Emerald ash borer | |
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Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Arthropoda |
Class: | Insecta |
Order: | Coleoptera |
Suborder: | Polyphaga |
Infraorder: | Elateriformia |
Family: | Buprestidae |
Genus: | Agrilus |
Species: | A. planipennis |
Binomial name | |
Agrilus planipennis Fairmaire, 1888 | |
Synonyms | |
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The emerald ash borer (Agrilus planipennis), also known by the acronym EAB, is a green buprestid or jewel beetle native to north-eastern Asia that feeds on ash species (Fraxinus spp.). Females lay eggs in bark crevices on ash trees, and larvae feed underneath the bark of ash trees to emerge as adults in one to two years. In its native range, it is typically found at low densities and does not cause significant damage to trees native to the area. Outside its native range, it is an invasive species and is highly destructive to ash trees native to Europe and North America. Before it was found in North America, very little was known about the emerald ash borer in its native range; this has resulted in much of the research on its biology being focused in North America. Local governments in North America are attempting to control it by monitoring its spread, diversifying tree species, and through the use of insecticides and biological control.
French priest and naturalist Armand David collected a specimen of the species during one of his trips through Imperial China in the 1860s and 1870s. He found the beetle in Beijing and sent it to France, where the first brief description of Agrilus planipennis by the entomologist Léon Fairmaire was published in the Revue d'Entomologie in 1888. [2] Unaware of Fairmaire's description, a separate description naming the species as Agrilus marcopoli was published in 1930 by Jan Obenberger. [2]
Adult beetles are typically bright metallic green and about 8.5 mm (0.33 in) long and 1.6 mm (0.063 in) wide. Elytra are typically a darker green, but can also have copper hues. Emerald ash borer is the only North American species of Agrilus with a bright red upper abdomen when viewed with the wings and elytra spread. The species also has a small spine found at the tip of the abdomen and serrate antennae that begin at the fourth antennal segment. [3] They leave tracks in the trees they damage below the bark that are sometimes visible. [4] Adult beetles of other species can often be misidentified by the public. [5] [6]
The emerald ash borer life cycle can occur over one or two years depending on the time of year of oviposition, the health of the tree, and temperature. [7]
After 400–500 accumulated degree-days above 10 °C (50 °F), adults begin to emerge from trees in late spring, and peak emergence occurs around 1,000 degree-days. After emergence, adults feed for one week on ash leaves in the canopy before mating, but cause little defoliation in the process. [8] Males hover around trees, locate females by visual cues, and drop directly onto the female to mate. Mating can last 50 minutes, and females may mate with multiple males over their lifespan. [9] A typical female can live around six weeks and lay approximately 40–70 eggs, but females that live longer can lay up to 200 eggs. [8]
Eggs are deposited between bark crevices, flakes, or cracks and hatch about two weeks later. Eggs are approximately 0.6 to 1.0 mm (0.02 to 0.04 in) in diameter, and are initially white, but later turn reddish-brown if fertile. [8] [7] After hatching, larvae chew through the bark to the inner phloem, cambium, and outer xylem where they feed and develop. [9] Emerald ash borer has four larval instars. By feeding, larvae create long serpentine galleries. Fully mature fourth-instar larvae are 26 to 32 mm (1.0 to 1.3 in) long. [7] In fall, mature fourth-instars excavate chambers about 1.25 cm (0.49 in) into the sapwood or outer bark where they fold into a J-shape. [9] These J-shaped larvae shorten into prepupae and develop into pupae and adults the following spring. To exit the tree, adults chew holes from their chamber through the bark, which leaves a characteristic D-shaped exit hole. Immature larvae can overwinter in their larval gallery, but can require an additional summer of feeding before overwintering again and emerging as adults the following spring. [7] This two-year life cycle is more common in cool climates, such as European Russia. [10]
The native range of the emerald ash borer is temperate north-eastern Asia, which includes Russia, Mongolia, northern China, Japan, and Korea. [11] [10]
