Forest disturbance by invasive insects and diseases in the United States

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An ash-dominant forest in decline from emerald ash borer damage The Plight of Ash Trees.jpg
An ash-dominant forest in decline from emerald ash borer damage

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 (see extended listing below). The loss of these tree species is typically rapid with both short and long-term impacts to the forest ecosystem. [1]

Contents

Spongy moth spread 1900-2007 Gypsy moth spread 1900-2007.gif
Spongy moth spread 1900-2007
Spongy moth damage in Harper's Ferry, WV Gypsy Moth damage Hapers Ferry.jpg
Spongy moth damage in Harper's Ferry, WV

Pathways and spread

Pathways

International trade is the primary pathway for introduction of invasive insects and pathogens into the United States. [2]

Live plants

The importation of live plants has been the pathway attributed to over 60 percent of the most damaging non-native pests to the United States. [2] Live plants are an especially effective pathway because they provide sustenance to the pest over long journeys on otherwise inhospitable cargo ships. Notable early examples of live plant harbored pests—such as beech scale, chestnut blight, and white pine blister rust—led to the Plant Quarantine Act of 1912 and regulations from the US Department of Agriculture in 1919. Despite these and subsequent regulations, insects and pathogens have continued to be introduced through live plants. [1]

Wood products

Wood products—including wood packaging material—are the second most common pathway, accounting for about 30 percent of the most damaging non-native pests to the United States. These low quality wood products, like crating and pallets, often retain the outer part of the tree (bark and phloem) where insect life can be harbored. [1]

Other

Other less common pathways include "hitch-hiking" directly on cargo ships and containers and in air passenger baggage. [1]

Spread

Once introduced into forests of the United States, invasive insects and pathogens can spread in two ways: natural and human-assisted dispersal. The rate of human-assisted dispersal can be erratic and cause the pest to invade distant, discrete locations. The rate of natural dispersal varies depending on the adaptations of the pest species but is more diffuse, spreading outward from the invaded area. A typical dispersal of invasive insects and pathogens is characterized by both means: diffuse spread coupled with long-distance "jumps" to discrete areas. [3]

Natural dispersal

Pathogens can spread through the air, water, or on insect and animal vectors (hosts). Dutch elm disease was spread by elm bark beetles, yet the tree mortality was caused by a pathogen. [4] Chestnut blight is a fungus spread through wind dispersal and rain splatter; the blight traveled up to 50 miles in a year by natural means. [5] Insect pests, once they reach the adult phase, have the ability to disperse by flight. Emerald ash borer beetles can travel over seven kilometers per day. [6] The Asian long-horned beetle, however, usually stays with the tree it matured on, maintaining localized natural spread. [7]

Human-assisted dispersal

Human-assisted spread can cause invasive insects or pathogens to "jump" to new and often distant locations. The transported pest may then spread by natural means, forming new, discrete populations. [3] A significant means of human-assisted spread is through transportation of infected firewood. For example, discrete populations of emerald ash borer have been traced to remote campsites where natural spread would not have been possible. [8]

Forest impacts

Non-native invasive species can disrupt ecosystems because they do not have natural predators, or other ecological checks-and-balances. Thus, with less competition from native species, non-native populations can explode. [9] Invasive insects and pathogens have eliminated entire tree species from forests of the United States in as little as decades. This rapid change in forest composition has consequences for the entire ecosystem. [1] The extent of the impact depends on characteristics of both the pest and host tree. [10]

Pests differ in their means of attack and extent of damage. For example, defoliating insect (spongy moth) may weaken trees but not directly cause mortality, while boring insects (emerald ash borer) can often cause serious damage. A pathogen may slowly weaken a tree (beech bark disease) or it may cause rapid decline (chestnut blight). A pest may attack a specific species (hemlock woolly adelgid) or it may attack several species (emerald ash borer) or species across many genera (Asian long-horned beetle). [10]

