Tree species richness and ash density have variable effects on emerald ash borer biological control by woodpeckers and parasitoid wasps in post-invasion white ash stands.

Emerald ash borer (EAB) (Agrilus planipennis Fairmaire) (Coleoptera: Buprestidae) is the most destructive insect to invade North American forests. Identifying habitat features that support EAB natural enemies is necessary to enhance EAB biological control. In many forest ecosystems, tree species diversity has been linked with reduced pest abundance and increases in natural enemy abundance. We assessed the influence of tree species richness, ash density, and proportion of total ash basal area on ash canopy condition, EAB larval densities, and biocontrol by woodpeckers and parasitoids in pairs of healthy and declining overstory (DBH > 10 cm) and recruit-sized ash (DBH 2-10 cm) in 4 post-invasion forests in Michigan, USA. Tree species richness and ash density were not significantly associated with EAB larval densities, ash canopy dieback and transparency, and woodpecker predation of EAB larvae. In declining and healthy overstory ash, woodpeckers killed 38.5 ±â€…3.9% and 13.2 ±â€…3.7% of larvae, respectively, while the native parasitoid Phasgonophora sulcata Westwood killed 15.8 ±â€…3.8% and 8.3 ±â€…3.0% and the introduced parasitoid Spathius galinae Belokobylskij & Strazanac killed 10.8 ±â€…2.5% and 5.0 ±â€…2.6% of EAB larvae. Parasitism by P. sulcata was inversely related to ash density while parasitism by S. galinae was positively associated with ash density. Ash density, but not tree diversity, appears to differentially influence biological control of EAB by parasitoids, but this effect is not associated with reduced EAB densities or improved canopy condition.

Hosts and impacts of elongate hemlock scale (Hemiptera: Diaspididae): A critical review.

Fiorinia externa Ferris, elongate hemlock scale, was inadvertently introduced to North America from Japan. This insect is particularly problematic on hemlock, Tsuga spp., though it has been reported in association with several other conifers. The evidence that other conifers might be hosts, capable of supporting growing populations of the insect, has not been previously reviewed. Our review confirms that F. externa is an oligophagous pest of members of Pinaceae. Although species of Cupressaceae and Taxaceae have been reported as hosts of F. externa, they seem unable to support population growth of this pest. Evidence of the tree-killing potential of the insect, even on suitable hosts, is remarkably scant. The degree of pest risk posed by F. externa with respect to tree mortality in areas beyond the geographic range of hemlock seems modest, but uncertain.

Successful establishment, spread, and impact of the introduced parasitoid Spathius galinae (Hymenoptera: Braconidae) on emerald ash borer (Coleoptera: Buprestidae) populations in postinvasion forests in Michigan.

Spathius galinae is a larval parasitoid native to the Russian Far East that was approved for release in the United States in 2015 for biological control of the emerald ash borer (EAB), Agrilus planipennis, an invasive beetle from Asia responsible for widespread mortality of ash trees (Fraxinus spp.) in North America. From 2015 to 2017, 1,340-1,445 females of S. galinae along with males were released into each release plot, paired with a nonrelease control plot (1-12.5 km apart), at 6 postinvasion forested sites containing abundant pole-sized ash trees in Michigan. By 2018, S. galinae had spread to all but one control plot. Based on the first year that S. galinae was found in trees in each control plot and the distances of those trees to the parasitoid release point within each site, we estimated that S. galinae spread at 3.7 (±1.9) km per year after its initial releases in 2015. The proportion of sampled trees with S. galinae broods, brood densities within sampled trees, and parasitism of EAB larvae increased sharply in both control and release plots after the last field releases in 2017, with the highest parasitism rates (42.8-60.3%) in 2020. Life table analysis showed that S. galinae alone reduced EAB's net population growth rate by 35-55% across sites from 2018 to 2020. These results demonstrate that S. galinae has established an increasing population in Michigan and now plays a significant role in reducing EAB populations in the area.

