Oregon ash in swamp in Ankey National Wildlife Refuge, Willamette Valley, Oregon; photo by Wyatt Williams, Oregon Department of Forestry
1) Funds still not released
As of the end of June, the Office of Management and Budget has not released funds to programs under the USDA Forest Service’ State, Private, and Tribal section. This includes many programs – grants, etc. – that support state and other entities’ efforts and operation of the Forest Health Protection program. Meanwhile, tree-killing insects, pathogens, and nematodes pursue their lives … killing trees in the process.
Congress has not yet acted on legislation that will determine the funding level for USFS FHP and Research programs in Fiscal Year 2026, which begins on October 1. I remind you that the Administration has proposed 0 funds for these programs. Take advantage of the Congressional delay – contact your Member of the House and Senators.
2) the Mediterranean oak beetle (MOB) Xyleborus monographus has now been detected in nine California counties, including Napa, Sonoma, Lake, Sacramento, El Dorado, Yolo, Mendocino, and as of last month, Marin. MOB is also present in Oregon – in Multnomah, Clakamass, Marion and Washington counties.
There is no treatment for infected trees. California authorities urge landowners to search for the insect and remove infected trees – and to avoid moving infested wood.
3) Oregon Department of Forestry has announced that thousand canker disease (TCD) of walnut is killing trees of the Juglans genus in the Willamette Valley. Where the insect has been found in traps, the majority of black walnut trees have since died. Black walnut (Juglans nigra) is not native in Oregon; its range east of the Great Plains. Apparently the range of northern California black walnut (Juglans hindsii) also does not extend into Oregon.
Cities in Oregon are preparing for the inevitable arrival of the emerald ash borer (EAB) (Agrilus planipennis) which was detected in Forest Grove in June 2022. The City of Salem inventoried all of its street & park trees in the last 5 years. It plans to inject a systemic pesticide into at least 550 trees this year and a similar number next year. These trees have been judged to be in good condition. Ash trees in poor condition along streets or in parks are gradually being removed and replaced.
Salem also plans to inventory Oregon ash growing in the city’s natural areas so as to understand where they will need to plant other native species. I blogged earlier about the threat EAB and MOB pose to western Oregon’s wetlands and oak savannahs.
4) Ann Hajek of Cornell and colleagues haves published a review of 20 years of research on entomopathogens that might contribute to control efforts targetting the Asian longhorned beetle (ALB) (Anoplophora glabripennis). The authors call for renewed efforts to find appropriate control agents and techniques. They conclude that various pathogens – especially fungi – can support ALB eradication efforts. They would be particularly helpful if ALB populations spread – or a new outbreak is detected. (Remember, ALB has been detected in seven locations in North America – some the result of more than one introduction; and nine locations in Europe.)
The article is open access!! See
Ann E. Hajek, A.E., E.H. Clifton, and L.F. Solter. 2025. Entomopathogens for control of Asian longhorned beetles (Coleoptera: Cerambycidae). Environmental Entomology, XX(XX), 2025, 1–10 https://doi.org/10.1093/ee/nvaf016
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 http://treeimprovement.utk.edu/FadingForests.htm
Several bioinvasion scientists have announced launch of a bi-annual Invasions Newsletter. It will be an open-access digital magazine intended to meet the growing need for effective communication across the diverse community of researchers, practitioners, & policymakers. It will offer accessible insights into current research; communication & management strategies; novel technologies; research centers, groups, journals, networks, projects & resources; emerging policy trends; & past & upcoming meetings & events.
The initial issue is available here Among the topics addressed:
invasive plant biocontrol efforts in Zimbabwe,
setting national-level invasive species priorities in Chile,
conservation actions to recover invaded endemic forests in Galápagos Islands,
protecting ground-nesting birds from introduced predators,
IUCN ISSG’s assistance in tackling invasive species in Europe.
The organizers — Ana Novoa, Susan Canavan, Katelyn T. Faulkner, Piero Genovesi, Deah Lieurance, Dan Simberloff, Hsiao-Hsuan Wang, Tsungai Zengeya, & Laura A. Meyerson – invite us to contribute to future editions. Inform your global colleagues about your efforts and findings: fieldwork, model development, designing or implementing management interventions at a local, national or continental scale, or crafting policy frameworks.
outreach
To submit a contribution, contact any of the organizers.
Let’s ensure that tree-killing critters (with or without legs) get the attention they deserve!
Northern red oak – one of the species planted in Europe & China as part of sentinel garden project; photo by F.T. Campbell
During the USDA Interagency Forum on Invasive Species, Dr. Eliana Torres Bedoya, from the Bonello lab at Ohio State, provided insights gained from a sentinel garden project operating for the last five years.
The gardens were established in six locations: two in China in the Nanjing area, one each in Italy, Sweden, Ohio, and New Hampshire. The network required collaboration among scientists in several countries, a difficult task in itself. (Jiri Hulcr of the University of Florida has also stressed the importance of mutually beneficial collaborations.)
The focus was on detecting and identifying novel fungal pathogens abroad before they ever enter a country, in an approach called ex patria sentinel plantings. Altogether, 32 tree species were planted in at least one location. For example, Chinese and European tree species were planted in the U.S. to identify potential threats to China and Europe. Conversely, North American species were planted in Europe and China to detect potential threats to the U.S. As noted, the reciprocity is crucial to establishing and maintaining a long-term relationship.
Key information gained to date:
While the scientists isolated several potential pathogens from symptomatic plants, analysis of all plants’ leaf microbiomes showed that asymptomatic plants harbored many more potential pathogens that had not been isolated.
