Both of these Verticillium species attack a wide range of plants besides woody ornamental trees and shrubs.Verticilium albo-atrum is adapted for the cooler soils in the world so is not usually found in tropical soils.Verticillium dahlia is more commonly found in most soils around the world. Avoid fields previously planted to potato or tomato; however, former peppermint fields may be lower risk. ... verticillium wilt, anthracnose, leaf spot, tar spot, several insects manganese chlorosis on high pH soils ; ... 'Silver Queen' - A common cultivar with upright habit that is rounded when mature (to 50' tall and wide). Symptoms canoccur at any time of the year but often show up when hot, dry weatherbegins.Sometimes a single branch or the foliage on one side of a tree will die. Obvious Verticillium Wilt on Crimson Queen Japanese maple. During the acute stage, maple trees will have leaves that curl, dry, appear abnormally red or yellow, and fall. Silver maple has a vase shape and is a rapidly growing, fairly weak-wooded tree that reaches a height of 60 to 80 feet with a 5- to 6- foot diameter trunk on a moist site. Effect was observed up to 4 years after planting. The trees may also have a more serious problem. They eventually turn brown and drop off. A fungal infection of the soil that penetrates a tree’s roots, verticillium wilt takes … As the disease spreads, vascular tissue is damaged, which prevents the transport of water and mineral to tree branches and leaves. In addition, extra watering can help to decrease the problems of wilt. The fungi that cause verticillium wilt affect the tree's vascular system and causes symptoms of wilting and yellowing leaves that are concentrated in one particular area. While maple trees can suffer from a number of general problems, verticillium wilt seems to be the most common and dangerous disease plaguing maple tree owners. I’ve had this happen to at least four Lace leaf weeping Japanese maples in my landscape and it’s happened over a period of time. Verticillium Wilt in the Pacific Northwest. It has a somewhat weeping, pyramidal growth form and the leaves are finely dissected, not resembling silver maple at all. Of the maples the silver maple seems most resistant while the sugar maple is the most disease prone. Additional Notes When planted in alkaline soils, this tree may exhibit chlorosis (yellowing leaves) due to a manganese deficiency. Silver maple has a vase shape and is a rapidly growing, fairly weak-wooded tree that reaches a height of 60 to 80 feet with a 5- to 6- foot diameter trunk on a moist site. Verticillium wilt (Verticillium albo-atrum) is a stem disease that can cause sudden death of silver maple trees. Infected trees may die slowly or suddenly. They al… In Iowa, it is most commonly seen on maple, ash, and catalpa trees, although it is also frequently found on smoke tree, viburnum, lilac, cherry, plum and several other trees and shrubs. Verticillium Wilt within Phloem. Evaluation of Phyton-27 for control of Verticillium wilt on Japanese Maple, 2006-2008. Many authorities recommend against planting silver maple. As with many fast growing trees, silver maple has weak brittle branches that are susceptible to breaking in high winds or when coated with ice/snow in winter. Weak wooded and is prone to storm damage. to the junction of the suspect branch with the main stem. Verticillium Wilt. It should be saved for planting in wet areas or where nothing else will thrive. Verticillium is a soil borne fungus that infects roots then settles into the vascular system of the tree blocking water movement. and van Bruggen, H.C. 2004. If an Autumn Blaze maple is suffering from chlorosis, or iron deficiency, the first sign will be yellow pigmentation between the normally deep-green leaf veins. Verticillium wilt susceptibility: susceptible Pest resistance: long-term health usually not affected by pests USE AND MANAGEMENT Silver Maple will grow in areas which have standing water for several weeks at a time. You may first notice these symptoms in spring or fall when temperatures are mild. The … A few chemicals are registered for tree injection as a preventative treatment. In Iowa, it is most commonly seen on maple, ash, and catalpa trees, although it is also frequently found on smoke tree, viburnum, lilac, cherry, plum and several other trees and shrubs. Frequently, it affects the bark as well. Symptoms: wilting of leaves on one limb, reduced growth, gray-green streaks in sapwood. In a hybrid-like autumn blaze maple, the condition may not be severe and the only symptom may be a thin, slow-growing crown. Over 300 plant species are affected by Verticillium wilt. Shallow roots may damage nearby sidewalks. Introduction 'Skinneri' silver maple is a fast-growing, weak-wooded tree that probably reaches a height of about 60 feet with a large trunk. Growth Rate: Slow to moderate. Once in the xylem, this fungus partially blocks water movement and produces toxins that result in wilt symptoms. 