ash trees.     

Occurrence of emerald ash borer (Coleoptera: Buprestidae) and biotic factors affecting its immature stages in the Russian Far East

Field surveys were conducted from 2008 to 2011 in the Khabarovsk and Vladivostok regions of Russia to investigate the occurrence of emerald ash borer, Agrilus planipennis Fairmaire, and mortality factors affecting its immature stages. We found emerald ash borer infesting both introduced North American green ash (Fraxinus pennsylvanica Marshall) and native oriental ashes (F. mandshurica Rupr. and F. rhynchophylla Hance) in both regions. Emerald ash borer densities (larvae/m(2) of phloem area) were markedly higher on green ash (11.3-76.7 in the Khabarovsk area and 77-245 in the Vladivostok area) than on artificially stressed Manchurian ash (2.2) or Oriental ash (10-59). Mortality of emerald ash borer larvae caused by different biotic factors (woodpecker predation, host plant resistance and/or undetermined diseases, and parasitism) varied with date, site, and ash species. In general, predation of emerald ash borer larvae by woodpeckers was low. While low rates (3-27%) of emerald ash borer larval mortality were caused by undetermined biotic factors on green ash between 2009 and 2011, higher rates (26-95%) of emerald ash borer larval mortality were caused by putative plant resistance in Oriental ash species in both regions. Little (<1%) parasitism of emerald ash borer larvae was observed in Khabarovsk; however, three hymenopteran parasitoids (Spathius sp., Atanycolus nigriventris Vojnovskaja-Krieger, and Tetrastichus planipennisi Yang) were observed attacking third - fourth instars of emerald ash borer in the Vladivostok area, parasitizing 0-8.3% of emerald ash borer larvae infesting Oriental ash trees and 7.3-62.7% of those on green ash trees (primarily by Spathius sp.) in two of the three study sites. Relevance of these findings to the classical biological control of emerald ash borer in newly invaded regions is discussed.     

Submergence of black ash logs to control emerald ash borer and preserve wood for American Indian basketmaking

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Evaluation of tree mortality and parasitoid recoveries on the contiguous western invasion front of emerald ash borer

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Impacts of the emerald ash borer (Agrilus planipennis Fairmaire) induced ash (Fraxinus spp.) mortality on forest carbon cycling and successional dynamics in the eastern United States

Invasive species are widely recognized as altering species and community dynamics, but their impacts on biogeochemical cycling and ecosystem processes are less understood. The emerald ash borer (Agrilus planipennis Fairmaire) is a phloem feeding beetle that was inadvertently introduced to the US in the 1990s and relies solely on ash trees (Fraxinus spp.) to complete its life cycle. Ash trees have a wide geographic distribution and are an important component of many different forest types in the US. The larval feeding behavior of the emerald ash borer (EAB) effectively girdles the tree’s phloem tissue resulting in tree mortality in as little as 2 years and stand mortality in as little as 5 years. Using the forest inventory and analysis database, we found that forest lands in the lower 48 states hold approximately 8.7 billion ash trees and saplings, which represent ~2.5 % of the aboveground forest carbon mass. Furthermore, we measured tree growth in 7 EAB impacted and 5 non-EAB impacted temperate forests in the Midwestern United States to quantify the impacts of EAB induced tree mortality on tree growth. We hypothesized that the initial C lost would be partly compensated for by the enhanced non-ash tree growth in EAB-impacted regions relative to non-EAB impacted regions. The EAB disturbance enhanced growth of non-ash trees in the EAB impacted region relative to the non-EAB impacted region. Results also indicate that in EAB impacted areas, growth of trees from the genera Acer and Ulmus responded most positively. Finally, we quantified annual biometric net primary productivity of the EAB impacted forests and compared these quantities to modeled growth of these forests in the absence of EAB and found that large scale ash tree mortality has reduced short term regional forest productivity. The loss of ash biometric net primary productivity is, in part compensated by enhanced growth of non-ash species. As expected, EAB disturbance severity was greater in forests with higher basal areas of ash. This study illustrates the ecosystem and regional scale impacts of invasive pest-induced disturbance on biogeochemical cycling and forest species composition.     

Dispersal of the emerald ash borer, Agrilus planipennis, in newly-colonized sites

• 1 Emerald ash borer Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) is an invasive forest insect pest threatening more than 8 billion ash (Fraxinus spp.) trees in North America. Development of effective survey methods and strategies to slow the spread of A. planipennis requires an understanding of dispersal, particularly in recently established satellite populations.
• 2 We assessed the dispersal of A. planipennis beetles over a single generation at two sites by intensively sampling ash trees at known distances from infested ash logs, the point source of the infestations. Larval density was recorded from more than 100 trees at each site.
• 3 Density of A. planipennis larvae by distance for one site was fit to the Ricker function, inverse power function, and the negative exponential function using a maximum likelihood approach. The prediction of the best model, a negative exponential function, was compared with the results from both sites.
• 4 The present study demonstrates that larval densities rapidly declined with distance, and that most larvae (88.9 and 90.3%) were on trees within 100 m of the emergence point of the adults at each site. The larval distribution pattern observed at both sites was adequately described by the negative exponential function.