Postrelease assessment of Oobius agrili (Hymenoptera: Encyrtidae) establishment and persistence in Michigan and the Northeastern United States.

The emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), is an invasive woodboring pest of ash trees (Fraxinus sp.) in North America. Among the Asiatic parasitoids being released for the management of EAB in North America, Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae) is the only EAB egg parasitoid. To date, more than 2.5 million O. agrili have been released in North America; however, few studies have examined its success as a biological control agent of EAB. We conducted studies to assess O. agrili establishment, persistence, spread, and EAB egg parasitism rates in Michigan at the earliest release sites (2007-2010), as well as at more recent release sites (2015-2016) in 3 Northeastern United States (Connecticut, Massachusetts, New York). In both regions, we documented successful O. agrili establishment at all but one release site. In Michigan, O. agrili has persisted at release sites for over a decade and spread to all control sites located 0.6-3.8 km from release sites. Overall, EAB egg parasitism ranged from 1.5% to 51.2% (mean of 21.4%) during 2016-2020 in Michigan and from 2.6% to 29.2% (mean of 16.1%) during 2018-2020 in the Northeastern states. Future research efforts should focus on factors affecting the spatiotemporal variation in EAB egg parasitism rates by O. agrili, as well as its potential range in North America.

An integrative phenology and climatic suitability model for emerald ash borer.

Introduction: Decision support models that predict both when and where to expect emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), are needed for the development and implementation of effective management strategies against this major invasive pest of ash (Fraxinus species) in North America and other regions such as Europe. We present a spatialized model of phenology and climatic suitability for EAB for use in the Degree-Days, Risk, and Phenological event mapping (DDRP) platform, which is an open-source decision support tool to help detect, monitor, and manage invasive threats. Methods: We evaluated the model using presence records from three geographic regions (China, North America, and Europe) and a phenological dataset consisting primarily of observations from the northeastern and midwestern United States. To demonstrate the model, we produced phenological event maps for a recent year and tested for trends in EAB's phenology and potential distribution over a recent 20-year period. Results: Overall, the model exhibited strong performance. Presence was correctly estimated for over 99% of presence records and predicted dates of adult phenological events corresponded closely with observed dates, with a mean absolute error of ca. 7 days and low estimates of bias. Climate stresses were insufficient to exclude EAB from areas with native Fraxinus species in North America and Europe; however, extreme weather events, climate warming, and an inability for EAB to complete its life cycle may reduce suitability for some areas. Significant trends toward earlier adult emergence over 20 years occurred in only some areas. Discussion: Near real-time model forecasts for the conterminous United States are available at two websites to provide end-users with decision-support for surveillance and management of this invasive pest. Forecasts of adult emergence and egg hatch are particularly relevant for surveillance and for managing existing populations with pesticide treatments and parasitoid introductions.

Mortality of Bark- and Wood-boring Beetles (Coleoptera: Buprestidae, Cerambycidae, and Curculionidae) in Naturally Infested Heat-treated Ash, Birch, Oak, and Pine Bolts.

Firewood and wood packaging material (WPM) are major pathways for moving bark- and wood-infesting insects (borers). Heat treatment regulations for interstate firewood movement vary among U.S. states: from 56°C for 30 min to 71.1°C for 75 min. Current WPM international standards (ISPM 15) require heating to a minimum of 56°C for 30 min throughout the profile of the wood. Using bolts from infested ash (Fraxinus), birch (Betula), oak (Quercus), and pine (Pinus) trees in Michigan, we assessed borer mortality at core temperatures of 50, 53, 56, and 60°C maintained for 30 min in chambers set to 60, 65, 70, or 75°C. After treatment, bolts were monitored for adult emergence and later dissected to determine borer mortality rates. Mortality was high to complete for all heat treatments and increased with both increasing core and chamber temperatures. For the Agrilus (Buprestidae) species tested, there was complete mortality of Agrilus anxius on birch and Agrilus planipennis on ash when core temperatures of 56°C or higher were targeted regardless of chamber temperature. However, on oak, a few Agrilus bilineatus and Agrilus sulcicollis survived in bolts heated to 56°C in chambers at 60 and 65°C, and some A. sulcicollis survived in chambers set at 70 and 75°C. Similarly, a few pine-infesting borers survived heating to 56°C at all chamber temperatures. However, there was complete mortality in all hosts when bolts were heated to 60°C for 30 min, regardless of chamber temperature. Results are discussed in terms of current treatment regulations for firewood and WPM.