Healthy plants tend to harbor larger and often more diverse microbial communities. This study found that asymptomatic plants supported a significantly more abundant, richer, and taxonomically diverse leaf-associated fungal community than symptomatic plants. Importantly, this pattern pertains also to the subset of taxa classified as potential plant pathogens.
Detection of the full range of fungal pathogens requires that samples must be collected both early and mid-to-late in the growing season because microbes present differ.
Core leaf microbiomes were associated with specific tree species, no matter where they were planted. However, the constituents of the core microbiome were outnumbered by other organisms driven primarily by the location of the planting. This had been expected.
Other contributing factors – in declining order – were geographic location, tree species, season, and host health status. In other words, the phylogenetic relationship.
The drivers of fungal community composition interact in complex ways. For instance, the effect of the plant’s health on pathogenic fungal communities might depend on the host species. This relationship can be further modulated by seasonal variation and geographic context.
European & Asian trees planted in Ohio as part of the sentinel garden program; photo by P (E) Bonello
Implications:
Sentinel gardens can facilitate identification of novel host-pathogen interactions in symptomatic and asymptomatic plants, so they should be adopted / supported by governmental and regional phytosanitary agencies.
The findings demonstrate the need to expand surveillance beyond symptomatic plants – at both sentinel gardens and plant health border inspection stations. Phytosanitary agencies should employ both full microbial community molecular characterization to detect threats in asymptomatic plants and traditional symptom-based approaches. These modern approaches are described in Munck and Bonello 2018 (full reference at end of the blog).
Enrico Bonello (pers. comm.) thinks it is likely that similar context-dependent interactions among host and fungus species, season, and geography also drive disease infection and virulence.
Eliana Torres Bedoya (pers. comm.) clarifies that the leaf microbiome is the community of microorganisms living on and within tree leaves. These microbes can contribute to protecting trees against pathogens, enhance tolerance to environmental stressors such as drought or pollution, and influence how trees interact with their surroundings. Because the composition of the leaf microbiome responds to changes in climate, location, and tree species, it also serves as a valuable indicator of forest health and environmental change.
There are several approaches to studying microbial communities in leaves. One is the traditional, culture-based method, which relies on isolating and cultivating microorganisms on nutrient media. While this approach is effective for recovering fast-growing and easily culturable taxa, it has a major limitation: the vast majority of environmental microbes are not readily culturable under standard laboratory conditions. As a result, full understanding requires use of culture-independent methods. One technique widely used is metabarcoding. This technique involves extracting total DNA from leaf tissue and amplifying a phylogenetically informative genetic marker specific to the microbial group of interest (for example, the internal transcribed spacer (ITS) region for fungi or the 16S rRNA gene for bacteria). The amplified regions are then sequenced using high-throughput sequencing platforms. After a series of processing steps, the sequences are clustered into Amplicon Sequence Variants (ASVs), which represent unique DNA sequences that can be used as proxies for microbial taxa present in the sample. Torres Bedoya and Bonello used the ASVs for comparative analysis.
Tilia cordata (linden) via Picaryl (seed wings make a great tea!)
In her presentation, Torres Bedoya provided examples of the complexities arising when trying to detect fungi associated with trees. Eleven potentially pathogenic fungal genera were found to be more abundant in asymptomatic Northern red oak (Quercus rubra) trees (a North American species) planted in both Europe and China. Five ASVs were more abundant in asymptomatic Fraxinus mandshurica trees (an Asian species) planted in Sweden. In this case, the season when the leaves were sampled explained a higher proportion of the variance in the community composition than did the health status of the host. Molecular methods detected 10 genera not revealed through isolation from little-leaf linden species (Tilia cordata) trees (a European species) planted in China and the US.
This “proof of concept” study considered only fungi associated with leaves. As shown above, learning the true plant health risk associated with any tree taxon’s leaves is already complicated and resource-demanding. To fully exploit the power of the ex patria sentinel plantings approach, phytosanitary officials must provide additional resources (land, people, equipment, money) to enable screening of all plant parts, above and below-ground, and all potentially pathogenic taxa, including nematodes, phytoplasmas, and viruses. These systems must be maintained over years.
Reference
Munck, I.A., Bonello, P. 2018. Modern approaches for early detection of forest pathogens are sorely needed in the United States. Forest Pathology 48 (5). doi:10.1111/efp.1445
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/
ash tree killed by emerald ash borer; photo courtesy of (then) Mayor of Ann Arbor John Hieftje
Since the 1990s, scientists have been trying to the determine costs imposed by invasive species. They hope that measuring monetary costs will motivate political decision-makers to take more assertive actions to counter this ecological treat. As Daigne et al. (2021) point out, too few countries are implementing effective control and mitigation strategies. They say this inaction stems, largely, from undervaluing bioinvasions’ impacts by the general public, stakeholders and decision-makers.
A major step in this effort was creation of the InvaCost database. The goal was to provide a reliable, comprehensive, standardized and easily updatable synthesis of bioinvasions’ monetary costs worldwide.
Several publications based on this database appeared. I have blogged about studies published in 2021 or 2022: a) the costs of bioinvasions generally (Cuthbert et al. 2022); b) the costs imposed by invasive species in protected areas (Moodleyet al. 2022; c) a focus on the “worst” 100 invasives (as determined by the IUCN) (Ahmedet al. 2022); and d) assessing costs associated with various pathways of introduction (Turbelin et al. 2022).