2009. Maple wilt, or Verticillium wilt, is a soil-borne fungus that primarily targets woody ornamental trees. Wilt (fungus – Verticillium albo-atrum); Sudden wilting and drying of leaves on individual branches, particularly on one side of the tree. Leaves that curl, wilt, discolor and die may mean that a plant is suffering from verticillium wilt. Even though V. albo-atrum is not as common as V. dahlia, it is more likely to be fatal to most plants. Maple wilt is also known as Verticillium wilt, because it is caused by fungi from the Verticillium genus, including Verticillium albo-atrum and Verticillium dahliae. Verticillium Wilt, often called Maple wilt, is a very common disease that attacks a large number of trees. Rachel Delp has been writing and editing for academic and medical publications since 1992. Leaf color is good -- bright green above, silvery beneath and yellowish in autumn. Long-term effect of biological soil disinfestation on Verticillium wilt. In Minnesota the most common species to be affected are the maple varieties. Note how the discoloration occurs along the direction of the grain and appears as a circular or semicircular pattern in cross section. Sapwood on trees infected with verticillium wilt will be stained dark green. Delp holds a Bachelor of Arts in Spanish with a minor in art from Moravian College. This post is about a common bacterial disease called Pseudomonas syringae, which frequently affects Japanese maples yet is relatively easy to control. A fungal infection of the soil that penetrates a tree’s roots, verticillium wilt takes … Both infect a very wide range of garden plants through the roots and then grow upwards in the water-conducting tissues, causing wilting of the upper parts due to water stress. Invasiveness: very invasive, especially in woods and natural areas due to production of lots of fruit/seed that creates a litter mess By late summer, the spots have become thick, shiny lesions up to 1/2 inch in diameter. Verticillium Wilt on Silver Maple. Variation in response of Norway maple cultivars to Verticillium dahliae. Silver maple will grow in areas that have standing water for several weeks at a time. how to tell if a silver maple is dying. It grows best on acid soil which remains moist, … Invasiveness: very invasive, especially in woods and natural areas due to production of lots of fruit/seed that creates a litter mess By late summer, the spots have become thick, shiny lesions up to 1/2 inch in diameter. Verticillium wilt is hard to eradicate and the best defense is to plant resistant cultivars. Both infect a very wide range of garden plants through the roots and then grow upwards in the water-conducting tissues, causing wilting of the upper parts due to water stress. Cottony maple scale, maple bladder gall (common but not serious), anthracnose (occasionally) and Verticillium wilt . Disease cycle of Verticillium Wilt The fungus Verticillium is found in the soil. There are over 300 plant species susceptible to this fungus. Verticillium wilt is a common fungus of plants that originates in the root system and multiplies in the living tissue of the host. Well, anyway, since it works very well on fungus, virus and bacteria I thought it was worth a try to use it on my verticillium wilt infected japanese maple. If branches abruptly wilt and die, verticillium wilt may be to blame. Plants are occasionally susceptible to verticillium wilt, leaf spots, root rots, anthracnose, collar rot and stem canker. Maple Box elder Tree-of - heaven Pecan Catalpa Carob Redbud Camphor tree Yellow wood Carrotwood ... 4 The VF number varieties of tomatoes are resistant to both Verticillium wilt and Fusarium wilt. Symptoms include yellowing and defoliation of leaves on a few branches or even the entire tree. Verticillium wilt is a soil borne fungus that can exist in the soil almost indefinitely. I believe my silver maple is infected with verticillium wilt. They’ll begin to develop yellow blotches on the lowest leaves and can end up killing off entire limbs of trees at once. Its root system clogs sewer lines and lifts sidewalks. Vascular discoloration of sugar maple caused by Verticillium wilt. "Plant pathogenic Verticillium species: how many of them are there?" This fungi attack more than 300 plants, shrubs and trees, causing a vascular disease that may result in plant death. Its large, vigorous, shallow-rooted root system can damage sidewalks and driveways, clog drain pipes, and penetrate septic systems and sewer pipes. Another species, Verticillium albo-atrum, is less common. Irregular, light