riparian ash killed by EAB; in this case, Mattawoman Creek in Maryland. Photo by Leslie A. Brice
Good news at the recent 33rd USDA Research Forum on Invasive Species. Scientists presented the first study that demonstrates significantly lower ash tree mortality in sites with high parasitism rates of two larval parasitoids, Tetrastichus planipennisi and Spathius galinae.
Their study area is the ash-dominated riparian area along the Connecticut River that flows north to south across the middle of Massachusetts. Knowing in advance that the emerald ash borer (Agrilus planipennis; EAB) would invade the area, scientists established monitoring plot that consisted of marked individual ash trees. EAB was first detected in the southern reach of the riparian area in 2015. It gradually moved north. By 2020 isolated mortality was observed at all sites. Meantime, they released three biocontrol agents – T. planipennis,S. galinae, and Oobius agrilii – early in the invasion at three of the six monitoring sites. These released occurred in 2018 – 2020 and again in 2022.
In 2021 and 2025, the scientists counted the numbers of biocontrol agents in the marked trees or sentinel logs. Thus the first evaluation occurred six years after EAB arrived, three years after the first releases of biocontrol agents.
They found that at southern Massachusetts sites, where EAB density was higher at the time of the biocontrol agents’ initial release, remaining ash grew more slowly than in the North. They believe the trees’ growth rate was suppressed by the trees having fewer resources. They also observed dieback. Smaller trees grew faster, perhaps responding to opening of the canopy as mature ash succumbed to EAB invasion.
The most important finding was that ash mortality at all sites was ~50% or less … not the 90% expected based on experience in the upper Midwest where the EAB invasion occurred before biocontrol agents were developed.
SOURCE
Ash survival and growth response to emerald ash borer invasion in Massachusetts riparian forests: impacts of biological control. Mitchell A. Reed, Jian Duan, Ryan S. Crandall, Roy G. van Driesche, Jeremy C. Anderson, Joseph S. Elkington. Presentation to the 33rd USDA Interagency Research Forum on Invasive Species, Annapolis, Maryland February 25-28, 2025 (The proceedings should be posted online before the end of the year.)
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at https://treeimprovement.tennessee.edu/
The pest alert system “PestLens” has again alerted us to plant pests in Europe or Asia that feed on species closely related to tree species native to North American forests. Two of the insects named in the alert apparently pose a hazard to icons of the forests of America’s Pacific coast forests, giant sequoia and redwood.
I hope APHIS is using this information to alert port and on-the-ground staff and perhaps initiating more in-depth risk assessments.
The posting on February 27, 2025 reported that cotton jassid, Jacobiasca lybica (Hemiptera: Cicadellidae), affects not just cotton and citrus but also Cupressus sempervirens (Mediterranean cypress) [Cupressaceae]. More than a dozen North American trees species are in this family, including
Sequoiadendron giganteum or giant sequoia. Giant sequoia is listed as an endangered species by the IUCN with fewer than 80,000 remaining in its native California.
Chamaecyparis thyoides and C. lawsoniana (Port-Orford cedar). Port-Orford cedar has been decimated in its native range by an introduced pathogen, Phytopthora lateralis. A major breeding effort has developed trees that are resistant to the pathogen; they are now available for people to plant.
Thuja occidentalis, also known as northern white-cedar, eastern white-cedar, or arborvitae,
Taxodium ascendens, also known as pond cypress
several Juniperus
Hesperocyparis macrocarpa also known as Cupressus macrocarpa, or the Monterey cypress. NatureServe ranks the cypress as GI – critically imperiled.
Cotton jassid been reported from several countries in Europe, Africa, and the Middle East.
China has reported the existence of a previously unknown bark beetle species, Phloeosinus metasequoiae (Coleoptera: Curculionidae). It was found infesting Metasequoia glyptostroboides (dawn redwood) trees in China. Affected trees exhibited reddened leaves and holes and tunnels in branches.
China has also discovered a several new hosts utilized by the fungus Pestalotiopsis lushanensis (Sordariomycetes: Amphisphaeriales). Formerly known to infect tea (Camellia sinensis) and several other plant species, P. lushanensis has now been found shoot causing blight and leaf drop on a conifer, deodar cedar (Cedrus deodara) and leaf spots on an angiospermwith congeners in North America — the rare Chinese species, Magnolia decidua. There are eight species of Magnolia native to North America.
Magnolia grandiflora; photo by DavetheMage via Wikimedia
APHIS’ ability to respond to alerts remains uncertain.
The agency’s probationary employees have been fired – just as at other agencies. APHIS staff were prohibited from participating in last week’s annual USDA Invasive Species Research Forum – the 33rd such meeting. The bird flu emergency is demanding all the attention and funds.
So – how can the rest of us fill in?
At the USDA Research Forum I again presented a poster urging greater attention to tree-killing pathogens. Scientists have made considerable progress in identifying factors that indicate whether a non-native insect might pose a significant threat (see blogs on conifer and deciduous species; more to come!). However, USDA had not funded a similar effort to improve understanding of pathogens. The most promising strategy so far are sentinel plantings. However, these systems have weaknesses; I will blog in the near future about another analysis.
I propose that APHIS start by working with independent scientists to determine the actual, current level of pathogens associated with various types of incoming goods. Contact me directly if you wish to read the text of my poster.
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at https://treeimprovement.tennessee.edu/
emerald ash borer; some of PPA grants are funding evaluation of biocontrol efficacy
USDA APHIS has released information about its most recent annual allocation of funds under the Plant Pest and Disease Management & Disaster Prevention Program under §7721 of the Plant Protection Act. (Also see Fading Forests II and III; links provided at the end of this blog.) These funds support both critical needs and opportunities to strengthen the nation’s infrastructure for pest detection, surveillance, identification, and threat mitigation. Since 2009, this USDA program has provided nearly $940 million to more than 5,890 projects.
For FY25 APHIS allocated $62.725 million to fund 339 projects, about 58% of the proposals submitted. About $10 million has reserved for responding to pest and plant health emergencies throughout the year.
According to APHIS’ press release, the highest amount of funds (almost $16 million) is allocated to the category “Enhanced Plant Pest/Disease Survey.” Projects on “Enhanced Mitigation Capabilities” received $13.6 million. “Targetting Domestic Inspection Efforts to Vulnerable Points” received nearly $6 million. “Improving Pest Identification and Detection Technology” was funded at $5 million. Outreach & education received $4 million. I am not sure why these do not total $63 million.
Funding for States and Specific Pests
Wood-boring insects received about $2.3 million. These included more than $869,800 to assess the efficacy of biocontrol for controlling emerald ash borer (EAB) Agrilus planipennis, $687,410 was provided for various detection projects, and $450,000 for outreach efforts related to various pests. Ohio State received $93,000 to optimize traps for the detection of non-native scolytines (bark beetles).
Biocontrol efficacy will also be assessed for hemlock woolly adelgid, invasive shot hole borers, cactus moth, and several invasive plants (including Brazilian pepper). (Contact me to obtain a copy of CISP’s comments on this biocontrol program.)
Opuntia basilaris in Anza Boreggo; one of flat-padded Opuntia vulnerable to the cactus moth; photo by F.T. Campbell
Funding for other pests exceeded $1 million for spotted lanternfly (nearly $1.4 million), Asian defoliators ($1.2 million) and box tree moth (just over $1 million).
$630,000 was provided for detection surveys and studies of the sudden oak death pathogen Phytophthora ramorum, especially how it infects nursery stock. Nursery surveys are funded in Alabama, Louisiana, North Carolina, Ohio, Oklahoma, Pennsylvania, South Carolina, Tennessee, Virginia, and West Virginia. Most of these states are in regions considered most at risk to SOD infection of wildland plants.
sudden oak mortality of tanoak trees in southern Oregon; photo by Oregon Department of Forestry
Oregon received much-deserved $41,000 to evaluate the threat of the NA2 and EU2 lineages of P. ramorum to nurseries and forests Oregon also received $104,000 to respond to the detection of Phytophthora austrocedri in nurseries in the state. The Oregon outbreak has been traced to Ohio, but I see no record of funds to assist that state in determining how it was introduced.
Asian defoliator (e.g., Lymantrid moths) surveys have been funded for several years. This year’s projects are in Alaska, Arkansas, California, Kentucky, Maryland, Massachusetts, Mississippi, Montana, Nevada, North Carolina, Oregon, Tennessee, Texas, Washington, and West Virginia. While I agree that the introduction risk is not limited to coastal states with maritime ports, I don’t what criteria were applied in choosing the non-coastal states which are funded to search for these insects
Spotted lanternfly surveys (including technological improvements) or related outreach are funded in Alabama, Connecticut, Delaware, Kentucky, New Hampshire, New Jersey, North Carolina, Oregon, Pennsylvania, and Tennessee. California’s project is focused on postharvest treatments.
The Don’t Move Firewood project continues to be funded by APHIS. Several states also direct attention specifically to the firewood pathway: Kentucky, Maine, and Michigan.
I applaud the precautionary funding of the Agriculture Research Service to generate of high-quality genomic resources for managing the causal agent of Japanese oak wilt Dryadomyces quercivorous
Florida Department of Agriculture, North Carolina State University, and West Virginia University each received more than $100,000 to improve detection and management of invasive hornets.
Tennessee State University got $100,000 to continue efforts to detect and understand Vascular Streak Dieback in redbud Cercis canadensis.
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at https://treeimprovement.tennessee.edu/
BLD symptoms; photo by Matt Borden, Bartlett Tree Experts
As beech leaf disease (BLD) is detected in an ever-expanding number of counties from Michigan to Maine south to Virginia, scientists are trying to clarify how the causal nematode — Litylenchus crenatae ssp. mccannii (Lcm) – spreads. One focus is on local spread from tree to tree. Mankanwal Goraya and colleagues set up an experiment in Stone Valley Forest, a recreation and research site managed by Penn State in Huntington County, Pennsylvania. BLD is present – although I have not been able to determine for how many years. [The full citation to Goraya et al. is provided at the end of this blog.]
Goraya et al. (2024) set up four stands, each bearing three funnels, at varying distances from naturally BLD-infected American beech (Fagus grandifolia) trees. Two stands were at 3.51 m from symptomatic trees of starkly different sizes: one of the trees had a dbh of 50 cm, the other of only 5.6 cm. A third close-up stand was set up at 2.20 m from another large tree, having a dbh of 46 cm. The fourth stand was set up at a significantly longer distance, 11.74 m from a symptomatic beech tree; this tree was also small, with a dbh of 5 cm. This arrangement allowed the scientists to detect influences of both distance from the source of infection and relative canopy size of the source tree. They consider dbh to be an adequate substitute for canopy size. There was apparently no other effort to determine or vary the height of “source” trees, although I think that might influence speed of the wind flowing through the canopy.
Goraya et al. also tested whether it is possible to detect the presence of Lcm in association with other invertebrates that live in beech forests. To do this, they counted numbers of nematodes in frass from six species of caterpillars that had been feeding on leaves of infected trees, and in two spider webs spun in the branches of symptomatic trees. They also determined whether these nematodes were alive (active) or inactive – presumably dead.
The study makes clear that Lcm’s life cycle and impact are not as surprising as initially thought. Several species in the family Anguinidae – to which Lcm belongs – are considered significant pests. These nematodes can parasitize aerial parts of the plants (leaves, stems, inflorescences and seeds), causing swellings and galls. Furthermore, they are migratory; they can move across the surface of host tissues using water films. Once they have penetrated the host tissues, they can induce host cell hyperplasia and hypertrophy, resulting in leaf or bulb deformities, shorter internodes, and neoplastic tissues. Furthermore, heavy rainfall and wind are known to play significant roles in the dissemination of plant-infecting nematodes. In their desiccated state on infected seeds, some species of this family can survive passage through animals’ gastrointestinal digestive tract (e.g., domestic livestock, insects, & birds).
A crucial factor is that Lcm can reach densities of thousands of nematodes per leaf by late summer or early fall, increasing the likelihood of their exposure to facilitating environmental conditions at the time they migrate from leaves to buds. And once established within the bud tissues, the nematodes feed on bud scales and newly forming leaves to develop & increase their pop #s. They also use the bud as protection from adverse environmental conditions.