The InvaCost database, as applied in these studies, demonstrated that bioinvasions impose tremendous costs –a minimum of US $1.288 trillion for the period 1970 – 2017. These costs increased on average three times per decade (Daigne et al. 2022).
Still, everyone has recognized that InvaCost data have significant limitations. First, three-quarters of the records in the original database came from North America, Oceania and Europe; and referred to animal taxa, even though plants are a major group of invaders. Also, a large proportion of total invasion costs – for all taxa – probably is undetected. Finally, the many non-market values of species and ecosystems are extremely difficult to calculate (Daigne et al. 2022).
As a result of these deficiencies, the earlier studies discussed in the blogs referenced above substantially underestimated the true costs associated with bioinvasion (Cuthbert et al. 2022).
Now a new study, led by Ismael Soto, finds that the underestimate is huge. Global costs associated with a subset of 162 species (17% of all the species in the InvaCost database) is nearly 17 times higher than reported in the InvaCost database.
Soto et al. (2025) applied species distribution models and macroeconomic data to interpolate these 162 species’ probable impacts in 172 countries
Japanese knotweed – one of the invasive plants proving very costly in Europe, according to I. Soto
The newly identified costs were greatest in Europe; second place fell to North America. This is because both higher damage costs and management expenditures are linked to higher gross domestic product and extent of agricultural area, in addition to environmental suitability. Analysis of monetary costs per unit area revealed that ‘cost hot spots’ are predominantly located in densely populated urban areas and locations hosting key industries. These tend to be in coastal zones, i.e., Europe, the east coast of China, and the east and west coasts of the US.
cypress aphid Cinara cupressi – a threat to both native & plantation trees in Africa; photo by Blackman & Eastop via Wikimedia
The authors found that the greatest increase in estimated costs for countries in Africa and Asia. These countries had not previously recorded any economic costs arising from invasions by these 162 species. I have blogged about forest pest threats in Africa.
The authors also significantly increased estimated costs linked to invasive plants. Daigne et al. found that invasive insects caused ~90% of reported costs in the InvaCost database as of 2022. Vertebrates ranked second, plants third. In contrast, Soto et al. determined that invasive plants had the highest average estimated damage costs (US $42.10 billion) and management expenditures ($0.81 billion).
Substantial total costs were also reported for arthropods, mammals and birds. Reported damage and management costs were much lower for molluscs, fish, reptiles and amphibians. Daigne et al. suggest this might be due to their lower (observable) damage to human infrastructure, research biases leading to fewer studies, or disparities resulting from the filtering process used in their own study.
Williams et al. (2023) focus on insects, which cause damage primarily to agriculture, human health, and forestry. Insects constitute the highest number of species introduced as ‘Contaminants’ (n = 74) and ‘Stowaways’ (n = 43). They also impose the highest costs among species using these two pathways.
Forest insects and pathogens account for less than 1% of the records in the InvaCost database. I believe that this figure reflects significant under-reporting of these invasion events. Even at this paltry level of reported invasions, forest insects and pathogens were responsible for causing 25% of total annual costs ($43.4 billion) (Williams et al. 2023). This discrepancy illustrates the huge economic cost associated with widespread mortality of trees. Yet authorities in most countries continue to provide completely inadequate resources to counter this threat.
The authors of these publications examining economic losses associated with bioinvasion all note that ecological damage is additional. Soto et al. note that bioinvasions contribute to 60% of already recorded global extinctions. Interestingly, the species ranked third using the criterion of monetary damage is the cactus moth Cactoblastis cactorum. This insect threatens flat-padded Opuntia cacti across the United States and in the center of endemism, Mexico.
a flat-padded Opuntia — vulnerable to the cactus moth; photo by F.T. Campbell
Soto et al. found a lag of ~46 years between first (reported) detection of an introduced species and the peak of damage costs. They suggest that the rising monetary cost reflects the species becoming more abundant or occupying a larger area. The authors also say this finding demonstrates the value of implementing mitigation measures as soon as possible. Their finding thus validates others’ advocacy for investing in prevention and rapid response measures (see Cuthbert et al. and Daigne et al.). Soto et al. were cheered by the fact that spending on management measures – when it was reported – often followed soon after a species’ detection – or even before (e.g., prevention).
But Decision-Makers Usually Delay – Why?
Prevention is a hard sell. Decision-makers find it difficult to justify management expenditures before impacts become obvious. By that time, of course, management of the invasion is extremely difficult and expensive – if it is possible at all. Ahmed et al. found particularly effective wording to describe this problem: bioinvasion costs can be deceitfully slow to accrue, so policy makers don’t appreciate the urgency of taking action. Another contributing factor is that when efficient proactive management succeeds in preventing any impact, it paradoxically undermines evidence of the value of this action!
Programs to minimize the economic and ecological consequences of bioinvasion are severely obstructed – if not doomed! – by the following difficulties:
Resources are in short supply. Experts find that demands to address other threats to agriculture or natural systems outcompete appeals to ramp up invasive species efforts.
Prediction is uncertain. Cuthbert et al. found that none of the species with the highest pre-invasion investment was among the top 10 costliest invaders in terms of damages. Cuthbert et al. do not discuss whether this is evidence that the prevention efforts were effective? Or, alternatively, that prevention efforts target the wrong species.
Harm is in the eye of the beholder. Stakeholders’ perceptions of whether an introduced species causes a detrimental impact vary. For example, Moodley et al. point out that species imposing the highest economic costs might not be the ones causing the greatest ecological harm.
Externalities. Those harmed by a bioinvasion often are different from those that decide whether to act. Ahmed et al. argue that this creates a moral dilemma.