A Phenology Model for Simulating Oobius agrili (Hymenoptera: Encyrtidae) Seasonal Voltinism and Synchrony With Emerald Ash Borer Oviposition.

In North America, the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), continues to spread, and its egg parasitoid, Oobius agrili Zhang and Huang (Hymenoptera: Encyrtidae), is being released for emerald ash borer biocontrol well beyond their endemic climatic ranges in China. We developed a multiple cohort rate summation model to simulate O. agrili F0, F1, and F2 generations, and emerald ash borer oviposition for examining host-parasitoid synchrony across a north-south gradient from Duluth, MN (latitude 46.8369, longitude -92.1833) to Shreveport, LA (latitude 32.4469, longitude -93.8242). Temporal occurrences of critical day length for O. agrili diapause induction were integrated into the model. We used O. agrili and emerald ash borer trapping data from south central and northwestern Lower Michigan for model validation. Simulations demonstrated that 1) F0 adult emergence consistently occurred 2-5 d before emerald ash borer oviposition began; 2) F1 adult emergence was most synchronized with peak emerald ash borer oviposition compared with other generations; and 3) emerald ash borer oviposition was complete, or near so, when F2 adult emergence was predicted across the north-south gradient. Comparison of O. agrili trap captures with model simulations demonstrated that primarily two adult O. agrili generations (F0 and F1) emerged per year in Michigan and almost all F2 larvae entered diapause despite day lengths longer than critical day length in south central Michigan. Critical day length varied temporally across the north-south gradient during emergence of O. agrili generations. Determining day lengths perceived by O. agrili larvae in the field should improve model realism for examining spatiotemporal variation in O. agrili population dynamics.

Effects of Extreme Low Winter Temperatures on the Overwintering Survival of the Introduced Larval Parasitoids Spathius galinae and Tetrastichus planipennisi: Implications for Biological Control of Emerald Ash Borer in North America.

Climate change has been linked to shifts in the distribution and phenology of species although little is known about the potential effects that extreme low winter temperatures may have on insect host-parasitoid interactions. In late January 2019, northern regions of the United States experienced a severe cold wave caused by a weakened jet stream, destabilizing the Arctic polar vortex. Approximately 3 mo later at six study sites in southern Michigan and three in southern Connecticut, we sampled the overwintering larvae of the emerald ash borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), and two larval parasitoids, Spathius galinae (Hymenoptera: Braconidae) and Tetrastichus planipennisi (Hymenoptera: Eulophidae), that are being introduced as emerald ash borer biocontrol agents in North America. At these nine study sites, emerald ash borer-infested ash trees and/or saplings were debarked and each overwintering emerald ash borer and parasitoid larva was then examined for cold-induced mortality, as indicated by a brown coloration, flaccid, and watery consistency. In early spring in Michigan, we found 4.5-26% of emerald ash borer larvae, 18-50% of S. galinae larvae, and 8-35% of T. planipennisi larvae were killed by cold. In Connecticut where temperatures were more moderate than in Michigan during the 2019 cold wave, <2% of the larval hosts and parasitoids died from cold injury. Our findings revealed that cold-induced mortality of overwintering larvae of emerald ash borer and its larval parasitoids varied by location and species, with higher mortality of parasitoid larvae in most Michigan sites compared to host larvae. The potential impacts of our findings on the management of emerald ash borer using biocontrol are discussed.