Goraya and colleagues collected samples every other day from September 9 to November 23, 2023 – the period when Lcm migrate from highly infected leaves to newly forming buds. [I note that it in the mid-Atlantic – where Lcm is spreading – we had an extensive drought in autumn 2024 – more than 30 days without any rain from early October into November. I hope scientists are monitoring BLD spread sufficient closely to see whether this drought affected dispersal.]
Nematodes dispersal linked to weather
Goraya and colleagues collected 324 samples from the funnels. Eighty-two percent (n =266) of the samples had nematodes; up to 92% were identified as Lcm. Non-Lcm nematodes were distributed across different genera, mostly classified as free-living nematodes. While several hundred nematodes were found in the funnels on most days, numbers peaked noticeably on some days in September and October. A startling 2,452 nematodes were recovered from a single funnel in October. Depending on the sample, up to 67% of Lcm recovered from the funnels were active.
Analysis of the environmental (weather) variables found that increases in wind speed, humidity, and precipitation (rainfall) coincided with higher numbers of Lcm being recovered from the funnels. However, the effect of wind speed becomes less positive as precipitation increases or vice versa. Goraya et al. suggest a pronounced negative interaction between wind and rain. At low precipitation levels, increased wind speed might facilitate Lcm dispersal. As rainfall increases, higher wind speeds might carry the Lcm nematodes farther away. Support is seen in the fact that fewer nematodes were found in the funnels closer to the BLD-infected trees during these periods. Really heavy rain might push a significant preponderance of nematodes to the ground. The scientists point to a very complex interplay between weather patterns and Lcm population dynamics and dispersal.
BLD symptoms on beech tree in Fairfax County, Virginia – a dozen miles from known infestation; photo by F.T. Campbell
The model did not show any significant influence of maximum temperature on nematode numbers in autumn. Goraya et al. do not speculate on whether temperatures might play a role during summer, as distinct from cooler autumn periods.
Goraya et al.’s findings differ from those of previous studies. Earlier documentation of wind dispersal of nematodes concerned primarily free-living species. It was unexpected to find consistently much higher numbers of Lcm – especially because Lcm is a plant-parasitic nematode. Another surprise is the high proportion of nematodes that are active.
Goraya et al. conclude that because Lcm is actively migrating in large numbers during autumn months, it is primed to take advantage of favorable weather. This nematode will likely survive and thrive in the environmental conditions of beech forests in northeastern North America.
Considering the effect of distance, some findings fit expectations: significantly more Lcm were recovered from funnels placed near symptomatic “source” trees than from those farther away. However, this was not a simple relationship. For example, in two cases the scenarios seemed nearly alike: both “source” trees were large (dbh 46 or 50 cm) and symptoms were “medium-high” (more than half of leaves presenting dark-green interveinal bands). Distance of funnels from the “source” tree differed minimally: 2.2 m versus 3.51 m. Still, the number of nematodes retrieved from the two sets of funnels differed significantly: one set of funnels recovered the highest number of Lcm nematodes obtained during the entire experiment – 2,452; the second contained only up to 600 nematodes. The authors do not offer an explanation.
I am not surprised by the apparently strong correlation between numbers and proximity to the disease source (a symptomatic tree). Nor am I surprised that Lcm nematodes were also found in funnels 11 meters away. I do wonder, however, why they are certain that no source was closer. Detecting early stage infections is notoriously difficult.
beech with large canopy; photo by F.T. Campbell
Goraya et al. also evaluated the effect of size of the source tree. They used dbh a substitute for larger canopies. Trees with larger canopies can host more nematodes, so are likely to contribute more to dispersal events. Two sets of funnels were equidistant from separate “source” trees – 3.51 m. One tree was small – 5.6 cm dbh, 11% as large as the other tree (50 cm). They collected many fewer Lcm nematodes from the smaller tree – the maximum was only 132 compared to 600 (a decrease of 78%).
Still, small trees can apparently support spread of the nematode to a reasonable distance. The fourth set of funnels was set up more than three times farther away (11.74 m) from an infected tree of a similar size (dbh = 5 cm) but recovered almost the same number of Lcm nematodes (0 – 119).
I find it alarming that both small trees in this part of the experiment had low BLD symptoms – only a few leaves were banded. Yet they apparently are the source of Lcm spread. The alternative, as I noted above, is that other “source” trees were in the vicinity but were not detected, possibly because they did not yet display symptoms?
Goraya et al. conclude that “source” tree size directly impacts the number of recovered nematodes. In addition, wind plays a pivotal role in their local distribution. This suggests a complex dispersal pattern in which proximity to the source leads to higher numbers of nematodes but longer-distance spread is possible.
Tussock moth; photo by Jon Yuschock via Bugwood
Nematodes’ association with other organisms
Goraya et al. (2024) collected one each of six caterpillar species from BLD-symptomatic trees. The frass of one – the tussock moth caterpillar (Halysidota tessellaris) — contained 12 nematode specimens — 10 of them Lcm. Two of the Lcm were alive and active. Their presence indicates that Lcm can survive passage through the caterpillar’s gastrointestinal tract. The authors conclude that caterpillars feeding on symptomatic leaves might contribute to local dispersal of Lcm.
Hundreds of Lcm were recovered from the two spider webs collected from the branches of a BLD-infected beech tree. From one web, 255 nematodes were captured; 58 were active. In the second web there were only 34 Lcm, but one-third — 10 – were active.
Goraya et al. (2024) hypothesized that any biotic form having the ability to move from a BLD-infected tree would be able to transport Lcm to other non-infected trees. Beyond caterpillars, they speculate that birds consuming these caterpillars might also disperse Lcm. Doug Tallamy has documented that many birds feed on caterpillars, link although he is focused on those that consume caterpillars in the spring, not the autumn. They note that others are studying that the bird species that feed on beech buds (e.g., finches) might transport nematodes. They note the need for additional research to clarify whether the nematode can survive birds’ digestive system.
Re: detection of live Lcm in spider webs, Goraya et al. suggest two possible interpretations: 1) this finding demonstrates that nematodes might fall from leaves, potentially spreading the infection to other trees beneath the canopy. (Supporting this idea is the fact that sub-canopy trees are often heavily infected with BLD and are frequently the first to exhibit BLD symptoms.) 2) Nematodes in spider webs are very likely to be transported by other “incidental organisms” (e.g., insects, birds, mammals) that feed on invertebrates trapped in webs — thereby potentially increasing the number and impact of nonspecific nematode vectors.
In conclusion, Goraya et al. found that many factors, e.g., distance & size of infected beech trees, wind speed, & humidity, contribute significantly to Lcm dispersal. The multitude of organisms interacting beneath the canopy also play a role.
They suggest that several major questions still need to be explored. These include how Lcm navigate environmental factors in their spread; and whether Lcm can survive – perhaps in a anhydrobioses state –transport over long distances, whether by abiotic or biotic vectors.
I remind my readers of the importance of beech in the hardwood forests in northeastern North America. Many wild animals, including squirrels, wild turkeys, white-tailed deer, and bears depend on beechnuts for fats and proteins. Moreover, some insects birds rely on beech tree canopies for shelter & nesting.
Other Hosts
Beech leaf disease attacks not just American beech (Fagus grandifolia). In North America, it has also attacked planted European beech(F. sylvatica), Chinese beech (F. engleriana), and Oriental beech (F. orientalis). Thus if it spreads it could have severe impacts across forests of much of the Northern Hemisphere.
range of European beech; from Royal Botanic Gardens, Kew
I appreciate that this project was funded by the USDA Forest Service International Program. I will pursue information concerning efforts by USFS Research and Development and the Forest Health Protection program.
SOURCE
Goraya, M., C. Kantor, P. Vieira, D. Martin, M. Kantor. 2024 Deciphering the vectors: Unveiling the local dispersal of Litylenchus crenatae ssp mccanni in the American beech (Fagus grandifolia) forest ecosystem PLOS ONE |https://doi.org/10.1371/journal.pone.0311830 November 8, 2024 1 / 16
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at https://treeimprovement.tennessee.edu/
ʻŌhiʻa trees killed by ROD; photo by Richard Sniezko, USFS
Several Hawaiian tree species are at risk due to introduced forest pests. Two of the Islands’ most widespread species are among the at-risk taxa. Their continuing loss would expose watersheds on which human life and agriculture depend. Habitats for hundreds of other species – many endemic and already endangered – would lose their foundations. These trees also are of the greatest cultural importance to Native Hawaiians.
I am pleased to report that Hawaiian scientists and conservationists are trying to protect and restore them.
Other tree species enjoy less recognition … and efforts to protect them have struggled to obtain support.
1) koa (Acacia koa)
Koa is both a dominant canopy tree and the second-most abundant native tree species in Hawai`i in terms of areas covered. The species is endemic to the Hawaiian archipelago. Koa forests provide habitat for 30 of the islands’ remaining 35 native bird species, many of which are listed under the U.S. Endangered Species Act. Also dependent on koa forests are native plant and invertebrate species and the Islands’ only native terrestrial mammal, the Hawaiian hoary bat. Finally, koa forests protect watersheds, add nitrogen to degraded soils, and store carbon [Inman-Narahari et al.]
Koa forests once ranged from near sea level to above 7000 ft (2100 m) on both the wet and dry sides of all the large Hawaiian Islands. Conversion of forests to livestock grazing and row-crop agriculture has reduced koa’s range. Significant koa forests are now found on four islands – Hawai’i, Maui, O‘ahu, and Kauaʻi. More than 90% of the remaining koa forests occur on Hawai`i Island (the “Big Island) [Inman-Narahari et al.]
In addition to its fundamental environmental role, koa has immense cultural importance. Koa represents strength and the warrior spirit. The wood was used traditionally to make sea-going canoes. Now Koa is widely used for making musical instruments, especially guitars and ukuleles; furniture, surfboards, ornaments, and art [Inman-Narahari et al.]
Koa timber has the highest monetary value of any wood harvested on the Islands. However, supplies of commercial-quality trees are very limited (Dudley et al. 2020). Harvesting is entirely from old-growth forests on private land. [Inman-Narahari et al.]
Koa forests are under threat by a vascular wilt disease caused by Fusarium oxysporum f. sp. koae (FOXY). This disease can kill up to 90% of young trees and – sometimes — mature trees in native forests. The fungus is a soil-dwelling organism that spreads in soil and infects susceptible plants through the root system (Dudley et al. 2020).
Conservation and commercial considerations have converged to prompt efforts to breed koa resistant to FOXY. Conservationists hope to restore native forests on large areas where agriculture has declined. The forestry industry seeks to enhance supplies of the Islands’ most valuable wood. Finally, science indicated that a breeding program would probably be successful. Field trials in the 1990s demonstrated great differences in wilt-disease mortality among seed sources (the proportion of seedlings surviving inoculation ranged from 4% to 91.6%) [Sniezko 2003; Dudley et al. 2009].
In 2003, Dudley and Sniezko outlined a long-term strategy for exploring and utilizing genetic resistance in koa. Since then, a team of scientists and foresters has implemented different phases of the strategy and refined it further (Dudley et al. 2012, 2015, 2017; Sniezko et al. 2016]
First, scientists determined that the wilt disease is established on the four main islands. Having obtained more than 500 isolates of the pathogen from 386 trees sampled at 46 sites, scientists tested more than 700 koa families from 11 ecoregions for resistance against ten of the most highly virulent isolates (Dudley et al. 2020).
The Hawaiian Agricultural Research Center (HARC), supported by public and private partners, has converted the field-testing facilities on Hawai`i, Maui, and Oahu into seed orchards. The best-performing tree families are being grown to maturity to produce seeds for planting. It is essential that the seedlings be not just resistant to FOXY but also adapted to the ecological conditions of the specific site where they are to be planted [Dudley et al. 2020; Inman-Narahari et al. ] Locally adapted, wilt-resistant seed has been planted on Kauaʻi and Hawai`i. Preparations are being made to plant seed on Maui and O‘ahu also. Scientists are also exploring methods to scale up planting in both restoration and commercial forests [R. Hauff pers. comm.].
koa; photo by David Eickhoff via Flickr
Restoration of koa on the approximately half of lands in the species’ former range that are privately owned will require that the trees provide superior timber. Private landowners might also need financial incentives since the rotation time for a koa plantation is thought to be 30-80 years. [Inman-Narahari et al.]