These decisions are political — influenced by citizens’ expressed wishes. Changing decision-makers’ perceptions of what is important is up to us!!! Start a parade!!!
SOURCES
Ahmed, D.A., E.J. Hudgins, R.N. Cuthbert, .M. Kourantidou, C. Diagne, P.J. Haubrock, B. Leung, C. Liu, B. Leroy, S. Petrovskii, A. Beidas, F. Courchamp. 2022. Managing biological invasions: the cost of inaction. Biol Invasions (2022) 24:1927–1946 https://doi.org/10.1007/s10530-022-02755-0
Diagne, C., B Leroy, A-C. Vaissière, R.E. Gozlan, D. Roiz, I. Jaric, J-M. Salles, C.A. Bradshaw, and F. Courchamp. 2021. High and rising econ costs of bioinvasions worldwide Published online: 31 March 2021
Moodley, D., E. Angulo, R.N. Cuthbert, B. Leung, A. Turbelin, A. Novoa, M. Kourantidou, G. Heringer, P.J. Haubrock, D. Renault, M. Robuchon, J. Fantle-Lepczyk, F. Courchamp, C. Diagne. 2022. Surprisingly high economic costs of bioinvasions in protected areas. Biol Invasions. https://doi.org/10.1007/s10530-022-02732-7
Soto, I., P. Courtois, A. Pili, E. Tordoni, E. Manfrini, E. Angulo, C. Bellard, E. Briski, M. Buric, R.N. Cuthbert, A. Kouba, M. Kourantidou, R.L. Macêdo, B. Leroy, P.J. Haubrock, F. Courchamp and B. Leung. 2025. Using species ranges and macroeconomic data to fill gap in costs of biological invasions. Nat Ecol Evol doi: 10.1038/s41559-025-02697-5
Turbelin, A.J., C. Diagne, E.J. Hudgins, D. Moodley, M. Kourantidou, A. Novoa, P.J. Haubrock, C. Bernery, R.E. Gozlan, R.A. Francis, F. Courchamp. 2022. Introduction pathways of economically costly invasive alien spp. Biol Invasions (2022) 24:2061–2079 https://doi.org/10.1007/s10530-022-02796-5
Williams, G.M., M.D. Ginzel, Z. Ma, D.C. Adams, F.T. Campbell, G.M. Lovett, M. Belén Pildain, K.F. Raffa, K.J.K. Gandhi, A. Santini, R.A. Sniezko, M.J. Wingfield, and P. Bonello. 2023. The Global Forest Health Crisis: A Public Good Social Dilemma in Need of International Collective Action. Annual Review of Phytopathology Vol. 61, 2023
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 Trump Administration’s budget for Fiscal Year 2026 [which begins at the end of September 2025] proposes to eliminate funding for nearly all USFS research & Forest Health Protection.
Proposed Cuts to USFS Research: Timber the Sole Aim
In a letter from Office of Management and Budget (OMB) to Senate Appropriations Committee Chair Susan Collins (R-Maine, Director Russell Vought says the Administration wants to manage National forests “for their intended purpose of producing timber” and that the research and development program “is out of step with the practical needs of forest management for timber production.” The Administration proposes to eliminate funding for USFS research projects other than the small portion covering Forest Inventory and Analysis.
I understand that the USFS Chief told various NGOs that his job is to run the National Forest System, increase timber production by 40%, and do nothing else.
This single aim conflicts with the 1897 legislation founding and authorizing the USFS. It also violates provisions of subsequent legislation such as the Multiple-Use Sustained-Yield Act of 1960 and the National Forest Management Act of 1976. It also departs from long-standing US Forest Service policy – which is the intention.
The “intended purpose” of establishing “forest reserves” [which were later renamed National forests] has never been solely for timber production. The “Organic Act” of 1897 provided that any new forest reserves would have to meet the criteria of forest protection, watershed protection, and timber production.
Specifically, theORGANIC ACT OF 1897 [PUBLIC–No.2.] says:
“[All public lands heretofore designated and reserved by the President of the US under the provisions of the Act [of] March 3rd 1891, the orders for which shall be and remains in full force and effect, unsuspended and unrevoked, and all public lands that may hereafter be set aside as public forest reserves under said act, [these were the “forest reserves,”predecessors of “National Forests]” shall be as far as practicable controlled and administered in accordance with the following provisions:
“No public forest reservation shall be established, except to improve and protect the forest within the reservation, or for the purpose of securing favorable conditions of water flows, and to furnish a continuous supply of timber for the use and necessities of [US] citizens; but it is not the purpose or intent of these provisions, or of the Act providing for such reservations, to authorize the inclusion therein of lands more valuable for the mineral therein, or for agricultural purposes, than for forest purposes.”
The Department of the Interior, which then managed these forest reserves, promptly issued implementing regulations. The regulations stated that the “object” of forest reservations was:
“2. Public forest reservations are established to protect and improve the forests for the purpose of securing a permanent supply of timber for the people and insuring conditions favorable to continuous water flow.”
Therefore, I think the Administration has exaggerated the emphasis on timber production by calling it “the” intended purpose of the original establishment of National forests. The Administration has also chosen to ignore subsequent legislation, such as the Multiple-Use Sustained-Yield Act of 1960 and the National Forest Management Act of 1976.
Sec. 13 of the NFMA limits the sale of timber from each national forest to a quantity equal to or less than a quantity which can be removed from such forest annually in perpetuity on a sustained-yield basis. This limit might be exceeded under certain circumstances, but such excess must still be consistent with the multiple-use management objectives of the land management plan. Further, Sec. 14 requires public input into any decision to raise timber allowances.
