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.     

The persistence of blue ash in the aftermath of emerald ash borer may be due to adult oviposition preferences and reduced larval performance

1. We examined the health of mature blue ash, Fraxinus quadrangulata, in two forests in southwestern Ohio in relation to that of mature white ash, F. americana, and examined the potential importance of oviposition preferences and larval resistance in the persistence of blue ash.
2. Both blue ash and white ash were largely unaffected by emerald ash borer in 2012. By 2018, nearly 90% of the blue ash trees observed in these forests had full or nearly full canopies, as opposed to less than 20% of the white ash encountered in our studies. In 2021, blue ash maintained a similar degree of canopy health as in 2018, but no standing live mature white ashes remained.
3. Bark removals revealed no current or past larval feeding attempts in blue ash in 2018 or 2021, except for one attacked and killed tree in 2018. All white ash trees examined were attacked. In a laboratory bioassay with cut stems, emerald ash borer larvae fed less and grew significantly more slowly on blue ash than on white ash.
4. Both reduced larval performance and reduced adult oviposition likely contribute to the persistence of blue ash in forests devastated by emerald ash borer.

     

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.     

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.     

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.     

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.     

Effects of trap type, placement and ash distribution on emerald ash borer captures in a low density site

Effective methods for early detection of newly established, low density emerald ash borer (Agrilus planipennis Fairmaire) infestations are critically needed in North America. We assessed adult A. planipennis captures on four types of traps in a 16-ha site in central Michigan. The site was divided into 16 blocks, each comprised of four 50- by 50-m cells. Green ash trees (Fraxinus pennsylvanica Marshall) were inventoried by diameter class and ash phloem area was estimated for each cell. One trap type was randomly assigned to each cell in each block. Because initial sampling showed that A. planipennis density was extremely low, infested ash logs were introduced into the center of the site. In total, 87 beetles were captured during the summer. Purple double-decker traps baited with a blend of ash leaf volatiles, Manuka oil, and ethanol captured 65% of all A. planipennis beetles. Similarly baited, green double-decker traps captured 18% of the beetles, whereas sticky bands on girdled trees captured 11% of the beetles. Purple traps baited with Manuka oil and suspended in the canopies of live ash trees captured only 5% of the beetles. At least one beetle was captured on 81% of the purple double-decker traps, 56% of the green double-decker traps, 42% of sticky bands, and 25% of the canopy traps. Abundance of ash phloem near traps had no effect on captures and trap location and sun exposure had only weak effects on captures. Twelve girdled and 29 nongirdled trees were felled and sampled in winter. Current-year larvae were present in 100% of the girdled trees and 72% of the nongirdled trees, but larval density was five times higher on girdled than nongirdled trees.     

Effects of rearing conditions on reproduction of Spathius agrili (Hymenoptera: Braconidae), a parasitoid of the emerald ash borer (Coleoptera: Buprestidae)

Spathius agrili Yang (Hymenoptera: Braconidae) can be successfully reared on emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), larvae feeding in chambers drilled in small ash twigs that are wrapped with floral tape. Females maintained in groups with males for one week can receive enough sperm for production of female progeny throughout their lives. Volatiles released by emerald ash borer adults feeding on ash foliage increased parasitoid fecundity over ash foliage alone or no stimulus. The temperature at which the parasitoids were reared ranged from 20 to 25 degrees C in a daily cycle; however, raising the daily maximum temperature to 28 degrees C did not affect parasitoid longevity or fecundity. Adult females lived between 12 and 127 d, with an average of 60.8 +/- 4.5 d. Males lived slightly longer, with an average of 66 +/- 4.5 d. The first clutch of eggs was laid when the female was between 2 and 42 d old, with the average preoviposition period lasting 11.4 +/- 1.4 or 19.5 +/- 2.0 d in 2007 and 2009 trials, respectively. A higher proportion of the emerald ash borer larvae were feeding and thus attractive to parasitoids in the 2009 trial, and female S. agrili laid an average of 9.5 +/- 1.0 clutches containing 5.4 +/- 0.2 eggs, for an average of 51.2 eggs per female. Approximately three quarters of the progeny were female. The number of eggs per clutch was significantly greater when deposited on larger emerald ash borer larvae, further highlighting the need for quality larvae in rearing. Chilling S. agrili pupae at 10 degrees C to stockpile them for summer release was not successful; chilling resulted in lower survival and lower fecundity of emerging progeny. Female S. agrili proved capable of attacking emerald ash borer larvae through even the thickest bark of an ash tree that was 30-cm diameter at breast height. Even emerald ash borer larvae that were creating overwintering chambers in the outer sapwood of the tree were successfully attacked, suggesting that S. agrili could be reared on field collected logs infested with emerald ash borer.     