The beetle is invasive in North America where it has a core population in Michigan and surrounding states and provinces. Populations are more scattered outside the core area, and the edges of its known distribution range north to Ontario, south to northern Louisiana, west to Nebraska, and east to New Brunswick. [12] [13] Satellite populations also exist in Colorado and in the Pacific Northwest in Oregon. [14] [15] [16] In eastern Europe, a population was found in Moscow in 2003. [10] From 2003 to 2016, this population has spread west towards the European Union at up to 40 km (25 mi) per year and is expected to reach central Europe between 2031 and 2036. [17] [18] [10] Although not recorded from the European Union as of 2019, it has already spread to far eastern Ukraine from neighboring Russia. [19] [20] [21] [22]
In its native range, emerald ash borer is only a nuisance pest on native trees, as population densities typically do not reach levels lethal to healthy trees. [23] In China, it infests native Fraxinus chinensis , F. mandshurica, and F. rhynchophylla; in Japan it also infests F. japonica and F. lanuginosa. [10]
Emerald ash borer primarily infest and can cause significant damage to ash species including green ash ( F. pennsylvanica ), black ash ( F. nigra ), white ash ( F. americana ), and blue ash ( F. quadrangulata ) in North America. [24] In Europe, F. excelsior is the main ash species colonized, which is moderately resistant to emerald ash borer infestation. [10] [25] Ash susceptibility can vary depending on the attractiveness of chemical volatiles to adults, or the ability of larvae to detoxify phenolic compounds. [9] Emerald ash borer has also been found infesting white fringe tree in North America, which is a non-ash host, but it is unclear whether the trees were healthy when first infested, or were already in decline because of drought. [9] [26] Another non-ash host has also been discovered, Olea europaea , albeit in a lab setting. [27]
Adults prefer to lay eggs on open grown or stressed ash but readily lay eggs on healthy trees amongst other tree species. Ashes that grow in pure stands, whether naturally occurring or in landscaping, are more prone to attack than isolated trees or ones located in mixed forest stands. Ashes used in landscaping also tend to be subjected to higher amounts of environmental stresses including compacted soil, lack of moisture, heating effects from urban islands, road salt, and pollution, which may also reduce their resistance to the borer. Furthermore, most ashes used in landscaping were produced from a handful of cultivars, resulting in low genetic diversity. [9] Young trees with bark between 1.5 mm (0.059 in) to 5 mm (0.20 in) are preferred. [10] Both males and females use leaf volatiles and sesquiterpenes in the bark to locate hosts. [9] Damage occurs in infested trees by larval feeding. The serpentine feeding galleries of the larvae disrupt the flow of nutrients and water, effectively girdling, thus killing the tree, as it is no longer able to transport sufficient water and nutrients to the leaves to survive. Girdled ashes will often attempt to regenerate through stump sprouting, and there is evidence that stressed trees may also generate higher than normal seed crops as an emergency measure. [8]
Outside its native range, emerald ash borer is an invasive species that is highly destructive to ash trees in its introduced range. [11] Before emerald ash borer was found in North America, very little was known about the insect in its native range aside from a short description of life-history traits and taxonomic descriptions, which resulted in focused research on its biology in North America. [8] The insect was first identified in Canton, Michigan (near Detroit [28] ), in 2002, [28] but it may have been in the U.S. since the late 1980s. [29] It is suspected that it was introduced from overseas in shipping materials such as packing crates. [28]
Without factors that would normally suppress emerald ash borer populations in its native range (e.g., resistant trees, predators, and parasitoid wasps), populations can quickly rise to damaging levels. [8] After initial infestation, all ash trees are expected to die in an area within 10 years without control measures. [8] Every North American ash species has susceptibility to emerald ash borer, as North American species planted in China also have high mortality from infestations, but some Asian ash species are resistant, including F. baroniana, F. chinensis, F. floribunda, F. mandshurica, and F. platypoda . [30] [31] [32]