During his period as Chief (1905 – 1910), Gifford Pinchot invented and applied the concept of “conservation” of natural resources. As a result “wise use” became accepted as the national goal.
Culminating more than a century of legislation and informed policy, the mission of the USDA Forest Service is to “sustain the health, diversity, and productivity of the nation’s forests and grasslands to meet the needs of present and future generations.”
Proposed Cuts to State, Private, and Tribal Forests
The budget also cuts $303 million from the State, Private, and Tribal Forests program. (I understand this zeroes out the entire program). The OMB Director alleges that the program has been “plagued by oversight issues, including allegation of impropriety by both the Agency and State governments.” I understand that this would eliminate the cooperative projects managed by the Forest Health Protection program, too.
Implications for Non-native Insects and Pathogens
Remember that USFS’s research and development program is intended to improve forest managers’ understanding of ecosystems, including human interactions and influences, thereby enabling improvements to the health and use of our Nation’s forests and grasslands. Most importantly to me, this program provides foundational knowledge needed to develop effective programs to prevent, suppress, mitigate, and eradicate the approximately 500 non-native insects and pathogens that are killing America’s trees.
The Forest Health Program provides technical and financial assistance to the states and other forest-management partners to carry out projects (designed based on the above research) intended to prevent, suppress, mitigate, and eradicate those non-native insects and pathogens. The program’s work on non-federal lands is crucial because introduced pests usually start their incursions near cities that receive imports (often transported in crates, pallets, or imported plants).
Eliminating either or both programs will allow these pests to cause even more damage to forest resources – including timber.
Both supporting research and on-the-ground management must address pest threats across all U.S. forests, including the more than 69% that are located on lands managed by others than the USFS. Already, the 15 most damaging of these pests threaten destruction of 41% of forest biomass in the “lower 48” states. This is a rate similar in magnitude to that attributed to fire (Fei et al. 2019). It is ironic that the Administration considers the fire threat to be so severe that it has proposed restructuring the government’s fire management structure.
I remind you that the existing USFS R&D budget allocates less than 1% of the total appropriation to studying a few of the dozens of highly damaging non-native pests. I have argued that this program should be expanded, not eliminated. Adequate funding might allow the USFS to design successful pest-management programs for additional pests (as suggested by Coleman et al.).
As a new international report (FAO 2025) notes, genetic resources underpin forests’ resilience, adaptability, and productivity. Funding shortfalls already undercut efforts to breed trees able to thrive despite introduced pests and climate change (the latter threat is still real, although the Administration disregards it). I have frequently urged the Congress to increase funding for USFS programs – which are sponsored primarily by the National Forest System and State, Private, and Tribal, although some are under the R&D program.
Please ask your Member of Congress and Senators to oppose these proposed cuts. Ask them to support continued funding for both USFS R&D and its State, Private, and Tribal Programs targetting non-native insects and pathogens. America’s forests provide resources to all Americans – well beyond only timber production and they deserve protection.
Contacting your Representative and Senators is particularly important if they serve on the Appropriations committees.
House Appropriations Committee members:
Republicans: AL: Robert Aderholt, Dale Strong; AR: Steve Womack; AZ: Juan Ciscomani; CA: Ken Calvert, David Valadao, Norma Torres; FL: Mario Diaz-Balart, John Rutherford, Scott Franklin; GA: Andrew Clyde; ID: Michael Simpson; IA: Ashley Hinson; KY: Harold Rogers; LA: Julia Letlow; MD: Andy Harris; MI: John Moolenaar; MO: Mark Alford; MS: Michael Guest; MT: Ryan Zinke; NC: Chuck Edwards; NV: Mark Amodei; NY: Nick LaLota; OH: David Joyce; OK: Tom Cole, Stephanie Bice; PA: Guy Reschenthaler TX: John Carter, Chuck Fleishmann, Tony Gonzales, Michael Cloud, Jake Ellzey; UT: Celeste Maloy; VA: Ben Cline; WA: Dan Newhouse; WV: Riley Moore
Democrats: CA: Pete Aguilar, Josh Harder, Mike Levin; CT: Rosa DeLauro; FL: Debbie Wasserman Schultz, Lois Frankel; GA: Sanford Bishop; HI: Ed Case IL: Mike Quigley, Lauren Underwood; IN: Frank Mrvan; MD: Steny Hoyer, Glenn Ivey; ME: Chellie Pingree; MN: Betty McCollum; NJ: Bonnie Watson Coleman NY: Grace Meng, Adriano Espaillat, Joseph Morelle; NV: Susie Lee; OH: Marcy Kaptur; PA: Madeleine Dean; SC: James Clyburn; TX: Henry Cuellar, Veronica Escobar; WA: Marie Gluesenkamp Perez; WI: Mark Pocan
Senate Appropriations Committee members:
Republicans: AK: Lisa Murkowski; AL: Katie Britt; AR: John Boozman (AR); KS: Jerry Moran; KY: Mitch McConnell; LA: John Kennedy; ME: Susan Collins; MS: Cindy Hyde-Smith; ND: John Hoeven; NE: Deb Fischer; OK: Markwayne Mullin; SC: Lindsey Graham; SD: Mike Rounds TN: Bill Hagerty; WV: Shelley Moore Capito;
Democrats: CT: Chris Murphy; DE: Chris Coons; GA: Jon Ossof; HI: Brian Schatz; IL: Richard Durbin; MD: Chris van Hollen; MI: Gary Peters; NH: Jeanne Shaheen; NM: Martin Heinrich; NY: Kirsten Gillibrand; OR: Jeff Merkley; RI: Jack Reed; WA: Patty Murray; WI: Tammy Baldwin
SOURCES
Coleman, T.W, A.D. Graves, B.W. Oblinger, R.W. Flowers, J.J. Jacobs, B.D. Moltzan, S.S. Stephens, R.J. Rabaglia. 2023. Evaluating a decade (2011–2020) of integrated forest pest management in the United States. Journal of Integrated Pest Management, (2023) 14(1): 23; 1–17
FAO. 2025. The Second Report on the State of the World’s Forest Genetic Resources. FAO Commission on Genetic Resources for Food and Agriculture Assessments, 2025. Rome.