to reddish-brown, or purplish-brown, dead areas occur on the leaves. is there any thing I... Home; About; Ask a Question; Sign in; verticillium wilt. Habit: multi-trunked Verticillium wilt is a soil borne fungus that can exist in the soil almost indefinitely. Lists of plants susceptible or resistant It is often misidentified as Verticillium wilt, as both cause tip dieback. Clean boots, equipment, and tools before leaving an infested area. Symptoms of leaf scorch or die-back of branches would indicate a possibility that these symptoms could be caused by Verticillium. Frequency. Rhytisma acerinum or R. punctatum: No control measures are necessary. Branches with maple wilt will also have a small amount of the scorched leaves. The fungi that cause verticillium wilt affect the tree's vascular system and causes symptoms of wilting and yellowing leaves that are concentrated in one particular area. Another preventive approach is to plant resistant trees and cultivars whenever possible. You may first notice these symptoms in spring or fall when temperatures are mild. Current-season sapwood may not be infected, and symptoms may not reappear; or infection may occur without foliar symptoms. Verticillium Wilt on Maple Trees. The fungus grows into the xylem where it colonizes the plant through mycelial growth and conidial production. Leaf scorch can also occur at leaf margins. May be weedy, spreading many seedlings in lawns and gardens. Verticillium wilt is caused by a soil fungus called Verticillium dahliae. Keep nitrogenous fertilizers to a minimum-enough only to produce normal, not succulent, growth. Verticillium Wilt on Maple Trees. 1990. Note Although Phyton 27 is registered as a spray it is not recommended based on a single trial where it was ineffective when used before or after infection. Verticillium wilt: Early symptoms of verticillium wilt include heavy seed production, leaves that are smaller than normal, and browning of the margins of leaves. Maples are among the tree species susceptible to Verticillium wilt. Leaves that curl, wilt, discolor and die may mean that a plant is suffering from verticillium wilt. Verticillium Wilt within Phloem . The … Wilting is mostly seen from spring until autumn. Relative Disease Susceptibility and Sensitivity to Sulfur, Diagnosis and Control of Phytophthora Diseases, APHIS List of Regulated Hosts and Plants Proven or Associated with Phytophthora ramorum, Plants Resistant or Susceptible to Armillaria Root Rot, Plants Resistant or Susceptible to Verticillium Wilt, Plants Susceptible to Sclerotinia sclerotiorum, Potential Impact of Cyanobacteria on Crop Plants, Management of the Cyanobacterium Nostoc in Horticultural Nurseries, Impatiens Necrotic Spot: New Name, Same Threat, Virus Certification Program for Fruit and Ornamental Trees, Care and Maintenance of Wood Shingle and Shake Roofs, Winter Injury of Landscape Plants in the Pacific Northwest, Recognizing Sapsucker Damage on your Trees, Key to Nutrient Deficiencies in Vegetable Crops, Key to Nutrient Deficiencies of Deciduous Fruit and Nuts, Use of Disinfestants to Control Plant Pathogens, Current Status of Biological Weed Control Agents in Idaho, Oregon, and Washington, Biological Control Agents and Their Roles, Restricted-use Herbicides in Idaho, Oregon, and Washington, Testing for and Deactivating Herbicide Residues, Herbicide Effectiveness on Weeds in Grass Seed Crops, Dry Bean East of the Cascades - Phaseolus spp. Verticillium wilt is a fungal disease that affects over 300 species of plants, including many common trees and shrubs. I’ve had this happen to at least four Lace leaf weeping Japanese maples in my landscape and it’s happened over a period of time. They eventually turn brown and drop off. Do not track soil from infested areas into clean areas. Silver maple trees suffering from verticillium wilt will display wilting leaves and dying branches. Verticillium wilt susceptibility: susceptiblePest resistance: resistant to pests/diseases Use and Management . Without proper and frequent pruning high winds and ice can cause limbs to break. They include marginal scorch and complete wilting of leaves on individual branches in the crowns of potential hosts. If branches abruptly wilt and die, verticillium wilt may be to blame. Buy Silver Maples Online Branches with maple wilt will also have a small amount of the scorched leaves. It is best to consult local nurserymen or ag extension departments as to silver maple problems common to your area. These include scorch, anthracnose, fungal leaf spots, powdery mildew, and Verticillium wilt. I’ve read all kinds of explanations of what exactly is going on in the soil to cause this, but quite honestly, in all four cases my soil conditions were different, never soggy, just different