Green ash and black ash trees are preferred by emerald ash borer. White ash is also killed rapidly but usually only after all green and black ash trees are eliminated. Blue ash is known to exhibit a higher degree of resistance to emerald ash borer, which is believed to be caused by the high tannin content in the leaves making the foliage unpalatable to the insect. While most Asian ashes have evolved this defense, it is absent from American species other than blue ash. Researchers have examined populations of so-called "lingering ash", trees that survived ash borer attack with little or no damage, as a means of grafting or breeding new, resistant stock. Many of these lingering ashes were found to have unusual phenotypes that may result in increased resistance. Aside from their higher tannin content, Asian ashes also employ natural defenses to repel, trap, and kill emerald ash borer larvae. Although studies of American ashes have suggested that they are capable of mustering similar defensive mechanisms, the trees do not appear to recognize when they are under attack. [33] Many of the specialized predators and parasitoids that suppressed emerald ash borer in Asia were not present in North America. Predators and parasitoids native to North America do not sufficiently suppress emerald ash borer, so populations continue to grow. Birds such as woodpeckers feed on emerald ash borer larva, although the adult beetles have not been used by any American fauna as food. [8] Emerald ash borer populations can spread between 2.5 to 20 km (1.6 to 12.4 mi) per year. [8] It primarily spreads through flight or by transportation of ash bark containing products such as firewood or nursery stock, which allows it to reach new areas and create satellite populations outside of the main infestation. [8] [10]
Other factors can limit spread. Winter temperatures of approximately −38 °C (−36 °F) limit range expansion, [34] [35] and overwintering emerald ash borer survive down to average temperatures of −30 °C (−22 °F) because of antifreeze chemicals in the body and insulation provided by tree bark. [10] Larvae can also survive high heat up to 53 °C (127 °F). Conversely, much like ashes grown in the nursery trade, the population of emerald ash borer in North America is believed to have originated from a single group of insects from central China and also exhibits low genetic diversity. [10]
North American predators and parasitoids can occasionally cause high emerald ash borer mortality, but generally offer only limited control. Mortality from native woodpeckers is variable. Parasitism by parasitoids such as Atanycolus cappaerti can be high, but overall such control is generally low. [8]
The United States Department of Agriculture's Animal and Plant Health Inspection Service published a rule on December 14, 2020—to take effect one month later, January 14, 2021—ending all EAB quarantine activities in the United States due to ineffectiveness so far. [36] [37] Other means will be used instead, especially biological controls (see §Biological control below). [36] [37]
These insects have managed to eliminate close to 300,000 Ash trees in the National Capital Region in only nine years. This leaves only 80,000 ash trees left standing either due to luck or to some amount of resistance to the beetles. These forests used to have an extremely dense Ash population, having 17-18 trees per Hectare. Now, there are only 5-6 trees per Hectare. This illustrates extremely well the overall destructive power of the Emerald Ash Borer and the relevance to the everyday person. Something extremely important to note about this severe loss of Ash trees is the effect that it has on the ecosystem of that area. Swamplands that used to be home many Ash forest have now become shrublands, completely changing the ecosystem of that area permanently. The impact this has on the wildlife is extreme because of all the animals that used every part of the tree as refuge, such as birds in the foliage, small rodents amongst the roots, etc. [38]
Emerald ash borer threatens the entire North American genus Fraxinus . It has killed tens of millions of ash trees so far and threatens to kill most of the 8.7 billion ash trees throughout North America. [12] Emerald ash borer kills young trees several years before reaching their seeding age of 10 years. [8] In both North America and Europe, the loss of ash from an ecosystem can result in increased numbers of invasive plants, changes in soil nutrients, and effects on species that feed on ash. [10]