Plantings on both private and public lands will need to be protected from grazing by feral ungulates and encroachment by competing plants. These management actions are intensive, expensive, and must be maintained for years.
Some additional challenges are scientific: uncertainties about appropriate seed zones, efficacy of silvicultural approaches to managing the disease, and whether koa can be managed for sustainable harvests. Human considerations are also important: Hawai`i lacks sufficient professional tree improvement or silvicultural personnel, a functioning seed distribution and banking network — and supporting resources. Finally, some segments of the public oppose ungulate control programs. Inman-Narahari et al.
Finally, scientists must monitor seed orchards and field plantings for any signs of maladaptation to climate change. (Dudley et al. 2020).
2) ʻŌhiʻa Metrosideros polymorpha)
ʻŌhiʻa lehua is the most widespread tree on the Islands. It dominates approximately 80% the biomass of Hawaii’s remaining native forest, in both wet and dry habitats. ʻŌhiʻa illustrates adaptive radiation and appears to be undergoing incipient speciation. The multitude of ecological niches and their isolation on the separate islands has resulted in five recognized species in the genus Metrosideros. Even the species found throughout the state, Metrosideros polymorpha, has eight recognized varieties (Luiz et al. (2023) (some authorities say there are more).
Loss of this iconic species could result in significant changes to the structure, composition, and potentially, the function, of forests on a landscape level. High elevation ‘ohi‘a forests protect watersheds across the state. ʻŌhiʻa forests shelter the Islands’ one native terrestrial mammal (Hawaiian hoary bat), 30 species of forest birds, and more than 500 endemic arthropod species. Many species in all these taxa are endangered or threatened (Luiz et al. 2023). The increased light penetrating interior forests following canopy dieback facilitates invasion by light-loving non-native plant species, of which Hawai`i has dozens. There is perhaps no other species in the United States that supports more endangered taxa or that plays such a geographical dominant ecological keystone role [Luiz et al. 2023]
For many Native Hawaiians, ‘ōhi‘a is a physical manifestation of multiple Hawaiian deities and the subject of many Hawaiian proverbs, chants, and stories; and foundational to the scared practice of many hula. The wood has numerous uses. Flowers, shoots, and aerial roots are used medicinally and for making lei. The importance of the biocultural link between ‘ōhi‘a and the people of Hawai`i is described by Loope and LaRosa (2008) and Luiz et al. (2023).
In 2010 scientists detected rapid mortality affecting ‘ōhi‘a on Hawai‘i Island. Scientists determined that the disease is caused by two recently-described pathogenic fungi, Ceratocystis lukuohia and Ceratocystis huliohia. The two diseases, Ceratocystis wilt and Ceratocystis canker of ʻōhiʻa, are jointly called “rapid ‘ōhi‘a death”, or ROD. The more virulent species, C. lukuohia, has since spread across Hawai`i Island and been detected on Kaua‘i. The less virulent C. huliohia is established on Hawai`i and Kaua‘i and in about a dozen trees on O‘ahu. One tree on Maui was infected; it was destroyed, and no new infection has been detected [M. Hughes pers. comm.] As of 2023, significant mortality has occurred on more than one third of the vulnerable forest on Hawai`i Island, although mortality is patchy.
[ʻŌhiʻa is also facing a separate disease called myrtle rust caused by the fungus Austropuccinia psidii; to date this rust has caused less virulent infections on ‘ōhi‘a.]
rust-killed ‘ōhi‘a in 2016; photo by J.B. Friday
Because of the ecological importance of ‘ōhi‘a and the rapid spread of these lethal diseases, research into possible resistance to the more virulent pathogen, C. lukiohia began fairly quickly, in 2016. Some ‘ōhi‘a survive in forests on the Big Island in the presence of ROD, raising hopes that some trees might possess natural resistance. Scientists are collecting germplasm from these lightly impacted stands near high-mortality stands (Luiz et al. 2023). Five seedlings representing four varieties of M. polymorpha that survived several years’ exposure to the disease are being used to produce rooted cuttings and seeds for further evaluation of these genotypes.
ʻŌhiʻa flowers
Encouraged by these developments, and recognizing the scope of additional work needed, in 2018 stakeholders created a collaborative partnership that includes state, federal, and non-profit agencies and entities, ʻŌhiʻa Disease Resistance Program (‘ODRP) (Luiz et al. 2023). The partnership seeks to provide baseline information on genetic resistance present in all Hawaiian taxa in the genus Metrosideros. It aims further to develop sources of ROD-resistant germplasm for restoration intended to serve several purposes: cultural plantings, landscaping, and ecological restoration. ‘ODRP is pursuing screenings of seedlings and rooted cuttings sampled from native Metrosideros throughout Hawai`i while trying to improve screening and growing methods. Progress will depend on expanding these efforts to include field trials; research into environmental and genetic drivers of susceptibility and resistance; developing remote sensing and molecular methods to rapidly detect ROD-resistant individuals; and support already ongoing Metrosideros conservation. If levels of resistance in wild populations prove to be insufficient, the program will also undertake breeding (Luiz et al. 2023).
To be successful, ‘ODRP must surmount several challenges (Luiz et al. 2022):
increase capacity to screen seedlings from several hundred plants per year to several thousand;
optimize artificial inoculation methodologies;
determine the effects of temperature and season on infection rates and disease progression;
find ways to speed up seedlings’ attaining sufficient size for testing;
develop improved ways to propagate ʻōhiʻa from seed and rooted cuttings;
establish sites for field testing of putatively resistant trees across a wide range of climatic and edaphic conditions;
establish seed orchard, preferably on several islands;
establish systems for seed collection from the wide variety of subspecies/varieties;
if breeding to enhance resistance is appropriate, it will be useful to develop high-throughput phenotyping of the seed orchard plantings.
Developing ROD-resistant ‘ōhi‘a is only one part of a holistic conservation program. Luiz et al. (2023) reiterate the importance of quarantines and education to curtail movement of infected material and countering activities that injure the trees. Fencing to protect these forests from grazing by feral animals can drastically reduce the amount of disease. Finally, scientists must overcome the factors there caused the almost complete lack of natural regeneration of ‘ōhi‘a in lower elevation forests. Most important are competition by invasive plants, predation by feral ungulates, and the presence of other diseases, e.g., Austropuccinia psidii.
Hawaii’s dryland forests are highly endangered: more than 90% of dry forests are already lost due to habitat destruction and the spread of invasive plant and animal species. Two tree species are the focus of species-specific programs aimed at restoring them to remaining dryland forests. However, support for both programs seems precarious and requires stable long-term funding; disease resistance programs often necessitate decades-long endeavors.
naio in bloom; photo by Forrest & Kim Starr via Creative Commons
1) naio (Myoporum sandwicense)
Naio grows on all of the main Hawaiian Islands at elevations ranging from sea level to 3000 m. While it occurs in the full range of forest types from dry to wet, naio is one of two tree species that dominate upland dry forests. The other species is mamane, Sophora chrysophylla. Naio is a key forage tree for two endangered honeycreepers, palila (Loxioides bailleui) and `akiapola`au (Hemignathus munroi). The tree is also an important host of many species of native yellow-face bees (Hylaeus spp). Finally, loss of a native tree species in priority watersheds might lead to invasions by non-native plants that consume more water or increase runoff.
The invasive non-native Myoporum thrips, Klambothrips myopori, was detected on Hawai‘i Island in December 2008 (L. Kaufman website). In 2018 the thrips was found also on Oahu (work plan). The Myoporum thrips feeds on and causes galls on plants’ terminal growth. This can eventually lead to death of the plant.
Aware of thrips-caused death of plants in the Myoporum genus in California, the Hawaii Department of Lands and Natural Resources Division of Forestry and Wildlife and the University of Hawai‘i began efforts to determine the insect’s distribution and infestation rates, as well as the overall health of naio populations on the Big Island. This initiative began in September 2010, nearly two years after the thrips’ detection. Scientists monitored nine protected natural habitats for four years. This monitoring program was supported by the USFS Forest Health Protection program. This program is described by Kaufman.
naio monitoring sites from L. Kaufman article
The monitoring program determined that by 2013, the thrips has spread across most of Hawi`i Island, on its own and aided by human movement of landscaping plants. More than 60% of trees being monitored had died. Infestation and dieback levels had both increased, especially at medium elevation sites. The authors feared that mortality at high elevations would increase in the future. They found no evidence that natural enemies are effective controlling naio thrips populations on Hawai`i Island.
Kaufman was skeptical that biological control would be effective. She suggested, instead, a breeding program, including hybridizing M. sandwicensis with non-Hawaiian Myoporum species that appear to be resistant to thrips. Kaufman also called for additional programs: active monitoring to prevent thrips from establishing on neighboring islands; and collection and storage of naio seeds.
Ten years later, in February 2024, DLNR Division of Forestry and Wildlife adopted a draft work plan for exploring possible resistance to the Myoporum thrips. Early steps include establishing a database to record data needed to track parent trees, associated propagules, and the results of tests. These data are crucial to keeping track of which trees show the most promise. Other actions will aim to hone methods and processes. Among practical questions to be answered are a) whether scientists can grow even-aged stands of naio seedlings; b) identifying the most efficient resistance screening techniques; and c) whether K. myopori thrips are naturally present in sufficient numbers to be used in tests, or – alternatively – whether they must be augmented. [Plan]
Meanwhile, scientists have begun collecting seed from unaffected or lightly affected naio in hotspots where mortality is high. They have focused on the dry and mesic forests of the western side of Hawai`i (“Big”) Island, where the largest number of naio populations still occur and are at high risk. Unfortunately, these “lingering” trees remain vulnerable to other threats, such as browsing by feral ungulates, competition with invasive plants, drought, and reduced fecundity & regeneration.
Hawai`i DLNR has secured initial funding from the Department of Defense’s REPI program to begin a pest resistance project and is seeking a partnership with University of Hawai`i to carry out tests “challenging” different naio families’ resistance to the thrips [R. Hauff pers. comm.]
wiliwili; photo by Forrest & Kim Starr
2) wiliwili (Erythrina sandwicensis)
Efforts to protect the wiliwili have focused on biological control. The introduced Erythrina gall wasp, Quadrastichus erythrinae (EGW) was detected on the islands in 2005. It immediately caused considerable damage to the native tree and cultivated nonnative coral trees.
A parasitic wasp, Eurytoma erythrinae, was approved for release in November 2008 – only 3 ½ years after EGW was detected on O‘ahu. The parasitic wasp quickly suppressed the gall wasp’s impacts to both wiliwili trees and non-native Erythrina. By 2024, managers are once again planting the tree in restoration projects.
However, both the gall wasp and a second insect pest – a bruchid, Specularius impressithorax – can cause loss of more than 75% of the seed crop. This damage means that the tree cannot regenerate. By 2019, Hawaiian authorities began seeking permission to release a second biocontrol gent, Aprostocitus nites.Unfortunately, the Hawai’i Department of Agriculture still has not approved the release permit despite five years having passed. Once they have this approval, the scientists will then need to ask USDA Animal and Plant Health Inspection Service (APHIS) for its approval [R. Hauff, pers. comm.]
SOURCES
www.RapidOhiaDeath.org
Dudley, N., R. James, R. Sniezko, P. Cannon, A. Yeh, T. Jones, & Michael Kaufmann. 2009? Operational Disease Screening Program for Resistance to Wilt in Acacia koa in Hawai`i. Hawai`i Forestry Association Newsletter August 29 2009
Dudley, N., T. Jones, K. Gerber, A.L. Ross-Davis, R.A. Sniezko, P. Cannon & J. Dobbs. 2020. Establishment of a Genetically Diverse, Disease-Resistant Acacia koa Seed Orchard in Kokee, Kauai: Early Growth, Form, & Survival. Forests 2020, 11, 1276; doi:10.3390/f11121276 www.mdpi.com/journal/forests
Friday, J. B., L. Keith, and F. Hughes. 2015. Rapid ʻŌhiʻa Death (Ceratocystis Wilt of ʻŌhiʻa). PD-107, College of Tropical Agriculture and Human Resources, University of Hawai‘i, Honolulu, HI. URL: https://www.ctahr.HI.edu/oc/freepubs/pdf/PD-107.pdf Accessed April 3, 2018.