soils. First, never plant maples in an area where previous plants or trees have died from disease. Ash, catalpa, elm, sumac, and maple are the most common hosts in the Rocky Mountain Region. © Copyright 2020 Hearst Communications, Inc. The tree is useful in wet areas, transplants easily, and can grow where few others can. In cross sections of the stem, vascular discoloration will appear in rings or arcs. Once the infected tissue decays, microsclerotia are released and can survive several years in soil. While maple trees can suffer from a number of general problems, verticillium wilt seems to be the most common and dangerous disease plaguing maple tree owners. Mycelia from the microsclerotia infect the roots. how to tell if a silver maple is dying. Lastly, good sanitation practices -- keeping the area around plants and trees free from debris and fallen branches -- helps prevent disease like maple wilt. Leaf edges roll inward prior to … Infected trees may die within a few weeks or live for years. Another species, Verticillium albo-atrum, is less common. They’ll begin to develop yellow blotches on the lowest leaves and can end up killing off entire limbs of trees at once. Trees may show yellowing, wilting, and drying of leaves in the top branches, or sometimes the entire crown. It should be saved for planting in wet areas or where nothing else will thrive. Protect from late frosts as foliage tends to leaf out early in spring and may be damaged. None of these control methods will cure the maple. After the roots of susceptible trees are invaded, the fungus moves into the vascular system. Because Verticillium persists in the soil for years, you will need to replace it with a tree that is Verticillium-resistant. The tree is useful in wet areas, transplants easily, and can grow where few others can. Its a large tree with dying off of the top branches. Verticillium wilt susceptibility: susceptible Pest resistance: long-term health usually not affected by pests USE AND MANAGEMENT Silver Maple will grow in areas which have standing water for several weeks at a time. Avoid planting maple in fields with a history of Verticillium wilt. Vascular discoloration is more likely to be found in larger branches and trunks. PDMR 3:OT024. Symptoms Leaves on one side of the tree or on just an individual branch suddenly wilt and die. As it enters the tree, it will cause the leaves to discolor and shrivel up. Nurseries may wish to test individual core samples to determine the distribution in a particular field. & Clewes, E. (2003). HERBACEOUS OR NAM ENTALS-Susceptible Botanical Name ... silver dollar A I … It is usually found growing in open sunlight along creeks and waterways. Verticillium Wilt Verticillium wilt (Verticillium albo-atrum) is a stem disease that can cause sudden death of silver maple trees. Verticillium wilt is caused by a soil fungus called Verticillium dahliae. Verticillium wilt is a fungal disease of over 300 host plants, including a wide range of garden and greenhouse crops in addition to woody ornamentals, most noticeably elms, magnolias, maples, redbud, and viburnums. In a hybrid-like autumn blaze maple, the condition may not be severe and the only symptom may be a thin, slow-growing crown. It is caused by a soil-inhabiting fungus called Verticillium. There are over 300 plant … Verticillium Wilt, often called Maple wilt, is a very common disease that attacks a large number of trees. The disease is particularly destructive to trees in landscape plantings. If your trees are infected, you’ll most likely see it on the oldest branches and leaves on the tree. The disease is particularly destructive to trees in landscape plantings. The Norway maple cultivars Jade Glen and Parkway are tolerant and have few symptoms. ... red and silver maples. Over 300 plant species are affected by Verticillium wilt. is there any thing I can do to save it? Many weeds are susceptible and can help the fungus survive and disperse. Autumn Blaze is a hybrid cross between the silver maple and red maple. This species has become over planted. Verticillium dahliae, a soilborne fungus that is almost impossible to eradicate once established in the soil. Its a large tree with dying off of the top branches. Varieties of maple tree susceptible to Verticilium wilt include amur maple (Acer ginnala), red maple (Acer rubrum) and silver maple (Acer saccharinum), which grow in U.S. Department of Agriculture plant hardiness zones 3 through 8 or 9, among others. Maple wilt is also known as Verticillium wilt, because it is caused by fungi from the Verticillium genus, including Verticillium albo-atrum and Verticillium dahliae. (See following list for a more complete list of susceptible plants.) I believe my silver maple is infected with verticillium