The impacts to the forest also differ depending on characteristics of the host tree. The decline of a more dominant tree (American chestnut) or one that fills a unique role (such as a nitrogen fixer) has a greater impact. A pioneer tree species, one that is able to quickly regenerate after a disturbance, may be able to reassert its role in the forest while a climax tree species may not. [10]

Forest impacts can be divided into two phases: short-term physical disturbance caused directly by the pest, and long-term changes in ecosystem functions in response to the disturbance. [1]

Short-term

Short-term disturbances are physical damages directly caused by the invasive pest and are measured on a time-scale of months to years. Short-term forest productivity is reduced by significant tree mortality. For example, productivity of mixed ash forests affected by emerald ash borer were measured at around 30 percent reduction after the disturbance. Forest-productivity can be reduced even without widespread mortality. During a spongy moth outbreak, entire forests can be defoliated but then recover. However, the energy required for new foliage weakens the trees. Oaks—a favorite food source for spongy moth larva—produce a reduced acorn crop for years after a defoliating event. Animals that rely on acorns are thus affected; but some animals benefit from the spongy moth, namely those that prey on the caterpillars (larva). Understory plants can also benefit from the increased sunlight and rainfall; or they may perish if left too exposed. Nitrogen levels in the soil increase from the insect excrement, dead caterpillars, and dropped foliage. [10]

Other short-term impacts include: changes in forest micro-climate from a more open canopy, which increases temperature and reduces humidity, decline of species in symbiotic relationships with the affected tree (Clark's nutcracker and whitebark pine), decline of specialist species (ash specialist arthropods), decline of species which rely on affected trees for habitat (black-throated green warbler and hemlock), changes in aquatic species composition from streams switching from shaded to more sun, increased erosion from loss root mass and tree cover. [1] [10]

Long-term

Long-term disturbances are changes in the way the forest ecosystem functions as a result of the disturbance event and are measured in decades to centuries. Changes in the forest ecosystem are primarily a result of changes in forest structure and species composition. [1] American chestnut used to compose up to 50 percent of the canopy in many eastern deciduous forest, [11] Ash species make up 7 percent of all Wisconsin forests (for example) [12] and up to 90 percent of some forests. [13] Eastern hemlock is the most common conifer in many northeastern forests. [10] The loss of these species from the forest represents a loss of the contributions they made to the function of the ecosystem. Co-existing tree and plant species will gain in the void left by the affected species, but often providing different functions. Eastern hemlock, for example, is commonly replaced by black birch in the northeast—a slow-growing, shade-tolerant and deep shade producing evergreen tree, replaced by a fast-growing, open, deciduous tree. While short-term forest productivity was discussed above, long-term productivity of ash forests will depend on the species which fill the void; a slow-growing species (maples) would reduce forest productivity while a fast-growing species (box-elder) might increase productivity. [10]

Limitations

The complete picture of long-term ecological effects from invasive pests is difficult to measure; and it is too early to determine the effects of the many new invasive pest introductions. Much of the research has focused of the biology of the pests—i.e. life cycle and host preferences—towards understanding how to contain their spread. [10] Recent and rapid outbreaks such as emerald ash borer, however, have invigorated interest in understanding the ecological consequences of invasive pests and how to restore affected forests. [14]

Survival and recovery

The American chestnut, virtually eliminated from eastern forests, survived in small isolated pockets. Some survivors have been cross-bred with the blight-resistant Chinese chestnut and introduced into National Forests in trial studies. [11]

Parasitoid wasps have been approved for release by the USDA to combat the emerald ash borer. Pesticide applications are also used to protect ash trees against borer invasion. Blue ash is the one native ash species which shows some natural resistance to emerald ash borer. [15]

Though the American elm remains susceptible to Dutch elm disease, it persists in forests—thus preserving genetic diversity—because it reaches maturity at a young age, produces seed that travels long distances through wind dispersal, can establish in disturbed areas and is tolerant of harsh conditions. [16] Cultivars of American elm have been produced, and are available in the nursery trade, which are Dutch elm disease resistant, such as 'Valley Forge' and 'Princeton'. [16] [17]

List of invasive insects and pathogens affecting tree species in North America

Insects

Note: Pathogens carried by insects are included under pathogens.