     

Measuring the impact of biotic factors on populations of immature emerald ash borers (Coleoptera: Buprestidae)

Cohorts of emerald ash borer larvae, Agrilus planipennis Fairmaire, were experimentally established in July of 2008 on healthy green ash (Fraxinus pennsylvanica) trees in two wooded plots at each of three sites near Lansing, MI, by caging gravid emerald ash borer females or placing laboratory-reared eggs on trunks (0.5-2 m above the ground) of selected trees. One plot at each site was randomly chosen for release of two introduced larval parasitoids, Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) and Spathius agrili Yang (Hymenoptera: Braconidae), whereas the other served as the control. Stage-specific mortality factors and rates were measured for all experimentally established cohorts and for associated wild (i.e., naturally occurring) emerald ash borer immature stages via destructive sampling of 2.5 m (above the ground) trunk sections of cohort-bearing trees in the spring and fall of 2009. Host tree defense was the most important mortality factor, causing 32.0 to 41.1% mortality in the experimental cohorts and 17.5 to 21.5% in wild emerald ash borer stages by spring 2009, and 16.1 to 29% for the remaining experimental cohorts, and 9.9 to 11.8% for wild immature emerald ash borer stages by fall 2009. Woodpecker predation was the second most important factor, inflicting no mortality in the experimental cohorts but causing 5.0 to 5.6% mortality to associated wild emerald ash borer stages by spring 2009 and 9.2 to 12.8% and 3.2 to 17.7%, respectively, for experimental cohorts and wild emerald ash borer stages by fall 2009. Mortality from disease in both the experimental and wild cohorts was low (<3%) in both the spring and fall sample periods. In the fall 2009 samples, ≈ 1.5% of experimental cohorts and 0.8% of the wild emerald ash borer stages were parasitized by T. planipennisi. While there were no significant differences in mortality rates because of parasitism between parasitoid-release and control plots, T. planipennisi was detected in each of the three release sites by the end of the study but was not detected in the experimental cohorts or associated wild larvae in any of the three control plots.     

Dramatic expansion of the range of the invasive ash pest,buprestid beetle Agrilus planipennis Fairmaire, 1888 (Coleoptera,Buprestidae) in European Russia

The emerald ash borer (Agrilus planipennis Fairmaire, 1888) is a dangerous invasive pest of ashes. It was previously believed that in European Russia it occurs only in Moscow, Moscow Province, and in the eastern regions of Smolensk Province. An examination of ash trees in 12 cities of European Russia has revealed a much wider area of the emerald ash borer invasion. The pest was found in Konakovo (Tver Province), Tula, Kaluga, Orel, and Voronezh. We have found that A. planipennis damages not only Fraxinus pennsylvanica (an American species which is commonly planted in cities), but also the aborigine European ash Fraxinus excelsior. Ashes in the European forests and in the protective forest belts are badly endangered.     

Progression of ash canopy thinning and dieback outward from the initial infestation of emerald ash borer (Coleoptera: Buprestidae) in southeastern Michigan

Our objective was to characterize the rate at which ash (Fraxinus spp.) trees decline in areas adjacent to the leading edge of visible ash canopy thinning due to emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). Trees in southeastern Michigan were surveyed from 2003 to 2006 for canopy thinning and dieback by comparing survey trees with a set of 11 standard photographs. Freeways stemming from Detroit in all directions were used as survey transects. Between 750 and 1,100 trees were surveyed each year. A rapid method of sampling populations of emerald ash borer was developed by counting emerald ash borer emergence holes with binoculars and then felling trees to validate binocular counts. Approximately 25% of the trees surveyed for canopy thinning in 2005 and 2006 also were sampled for emerald ash borer emergence holes using binoculars. Regression analysis indicates that 41-53% of the variation in ash canopy thinning can be explained by the number of emerald ash borer emergence holes per tree. Emerald ash borer emergence holes were found at every site where ash canopy thinning averaged > 40%. In 2003, ash canopy thinning averaged 40% at a distance of 19.3 km from the epicenter of the emerald ash borer infestation in Canton. By 2006, the point at which ash trees averaged 40% canopy thinning had increased to a distance of 51.2 km away from Canton. Therefore, the point at which ash trees averaged 40% canopy thinning, a state of decline clearly visible to the average person, moved outward at a rate of 10.6 km/yr during this period.     