Fei, S., R.S. Morin, C.M. Oswalt, and A.M. 2019. Biomass losses resulting from insect and disease invasions in United States forests. PNAS August 27, 2019. Vol. 116 No. 35 17371–17376
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/
Guest blog by Kristy M. McAndrew, Department of Forestry, Mississippi State University
Virginia juniper (Juniperus virginiana) preforming its ecological role: succession in a field (in Ohio); photo by Greg Hume via Wikimedia
Spread of non-native species is a facet of global change that is an unintended consequence of the modern global trade network. Despite efforts put in place to limit such transport, such as International Standards for Phytosanitary Measures (ISPMs), unintentional spread of species continues, and thus, an important part of forest health research and management includes non-native monitoring and control efforts. As other aspects of global change, such as climate and weather patterns, shift, the dynamics between native landscapes and introduced pests may unexpectedly shift as well. For example, increased climate stress of tree hosts may weaken tree defenses, allowing species that historically have not been pests of concern to reach pest status.
Japanese cedar longhorned beetle (Callidiellum rufipenne; JCLB) is a wood boring beetle in the longhorned beetle family, Cerambycidae. The adults are reddish brown in color, and relatively small for longhorned beetles, at only around 1 cm in length. Japanese cedar longhorned beetle has a long history of establishing outside of its native range but has largely been considered a non-issue. It has long been disregarded as a pest because it feeds primarily on dead or dying trees in both the native and invaded ranges. However, there are more examples of these beetles feeding on stressed, but alive, trees in North America. Therefore, I think it is an important insect to take a closer look at.
Life cycle
These beetles have a one-year life cycle, most of which is spent inside a host tree. Adults emerge from host trees in the early spring and seek out other adults to mate with and trees to lay eggs on. Eggs are laid on thin parts of bark or in bark crevices, and when the eggs hatch larvae chew beneath the bark where they feed on the phloem until they have completed larval development. Once larvae are fully developed, they burrow further into the tree, into the xylem tissue, where they pupate, overwinter as fully formed adults, and continue the cycle the following spring.
Native range
The native range of JCLB is eastern Asia. It is common throughout the Korean peninsula and across the islands of Japan. It is also considered native to Eastern China and Russia. Within the native range JCLB is found primarily on dead and/or dying trees and is thus considered a secondary pest. On dead trees they can be found on any diameter of dead woody material, but on declining trees they will likely be in the small diameter branches and stems.
Arborvitae (Thuja occidentalis); photo by James St. John via Flickr
Invasion history
Japanese cedar longhorned beetle was first documented as an invasive pest in the early 1900s in France, and since then has established in at least fifteen countries (Clément 2023). Most of these countries are in Europe, but the United States and Argentina also have established populations. As with most woodboring insects, the invasion pathway is believed to have been wood packaging material being transported via global trade routes. Between 1914 and 2022 it was intercepted over 700 times (reviewed by KM). Since the implementation of ISPM No. 15, only six interceptions have been reported up to 2022 (USDA APHIS data reviewed by K.M.). [For Faith’s view on the regulation of wood packaging, see Fading Forests II and III (links provided at the end of this blog) and earlier blogs posted here under the category “wood packaging”. esp. 1 from 2015].
A USDA risk assessment completed in 2000 suggested other possible pathways of introduction, including balled nursery stock, green logs, and pruned branches (USDA APHIS and Forest Service, 2000).
In terms of establishments in North America, JCLB was first detected in natural forests in North Carolina in 1997. It was soon discovered in Connecticut in 1998; in neighboring New York in 1999; and in Massachusetts, New Jersey, and Rhode Island in 2000. It was quickly discovered feeding on live arborvitae (also called northern white cedar; Thuja occidentalis) in these invaded regions. JCLB has since been found in Pennsylvania (in 2010) and Maryland (in 2011). It is important to note that it is not clear when this species truly established, because of its previously discussed long history of being intercepted in ports of entry.
Most introduced populations of JCLB are found in either dead hosts or in the damaged/dead limbs of live hosts. In Buenos Aires, for example, storm-damaged trees with broken limbs are often where beetles are collected (Turienzo 2007). In the United States, eastern red cedar (Juniperus virginiana) and common juniper (Juniperus communis ) are the two native species most commonly affected, but so far there is no evidence of live trees of these species being infested (Maier 2007). However, a growing concern in the United States is that JCLB has been documented on live trees – particularly in urban environments. These trees are typically arborvitae, and they are typically stressed urban trees that have been overwatered and often show signs and symptoms of other health issues.