The overwintering physiology of the emerald ash borer, Agrilus planipennis fairmaire (coleoptera: buprestidae)

Ability to survive cold is an important factor in determining northern range limits of insects. The emerald ash borer (Agrilus planipennis) is an invasive beetle introduced from Asia that is causing extensive damage to ash trees in North America, but little is known about its cold tolerance. Herein, the cold tolerance strategy and mechanisms involved in the cold tolerance of the emerald ash borer were investigated, and seasonal changes in these mechanisms monitored. The majority of emerald ash borers survive winter as freeze-intolerant prepupae. In winter, A. planipennis prepupae have low supercooling points (∼−30 °C), which they achieve by accumulating high concentrations of glycerol (∼4 M) in their body fluids and by the synthesis of antifreeze agents. Cuticular waxes reduce inoculation from external ice. This is the first comprehensive study of seasonal changes in cold tolerance in a buprestid beetle.     

Dieback and recovery in poplar and attack by the hornet clearwing moth, Sesia apiformis (Clerck) (Lepidoptera: Sesiidae)

A survey of 801 poplar trees in central east England in 1999 demonstrated a correlation between crown dieback and infestation by hornet clearwing moth (Sesia apiformis), induced by a period of drought in 1995-1996. To determine whether trees colonised by S. apiformis would subsequently deteriorate and die or whether they could recover despite infestation and damage to the stem, all trees in the original survey were re-assessed in 2001, 2003 and 2005. The repeat surveys showed that trees with 70% or less crown dieback in 1999 replaced their canopy and generally improved by 2005, irrespective of the numbers of S. apiformis in the stem, whereas trees that had 75% or more crown dieback in 1999 either died or declined further. The presence of S. apiformis did not prevent tree recovery, and there was little evidence that infestation slowed the rate of recovery. Populations of S. apiformis, measured in terms of the numbers of adult emergence holes visible in the base of the trees, decreased between 2001 and 2005 at the same time as the amount of dieback visible in the canopy of the poplars markedly decreased. However, the fall in numbers of emergence holes at this time reflected a decline in larval establishment 2-3 years earlier, and indicated that the moth population had responded to a more rapid restoration in the internal state of the trees, which was not reflected immediately by the gradual replacement of dead branches and reduction in dieback symptoms.     

Ashes in Europe are in danger: the invasive range of Agrilus planipennis in European Russia is expanding

The emerald ash borer, Agrilus planipennis (Coleoptera, Buprestidae), is a pest of ash native to Asia. This major stem borer has killed millions of ash trees in North America. It was first found in Europe in 2003 in the city of Moscow. Now it is rapidly spreading in European Russia. In 2012 A. planipennis was found in the Tula, Kaluga, and Smolensk regions. A survey of green plantations in 22 localities in 2013 has revealed that A. planipennis occurs also in the Tver, Orel, Voronezh, Tambov, and Yaroslavl regions. It occurs 230 km northeast, 350 km southeast, and 460 km south from Moscow. Most ashes in the Moscow region, both alien American Fraxinus pennsylvanica and the indiginous European ash Fraxinus excelsior, are dying or already dead. Thousands of trees in other regions are seriously damaged. The pest will cross the western border of Russia soon. It represents a serious threat for ashes in other European countries.     