Friday, J.B. 2018. Rapid ??hi?a Death Symposium -West Hawai`i (“West Side Symposium”) March 3rd 2018, https://vimeo.com/258704469 Accessed April 4, 2018 (see also full video archive at https://vimeo.com/user10051674)
Inman-Narahari, F., R. Hauff, S.S. Mann, I. Sprecher, & L. Hadway. Koa Action Plan: Management & research priorities for Acacia koa forestry in Hawai`i. State of Hawai`i Department of Land & Natural Resources Division of Forestry & Wildlife no date
Kaufman, L.V, J. Yalemar, M.G. Wright. In press. Classical biological control of the erythrina gall wasp, Quadrastichus erythrinae, in Hawaii: Conserving an endangered habitat. Biological Control. Vol. 142, March 2020
Loope, L. and A.M. LaRosa. 2008. ‘Ohi’a Rust (Eucalyptus Rust) (Puccinia psidii Winter) Risk Assessment for Hawai‘i.
Luiz, B.C. 2017. Understanding Ceratocystis. sp A: Growth, morphology, and host resistance. MS thesis, University of Hawai‘i at Hilo.
Luiz, B.C., C.P. Giardina, L.M. Keith, D.F. Jacobs, R.A. Sniezko, M.A. Hughes, J.B. Friday, P. Cannon, R. Hauff, K. Francisco, M.M. Chau, N. Dudley, A. Yeh, G. Asner, R.E. Martin, R. Perroy, B.J. Tucker, A. Evangelista, V. Fernandez, C. Martins-Keli’iho.omalu, K. Santos, R. Ohara. 2023. A framework for establishlishing a rapid ‘Ohi‘a death resistance program New Forests 54, 637–660. https://doi.org/10.1007/s11056-021-09896-5
Sniezko, R.A., N. Dudley, T. Jones, & P. Cannon. 2016. Koa wilt resistance & koa genetics – key to successful restoration & reforestation of koa (Acacia koa). Acacia koa in Hawai‘i: Facing the Future. Proceedings of the 2016 Symposium, Hilo, HI: www.TropHTIRC.org , www.ctahr.HI.edu/forestry
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at https://treeimprovement.tennessee.edu/
The number of introduced forest pathogens are increasing – creating a crisis that is recognized by more scientists. These experts say tree diseases are reshaping both native and planted forests around the globe. The diseases are threatening biodiversity, ecosystem services, provision of products, and related human wellbeing. Some suggest that bioinvasions might threaten forests as much as climate change, while also undermining forests’ role in carbon sequestration.
Unfortunately, I see little willingness within the plant health regulatory community to tackle improving programs to slow introductions. Even when the scientists documenting the damage work for the U.S. Department of Agriculture – usually the U.S. Forest Service — USDA policy-makers don’t act on their findings. [I tried to spur a conversation with USDA 2 years ago. So far, no response.]
counties where beech leaf disease has been detected
What the scientists say about these pests’ impacts
Andrew Gougherty (2023) – one of the researchers employed by the USDA Forest Service – says that emerging infectious tree diseases are reshaping forests around the globe. Furthermore, new diseases are likely to continue appearing in the future and threaten native and planted forests worldwide. [Full references are provided at the end of the blog.] Haoran Wu (2023/24) – a Master’s Degree student at Oxford University – agrees that arrival of previously unknown pathogens are likely to alter the structure and composition of forests worldwide. Weed, Ayers, and Hicke (2013) [academics] note that forest pests — native and introduced — are the dominant sources of disturbance to North American forests. They suggest that, globally, bioinvasions might be at least as important as climate change as threats to the sustainability of forest ecosystems. They are concerned that recurrent forest disturbances caused by pests might counteract carbon mitigation strategies.
Scientists have proclaimed these warnings for years. Five years ago, Fei et al. (2019) reported that the 15 most damaging pests introduced to the United States — cumulatively — had already caused tree mortality to exceed background levels by 5.53 teragrams of carbon per year. As these 15 pests spread and invasions intensify, they threaten 41.1% of the total live forest biomass in the 48 coterminous states. Poland et al. (2019) (again – written by USFS employees) document the damage to America’s forest ecosystems caused by the full range of invasive species, terrestrial and aquatic.
Fei et al. and Weed, Ayers, and Hicke (2013) also support the finding that old, large trees are the most important trees with regard to carbon storage. This understanding leads them to conclude that the most damaging non-native pests are the emerald ash borer, Dutch elm disease fungi, beech bark disease, and hemlock woolly adelgid. As I pointed out in earlier blogs, other large trees, e.g., American chestnut and several of the white pines, were virtually eliminated from much of their historical ranges by non-native pathogens decades ago. These same large, old, trees also maintain important aspects of biological diversity.
It is true that not all tree species are killed by any particular pest. Some tree genera or species decrease while others thrive, thus altering the species composition of the affected stands (Weed, Ayers, and Hicke). This mode of protection is being undermined by the proliferation of insects and pathogens that cumulatively attack ever more tree taxa. And while it is true that some of the carbon storage capacity lost to pest attack will be restored by compensatory growth in unaffected trees, this faster growth is delayed by as much as two or more decades after pest invasions begin (Fei et al.).
ash forest after EAB infestation; Photo by Nate Siegert, USFS
Still, despite the rapid rise of destructive tree pests and disease outbreaks, scientists cannot yet resolve critical aspects of pathogens’ ecological impacts or relationship to climate change. Gougherty notes that numerous tree diseases have been linked to climate change or are predicted to be impacted by future changes in the climate. However, various studies’ findings on the effects of changes in moisture and precipitation are contradictory. Wu reports that his study of ash decline in a forest in Oxfordshire found that climate change will have a very small positive impact on disease severity through increased pathogen virulence. Weed, Ayers, and Hicke go farther, making the general statement that despite scientists’ broad knowledge of climate effects on insect and pathogen demography, they still lack the capacity to predict pest outbreaks under climate change. As a result, responses intended to maintain ecosystem productivity under changing climates are plagued by uncertainty.
Clarifying how disease systems are likely to interact with predicted changes in specific characteristics of climate is important — because maintaining carbon storage levels is important. Quirion et al. (2021) estimate that, nation-wide, native and non-native pests have decreased carbon sequestration by live forest trees by at least 12.83 teragrams carbon per year. This equals approximately 9% of the contiguous states’ total annual forest carbon sequestration and is equivalent to the CO2 emissions from more than 10 million passenger vehicles driven for one year. Continuing introductions of new pests, along with worsening effects of native pests associated with climate change, could cause about 30% less carbon sequestration in living trees. These impacts — combined with more frequent and severe fires and other forest disturbances — are likely to negate any efforts to improve forests’ capacity for storing carbon.
Understanding pathogens’ interaction with their hosts is intrinsically complicated. There are multiple biological and environmental factors. What’s more, each taxon adapts individually to the several environmental factors. Wu says there is no general agreement on the relative importance of the various environmental factors. The fact that most forest diseases are not detected until years after their introduction also complicates efforts to understand factors affecting infection and colonization.
The fungal-caused ash decline in Europe is a particularly alarming example of the possible extent of such delays. According to Wu, when the disease was first detected – in Poland in 1992 – it had already been present perhaps 30 years, since the 1960s. Even then, the causal agent was not isolated until 2006 – or about 40 years after introduction. The disease had already spread through about half the European continent before plant health officials could even name the organism. The pathogen’s arrival in the United Kingdom was not detected until perhaps five years after its introduction – despite the country possessing some of the world’s premier forest pathologists who by then (2012) knew what they to look for.
Clearly, improving scientific understanding of forest pathogens will be difficult. In addition, effective policy depends on understanding the social and economic drivers of trade, development, and political decisions are primary drivers of the movement of pathogens. Wu calls for collaboration of ecologists, geneticists, earth scientists, and social scientists to understand the complexity of the host-pathogen-surrounding system. Bringing about this new way of working and obtaining needed resources will take time – time that forests cannot afford.
However, Earth’s forests are under severe threat now. Preventing their collapse depends on plant health officials integrating recognition of these difficulties into their policy formulation. It is time to be realistic: develop and implement policies that reflect the true level of threat and limits of current science.
Background: Rising Numbers of Introductions
Gougherty’s analysis of rising detections of emerging tree diseases found little evidence of saturation globally – in accord with the findings of Seebens et al. (2017) regarding all taxa. Relying on data for 24 tree genera, nearly all native to the Northern Hemisphere, Gougherty found that the number of new pests attacking these tree genera are doubling on average every 11.2 years. Disease accumulation is increasing rapidly in both regions where hosts are native and where they are introduced, but more rapidly in trees’ native ranges.This finding is consistent with most new diseases arise from introductions of pathogens to naïve hosts.
Gougherty says his estimates are almost certainly underestimates for a number of reasons. Countries differ in scientific resources and their scientists’ facility with English. Scientists are more likely to notice and report high-impact pathogens and those in high-visibility locations. Where national borders are closer, e.g., in Europe, a minor pest expansion can be reported as “new” in several countries. New pathogens in North America appear to occur more slowly, possibly because the United States and Canada are very large. He suggests that another possible factor is the U.S. (I would add Canada) have adopted pest-prevention regulations that might be more effective than those in place in other regions. (See my blogs and the Fading Forest reports linked to below for my view of these measures’ effectiveness.)
ash dieback in the UK
Wu notes that reports of tree pathogens in Europe began rising suddenly after the 1980s. He cites the findings by Santini et al. (2012) that not only were twice as many pathogens detected in the period after 1950 than in the previous 40 years, the region of origin also changed. During the earlier period, two-thirds of the introduced pathogens came from temperate North America. After 1950, about one-third of previously unknown disease agents were from temperate North America. Another one-third was from Asia. By 2012, more than half of plant infectious diseases were caused by introduction of previously unknown pathogens.
What is to be done?
Most emerging disease agents do not have the same dramatic effects as chestnut blight in North America, ash dieback in Europe, or Jarrah dieback in Australia. Nevertheless, as Gougherty notes, their continued emergence in naïve biomes increases the likelihood of especially damaging diseases emerging and changing forest community composition.
Gougherty calls for policies intended to address both the agents being introduced through trade, etc., and those that emerge from shifts in virulence or host range of native pathogens or changing environmental conditions. In his view, stronger phytosanitary programs are not sufficient.
Wu recommends enhanced monitoring of key patterns of biodiversity and ecosystem functioning, He says these studies should focus on the net outcome of complex interactions. Wu also calls for increasing understanding of key “spillover” effects – outcomes that cannot be currently assessed but might impact the predicted outcome. He lists several examples:
the effects of drought–disease interactions on tree health in southern Europe,
interaction between host density and pathogen virulence,
reproductive performance of trees experiencing disease,
effect of secondary infections,
potential for pathogens to gain increased virulence through hybridization.
potential for breeding resistant trees to create a population buffer for saving biological diversity. Wu says his study of ash decline in Oxfordshire demonstrates that maintaining a small proportion of resistant trees could help tree population recovery.
Quirion et al. provide separate recommendations with regard to native and introduced pests. To minimize damage from the former, they call for improved forest management – tailored to the target species and the environmental context. When confronting introduced pests, however, thinning is not effective. Instead, they recommend specific steps to minimize introductions via two principal pathways, wood packaging and imports of living plants. In addition, since even the most stringent prevention and enforcement will not eliminate all risk, Quirion et al. advocate increased funding for and research into improved strategies for inspection, early detection of new outbreaks, and strategic rapid response to newly detected incursions. Finally, to reduce impacts of established pests, they recommend providing increased and more stable funding for classical biocontrol, research into technologies such as sterile-insect release and gene drive, and host resistance breeding.
USDA HQ
Remember: reducing forest pest impacts can simultaneously serve several goals—carbon sequestration, biodiversity conservation, and perpetuating the myriad economic and societal benefits of forests. See Poland et al. and the recent IUCN report on threatened tree species.