wilt. Verticillium wilt is caused by a fungus and is a challenging one to deal with as it’s hard to diagnose and cannot be cured. These microsclerotia may lie dormant in the soil for years. Maple Wilt. Of the maples the silver maple seems most resistant while the sugar maple is the most disease prone. Obvious Verticillium Wilt on Crimson Queen Japanese maple. It was verticillium wilt even though it did not have the brown streaks in the cambian layer that is often a symptom. Leaves are yellowish and smaller than normal. The Verticillium fungus that cause maple wilt can live in the soil for years undetected. sometimes isn’t visible at all Verticillium wilt is a wilt disease affecting over 350 species of eudicot plants.It is caused by six species of Verticillium fungi: V. dahliae, V. albo-atrum, V. longisporum, V. nubilum, V. theobromae and V. tricorpus. Maples in the landscape and in natural woodlots and forests are commonly affected by several physiological disorders and diseases. Hosts—Verticillium wilt is a vascular wilt of hardwoods. Its brittle wood and poor branch development result in severe breakage during ice and wind storms. I believe my silver maple is infected with verticillium wilt. Roots often grow on the surface of the soil, making mowing grass difficult under the canopy. References Goud, J.C., Termorshuizen, A.J., Blok, W.J. It is best to consult local nurserymen or ag extension departments as to silver maple problems common to your area. It was verticillium wilt even though it did not have the brown streaks in the cambian layer that is often a symptom. If an Autumn Blaze maple is suffering from chlorosis, or iron deficiency, the first sign will be yellow pigmentation between the normally deep-green leaf veins. Symptoms include yellowing and defoliation of leaves on a few branches or even the entire tree. Plant Disease 74:44-46. how to tell if a silver maple is dying. The disease is caused by the common soil-borne fungi New leaves may be reduced in size and turn yellow. Maple wilt can show up as both acute and chronic signs of Verticillium infection. It forms microsclerotia that germinate in response to exudates from roots that are growing nearby. Most frequently, Verticillium spores enter the root system at the site of an earlier wound, but weakened trees are susceptible to direct infection. 5 . Plant Disease 88:688-694. This fungus lives in soil as small, darkened structures called microsclerotia. Plant-parasitic nematodes also can increase Verticillium wilt incidence and severity. Silver Maple Problems: The silver maple tree is susceptible to many insects and diseases including Verticillium wilt and other canker diseases. Her articles can be found on various websites. A cross section of the stem will show vascular discoloration in rings or arcs as can be seen in the stem on the right. Also susceptible to scale and borers. Wilting is mostly seen from spring until autumn. Verticillium wilt is a fungal disease of over 300 host plants, including a wide range of garden and greenhouse crops in addition to woody ornamentals, most noticeably elms, magnolias, maples, redbud, and viburnums. Leaves die and fall or hang on dead branches. It grows best on acid soil, which remains moist, but adapts to very dry, alkaline soil. As the fungus grows in the soil it can really only be handled by preventing its growth. Verticillium wilt symptoms mimic those of other plant diseases and environmental problems, and this makes it hard to diagnose. Expand. Relatively soft wood renders silver maple susceptible to a number of wood rotting fungi and it is susceptible to various leaf molds and wilts (e.g., anthracnose, verticillium wilt, leaf spot, tar spot). Read on to find out how to distinguish verticillium wilt from other plant diseases and what to do about it. The host range includes trees, shrubs, ground covers and vines, vegetables, field crops, fruits, herbaceous ornamentals, and many weeds. Leaf edges roll inward prior to turning brown if the tree is severely infected. Disease, pests, and problems. If the tree dies and/or is removed, replace it with a nonsusceptible host such as any conifer, birch, dogwood, or sycamore. Hosts—Verticillium wilt is a vascular wilt of hardwoods. Leaf Spots on a Japanese Maple Bloodgood Tree, University of Illinois Extension: Verticillium Wilt (Shrubs and Trees), Ohio State University Extension: Verticillium Wilt of Landscape Trees and Shrubs, New Mexico State University: Verticillium Wilt of Catalpa, Maple, and Elm, University of Missouri Extension: Verticillium Wilt in Maples, University of Illinois Extension: Verticillium Wilt, University of Minnesota Extension: Verticillium Wilt of Trees and Shrubs, Maple Tree Problems: The Leaves Turn Yellow in Summer. Photos courtesy of Jerry E. Weiland (USDA-ARS). how to tell if a silver maple is dying. Has a vigorous root system that can invade sewer pipes. Plants Resistant to Verticillium Wilt Botanical Name Arctostaphylos species Betula species Buxus species Carpinus species Ceanothus species Cercidiphyllum j. Cistus corbariensis ... Lunaria annua Honesty, silver dollar Mimulus species Monkey flower Nemesia strumosa Nemesia Nemophila menziesii Baby … If only one branch has symptoms you may miss it, but if the whole tree is affected, you may mistake the symptoms for premature fall leaf drop. The fungus first attacks the tree through the roots, which can spread the infection through the maple's vast vascular system. Once planted, keep trees healthy, including proper watering, especially during droughts, because healthy trees are better able to resist disease. Greenish streaks or bands that follow the grain can be found in sapwood but perhaps not in the earliest stages of infection. Incorporating freshly mown Italian ryegrass followed by covering the soil with plastic for 3 months in the late summer was effective in the Netherlands at reducing disease incidence in a nursery crop of Norway maple planted 6 months later. 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The leaves wilt and curl, and turn yellow or red. Fungicides are not effective for controlling or eliminating Verticillium or maple wilt. Silver Maple Problems: The silver maple tree is susceptible to many insects and diseases including Verticillium wilt and other canker diseases. This fungus lives in soil as small, darkened structures called microsclerotia. The cambium may die, resulting in an elongate canker, which can be colonized by other pathogens such as Nectria sp. Susceptible to verticillium wilt, anthracnose and canker. Oval to irregularly shaped, shiny, black spots up to ½ inch in diameter form on the leaves of silver or red maples. Areas often enlarge killing the entire leaf. (See, for example, Barbara, D.J. Other diseases, such as bacterial dieback, can also increase in incidence and severity on trees with Verticillium wilt. It grows best on acid soil which remains moist, … Its a large tree with dying off of the top branches. Verticillium wilt symptoms mimic those of other plant diseases and environmental problems, and this makes it hard to diagnose. Treescan go through years where no symptoms are present and then the symptoms sho… Pruning and burning or disposing of dead or affected branches will help limit the infection, but it will not eliminate the disease because the fungus has spread through the roots and vascular system. This fact sheet discusses the symptoms, causal factors, and management strategies to minimize their impact on tree health. In diseased branches, the wood under the maple's bark has olive or gray-green streaks. Young trees generally succumb to maple wilt within a year. Verticillium wilt of maple. Infected leaves fall to the ground, returning the fungus to the soil, and wind can aid in spreading spores farther, possibly infecting additional plants and trees. Roots often grow on the surface of the soil, making mowing grass difficult under the canopy. Anthracnose (fungus – Gloeosporium apocryptum): In rainy seasons this disease may be serious on silver maples and Japanese maples. Other fungi can also cause this discoloration. How to Reduce Bee Poisoning from Pesticides, Alfalfa Leafcutting Bee (Megachile rotunda) Pests, Horticultural, Landscape, and Ornamental Crops, Field Characteristics of Fruit-Tree-Attacking Spider Mites in the Pacific Northwest, Pests of Cabbage and Mustard Grown for Seed, Pests of Rutabaga and Turnip Grown for Seed, Biology and Control of the Garden Symphylan, Bean, Dry Cultivar Resistance to Bean Common Mosaic, Grape (Vitis spp.) Verticillium wilt in trees can be hard to diagnose as symptoms are often confused with other causes such as compacted soil, drought stress etc. (See following list for a more complete list of susceptible plants.) Verticillium wilt is caused by the soil-borne fungi Verticillium dahliae and V. albo-atrum. Cause Samples with Verticillium wilt have dominated the maple problem diagnoses sent to the OSU Plant Clinic. Pscheidt, J.W., and J.P. Bassinette. Verticillium Wilt in the Pacific Northwest Cause Samples with Verticillium wilt have dominated the maple problem diagnoses sent to the OSU Plant Clinic. It is caused by a soil-inhabiting fungus called Verticillium.The disease fungus can be spread by many methods including from plant-to-plant, through the soil, groundwater and often by infected pruning equipment that has not been properly sanitized. Later, other limbs wilt and die. Verticillium wilt is a common fungus of plants that originates in the root system and multiplies in the living tissue of the host.