Biology and damage traits of emerald ash borer planipennis Fairmaire）in China

Emerald ash borer （Agrilus planipennis Fairmaire） （Coleoptera： Buprestidae） is a major stem borer of ash （Fraxinus spp.）. It is univoltine in Tianjin, while it is semivoltine in Heilongjiang Province, and both univoltine and semivoltine in Changchun, Jilin Province, where the majority is univoltine. The longevity of emerald ash borer adults is 17.2 ± 4.6 days （n = 45）, eggs 9.0 5：1.1 days （n = 103）, univoltine larvae 308 days, semivoltine larvae 673 days, and pupae 61.2 ± 1.6 days （n = 45）. It takes about 100 days from the time larvae bore into the phloem to when they complete the pupal cell. In a 10-year-old velvet ash （Fraxinus velutina Tort.） plantation in Tianjin, emerald ash borer preferred to oviposit on the regions of boles from 50-150 cm above ground, accounting for 76.7% of the total girdling. Girdling on the south side of the tree boles accounted for 43.40% of the total girdling. The emerald ash borer population density is higher at the edge of the plantation compared with the center.     

Role of emerald ash borer (Coleoptera: Buprestidae) larval vibrations in host-quality assessment by Tetrastichus planipennisi (Hymenoptera: Eulophidae)

The biological control agent Tetrastichus planipennisi Yang (Hymenoptera: Eulophidae) is a gregarious larval endoparasitoid of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), an invasive cambium-feeding species responsible for recent, widespread mortality of ash (Fraxinus spp.) in North America. T. planipennisi is known to prefer late-instar emerald ash borer, but the cues used to assess host size by this species and most other parasitoids of concealed hosts remain unknown. We sought to test whether vibrations produced by feeding emerald ash borer vary with larval size and whether there are any correlations between these cues and T. planipennisi progeny number (i.e., brood size) and sex ratio. The amplitudes and rates of 3-30-ms vibrational impulses produced by emerald ash borer larvae of various sizes were measured in the laboratory before presenting the larvae to T. planipennisi. Impulse-rate did not vary with emerald ash borer size, but vibration amplitude was significantly higher for large larvae than for small larvae. T. planipennisi produced a significantly higher proportion of female offspring from large hosts than small hosts and was shown in previous work to produce more offspring overall from large hosts. There were no significant correlations, however, between the T. planipennisi progeny data and the emerald ash borer sound data. Because vibration amplitude varied significantly with host size, however, we are unable to entirely reject the hypothesis that T. planipennisi and possibly other parasitoids of concealed hosts use vibrational cues to assess host quality, particularly given the low explanatory potential of other external cues. Internal chemical cues also may be important.     

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.     

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.     

中国白蜡窄吉丁研究进展

白蜡窄吉丁Agrilus planipennis Fairmaire是近年来发生和影响比较严重的国际性检疫害虫,主要危害白蜡属(Fraxinus spp.)树木。它以幼虫在树木的韧皮部、形成层和木质部浅层蛀食,从而切断树木的营养输导组织,导致树木衰弱死亡。为了深入了解该害虫,探索合理的治理策略,本文对国内外有关白蜡窄吉丁的生物学特性,包括生活习性、生活史、分布与危害、寄主范围等方面以及防治技术的研究进展进行了系统总结,并讨论了白蜡窄吉丁今后的研究方向和研究趋势。     

Decreased emergence of emerald ash borer from ash treated with methyl jasmonate is associated with induction of general defense traits and the toxic phenolic compound verbascoside

The emerald ash borer (EAB; Agrilus planipennis Fairmaire) is causing widespread mortality of ash (Fraxinus spp.) in North America. To date, no mechanisms of host resistance have been identified against this pest. Methyl jasmonate was applied to susceptible North American and resistant Asian ash species to determine if it can elicit induced responses in bark that enhance resistance to EAB. In particular, phenolic compounds, lignin, and defense-related proteins were quantified, and compounds associated with resistance were subsequently tested directly against EAB larvae in bioassays with artificial diet. MeJA application decreased adult emergence in susceptible ash species, comparable to levels achieved by insecticide application. Concentration of the phenolic compound verbascoside sharply increased after MeJA application to green and white ash. When incorporated in an artificial diet, verbascoside decreased survival and growth of EAB neonates in a dose-dependent fashion. Lignin and trypsin inhibitors were also induced by MeJA, and analogs of both compounds reduced growth of EAB larvae in artificial diets. We conclude that the application of MeJA prior to EAB attack has the ability to enhance resistance of susceptible ash trees by inducing endogenous plant defenses, and report evidence that induction of verbascoside is a mechanism of resistance to EAB.     

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

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

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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.