Host breadth
The host breadth of JCLB encompasses much of the family Cupressaceae. Maier (2007) identified 19 potential hosts from the literature and research, with the vast majority (14) of the hosts being Cupressaceae species, which is indicative of JCLB being a relative generalist, especially when considering species in the cypress family. This is important, because there are over 130 species within Cupressaceae worldwide that could be suitable hosts for JCLB, meaning host will not be a limiting factor in many invasion scenarios for this insect. Most often trees infested by JCLB need to be either stressed or dead, which limits suitability to an extent. However, many landscape trees are inherently stressed, whether it be from a history of roots being balled and wrapped in burlap, being planted in less than ideal scenarios, or being overwatered.
A few reports from research in Japan record JCLB feeding on plants in Pinaceae, primarily Pinus and Abies species. One article reports use of Larix kaempferi; another documented JCLB on the Taxaceae species, Taxus cuspidata. North American pine (Pinus spp.) and fir (Abies spp.) species have not been tested, but if they are revealed as suitable that would increase the availability of hosts in North America significantly.
In southern New England at least nine species have been confirmed as suitable, all of which are in the family Cupressaceae. Native and abundant junipers, such as Juniperus virginiana, appear to be highly suitable hosts. Additional host testing would be beneficial – especially Cupressaceae species that are either threatened or have a limited range. Within the United States there are a total of 28 native Cupressaceae species. Thus the suitable range (in terms of hosts) covers the entire Eastern half of North America through central Texas, most of the Pacific Coast, and widespread but spotty/disjunct areas throughout the Intermountain West and High Plains regions.
Atlantic white cedar swamp (Chamaecyparis thyoides) in Brendan Byrne State Forest, New Jersey; photo by Famartin via Wikimedia
Suitability
Tools such as environmental niche models can give helpful estimates of suitability. For species that are typically secondary pests, such as JCLB, it can be difficult to obtain non-biased data with good coverage to make reliable predictions. Preliminary research (unpublished) has been completed to estimate suitable habitat with limited occurrence records from the native range. Despite limited occurrences, models performed well and estimated moderate to high suitability in most temperate regions globally. These preliminary models are still being optimized by working with collaborators within the native range of JCLB to increase the number of occurrences. It is also important to note that these models are only accounting for climate data. Host data was not included, but Cupressaceae species are abundant globally, and therefore host availability is not likely a limiting factor for JCLB in establishing in regions.
Importance of monitoring species
While JCLB is still mostly limited to dead, dying trees, many of the species it may affect in the Eastern United States are already of heightened conservation concern. Wetland Cupressaceae, such as bald cypress (Taxodium distichum) and Atlantic White Cedar (Chamaecyparis thyoides), are valuable in terms of ecosystem services they provide in coastal, and inland, wetlands. These wetlands are encountering heightened stress in the form of increasing saltwater intrusion, increased storm strength, and changing landscapes, all of which may predispose trees to insect attack. Japanese cedar longhorned beetle has been successfully reared out of logs of Atlantic White Cedar, but thankfully has not been documented on live trees of this species (Maier 2009)[Ma1] . Bald cypress has not yet been tested for suitability. It is unknown if the stressors these trees are facing and will continue to face will impact JCLB’s ability to infest these landscapes, or if they will remain restricted to dead trees in these coastal forests. Regardless, given JCLB already has an established foothold in the Eastern United States, it is important to better understand the potential impacts of this insect.
First steps to understanding those impacts include 1) better documenting the host range in the regions and 2) determining the climate that may support the species. Hopefully we can continue research in these areas to best manage this non-native pest.
Much of the research conducted on JCLB in North America took place almost 20 years ago (Maier 2007, 2009), so updated sampling has potential to provide a wealth of information regarding spread rate, suitable climate, and establishment patterns.
bald cypress(Taxodium distichum); photo by Kej605 via Wikimedia; it is unknown whether this species is vulnerable to the Japanese cedar longhorned beetle
Sources
Clément F. 2023. Le point sur la distribution en France et en Europe de Callidiellum rufipenne (Motschulsky, 1861)(Coleoptera, Cerambycidae, Cerambycinae, Callidiini). Le Coléoptériste. 26(3):188–203.
Maier CT. 2007. Distribution and Hosts of Callidiellum rufipenne (Coleoptera: Cerambycidae), an Asian Cedar Borer Established in the Eastern United States. JOURNAL OF ECONOMIC ENTOMOLOGY. 100(4).
Maier CT. 2009. Distributional and host records of Cerambycidae (Coleoptera) associated with Cupressaceae in New England, New York, and New Jersey. Proceedings of the Entomological Society of Washington. 111(2):438–453. https://doi.org/10.4289/0013-8797-111.2.438
Turienzo P. 2007. New records and emergence period of Callidiellum rufipenne (Motschulsky, 1860) [Coleoptera:Cerambycidae: Cerambycinae: Callidiini] in Argentina. Boletín de Sanidad Vegetal, Plagas. 33:341–349.
United States Department of Agriculture Animal and Plant Health Inspection Service and Forest Service 2000. (Pasek, J.E., H.H. Burdsall, J.F. Cavey, A. Eglitis, R.A. Haack, D.A. Haugen, M.I. Haverty, C.S. Hodges, D.R. Kucera, J.D. Lattin, W.J. Mattson, D.J. Nowak, J.G. O’Brien, R.L. Orr, R.A. Sequeira, E.B. Smalley, B.M. Tkacz, W.W. Wallner) Pest Risk Assessment for Importation of Solid Wood Packing Materials into the United States. USDA APHIS and Forest Service. August 2000.