Two new species of Oobius Trjapitzin (Hymenoptera, Encyrtidae) egg parasitoids of Agrilus spp. (Coleoptera, Buprestidae) from the USA, including a key and taxonomic notes on other congeneric Nearctic taxa.

Oobius Trjapitzin (Hymenoptera, Encyrtidae) species are egg parasitoids that are important for the biological control of some Buprestidae and Cerambycidae (Coleoptera). Two species, Oobiusagrili Zhang & Huang and Oobiuslongoi (Siscaro), were introduced into North America for classical biocontrol and have successfully established. Two new native North American species that parasitize eggs of Agrilus spp. (Buprestidae) are described and illustrated from the USA: Oobiusminusculus Triapitsyn & Petrice, sp. n. (Michigan), an egg parasitoid of both Agrilussubcinctus Gory on ash (Fraxinus spp.) and Agrilusegenus Gory on black locust (Robiniapseudoacacia L.) trees, and Oobiuswhiteorum Triapitsyn, sp. n. (Pennsylvania), an egg parasitoid of Agrilusanxius Gory on European white birch (Betulapendula Roth). A taxonomic key and notes on the Nearctic native and introduced Oobius species are also included.

Incidence of bark- and wood-boring insects in firewood: a survey at Michigan's Mackinac Bridge.

Firewood is a major pathway for the inadvertent movement of bark- and wood-infesting insects. After discovery of Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) in southeastern Michigan in 2002, quarantines were enacted including prohibition of transporting firewood across the Mackinac Bridge between Michigan's Lower and Upper peninsulas. Drivers are required to surrender firewood before crossing the bridge. We surveyed recently surrendered firewood in April, July, and September 2008 and categorized it by genus, cross-sectional shape (whole, half, or quarter), approximate age (years since it was a live tree), presence of bark, and evidence of bark- and wood-boring insects. The 1045 pieces of firewood examined represented 21 tree genera: primarily Acer (30%), Quercus (18%), Fraxinus (15%), Ulmus (12%), Betula (5%), and Prunus (5%). Live borers (Bostrichoidea, Brentidae, Buprestidae, Cerambycidae, Cossidae, Curculionidae [Scolytinae and non-Scolytinae], and Siricidae) were found in 23% of the pieces and another 41% had evidence of previous borer infestation. Of the 152 Fraxinus firewood pieces, 13% had evidence of past A. planipennis infestation, but we found no live A. planipennis. We discuss national "don't move firewood" campaigns and U.S. imports of fuelwood. During 1996-2009, the United States imported fuelwood valued at > dollars U.S. 98 million from 34 countries.

Bark- and wood-borer colonization of logs and lumber after heat treatment to ISPM 15 specifications: the role of residual bark.

Wood packaging material (WPM) is a major pathway for international movement of bark- and wood-infesting insects. ISPM 15, the first international standard for treating WPM, was adopted in 2002 and first implemented in the United States in 2006. ISPM 15 allows bark to remain on WPM after treatment, raising concerns that insects could infest after treatment, especially if bark were present. We conducted field studies to evaluate insect infestation of green logs and lumber with varying amounts of bark after heat treatment. In a log study, Cerambycidae and Scolytinae (ambrosia beetles and bark beetles) readily infested and developed in logs with bark after heat treatment. In a lumber study, Cerambycidae and bark beetles laid eggs in all sizes of bark patches tested (approximately 25, 100, 250, and 1,000 cm2) after heat treatment but did not infest control or heat-treated lumber without bark. Cerambycidae completed development only in boards with bark patches of 1,000 cm2, whereas bark beetles completed development on patches of 100, 250, and 1,000 cm2. Survival of bark beetles was greater in square patches (10 by 10 cm) versus rectangular patches (2.5 by 40 cm) of the same surface area (100 cm2). In surveys at six U.S. ports in 2006, 9.4% of 5,945 ISPM 15-marked WPM items contained bark, and 1.2% of 564 ISPM 15-marked WPM items with bark contained live insects of quarantine significance under the bark. It was not possible to determine whether the presence of live insects represented treatment failure or infestation after treatment.