Damage and efforts to control the spread of emerald ash borer have affected businesses that sell ash trees or wood products, property owners, and local or state governments. [8] Quarantines can limit the transport of ash trees and products, but economic impacts are especially high for urban and residential areas because of treatment or removal costs and decreased land value from dying trees. [39] Costs for managing these trees can fall upon homeowners or local municipalities. For municipalities, removing large numbers of dead or infested trees at once is costly, so slowing down the rate at which trees die through removing known infested trees and treating trees with insecticides can allow local governments more time to plan, remove, and replace trees that would eventually die. This strategy saves money as it would cost $10.7 billion in urban areas of 25 states over 10 years, while removing and replacing all ash trees in these same areas at once would cost $25 billion [39] [40] (with another estimate putting the removal alone at $20–60 billion). [28] Some urban areas such as Minneapolis have large amounts of ash with slightly more than 20% of their urban forest as ash. [41]
In areas where emerald ash borer has not yet been detected, surveys are used to monitor for new infestations. Visual surveys are used to find ash trees displaying emerald ash borer damage, and traps with colors attractive to emerald ash borer, such as purple or green, are hung in trees as part of a monitoring program. [8] These traps can also have volatile pheromones applied to them that attract primarily males. [9]
Sometimes trees are girdled to act as trap trees to monitor for emerald ash borer. The stressed tree attracts egg-laying females in the spring, and trees can be debarked in the fall to search for larvae. [8] If detected, an area is often placed under a quarantine to prevent infested wood material from causing new infestations. [29] [8] Further control measures are then taken within the area to slow population growth by reducing beetle numbers, preventing them from reaching reproductive maturity and dispersing, and reducing the abundance of ash trees. [8]
Government agencies in both the U.S. and Canada have utilized a native species of parasitoid wasp, Cerceris fumipennis , as a means of detecting areas to which emerald ash borer has spread. The females of these wasps hunt other jewel beetles and emerald ash borer if it is present. The wasps stun the beetles and carry them back to their burrows in the ground where they are stored until the wasps’ eggs hatch and the wasp larvae feed on the beetles. Volunteers catch the wasps as they return to their burrows carrying the beetles to determine whether emerald ash borer is present. This methodology is known as biological surveillance, as opposed to biological control, because it does not appear that the wasps have a significant negative impact on emerald ash borer populations. [42]
In areas where emerald ash borer is non-native and invasive, quarantines, infested tree removal, insecticides, and biological control are used to reduce damage to ash trees.
Once an infestation is detected, quarantines are typically imposed by state, or previously, national government agencies disallowing transport of ash firewood or live plants outside of these areas without permits indicating the material has been inspected or treated (i.e., heat treatment or wood chipping) to ensure no live emerald ash borer are present in the bark and phloem. [29] [43] In urban areas, trees are often removed once an infestation is found to reduce emerald ash borer population densities and the likelihood of further spread. Urban ash are typically replaced with non-ash species such as maple, oak, or linden to limit food sources. [44] In rural areas, trees can be harvested for lumber or firewood to reduce ash stand density, but quarantines may apply for this material, especially in areas where the material could be infested. [45]
Kentucky Extension specialists suggest selecting uncommon species to replace removed ashes in the landscape. [46] Previous generations created monocultures by planting ash trees in an overabundance, a factor in the extent of the devastation caused by the emerald ash borer. Favoring instead a diversity in species helps keep urban forests healthy. University of Kentucky scientists suggest choosing monotypic species such as the pawpaw, yellowwood, Franklin tree, Kentucky coffeetree, Osage orange, sourwood, and bald cypress.