Pathogens

Related Research Articles

<span class="mw-page-title-main">Chestnut blight</span> Fungus disease of chestnut trees

The pathogenic fungus Cryphonectria parasitica is a member of the Ascomycota. This necrotrophic fungus is native to East Asia and South East Asia and was introduced into Europe and North America in the early 1900s. The fungus spread rapidly and caused significant tree loss in both regions.

<i>Fraxinus americana</i> Species of ash

Fraxinus americana, the white ash or American ash, is a fast-growing species of ash tree native to eastern and central North America.

<span class="mw-page-title-main">Biological pest control</span> Controlling pests using other organisms

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.

<span class="mw-page-title-main">Pest (organism)</span> Organism harmful to humans/our concerns

A pest is any organism harmful to humans or human concerns. The term is particularly used for creatures that damage crops, livestock, and forestry or cause a nuisance to people, especially in their homes. Humans have modified the environment for their own purposes and are intolerant of other creatures occupying the same space when their activities impact adversely on human objectives. Thus, an elephant is unobjectionable in its natural habitat but a pest when it tramples crops.

<span class="mw-page-title-main">Asian long-horned beetle</span> Species of beetle

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.

<span class="mw-page-title-main">Emerald ash borer</span> Species of beetle

The emerald ash borer, also known by the acronym EAB, is a green buprestid or jewel beetle native to north-eastern Asia that feeds on ash species. 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 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.

<span class="mw-page-title-main">Bark beetle</span> Subfamily of beetles

A bark beetle is the common name for the subfamily of beetles Scolytinae. Previously, this was considered a distinct family (Scolytidae), but is now understood to be a specialized clade of the "true weevil" family (Curculionidae). Although the term "bark beetle" refers to the fact that many species feed in the inner bark (phloem) layer of trees, the subfamily also has many species with other lifestyles, including some that bore into wood, feed in fruit and seeds, or tunnel into herbaceous plants. Well-known species are members of the type genus Scolytus, namely the European elm bark beetle S. multistriatus and the large elm bark beetle S. scolytus, which like the American elm bark beetle Hylurgopinus rufipes, transmit Dutch elm disease fungi (Ophiostoma). The mountain pine beetle Dendroctonus ponderosae, southern pine beetle Dendroctonus frontalis, and their near relatives are major pests of conifer forests in North America. A similarly aggressive species in Europe is the spruce ips Ips typographus. A tiny bark beetle, the coffee berry borer, Hypothenemus hampei is a major pest on coffee plantations around the world.

<span class="mw-page-title-main">Oak wilt</span> Plant disease

Oak wilt is a fungal disease caused by the organism Bretziella fagacearum that threatens Quercus spp. The disease is limited to the eastern half of the United States; first described in the 1940s in the Upper Mississippi River Valley. The pathogen penetrates xylem tissue, preventing water transport and causing disease symptoms. Symptoms generally consist of leaf discoloration, wilt, defoliation, and death. The disease is dispersed by insect vectors and to adjacent trees through underground root networks. However, human spread is the most consequential dispersal method. Moving firewood long distances can potentially transport diseases and invasive species.

<i>Fraxinus pennsylvanica</i> Species of ash

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 Europe from Spain to Russia.

<span class="mw-page-title-main">Hemlock woolly adelgid</span> Species of true bug

Adelges tsugae, the hemlock woolly adelgid or HWA, is an insect of the order Hemiptera native to East Asia. It feeds by sucking sap from hemlock and spruce trees. In its native range, HWA is not a serious pest because populations are managed by natural predators and parasitoids and by host resistance. In eastern North America it is a destructive pest that threatens the eastern hemlock and the Carolina hemlock. HWA is also found in western North America, where it has likely been present for thousands of years. In western North America, it primarily attacks western hemlock Tsuga heterophylla and has only caused minor damage due to natural predators and host resistance. Accidentally introduced to North America from Japan, HWA was first found in the eastern United States near Richmond, Virginia, in 1951. The pest is now found from northern Georgia to coastal Maine and southwestern Nova Scotia as well as areas of western Michigan near the eastern Lake Michigan shoreline. As of 2015, HWA has affected 90% of the geographic range of eastern hemlock in North America.