SOURCES
Barrett, T.M. and G.C. Robertson, Editors. 2021. Disturbance and Sustainability in Forests of the Western United States. USDA Forest Service Pacific Northwest Research Station. General Technical Report PNW-GTR-992. March 2021
Clark, P.W. and A.W. D’Amato. 2021. Long-term development of transition hardwood and Pinus strobus – Quercus mixedwood forests with implications for future adaptation and mitigation potential. Forest Ecology and Management 501 (2021) 119654
Fei, S., R.S. Morin, C.M. Oswalt, and A.M. 2019. Biomass losses resulting from insect and disease invasions in United States forests. Proceedings of the National Academy of Sciences. www.pnas.org/cgi/doi/10.1073/pnas.1820601116
Gougherty AV (2023) Emerging tree diseases are accumulating rapidly in the native and non-native ranges of Holarctic trees. NeoBiota 87: 143–160. https://doi.org/10.3897/neobiota.87.103525
Lovett, G.M., C.D. Canham, M.A. Arthur, K.C. Weathers, and R.D. Fitzhugh. 2006. Forest Ecosystem Responses to Exotic Pests and Pathogens in Eastern North America. BioScience Vol. 56 No. 5 May 2006
Lovett, G.M., M. Weiss, A.M. Liebhold, T.P. Holmes, B. Leung, K.F. Lambert, D.A. Orwig, F.T. Campbell, J. Rosenthal, D.G. MCCullough, R. Wildova, M.P. Ayres, C.D. Canham, D.R. Foster, S.L. Ladeau, and T. Weldy. 2016. Nonnative forest insects and pathogens in the United States: Impacts and policy options. Ecological Applications, 26(5), 2016, pp. 1437-1455
Poland, T.M., Patel-Weynand, T., Finch, D., Miniat, C. F., and Lopez, V. (Eds) (2019), Invasive Species in Forests and Grasslands of the United States: A Comprehensive Science Synthesis for the United States Forest Sector. Springer Verlag.
Quirion, B.R., G.M. Domke, B.F. Walters, G.M. Lovett, J.E. Fargione, L. Greenwood, K. Serbesoff-King, J.M. Randall, and S. Fei. 2021 Insect and Disease Disturbance Correlate With Reduced Carbon Sequestration in Forests of the Contiguous US. Front. For. Glob. Change 4:716582. [Volume 4 | Article 716582] doi: 10.3389/ffgc.2021.716582
Weed, A.S., M.P. Ayers, and J.A. Hicke. 2013. Consequences of climate change for biotic disturbances in North American forests. Ecological Monographs, 83(4), 2013, pp. 441–470
Wu, H. 2023/24. Modelling Tree Mortality Caused by Ash Dieback in a Changing World: A Complexity-based Approach MSc/MPhil Dissertation Submitted August 12, 2024. School of Geography and the Environment, Oxford University.
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at https://treeimprovement.tennessee.edu/
tanoak killed by P. ramorum; photo by F.T. Campbell
I am belatedly catching up on the situation with regard to Phytophthora ramorum – sudden oak death – in the US and other countries.
For a general factsheet on this plant disease, see profile here. Here, I’m summarizing more detailed information contained in the February, May, and August 2024 newsletters of the California Oak Mortality Task Force (COMTF) (Newsletters for earlier months are posted here.)
To obtain the most recent information, you can attend the Fall 2024 virtual meeting of the Task Force on Tuesday, October 29, 2024, from 1 pm to 3 pm PDT. Speakers will focus on the status of P. ramorum in California and Oregon wildlands.
On the next day, Wednesday, October 30, the Phytophthoras in Native Habitats Work Group will discuss “Threats to California Native Plants” including from viruses and excessive heat, along with other concerns.
Participation is free, but registration is required. Complete agendas and more information will be available soon here. Sessions will be recorded and posted to the same site. Questions? Contact Janice Alexander.
More in-depth information à Matteo Garbelotto’s UC Berkeley class, “Ecology and Impacts of Emergent Forest Diseases in California,” is now available free and online. Recommended reading, lecture recordings, slides, even essay topic suggestions are posted. Subjects covered include several high impact forest diseases, molecular diagnostics, disease control, and prevention.
I note that the recent detections of new outbreaks in forests and nurseries support the importance of weather in promoting or hindering establishment and spread of Phytophthora ramorum.
Phytophthora ramorum in North American Forests
In Oregon, P. ramorum continued to spread in 2023 and the first half of 2024.
These outbreaks were detected through extensive surveillance. Aerial monitoring (in cooperation with the USDA Forest Service) and high-resolution imagery covered more than half a million acres in Curry County — the region between the California border and the Coos County line. Ground surveys covered 860 acres. Sampling included 518 trees; 117 tested were positive for the fungus. Stream baits were deployed to 63 sites; 26 tested positive at least once (COMTF newsletter, February 2024; includes maps).
By summer 2024, 23 new P. ramorum infestations had been detected at or beyond the Generally Infested Area (GIA; the area where the pathogen is most commonly found) since 2021. Some of these involve one of the newly detected clonal lineages. Oregon officials are expecting to expand the state’s quarantine area to 901 square miles – 45% of Curry County. The designated GIA would also be enlarged to 178 square miles(COMTF newsletter, August 2024; contains maps).
Oregon continues trying to treat high-priority infestations. In 2023, the state treated 165 acres infested by one of the newly detected clonal lineages, NA2, in the Humbug Mountain area and 347 acres in the Port Orford infestation. Since 2001, Oregon’s Department of Forestry has completed eradication treatments on more than 9,000 acres at an estimated cost of over $37 million. Federal lands comprised 28% of treated acres; the remainder were private and state lands. Still, more than 1,000 high-priority acres have not been treated because neither state nor federal agencies could provide sufficient funds (COMTF newsletter, February 2024).
The stream baiting program in 64 stream drainages has – so far – detected six positive streams. Ground surveys are planned for the new positive drainages along the north bank of the Rogue River and a stream that drains into the Elk River east of Port Orford (COMTF newsletter, August 2024).
In California, recent wet winters have prompted several new detections. The first was in Del Norte County near previously detected sites. The UC Berkeley-coordinated “SOD Blitz” plans intensive surveys in this region in coming months (COMTF newsletter May 2024; contains map).
Somewhat later, new infestations were detected farther south, in Humboldt Redwoods State Park. The new sites were outside the formerly detected sites, on the north side of the creek and up to the top of the ridge (COMTF newsletter, August 2024).
Scientists have realized another concern: several other pathogens cause symptoms on bay laurel, tanoak, and madrone that are almost indistinguishable from SOD. This development will complicate monitoring (COMTF newsletter for August 2024; see below for more details).
Meanwhile, scientists determined that sites where the P. ramorum epidemic is driven by higher bay laurel (Umbellularia californica) densities sustained a higher genotypic diversity of P. ramorum. While tanoak (Notholithocarpus densiflorus) doesn’t contribute much to infection of true oaks (Quercus spp.) it can infect bay laurel, thus perpetuating the infection. Infected oaks and tanoaks maintain host-specific pathogen genotypes (Kozanitas et al. 2024)
The USDA Forest Service program that monitors streams in the East to detect P. ramorum placed baits in 63 streams in 10 eastern states: Alabama, Florida, Georgia, Illinois, Maryland, Mississippi, North Carolina, Pennsylvania, South Carolina, and Texas. In 2023, positive findings for P. ramorum were detected from two streams in Alabama, and one each in Mississippi and North Carolina. All sites are associated with nurseries that had previously tested positive for P. ramorum. Over the last five years – since 2019 – eight streams in four states have tested positive at least once: five in Alabama, and one each in Mississippi, North Carolina, and South Carolina. The detection in South Carolina is new. Vegetation in the watershed has been sampled multiple times; all samples collected so far — plant, soil, and run-off water – have been negative. The pathogen belongs to the NA1 lineage – the one established in forests in West Coast states. [COMTF newsletter February 2024]
from D.J. Haller & M.C. Wimberly. 2020
Situation in Europe
The February 2024, the COMTF newsletter summarized the situation in Great Britain. In England, aerial surveillance covered more than 31,000 ha of larch (Larix kaempferi)plantations. Follow-up investigations detected considerably fewer infested sites than the approximately 200 detected in 2022. Most remain in the southwest and northwest of the country. Weather conditions in 2023 were less conducive for sporulation in 2021 and 2022, which seemed to lead to a reduced level of disease in 2022 and 2023.
In Scotland, widespread aerial and ground surveillance detected a number of sites similar to those found since 2018. Scottish authorities note that where positive findings are not quickly followed by tree removal, localized spread occurred.
In Wales, four helicopter surveillance flights identified around 150 sites deserving further investigation. About 60 of these sites held infected trees, mainly larch, but some noble fir (Abies procera). The COMTF newsletter contains a map showing infested locations. This year’s infection level might be less than in previous years, but this might reflect the fact that the infections are in smaller forest blocks. However, the wet and mild weather in autumn/winter 2023 provided optimal conditions for sporulation, so the scientist expected higher infection rates in 2024. The Welsh Government is working on a new strategy for managing P. ramorum.
In Northern Ireland, P. ramorum was described as still active and spreading. Only two surveys were flown. They identified 49 locations for follow-up, many in forests where the pathogen had been found previously. At two locations, follow-up inspections and sampling of larch confirmed infection by a different pathogen, Phytophthora pseudosyringae. So in 2024, larch samples will be tested for both P. ramorum and P. pseudosyringae.
Other Phytopthoras in Europe
English scientists are trying to determine how damaging P. pseudosyringae is on larch. Infections have been observed at several locations in the north of England, as well as in Northern Ireland (COMTF newsletter February 2024).
Mullet et al. (2024) report that P. pseudosyringae is a self-fertile pathogen of woody plants, especially tree species in the genera Fagus, Notholithocarpus, Nothofagus and Quercus. It is found across Europe and in parts of North America and Chile. Genetic studies show that the North American population originated from Europe. P. pseudosyringae can infect roots; the stem collar region; bark; twigs and stems; as well as leaves. They report it is causing particular damage in Great Britain and western North America. Mullet et al. call for investigation of differences in life history traits between the two main population clusters, including their virulence and host ranges.
Nothofagus obliqua; photo by Line1 via Wikimedia
Chile (COMTF newsletter May 2024)
Concerned about decades of mortality of Nothofagus trees in native forests in Chile, González et al. 2024 sought to understand which other native plants might be reservoirs of inoculum of the pathogen Phytophthora pseudosyringae — which is a documented causal agent of partial defoliation and bleeding cankers on two native tree species,Nothofagus obliqua and N. alpina. P. pseudosyringae can sporulate on lesions on Cryptocarya alba, Nothofagus dombeyi and N. obliqua leaves. On Sophora macrocarpa, sporulation occurs on both asymptomatic tissues and on lesions. S. macrocarpa is a common understory species in Nothofagus forests, so it might be an inoculum reservoir for epidemic events in them.
Look-alikes on California Bay Laurel (COMTF newsletter May 2024)
Similar symptoms from a wide variety of pathogenic organisms were detected on bay laurels after last year’s wet winter. Among the pathogens — the list is not exhaustive — includes P. cinnamomi, Neofusicoccum nonquaesitum, Ganoderma brownie, P. pseudosyringae, P. nemorosa, Botryosphaeria dothidea, Armillaria gallica, Diplodia corticola, and others.
Foliar symptoms tend to look identical on bay laurel leaves. Two foliar pathogens cause particular concern. The first is an “anthracnose” disease of bay laurel caused by a species of Kabatiella. Although known to be present for ~80 years, this organism did not seem to cause problems until 2023. In multiple locations around the San Francisco Bay area, it has caused extensive browning defoliation of bay laurel crowns. Whether the trees will die is uncertain.
The second focus is on a recently named species, Calonectria californiensis. This organism produces P. ramorum-like similar symptoms on a wide variety of native plants, including bay laurel, tanoak, salal, mock-orange, Oregon-grape, and rhododendron. On most of these plants this fungus causes black spots that can grow to kill entire leaves, but apparently C. californiensis is not a pathogen of woody plant parts. Initial symptoms of infection on bay laurel appear identical to those caused by the SOD pathogen (Phytopthora ramorum). C. californiensis does not appear (yet) to lead to lasting debilitating disease or tree mortality.
Nurseries and Managed Landscapes
In administering APHIS’ cooperative program aimed at minimizing spread of P. ramorum via interstate trade in plants, California’s Department of Agriculture (CDFA) relies – at least in part – on funds from USDA. CDFA received $1,308,771 from APHIS in 2023. More than 300 establishments in California are regulated under the program. They submitted ~ 7,400 P. ramorum regulatory samples to the CDFA in 2023. Seventy-eight of the samples were positive (COMTF newsletter February 2024).