Insecticides with active ingredients such as azadirachtin, imidacloprid, emamectin benzoate, and dinotefuran are currently used. Dinotefuran and imidacloprid are systemic (i.e., incorporated into the tree) and remain effective for one to three years depending on the product. [8] [47] [48] Insecticides are typically only considered a viable option in urban areas with high value trees near an infestation. [47] Ash trees are primarily treated by direct injection into the tree or soil drench. Some insecticides cannot be applied by homeowners and must be applied by licensed applicators. Damage from emerald ash borer can continue to increase over time even with insecticide applications. [8] Insecticide treatments are not feasible for large forested areas outside of urban areas. [8]
The native range of emerald ash borer in Asia was surveyed for parasitoid species that parasitize emerald ash borer and do not attack other insect species in the hope they would suppress populations when released in North America. [49] Three species imported from China were approved for release by the USDA in 2007 and in Canada in 2013: Spathius agrili , Tetrastichus planipennisi , and Oobius agrili , while Spathius galinae was approved for release in 2015. [50] [51] Excluding Spathius galinae, which has only recently been released, the other three species have been documented parasitizing emerald ash borer larvae one year after release, indicating that they survived the winter, but establishment varied among species and locations. [51] Tetrastichus planipennisi and Oobius agrili established and have had increasing populations in Michigan since 2008; Spathius agrili has had lower establishment success in North America, which could be caused by a lack of available emerald ash borer larvae at the time of adult emergence in spring, limited cold tolerance, and better suitability to regions of North America below the 40th parallel. [51]
The USDA is also assessing the application of Beauveria bassiana , an insect fungal pathogen, for controlling emerald ash borer in conjunction with parasitoid wasps. [52]
Fraxinus, commonly called ash, is a genus of plants in the olive and lilac family, Oleaceae, and comprises 45–65 species of usually medium-to-large trees, most of which are deciduous trees, although some subtropical species are evergreen trees. The genus is widespread throughout much of Europe, Asia, and North America.
Fraxinus quadrangulata, the blue ash, is a species of ash native primarily to the Midwestern United States from Oklahoma to Michigan, as well as the Bluegrass region of Kentucky and the Nashville Basin region of Tennessee. Isolated populations exist in Alabama, Southern Ontario, and small sections of the Appalachian Mountains. It is typically found over calcareous substrates such as limestone, growing on limestone slopes and in moist valley soils, at elevations of 120–600 m (390–1,970 ft).
Fraxinus americana, the white ash or American ash, is a fast-growing species of ash tree native to eastern and central North America.
Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or other natural mechanisms, but typically also involves an active human management role. It can be an important component of integrated pest management (IPM) programs.
The Asian long-horned beetle, also known as the starry sky, sky beetle, or ALB, is native to the Korean Peninsula, northern and southern China, and disputably in northern Japan. This species has now been accidentally introduced into the eastern United States, where it was first discovered in 1996, as well as Canada, and several countries in Europe, including Austria, France, Germany, Italy and UK.
Fraxinus pennsylvanica, the green ash or red ash, is a species of ash native to eastern and central North America, from Nova Scotia west to southeastern Alberta and eastern Colorado, south to northern Florida, and southwest to Oklahoma and eastern Texas. It has spread and become naturalized in much of the western United States and also in Argentina and Europe, from Spain to Russia.
Fraxinus nigra, the black ash, is a species of ash native to much of eastern Canada and the northeastern United States, from western Newfoundland west to southeastern Manitoba, and south to Illinois and northern Virginia. Formerly abundant, as of 2017 the species is threatened with near total extirpation throughout its range within the next century as a result of infestation by an invasive parasitic insect known as the emerald ash borer.
Chionanthus virginicus is a tree native to the savannas and lowlands of the northeastern and southeastern United States, from Massachusetts south to Florida, and west to Oklahoma and Texas.
Fraxinus mandshurica, the Manchurian ash, is a species of Fraxinus native to northeastern Asia in northern China, Korea, Japan and southeastern Russia.
Fraxinus profunda, the pumpkin ash, is a species of ash (Fraxinus) native to eastern North America, where it has a scattered distribution on the Atlantic coastal plain and interior lowland river valleys from the Lake Erie basin in Ontario and New York west to Illinois, southwest to Missouri and southeast to northern Florida. It grows in bottomland habitats, such as swamps, floodplains and riverbanks. It is threatened by the emerald ash borer, an invasive insect which has caused widespread destruction of ash trees in eastern North America.