<i>Fraxinus nigra</i> Species of ash

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.

<span class="mw-page-title-main">Woodboring beetle</span> Common name for a beetle that eats wood

The term woodboring beetle encompasses many species and families of beetles whose larval or adult forms eat and destroy wood. In the woodworking industry, larval stages of some are sometimes referred to as woodworms. The three most species-rich families of woodboring beetles are longhorn beetles, bark beetles and weevils, and metallic flat-headed borers. Woodboring is thought to be the ancestral ecology of beetles, and bores made by beetles in fossil wood extend back to the earliest fossil record of beetles in the Early Permian (Asselian), around 295-300 million years ago.

<span class="mw-page-title-main">Christmas tree pests and weeds</span>

Pine and fir trees, grown purposely for use as Christmas trees, are vulnerable to a wide variety of pests, weeds and diseases. Many of the conifer species cultivated face infestations and death from such pests as the balsam woolly adelgid and other adelgids. Aphids are another common insect pest. Christmas trees are also vulnerable to fungal pathogens and their resultant illnesses such as root rot, and, in the U.S. state of California, sudden oak death. Douglas-fir trees in particular are vulnerable to infections from plant pathogens such as R. pseudotsugae.

<i>Lymantria dispar</i> in the United States Spread of an invasive species

The spongy moth, formerly known as the gypsy moth, was introduced in 1868 into the United States by Étienne Léopold Trouvelot, a French scientist living in Medford, Massachusetts. Because native silk-spinning caterpillars were susceptible to disease, Trouvelot imported the species in order to breed a more resistant hybrid species. Some of the moths escaped, found suitable habitat, and began breeding. The gypsy moth is now a major pest of hardwood trees in the Eastern United States.

<span class="mw-page-title-main">Invasive species in the United States</span>

Invasive species are a crucial threat to many native habitats and species of the United States and a significant cost to agriculture, forestry, and recreation. An invasive species refers to an organism that is not native to a specific region, this poses significant economic and environmental threats to its new habitat. The term "invasive species" can also refer to feral species or introduced diseases. Some introduced species, such as the dandelion, do not cause significant economic or ecologic damage and are not widely considered as invasive. Economic damages associated with invasive species' effects and control costs are estimated at $120 billion per year.

Forest pathology is the research of both biotic and abiotic maladies affecting the health of a forest ecosystem, primarily fungal pathogens and their insect vectors. It is a subfield of forestry and plant pathology.

<i>Agrilus auroguttatus</i> Species of beetle

Agrilus auroguttatus is a species of jewel beetle in the United States, known by the common name goldspotted oak borer. It is not native to California, but is native to Arizona. It is a woodboring beetle best known for destroying stands of oak trees in the Cleveland National Forest in San Diego County, California, in the United States. The distribution of this insect in California can be found at CalInvasive website. 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.

<i>Lymantria dispar dispar</i> Subspecies of moth (gypsy moth)

Lymantria dispar dispar, commonly known as the gypsy moth, European gypsy moth, LDD moth, or North American gypsy moth or spongy moth, is a species of moth in the family Erebidae. It has a native range that extends over Europe and parts of Africa, and is an invasive species in North America.

Trunk injection or endotherapy also known as vegetative endotherapy, is a method of target-precise application of pesticides, plant resistance activators, or fertilizers into the xylem vascular tissue of a tree with the purpose of protecting the tree from pests, or to inject nutrients to correct for nutrient deficiencies. This method largely relies on harnessing the tree's vascular system to translocate and distribute the active compounds into the wood, canopy and roots where protection or nutrition is needed.

<i>Xyleborus dispar</i> Species of beetle

Xyleborus dispar is a species of bark beetle commonly called the Pear blight beetle, or the European shothole borer. It is an invasive species in North America, and can be a pest in orchards and forests throughout its range.

References

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