At the end of 2023, seven California nurseries that had tested positive for the presence of P. ramorum were operating under the APHIS regulation governing positive nurseries. This was an increase over previous years; zero in 2022, three in 2021 (COMTF newsletter February 2024 Table 4). During 2024 five nurseries were confirmed as positive. Three of these had tested positive in previous years. Two retail nurseries were newly positive; one of these was apparently infected when it brought in plants from another nursery (COMTF newsletter August 2024). I wonder whether the very wet winters California has experienced lately have enhanced the pathogen’s ability to grow – and be detected.
In Oregon, in 2023 the Department of Agriculture regulated five interstate shippers under federal compliance agreements and a sixth intrastate shipper regulated under state requirements (COMTF newsletter February 2024). Spring compliance surveys tested 1,228 foliar samples; ten were positive. After this nursery incinerated all nearby plants, none of the 1,664 foliar samples tested in the fall was positive.
In 2023, the Washington State Department of Agriculture processed more than 300 plant, soil, and water samples; all were negative. Washington also inspected five of the nine nurseries that had ‘opted-out’ of the Federal program so they can no longer ship interstate. Host material appeared free of symptoms so no samples were collected (COMTF newsletter February 2024).
Washington nurseries and regulators frequently encounter the problem of infected plants being shipped into the state from outside. (P. ramorum has been found in 33 Washington nurseries since 2003.) During 2023, the Washington State Department of Agriculture conducted three trace-forward investigations. Fortunately no infestations were detected (COMTF newsletter February 2024). In March 2024, Washington faced another trace-forward involving plants sold to homeowners (COMTF newsletter May 2024). Thirteen tissue samples and two soil samples all tested negative (COMTF newsletter August 2024)
Finally, Washington conducted stream baiting. In 2023, none of the 66 samples was positive (COMTF newsletter February 2024)
Infested Plants
Most of the plant species on which P. ramorum was detected during these years are the usual ones: Rhododendron, Viburnum, Pieris, Arbutus, Prunus, Camellia, Loropetalum. I think the several Cornus species might be somewhat unusual. Disease was confirmed on a new Cornus species, C. capitata (evergreen dogwood). One taxon — Arbutus x ‘Marina’ — is not yet listed by APHIS as a host because Koch’s postulates have not been completed (COMTF newsletters for February 2024 and August 2024).
Research (summarized in the February 2024 newsletter)
Two studies found evidence of seasonal and weather factors influenced establishment of P. ramorum. One study found a clear seasonal pattern of pathogen incidence in the western US, plus a link to the El Niño-Southern Oscillation (ENSO) (Xuechung et al. 2024. The second study looked at a Japanese larch plantation in Scotland (Dun et al. 2024).
In both Scotland (above) and France (Beltran et al. 2024 2024), scientists demonstrated that prompt action helps to suppress P. ramorum establishment.
APHIS Updates its Regulations
In March 2024, APHIS revised the P. ramorum “Domestic Regulatory Program Manual.” The agency said it updated figures and definitions, clarified operational steps, and revised the Retail Nursery Dealer Protocol (COMTF newsletter for May 2024).
Funding
In Fiscal Year 2024, under the Plant Protection Act Section 7721 program, APHIS funded $1 million worth of projects focused on P. ramorum and related species. This was out of a total $62 million in funds dispersed for pest survey, research, mitigation, and outreach programs. This money funded nursery surveys in 11 states. Also, it paid for a project to evaluate the threat of the NA2 & EU2 lineages to nurseries and forests (COMTF newsletter May 2024).
SOURCES
Beltran, A.; Laubray, S.; Ioos, R.; Husson, C.; Marçais, B. 2024. Low persistence of Phytophthora ramorum in western France after implementation of eradication measures. Annals of Forest Science. 81: 7. https://doi.org/10.1186/s13595-024-01222-1
Dun, H.F.; MacKay, J.J. & Green, S. 2024. Expansion of natural infection of Japanese larch by Phytophthora ramorum shows trends associated with seasonality & climate. Plant Pathology. 73(2): 419-430).
González, M.P.; Mizubuti, E.S.G.; Gonzalez, G.; Sanfuentes, E. 2024. Uncovering the hidden hosts: Identifying inoculum reservoirs for Phytophthora pseudosyringae in Nothofagus forests in Chile. Plant Pathology. 73(4): 937-947. https://doi.org/10.1111/ppa.13855. (Summarized in COMTF newsletter February 2024.)
Kozanitas, M.; Knaus, B.J.; Tabima, J.F.; Grünwald, N.J.; Garbelotto, M. 2024. Climatic variability, spatial heterogeneity & the presence of multiple hosts drive the population structure of the pathogen P ram & the epidemiology of Sudden Oak Death. Ecogeography. https://doi.org/10.1111/ecog.07012. (Summarized in COMTF newsletter May 2024.)
Mullet, M.S.; Harris, A.R.; Scanu, B. [and others]. 2024. Phylogeography, origin & population structure of the self-fertile emerging plant pathogen Phytophthora pseudosyringae. Molecular Plant Pathology. https://doi.org/10.1111/mpp.13450. (Summarized in COMTF newsletter for May 2024.)
Xuechung, K.; Wei, C.; Siliang, L.; Tiejun, W.; Le, Y. & Singh, R. 2024. Spatiotemporal distribution of sudden oak death in the US & Europe. Agricultural & Forest Meteorology. 346: 109891)
beech leaf disease symptoms; photo by Matthew Borden via Flickr
Beech leaf disease (BLD) came to attention in 2012 near Cleveland. It has since spread to the Atlantic – Maine to New Jersey and northern Delaware; south into Virginia; north in Ontario; and west to eastern Michigan.
Scientists have scrambled to understand the disease – how it hijacks the tree’s metabolism; & here its impacts on seedlings, saplings, and mature trees; how it spreads, locations at greatest risk.
Now Bartlett Tree Research Laboratories – the research arm of Bartlett Tree Experts – has announced development of Integrated Pest Management (IPM) strategies to treat individual trees – sadly not yet beech in the forest. The project is led by Dr. Andrew Loyd and Dr. Matthew Borden.
Seeing the disease’s impacts on a tree species with aesthetic and ecological values not easily replaced, and its rapid spread, scientists at Bartlett Tree Research Laboratories began testing fungicides and nematicides registered under the Federal Insecticide, Fungicide and Rodenticide Act (FIFRA) by the U.S. Environmental Protection Agency (EPA) to see whether they might be effective against the causal nematode Litylenchus crenatae ssp mccannii.
As Drs. Loyd and Borden note, managing BLD presents numerous challenges:
1. The disease was discovered recently, so there were many unknowns, including how it spreads and the causal organism’s novel life cycle.
2. The damage occurs in leaf buds during winter dormancy. There has been little previous research on such a system. It is difficult for chemicals to reach the tissues.
3. Mature trees are large, so reaching the vulnerable leaves in the canopy is difficult.
4. Treatment efficacy cannot be evaluated until nearly a year after application.
5. Few chemicals are registered for use against foliar nematodes or for trees in forest, nursery, or landscape settings.
6. Obtaining funding is difficult because protecting beech is a low priority among many of the usual sources.
Fortunately, the leadership at Bartlett – the company’s research department, the New England Division leadership, and especially Robert A. Bartlett, Jr. (head of the family-owned company) – saw the importance of protecting beech and have supported this research. The USDA Forest Service has also funded some of studies exploring soil drenches. Cameron McIntire reports that these studies do not yet have results.
Furthermore, Bartlett has chosen to make the science easily available to all interested parties. Three posters explaining experiments to date are available at ResearchGate. They have also published a study on the early tests of Fuopyram as a foliar spray. It is open-access. Additional publications presenting data on experiments with both spray (Fluopyram) and injection (Thiabendazole/Arbotect) are in preparation.
I summarize briefly here their findings as of August 2024.
In all the trials, the scientists judged efficacy of treatments by counting the number of viable nematodes in leaves, viable nematodes in overwintering buds, and BLD symptom severity at appropriate times before and after treatment (spray or injection).
Tests of foliar sprays on small to medium sized trees
The first tests of foliar applications that resulted in BLD suppression were carried out in Ohio starting in 2021, then expanded to other field sites in Ohio and several states in New England in 2022 and 2023 seasons. In early trials, trees were sprayed four times starting in mid to late July at 21-day intervals. The scientists say that recent trials focus on application timing and rate. They hope that optimizing these factors will help generate new recommendations that are more sustainable while maintaining efficacy.
At the annual meeting of the American Phytopathological Society in July 2023, Bartlett announced that Fluopyram is an effective management tool to combat BLD – on smaller trees that can be treated using foliar application. There are several EPA-registered products, though only one, Broadform, has been so far been granted a section 2(EE) recommendation “For Control of Beech Leaf Disease on Beech Trees.”
Treatments are less effective in situations where the inoculum load is very high (for example, a very dense stand of infected trees); or where mature, untreated canopies hang over treated understory beech.
They suggest that managers focus treatments on high-value specimen beech, collection preservation, and potentially uncrowded mixed natural stands.
Treatments should be made by certified pesticide applicators who are familiar with the disease and treatment specifications. For the injection treatment, technical training and specialized equipment is needed. Bartlett arborists and plant health care specialists in locations affected by BLD have all been trained to perform the treatments, and some other arborists are doing BLD treatments as well using the same products.
Soil drench
Matt Borden said that they tested drenches with three different chemicals. The approach did not reduce nemtatode numbers sufficiently. However, as noted above, the Forest Service is funding additional tests exploring possible combinations of drenches with other actions, such as thinning. Discovering management options across a range of application methods (e.g., foliar, injection, drench) and modes of action is vital for a disease that covers such a broad range of locations and tree sizes and forms.
a macroinjection demonstration; photo by Matthew Borden via Flickr
Injections
Scientists injected Thiabendazole (TBZ) into beech on private land in three locations in Ohio and New Jersey. They tested two application rates and three application timings. They have two years of follow-up data for one site, one year for the others.
Key findings:
nematode numbers in buds in late winter consistently reflected foliar symptoms when the leaves opened.
Injections made before mid-July provided the greatest reduction in nemtatode numbers and best canopy improvement. Trees injected late in the season (30 August), after the nematode has begun dispersing from leaves to buds, exhibited some BLD symptoms the next year, but suffered less canopy dieback than controls.
Margery Daughtrey of Cornell said during a discussion of these finding that the trees’ persistence suggests that trees can tolerate some level of symptoms. Among other things, it might be possible to treat the trees less frequently than annually.
TBZ appears to provide at least two seasons of nematode suppression
Bartlett continues to monitor these trees to see how long the injected chemical suppresses nematode numbers and how long the tree remains healthy. They are also establishing new field sites to further optimize rate and timing.
TBZ – in a product called Arbotect 20-S – has been used to manage Dutch elm disease and sycamore anthracnose since the 1970s. However, it is also a well-known nematicide, previously used as an anti-parasitic drug in human and veterinary medicine. Once injected, TBZ protects the tree for more than one season. The injection technology (MACRO-Injection) has also been used for decades. It infuses the chemical directly into the tree’s vascular system; it does not rely on root uptake. Matt says injection does require take technical skill and the right equipment. To minimize the risk of the wound cracking and weeping, the injection should be done low on the side of the root flare, not on top.
While Arbotect 20-S has been registered for use in 48 states for many years, new labeling is required for its use in beech trees and against BLD. Special Local Needs labels, 24(C)s, have been granted by eight states – Connecticut, Massachusetts, Maine, New Jersey, New York, Pennsylvania, and Virginia. Registration in a ninth – Maryland – is in progress and Bartlett scientists are prepared to apply for several more. The problem is that only a limited number of these “special needs” labels may be issued, and BLD has expanded so far, and so rapidly, that it is already infesting beech in more states than may be covered by 24(C)s. Furthermore, 24(C) labels expire if not renewed. Most current 24(C)s will be active through 2028 – not ideal for a disease that will likely be with us long into the future. The product manufacturer (Syngenta) and distributor (Rainbow Ecoscience) are drafting a change to the main Arbotect 20-S label to add beech and the new nematode pest, but warn that EPA review and approval of amendments can take a very long time. Until then, we must resort to limited special local needs labels, and some states will miss out.
contrasting canopy transparency in beech treated with TBZ v. untreated controls; photo by Matthew Borden
One of the key scientists who developed these treatments for Dutch elm disease, R. Jay Stipes, professor emeritus at Virginia Tech, is quoted by Bartlett rejoicing that his work might help protect another tree species.