Cerceris fumipennis, the only species of buprestid-hunting Philanthidae occurring in eastern North America, is found throughout the continental United States east of the Rockies: from Texas and Florida north to Maine, Wyoming, and into Canada. The wasps most often nest in open areas of hard-packed sandy soil surrounded by woody habitat suitable for their buprestid beetle prey.
Tetrastichus planipennisi is a parasitic non-stinging wasp of the family Eulophidae which is native to North Asia. It is a parasitoid of the emerald ash borer, an invasive species which has destroyed tens of millions of ash trees in its introduced range in North America. As part of the campaign against the emerald ash borer (EAB), American scientists in conjunction with the Chinese Academy of Forestry searched since 2003 for its natural enemies in the wild leading to the discovery of several parasitoid wasps, including Tetrastichus planipennisi which is a gregarious endoparasitoid of EAB larvae on Manchurian Ash and has been recorded to attack and kill up to 50 percent of EAB larvae.
Oobius agrili is a parasitic non-stinging wasp of family Encyrtidae which is native to North Asia. It is a parasitoid of the emerald ash borer, an invasive species which has destroyed tens of millions of ash trees in its introduced range in North America. As part of the campaign against the emerald ash borer (EAB), American scientists in conjunction with the Chinese Academy of Forestry searched since 2003 for its natural enemies in the wild leading to the discovery of several parasitoid wasps, including Oobius agrili, which is a solitary egg parasitoid of EAB found on ash trees in Jilin province in 2004; it has been recorded to kill up to 60 percent of EAB eggs.
Spathius agrili is a parasitic non-stinging wasp of family Braconidae which is native to North Asia. It is a parasitoid of the emerald ash borer, an invasive species which has destroyed tens of millions of ash trees in its introduced range in North America. As part of the campaign against the emerald ash borer (EAB), American scientists in conjunction with the Chinese Academy of Forestry began searching in 2003 for its natural enemies in the wild, leading to the discovery of several parasitoid wasp species, including Spathius agrili. S. agrili was discovered in Tianjin, China where it is a prevalent parasitoid of EAB larvae in stands of an introduced ash species, and an endemic ash species. S. agrili has been recorded to attack and kill up to 90 percent of EAB larvae.
Agrilus auroguttatus is a species of jewel beetle in the United States, known by the common name goldspotted oak borer. Native to southeastern Arizona, it is best known for its probably invasive presence in California, destroying stands of oak trees in the Cleveland National Forest in San Diego County. It was originally considered a subspecies of the Central American species Agrilus coxalis, and much of the literature refers to it by this name, but now it is regarded as a separate species, known only from Arizona and California.
Agrilus biguttatus is a species of beetle in the family Buprestidae, the jewel beetles. Common names include oak splendour beetle, oak buprestid beetle, and two-spotted oak borer. This beetle is known as a pest that causes damage to oak trees and is a major factor in oak decline.
Spathius galinae is a parasitoid of the emerald ash borer. The known range of S. galinae extends from the Russian Far East to South Korea. It is currently approved for release in some areas of North America as part of a biological control program against the emerald ash borer.
Platypus cylindrus, commonly known as the oak pinhole borer, is a species of ambrosia beetle in the weevil family Scolytinae. The adults and larvae burrow under the bark of mature oak trees. It is native to Europe.
Species which are not native to a forest ecosystem can act as an agent of disturbance, changing forest dynamics as they invade and spread. Invasive insects and pathogens (diseases) are introduced to the United States through international trade, and spread through means of natural and human-dispersal. Invasive insects and pathogens are a serious threat to many forests in the United States and have decimated populations of several tree species, including American chestnut, American elm, eastern hemlock, whitebark pine, and the native ash species. The loss of these tree species is typically rapid with both short and long-term impacts to the forest ecosystem.
Phasgonophora sulcata is a species of chalcidid wasp in the family Chalcididae. It is a endoparasitoid of Agrilus beetles, and thus used in biological pest control of the emerald ash borer.