Effects of cutting date, outdoor storage conditions, and splitting on survival of Agrilus planipennis (Coleoptera: Buprestidae) in firewood logs.

The emerald ash borer Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) is an exotic pest of ash (Fraxinus spp.) in North America. We conducted studies in Michigan to evaluate how different tree cutting dates, outdoor storage conditions, and splitting affected A. planipennis survival in firewood logs. In 2002-2003, we cut logs from A. planipennis-infested ash trees each month, from July to October, and stored half of the logs in shade and half in full sun. In 2003-2004, we tested logs cut July-December; stored in sun versus shade; tarped versus untarped; and whole logs versus split logs. For both years, A. planipennis successfully emerged the summer after cutting from logs that represented all treatments and all cutting dates tested. Adult emergence density was significantly lower in logs cut in July and August. In 2003-2004, A. planipennis adult length was significantly shorter, and percentage of mortality was significantly higher for logs cut in August compared with later months. Emergence density was significantly lower for split logs compared with whole logs for all cutting months except for December. There was no significant difference in adult emergence density between logs stored in full sun versus shade in 2002-2003. In 2003-2004, untarped logs in full sun or shade had significantly lower adult emergence densities than tarped logs in the sun or shade. In conclusion, emergence, survival, and size of A. planipennis was significantly reduced if logs were cut early during larval development (July or August); splitting logs and storing them untarped in full sun or shade further reduced adult emergence. No treatment was 100% effective in preventing adult emergence.

Field evaluations of systemic insecticides for control of Anoplophora glabripennis (Coleoptera: Cerambycidae) in China.

Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae), a pest native to China and Korea, was discovered in North America in 1996. Currently, the only reliable strategy available for eradication and control is to cut and chip all infested trees. We evaluated various doses of the systemic insecticides azadirachtin, emamectin benzoate, imidacloprid, and thiacloprid for control of A. glabripennis in naturally infested elms (Ulmus spp.), poplars (Populus spp.), and willows (Salix spp. ) in China between 2000 and 2002. Significantly more dead A. glabripennis adults were found beneath elm and poplar trees treated with imidacloprid (in 2000 and 2001) or thiacloprid (in 2001) and beneath willow trees injected with imidacloprid or thiacloprid (in 2002) compared with control trees. In 2000, 4 mo after injection, the density of live A. glabripennis was significantly reduced in poplar trees treated with imidacloprid (90%) and in willow trees treated with imidacloprid (83%) or emamectin benzoate (71%) compared with controls. In 2001, 9 mo after injection, the density of live A. glabripennis was significantly reduced in poplar (76%) and willow (45%) trees treated with imidacloprid compared with control trees. Similarly, percentage mortality of all life stages of A. glabripennis feeding within trees was significantly higher on poplar trees 4 mo after injection with imidacloprid (64%) in 2000 and on elms (55%) and poplars (63%) 9 mo after injection with imidacloprid in 2001 compared with control trees. Imidacloprid residue levels in leaves and twigs collected at various times from 1 d to 9 mo after injection ranged from 0.27 to 0.46 ppm. Injecting A. glabripennis-infested trees with imidacloprid can result in significant mortality of adults during maturation feeding on leaves and twigs and of all life stages feeding within infested trees. Imidacloprid is translocated rapidly in infested trees and is persistent at lethal levels for several months. Although, injection with imidacloprid does not provide complete control of A. glabripennis, systemic insecticides may prove useful as part of an integrated eradication or management program.

Laboratory evaluation of the toxicity of systemic insecticides for Control of Anoplophora glabripennis and Plectrodera scalator (Coleoptera: Cerambycidae).