Interspecific proteomic comparisons reveal ash phloem genes potentially involved in constitutive resistance to the emerald ash borer

The emerald ash borer (Agrilus planipennis) is an invasive wood-boring beetle that has killed millions of ash trees since its accidental introduction to North America. All North American ash species (Fraxinus spp.) that emerald ash borer has encountered so far are susceptible, while an Asian species, Manchurian ash (F. mandshurica), which shares an evolutionary history with emerald ash borer, is resistant. Phylogenetic evidence places North American black ash (F. nigra) and Manchurian ash in the same clade and section, yet black ash is highly susceptible to the emerald ash borer. This contrast provides an opportunity to compare the genetic traits of the two species and identify those with a potential role in defense/resistance. We used Difference Gel Electrophoresis (DIGE) to compare the phloem proteomes of resistant Manchurian to susceptible black, green, and white ash. Differentially expressed proteins associated with the resistant Manchurian ash when compared to the susceptible ash species were identified using nano-LC-MS/MS and putative identities assigned. Proteomic differences were strongly associated with the phylogenetic relationships among the four species. Proteins identified in Manchurian ash potentially associated with its resistance to emerald ash borer include a PR-10 protein, an aspartic protease, a phenylcoumaran benzylic ether reductase (PCBER), and a thylakoid-bound ascorbate peroxidase. Discovery of resistance-related proteins in Asian species will inform approaches in which resistance genes can be introgressed into North American ash species. The generation of resistant North American ash genotypes can be used in forest ecosystem restoration and urban plantings following the wake of the emerald ash borer invasion.     

Effects of the emerald ash borer invasion on the community composition of arthropods associated with ash tree boles in Maryland,U.S.A.

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Feeding by emerald ash borer larvae induces systemic changes in black ash foliar chemistry

The exotic wood-boring pest, emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), has been threatening North American ash (Fraxinus spp.) resources, this being recognized since its first detection in Michigan, USA and Ontario, Canada in 2002. Ash trees are killed by larval feeding in the cambial region, which results in disruption of photosynthate and nutrient translocation. In this study, changes in volatile and non-volatile foliar phytochemicals of potted 2-yr-old black ash, Fraxinus nigra Marshall, seedlings were observed in response to EAB larval feeding in the main stem. EAB larval feeding affected levels of six compounds [hexanal, (E)-2-hexenal, (Z)-3-hexenyl acetate, (E)-β-ocimene, methyl salicylate, and (Z,E)-α-farnesene] with patterns of interaction depending upon compounds of interest and time of observation. Increased methyl salicylate emission suggests similarity in responses induced by EAB larval feeding and other phloem-feeding herbivores. Overall, EAB larval feeding suppressed (Z)-3-hexenyl acetate emission, elevated (E)-β-ocimene emission in the first 30 days, but emissions leveled off thereafter, and generally increased the emission of (Z,E)-α-farnesene. Levels of carbohydrates and phenolics increased overall, while levels of proteins and most amino acids decreased in response to larval feeding. Twenty-three amino acids were consistently detected in the foliage of black ash. The three most abundant amino acids were aspartic acid, glutamic acid, glutamine, while the four least abundant were α-aminobutyric acid, β-aminoisobutyric acid, methionine, and sarcosine. Most (16) foliar free amino acids and 6 of the 9 detected essential amino acids decreased with EAB larval feeding. The ecological consequences of these dynamic phytochemical changes on herbivores harbored by ash trees and potential natural enemies of these herbivores are discussed.     

Economic analysis of emerald ash borer (Coleoptera: Buprestidae) management options

Emerald ash borer, Agrilus planipennis (Fairmaire) (Coleoptera: Buprestidae), plays a significant role in the health and extent of management of native North American ash species in urban forests. An economic analysis of management options was performed to aid decision makers in preparing for likely future infestations. Separate ash tree population valuations were derived from the i-Tree Streets program and the Council of Tree and Landscape Appraisers (CTLA) methodology. A relative economic analysis was used to compare a control option (do-nothing approach, only removing ash trees as they die) to three distinct management options: 1) preemptive removal of all ash trees over a 5 yr period, 2) preemptive removal of all ash trees and replacement with comparable nonash trees, or 3) treating the entire population of ash trees with insecticides to minimize mortality. For each valuation and management option, an annual analysis was performed for both the remaining ash tree population and those lost to emerald ash borer. Retention of ash trees using insecticide treatments typically retained greater urban forest value, followed by doing nothing (control), which was better than preemptive removal and replacement. Preemptive removal without tree replacement, which was the least expensive management option, also provided the lowest net urban forest value over the 20-yr simulation. A "no emerald ash borer" scenario was modeled to further serve as a benchmark for each management option and provide a level of economic justification for regulatory programs aimed at slowing the movement of emerald ash borer.     

Submersion as a tactic to prevent emergence of emerald ash borer Agrilus planipennis from black ash logs

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