Matt believes the treatments will be effective if applied every 2-3 years. This approach would also spread out the cost – which will depend on the arborist but Dave Anderson of Rainbow Ecoscience estimated to be about $25 / inch of dbh.
It is always best to obtain an accurate diagnosis before treatment. The next step is talking through your options with a certified arborist or tree disease specialist. The “good” thing about BLD is that it is a progressive disease and will not kill a tree in a single year. Therefore, waiting until you know the disease is present or active locally is generally recommended.
Tree injection is better than foliar application where the latter is impractical (e.g., the tree is tall) or to reduce runoff, particularly near streams. Bartlett recommends treating any beech larger than 10 cm dbh by injection; smaller trees by foliar spray.
Treated trees should be sound, without serious decay, girdling roots, or other conditions that curtail uptake. Based on research results to date, they recommend treating the tree before mid-July. Bartlett is testing the results of injecting the shortly after full leaf expansion – early to mid-June. Bartlett scientists are testing several application rates to determine how long a single injection will suppress BLD. So far they have had good results from both low and moderate label rates (0.4-1.6 fl oz/inch DBH).
All the technical information re: research into treatments and recommendations for applying either the foliar or injection treatments has been provided by Dr. Matthew Borden of Bartlett Tree Research Laboratories. He can be reached at
Dr. Borden says he is immensely grateful for the support that allows him and Dr. Loyd to travel widely to establish the BLD research sites and spend weeks collecting data each year with their team. Company founder Francis A. Bartlett established the Bartlett Tree Research Laboratories as a separate entity within the company, where capital is reinvested directly into stable, long-term support of scientific tree research and preservation. The model is well-suited to provide the flexibility and freedom needed to rapidly respond to emerging invasive species issues.
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at https://treeimprovement.tennessee.edu/
The National Plant Board’s members are the lead plant health officials of the states and territories. Many federal officials also attend – from APHIS and DHS Bureau of Customs and Border Protection. Representatives of other North American phytosanitary entities participate – i.e., Canada, Mexico, and the North American Plant Protection Organization (NAPPO). Some stakeholder groups participate, especially the nursery industry. I have attended these meetings for over a decade because they provide an overview of pest issues and programs plus an unparalleled opportunity to network. The Nature Conservancy’s Leigh Greenwood also attends. We are the only representatives of the species conservation community to attend – others are missing great opportunities.
Here, I’ve listed 10 items that are among the most important the group discussed.
1) The funding situation for APHIS is worse than I realized
APHIS Administrator Mike Watson and Deputy Administrator (for plants) Mark Davidson both spoke about the need to cut programs to stay within the limits set by congressional appropriations. Funding for APHIS, as a whole, was cut only 1% for the current year (Fiscal Year 2024), cost-of-living salary increases mean less money for programs. (I believe Dr. Watson said $41 million less for FY24). If FY25 funding is the same, Congressionally mandated additional payraises will mean an another $20 million decrease in program funding.
Dr. Davidson said that the plant programs (Plant Protection and Quarantine) had been cut by 5% in FY24. However, Congress did not finalize the funding levels until about half-way through the fiscal year – so staying within the limits required even more severe cuts to programs in the remainder of FY24. To stay within these limits, APHIS cut several programs, among them a $3.6 million cut from the “tree and wood pest” program. This meant loss of funds to manage the polyphagous and Kuroshio shot hole borers, smaller cuts for programs managing Asian longhorned beetle and emerald ash borer, and perhaps the Asian flighted spongy moth. They anticipate additional cuts in these programs in FY25. The one bright light is the Section 7721 Plant Pest & Disease Management & Disaster Prevention Program. It provides steady funding for a range of plant health programs. The House version of the still-pending Farm Bill calls for increasing funding for this program by $15 million each year.
Nearly 100% ash trees in Oregon wetland — exposed to spreading EAB. Photo by Wyatt Williams, Oregon Department of Forestry
Remember this when I ask you to lobby for appropriations! If we don’t advocate for funding the programs dealing with “our” pests, they will shrink.
Watkins and Davidson also worry that whoever is the next secretary of USDA might not support the agency when it seeks to withdraw funds to cover emergencies from the Commodity Credit Corporation – as Secretary Vilsack has.
APHIS and the DHS Customs and Border Protection (CBP) both praised a recent regulatory action that increases user fees for importers having goods cleared at ports. Kevin Harriger, CPB official in charge of agriculture programs, said the new funds would allow CBP to hire 700 new agricultural inspectors (currently there are 2,800 agricultural officials). That sounds great, but … when trade and passenger volumes crashed early in the COVID pandemic, things looked dicey for a while. Plus – as I have argued repeatedly – real protection against pest introductions will come from stronger policies, not ramped-up inspections.
Pathologist Bruce Moltzan reported on the USFS Forest Health Protection program. He pointed out that the USFS has a very limited toolbox. In this fiscal year, the program has about $48 million, after salaries, to support its activities. Projects targetting insects receive 70% of the funding; those targetting pathogens 15%.
2) Invasive hornets
Washington State has not found any new nests of the Northern (formerly Asian) giant hornet (Vespa mandarinia). Miraculous!
However, Georgia detected another species, the yellow-legged hornet (Vespa velutina), near Savannah in August 2023. The Georgia Department of Agriculture, APHIS, and the University of Georgia are working to find nests – which are located at the top of tall pine trees in residential areas. Five nests were found in 2023; another four so far in 2024. Georgia hopes to place traps 100 miles out from each detection site. Like the northern hornet, V. velutina preys on honey bees. It was probably transported by ship or with its cargo.
A third species, V. tropica, has been introduced on Guam.
3) Better Federal-State Cooperation — Sometimes
APHIS and the state phytosanitary officials have set up structures – e.g., Strategic Alliance/Strategic Initiative, or SASI – to work together more closely. CBP joins the coordinating meetings. One program described at the meeting is the effort to contain spread of the box tree moth (Cydalima perspectalis). This effort came out of discussions at last year’s Plant Board meeting, with follow-up gatherings of APHIS, the states, and the nursery industry. The moth is known to be present in New York, Massachusetts, Michigan, Ohio, and now Delaware – plus several Canadian provinces.
A second project concerns how much data to share about state detections of pests – which are recorded in the National Plant diagnostic Network database. These data have accrued over 20 years … and are sought by both other states and academic researchers. States are often reluctant to allow public review of detection data because they fear it will cause other states or private parties to avoid buying plants or other goods from the infested area. While the project team has not yet decided how to deal with these conflicts, they said they were more inclined to share “nonconsequential data” – meaning data on a pest everyone already knows is present, not a pest under regulation or a new detection. In other words, “consequential” seems to pertain to industry profits, not damage to agricultural or natural resources.
EAB-killed ash along Mattawoman Creek, Maryland. Photo by Leslie A. Brice
Craig Kellogg, APHIS’ chief plant health representative in Michigan, reviewed 20 years of dealing with the emerald ash borer (EAB). He is optimistic about the impact of the biocontrol agents that have now been released in 32 states and four provinces. The larval parasitoids are dispersing and EAB densities are coming down. He conceded that over-story and mature ash are still dying, but says ash in long-infested areas are regenerating well. Scientists agree (see Wilson et al. 2024; full citation at end of the blog). Woodpeckers are still the most effective biocontrol agent of EAB for over-story ash, especially in locations where introduced parasitoids are not established. So far, the growing numbers of biocontrol agents are still parasitizing too few EAB larvae to prevent decline of over-story ash trees.
APHIS reported on recent detections of flighted spongy moth from Asia on ships coming to U.S. ports. The program covers four closely related species of Lymantria. All have much broader host ranges than Lymantria dispar, plus the females are capable of sustained flight, so they spread more rapidly.
The principal strategy to prevent their introduction is to require ships that call at ports along the Pacific coast in Russia, China, Japan, and North and South Korea to ensure that the ships’ superstructures and cargo are clean. Before arriving at U.S. ports, the ship’s captain must inform CBP where it has called over the last 24 months. Then, CBP conducts an inspection. If CBP inspectors find a small number of egg masses, they remove the eggs and spray pesticide. If the inspectors detect a large number of egg masses, the ship is ordered to leave port, clean itself, and undergo re-inspection before it can return.
Four countries in the Americas – the U.S., Canada, Chile, and Argentina – and also New Zealand have very similar programs.
Detections follow natural changes in population levels in the exporting regions. APHIS’ program leader, Ingrid Asmundsson, reported on an unfortunate coincidence in 2014. A huge moth population outbreak occurred simultaneously with very low fuel prices in Russia. The latter attracted many ships to call there. An even bigger population surge occurred in 2019. Asmundsson expects another high-moth period this year.
flighted spongy moths infesting a ship superstructure
APHIS is working on putting this program on a regulatory foundation; this would allow the agency to be more specific in its requirements and to impose penalties (other than expulsions from ports). I’ll let you know when the proposed rule is published for comment.
6) Regional Reports: Old Pests, New Pests
Representatives of the regional plant boards informed us of their “new pest” or other concerns.
Gary Fish, president of the Eastern Plant Board, mentioned
awareness that laurel wilt is moving into Virginia and maybe farther north.
elm zig-zag sawfly; photo by Gyorgy Csoka via Bugwood
There was a more general discussion of beech leaf disease. What can be done, given that the disease is so widespread that no one is regulating movement of beech. Gary Fish advised outreach and efforts to reach agreement on management approaches. Chris Benemann, of Oregon, suggested informing other states so that they can decide whether to take regulatory action. A representative of CBP urged engaging stakeholders by asking for their help.
Chris Benemann, President of the Western Plant Board, expressed concern about APHIS’ reduced funding for spongy moth detection and control efforts. She also worries about the recently detected population of Phytophthora austrocedriiin an Oregon nursery. The western states are also focused on several longstanding pest problems – grasshoppers, Japanese beetle; and a new beetle from Australia that is attacking almonds, pistachios, and walnuts.
tree infested by hemlock woolly adelgid; photo by F.T. Campbell
Megan Abraham of Indiana reported that members of the Central Plant Board are concerned about
funding decrease for APHIS and USFS efforts to slow the spread of spongy moth and manage Asian longhorned beetle and emerald ash borer;
She noted that nursery stock is increasingly coming from more distant – and cheaper – producers, raising the risk of new pests being introduced.
Finally, Abraham expressed concern about decreased funding at the same time as the need is growing – and asked with whom states should collaborate in order to reverse this trend.
Kenny Naylor of Oklahoma, Vice President of the Southern Plant Board, concurred that funding levels are a major concern. He mentioned specifically the spongy moth Slow the Spread program and eradication of the Asian longhorned beetle outbreak in South Carolina. Another concern is the Georgia hornet outbreak.
7) Phasing Out Post-Entry Quarantine
APHIS and the NPB have agreed to phase out the post-entry quarantine (PEQ) program. A program review revealed several problems, some of which astound me. When examining plants in quarantine the scientists still relying on visual inspection! And they are looking for pests identified 45 years ago (1980)! While I think PEQ programs can be valuable in preventing introduction of disease agents, as implemented in recent decades it does not. Twenty years ago, citrus longhorned beetles escaped from a “quarantine” area in a commercial nursery in Washington state. These Cerambycids are more than an inch long!
citrus longhorned beetle; photo by Art Wagner, USDA via Bugwood
Part of this phase-out is to transfer plant species harboring pests of concern to the Not Authorized for Importation Pending Pest Risk Assessment (NAPPRA). While the APHIS speaker said that NAPPRA allows the agency to act quickly when it detects evidence of pest risk, I have found lengthy delays. The third round of proposals was published in December 2019! The fourth round of species proposed for NAPPRA listing should be published soon; a fifth round is now in draft inside the agency.
8) Christmas Greens – Spreading Pests
Officials from Oregon, Maine, and Illinois described their concerns about pests being spread by shipments of various forest or plant products, especially Christmas greens. Mentioned were spongy moths, link hemlock woolly adelgid, link elongate hemlock scale, balsam woolly adelgid, link and box wood moth. Part of the challenge is that the vectoring items are often sold by unregulated outlets – multiple stores, Christmas tree lots – and through on-line or catalog outlets. There are also extreme demands on the regulatory enforcement staff during the brief holiday sales season. Several states are unsure whether they have authority over decorative products – although others pointed out that they are regulating the pest, regardless of the object for sale or type of store.