Anoplophora glabripennis (Motschulsky) (Coleoptera: Cerambycidae) is one of the most serious nonnative invasive forest insects discovered in North America in recent years. A. glabripennis is regulated by federal quarantines in the United States and Canada and is the subject of eradication programs that involve locating, cutting, and chipping all infested trees. Other control methods are needed to aid in eradication and to form an integrated management program in the event eradication fails. We conducted laboratory bioassays to determine the toxicity of two systemic insecticides, azadirachtin and imidacloprid, for potential control of A. glabripennis and the cottonwood borer, Plectrodera scalator (F.) (Coleoptera: Cerambycidae), a closely related native cerambycid. Larvae of both cerambycid species were fed artificial diet with dilutions of azadirachtin or imidacloprid for 14 wk. Both insecticides exhibited strong antifeedant effects and some toxicity against A. glabripennis and P. scalator larvae. For A. glabripennis, the highest larval mortality at the end of the bioassay was 60% for larvae fed artificial diet treated with azadirachtin (50 ppm) or imidacloprid (1.6 ppm). For P. scalator, the highest larval mortality at the end of the bioassay was 100% for larvae fed artificial diet treated with azadirachtin (50 ppm) or imidacloprid (160 ppm). At 14 wk, the LC50 values for P. scalator were 1.58 and 1.78 ppm for azadirachtin and imidacloprid, respectively. Larvae of both species gained weight when fed diet treated with formulation blanks (inert ingredients) or the water control but lost weight when fed diet treated with increasing concentrations of either azadirachtin or imidacloprid. In a separate experiment, A. glabripennis adults were fed maple twigs treated with high and low concentrations of imidacloprid. A. glabripennis adult mortality reached 100% after 13 d on twigs treated with 150 ppm imidacloprid and after 20 d on twigs treated with 15 ppm imidacloprid. There was no visible feeding by A. glabripennis adults on twigs treated at the higher imidacloprid rate, and feeding was significantly reduced for adults placed on twigs treated at the low imidacloprid rate compared with adults on untreated twigs. In summary, imidacloprid and azadirachtin had both antifeedant and toxic effects against A. glabripennis and P. scalator and have potential for use in management programs. Based on our results, the delivery of high and sustained insecticide concentrations will be needed to overcome the antifeedant effects and lengthy lethal time for both larvae and adults exposed to these insecticides.

Tomicus piniperda (Coleoptera: Scolytidae) initial flight and shoot departure along a north-south gradient.

The exotic pine shoot beetle, Tomicus piniperda (L.) (Coleoptera: Scolytidae), established in the north central and northeastern United States (U.S.) and adjacent regions in Canada, is regulated by a federal quarantine that restricts movement of pine material during specific times of the year based on the beetle's life history. Although climatic variation occurs across T. piniperda's range, a single set of dates is used for timing the movement of pine logs. We monitored T. piniperda spring flight, fall shoot departure, and air and internal tree temperatures at three sites along a 300-km north-south gradient in Michigan and Indiana. We also estimated dates for initial spring flight (12 degrees C threshold) and fall shoot departure (0 degrees C threshold) across an 850-km gradient using historical temperature records (1901 to 1999). Average daily temperatures in fall 1997 (8 October to 12 December) and spring 1998 (20 February to 21 April) were 1.8 to 2.4 degrees C colder, respectively, at the northern field site than at the southern field site. Fall shoot departure began at approximately the same time (day 289 to 290) at all three field sites, but complete shoot departure was extended by 3 wk at the southern site (day 336) compared with the northern site (day 317). T. piniperda adults were first captured in funnel traps on calendar day 86 at the northern site and on day 59 at the central and southern field sites. Peak flight occurred at approximately the same time (day 86) at all three sites. Within-shoot temperatures were very similar to air temperatures in the fall and aboveground inside-bark temperatures were similar to air temperatures in the spring. Average predicted dates based on historical temperature records varied by 31 d for initial shoot departure and 84 d for initial spring flight between northern Michigan and southern Indiana. Because considerable variation can occur in T. piniperda behavior across a broad geographic range, dates specified in the U.S. Federal quarantine should be adjusted according to local temperatures.