9) Pests in Firewood
Leigh Greenwood of The Nature Conservancy noted that the state agencies that issue firewood regulations – often the plant protection organization (state department of agriculture) — do a good job alerting the public about the risks and rules. However, the public looks first to state parks agencies for information about camping – and those agencies have less robust educational efforts. It is important to put the message where the public can find it when they don’t know it exists – before they include firewood from home in their camping gear.
10) Projects of the North American Plant Protection Organization
The North American Plant Protection Organization (NAPPO) is working on several projects of interest to those of us concerned about tree-killing pests. One project is evaluating risks associated with wood products, especially how well one international regulation, ISPM#15 is working for dunnage. Another projects is testing the efficacy of the heat treatment specified by ISPM#15 (50o C for 30 minutes). A third project — almost completed – is evaluating fumigants that can be alternatives to methyl bromide.
In conclusion, each annual meeting of the National Plant Board is packed with new information, updates on current pests, and comments on by the state agencies who suggest new approached to APHIS and hold the agency to account. It is well worth attending. Information about upcoming meetings of both the national and four regional plant boards is posted on the NPB website, https://www.nationalplantboard.org/
Signatories to the APHIS-NPB strategic alliance
SOURCE
Wilson, C.J., T.R. Petrice, T.M. Poland, and D.G. McCullough. 2024. Tree species richness and ash density have variable effects on emerald ash borer biological control by woodpeckers & parasitoid wasps in post-invasion white ash stands. Environmental Entomology.
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at https://treeimprovement.tennessee.edu/
Flighted spongy moths infesting a ship’s superstructure
USDA’s Animal and Plant Health Inspection Service (APHIS) has issued its annual report for Fiscal Year 2023. The report is part of an enhanced outreach effort that I believe is an effort to persuade the Congress to provide additional funds. However, as I describe below, at this summer’s annual meeting of the National Plant Board, link APHIS’ leadership stated that funding shortfalls are forcing them to curtail many programs. These include ones important to those of us concerned about threats to North American trees. I applaud this action and hope it succeeds!
The report contains some good news but I consider the overall approach depressing. Tree-killing pests continue to receive little attention. The report also emphasizes APHIS’ efforts to facilitate export of agricultural products – an understandable stance given American politics.
The opening summarizes the agency’s activities includes:
Examples of programs targetting pests abroad, before they can reach the U.S. All are fresh fruits and vegetables;
APHIS or staff at U.S. borders:
Approved (cleared) 27,235 shipmentscontaining over 1.87 billion plant units (e.g., a single plant or cutting, or vial of tissue culture plantlets) and 670,811 kilograms of seeds. They intercepted 2,176 quarantine pests. (APHIS carry out these inspections at Plant Inspection Stations – separate from the port environment where DHS Customs and Border Protection (CBP) staff inspects other cargo.)
Identified approximately 92,000 pestsfound during CBP inspections of cargo, mail, and express carrier shipments and took quick action to prevent those of concern from entering the U.S.
Facilitated entry of regulated agricultural cargo by monitoring more than 62,000 treatments of various kinds, that is, fumigations, cold or heat treatments, and irradiation.
Examples of APHIS’ efforts to slow pests’ spread within the country cited plant pest surveys — with coordinated responses — for approximately 45 pests. Also APHIS described funding to help citrus growers combat citrus greening.
The report has separate subreports on the following programs: risk analysis, pest detection, “specialty crop” pests, and tree and wood pests. The last two contain information specific to our interests.
Tree and Wood Pests
This program protects forests, private working lands, and natural resources. It targets specific pests: the Asian longhorned beetle, emerald ash borer, spongy moth, and most recently the invasive shot hole borers. The report notes that numerous native, widespread hardwood tree species are vulnerable to these pests. APHIS asserts an economic justification for the program: conserving forests enhances rural communities’ economic vitality, supports forest-related industries, and maintains the ecosystem services provided by urban trees.
Unfortunately, at this summer’s annual meeting of the National Plant Board APHIS leadership said funding shortfalls forced them to pull back on all these programs.
ALB eradication aims to protect the 30% of U.S. trees that are ALB hosts. These trees support multi-billion-dollar maple syrup, timber, tree nursery, trade, and tourism industries. After reviewing the history of ALB detections, starting in Brooklyn in August 1996, the report describes APHIS’ eradication strategy as comprising surveys, regulatory inspections and quarantine restrictions, removal of infested and high-risk trees, and chemical treatment applications. In FY 2023, the program surveyed more than 763,000 trees across the four regulated areas: New York, Massachusetts, Ohio, and South Carolina. Each program is summarized.
Good news at two locations. On Long Island: only 11 new infested trees were found after a survey of 43,480 trees. In Worcester County, Massachusetts, no new infested trees were found after surveying nearly 360,000 trees. However, in Tate Township, Ohio, surveys detected 163 new infested trees. And in
South Carolina, the program is at an earlier stage — surveying a portion of the quarantine area. The program surveyed nearly 140,000 trees and removed 1,700 in FY 2023.
At the National Plant Board Meeting, Deputy Administrator Mark Davidson explained that the FY2024 appropriation cut $3.6 million from the “tree and wood pests” account. This required the agency to reduce funding for the ALB eradication program.
The report summarizes the spread of EAB since its first detection in 2002 in Michigan to 37 states and the District of Columbia (APHIS does not mention EAB’s presence in five Canadian provinces.)
Saying that EAB has spread beyond what a regulatory program can control, the report notes that APHIS ended the regulatory program in FY 2021. In FY 2023 the agency continued the transition to a program relying primarily on biocontrol. In FY2023, APHIS provided parasitoids to 155 release sites – three in Canada, the rest in 122 counties in 25 states. APHIS and cooperators continue to assess their impacts on EAB populations and tree health at release sites and nearby areas. Field evaluations indicate the EAB parasitoid wasps and other EAB natural enemies (woodpeckers) are protecting regenerating sapling ash from EAB.
At the National Plant Board Meeting, Deputy Administrator Mark Davidson explained that the FY2024 appropriation cut $3.6 million from the “tree and wood pests” account. This required the agency to reduce funding for the EAB containment program – probably the biocontrol component.
Spongy moths (the species formerly called European gypsy moths) are established in all or parts of 20 eastern and midwestern states, plus the District of Columbia. APHIS and state cooperators regulate activities in the quarantine area to prevent the moths’ human-assisted spread to non-quarantine (non-infested) areas – primarily West coast states. To address the moths’ natural spread, APHIS PPQ monitors the 1,200-mile-long border of the quarantine area and adds newly infested areas to the regulated area. The USDA Forest Service – APHIS – Slow-the-Spread Foundation program has greatly reduced the moth’s rate of spread and has eradicated isolated populations.
Another component of the program aims to prevent introduction of members of the flighted spongy moth complex link from Asia. The Asian species have broader host ranges and the females can fly, so they could spread faster. A multi-nation cooperative program is designed to prevent the moths’ hitchhike on vessels coming from Asia. link APHIS supports this program through negotiations and support of CBP’s offshore vessel inspection, certification, and cleaning requirements. Canada participates in the same program.
In FY 2023, APHIS and state cooperators continued efforts to delimit possibly introductions in Washington State (no additional moths detected); and California and Oregon (initial detections in FY 2020).
At the National Plant Board Meeting, Deputy Administrator Mark Davidson explained that the FY2024 appropriation cut $3.6 million from the “tree and wood pests” account. This required the agency to reduce funding for the flighted spongy moth program.
California sycamore infested by polyphagous shot hole borer; photo by Beatriz Nobua-Behrmann UC Cooperative Extension
The report notes that various non-native shot hole borers have been detected in several states. Their hosts include trees in forests and urban landscapes, tea plantations, and avocado orchards. The program’s focus was apparently on the polyphagous and Kuroshio shot hole borers devastating riparian habitats in southern California and urban areas in other parts of California. At California’s request, APHIS and the USDA Forest Service helped establish a working group, led by USFS, with the goal of strategically addressing both shot hole borers in California. In FY 2023, APHIS’ helped with foreign explorations for possible biocontrol agents, as well as host specificity testing.
APHIS leadership told the National Plant Board in July 2024 that it had dropped this entire program due to funding shortfalls.
Specialty Crop Pests
While much of this report concerns pests of agricultural crops (e.g., grapes, citrus, potatoes), it also summarized efforts re: Phytophthora ramorum (sudden oak death) and spotted lanternfly. APHIS says its efforts protected nursery stock production worth approximately $1.3 billion as of 2019, and tree fruit production worth approximately $1.7 billion in 2021.
map showing areas of the Eastern United States at risk to P. ramorum – developed by Gilligan of Cambridge University
The report states that APHIS seeks to limit P. ramorum’s spread from affected nurseries. The agency does this via regulatory strategies. During FY 2023, 16 nurseries were governed by more stringent rules under the federal program which are imposed on nurseries that have been determined in past years to harbor P. ramorum-infected plants.
In addition, Oregon officials continued surveys of an area outside its quarantine zone because of a detection the previous year. APHIS will adjust the federal quarantine depending on the state’s findings.
The APHIS report does not discuss several pertinent events that occurred in FY2023. [For more details, read the California Oak Mortality Task Force newsletters for 2023 – posted here.
First, APHIS does not mention or discuss the implications of detection of two new strains of P. ramorum — EU1 & NA2 — in west coast forests. The presence of EU1 in a new California county (Del Norte) was confirmed in Feb 2023.
Second, the report said that Oregon is trying to determine the extent of the P. ramorum infection detected outside the state’s quarantine zone. However, it does not mention that this outbreak involves the new NA2 lineage – and that NA2 was known to be present in nurseries in the region since 2005.
The report also does not clarify that three nurseries to added to the more stringent program were so treated because SOD-infected plants were found on their premises.
Nor does the report note that at least two new naturally-infected hosts of P. ramorum were identified: Western sword fern (Polystichum munitum) and Arbutus x ‘Marina’.Koch’s postulates need to be completed on the latter so it has not yet been added to APHIS’ official host list.
Spotted Lanternfly (SLF) was found in 16 states in FY 2023. APHIS’ program enjoyed funding provided through Specialty Crop Pests and from the Plant Protection Act’s Section 7721 link ($6 million from the latter).
The report notes that APHIS still does not have enough data to determine SLF’s impacts on agriculture. Thus far, vineyards have been the most adversely affected agricultural segment, mostly due to SLF acting as a stressor to vines. Also, the sticky, sugary “honeydew” produced by SLF attracts other insects and promotes sooty mold growth. These can ruin the fruit and further damage the plant.
SLF populations are strongly linked to major transportation pathways, such as railroads and interstate highways. APHIS targets treatments and, in some areas, removes SLF’s preferred host plant (tree of heaven [Ailanthus]), from transportation hubs. The aim is to reduce the risk of SLF’ spread to new areas and to eradicate isolated infestations. In FY 2023, APHIS and cooperators treated 4,637 properties covering 6,455 acres in affected areas. However, during the National Plant Board meeting both state and APHIS officials complained to me that managers of these transportation hubs raise many barriers to their access, sharply limiting the program’s chance of success.
Ailanthus altissima – drive of spotted lanternfly invasion
The program was expanded after National Environmental Policy Act-mandated environmental review. This allowed APHIS to conduct treatments in four additional states—Indiana, Massachusetts, Michigan, and Rhode Island. In addition, program cooperators identified three potential biological control organisms, one that targets the tree of heaven and two that target SLF. APHIS will continue to evaluate them and develop methods to rear them in the laboratory.
Finally, in fiscal year 2023, APHIS joined the National Association of State Departments of Agriculture and the National Plant Board to develop a national strategic plan outlining the future direction of the SLF program. With the strategic plan, PPQ aims to harmonize the approach across states to slow SLF’s spread, develop consistent outreach messaging for a nationwide audience, and more effectively use existing state and Federal resources. Continued spread of SLF despite the huge effort, rising costs of the program, and new scientific findings spurred reconsideration of the strategy.
To summarize, I hope that APHIS’ annual report will – in the future – help members of Congress and their staff understand the agency’s programs’ purpose and past successes. This increased understanding might make it easier to advocate for more funding. I am troubled, however, by the agency’s glossing over significant problems.
Posted by Faith Campbell
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For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at https://treeimprovement.tennessee.edu/