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/
results of invasion by emerald ash borer (photo courtesy of Nathan Siegert, USFS ); one of the woodboring beetles found to be so damaging to hardwood trees
Over the last nine years scientists have made significant progress in identifying aspects of insect-plant host relationships that play important roles in determining how much damage an introduced, non-native pest is likely to cause within forest ecosystems in the United States. Predicting which introductions will probably cause the greatest damage is vitally important because scientists, phytosanitary officials, and resource managers cannot address all the hundreds of established insects, much less the thousands which might be introduced. This shortfall increases with each surge in import volumes (see my previous blogs about wood packaging by scrolling down the website below the “Archives” to “Categories”, then find “SWPM”), proliferation of goods types and source areas, and cutbacks in funding.
I hope USDA APHIS and Forest Service are adjusting their procedures to apply the scientists’ path-breaking findings.
Their progress will help protect our forests. I apologize if I seem ungrateful — but we need similar progress in managing plant pathogens. Consider the damage caused by chestnut blight, white pine blister rust, Dutch elm disease, sudden oak death, beech leaf disease … (All these and other pathogens are described briefly here.) Understanding the universe of introduced fungi, water molds, nematodes, viruses, etc., is per se much more challenging. Ashley Schulz points out that among the complications are pathogens’ complex life cycles, and possible new relationships with vectors.
Undertaking this analysis will be set back decades if agencies’ resources – funds and staffs – are decimated during the current “downsizing” of government. We must speak up!!
At least regarding non-native insects that attack North American tree species, scientists’ analyses promise a new ability to set priorities. This should improve the efficacy of phytosanitary programs – if government downsizing is not allowed to destroy USDA’s scientific, regulatory, and resource management programs.
We must speak up!!
What science tells us now
Schulz et al. (2025) summarize current findings. (Full citation to all references appear at the end of the blog.)
Earlier, scientists sought to find commonalities associated with introduced insects that caused high impacts on North American conifer trees [Mech et al. (2019)] and hardwoods (angiosperms) [Schulz et al. (2021)] (Full citations at the end of the blog; earlier blogs posted here and here.) Both studies found that the time elapsed since tree species in North America diverged from the host plants of the insects in their native range (i.e., host evolutionary history) is a diagnostic factor. This factor best predicted non-native insect impact compared to the other factors that were significant for conifer and hardwood specialists. For conifers, the other significant factors included the shade and drought tolerance of the North American host plants and whether there was a related insect native to North America on the same hosts that the non-native insect impacted. For hardwoods, another important factor explaining a specialist insect’s impact is if the insect is a wood borer, especially a scolytine beetle. The wood density of the North American host plant was also considered a significant factor when predicting impact of the non-native insect.
In 2022, Uden et al. applied the divergence time method to insect species not yet introduced to North America that might attack conifer species. They hoped to identify both insects posing the greatest hazard and tree species most vulnerable to introduced pests.
Now, a new team again led by Ashley Schulz and Angela Mech (see Schulz et al. 2025) has applied a similar approach to a more comprehensive range of pest-host relationships, including the pests that specialize on host plants and pests that feed on a broader array of hosts. Some feed on both conifers and hardwoods. They found that:
It is possible to quantify insect host breadth and identify the cutoff where “specialists” and “generalists” diverge. Specifically, the split occurs around 2,250 cumulative million years, where insects that feed on hosts that add up to less than that have narrow host breadth (i.e., “specialists”) and insects that feed on hosts that add up to more than that have broad host breadth (i.e., “generalists”). This technique also helps categorize insects that fall within the middle range of host breadth and are traditionally difficult to classify as either specialists or generalists based on differing qualitative definitions of the terms.
Insects that use more hosts in their native range also tend to use more hosts in the introduced range (North America). However, many of these insects utilized fewer hosts in the introduced range compared to the native range. This shrinkage was not universal, however; about 30% of insects increased their host breadth in the introduced range. Most of these fed on a single species in their native range but attacked additional species in the same family in North America. The corresponding i-Tree Pest Predictor tool uses the list of hosts in the insect’s native range and these models to determine the insect’s likelihood that it would cause high impact, as well as each North American tree species’ susceptibility to the insect entered into the tool.
Certain feeding guilds had – on average — a significantly narrower host breadth in North America than in their native ranges. These were gall makers (13 species analyzed); sap feeders (120 species); and wood borers (35 species). In contrast, host ranges did not differ for folivores (68 species), reproductive feeders (7 species), and root feeders (5 species). Still, we know that wood borers, as a group, have caused enormous damage to a range of North American tree taxa (see emerald ash borer, redbay ambrosia beetle, invasive shot hole borers (all described briefly here). Again, the i-Tree Pest Predictor tool can help identify the threat to particular tree species.
Of course, APHIS should not disregard pests with narrow host ranges; several have caused enormous damage.
red spruce (Pikea rubens) in Great Smoky Mountains National Park; photo by famartin via Wikipedia. Red spruce is the species found to be most vulnerable by the Uden et al. study
Schulz et al. (2025) developed models for three groups of introduced herbivorous insects that feed on trees:
1) conifer specialists (based on analysis of 69 species);
2) hardwood specialists (based on analysis of 141 species);
3) hardwood generalists (based on analysis of 30 species).
Because of their quantification of host breadth, they defined the “specialist” group more broadly than is commonly done, e.g., an insect that feeds on the three families Betulaceae, Fagaceae, and Juglandaceae would be considered “specialists” because all three host families are in the Fagales clade.
Tree relatedness was the only significant explanatory factor for all three host breadth categories. As determined in the previous studies, North American host tree species that were too closely or distantly related to the insect’s hosts in its native range were less impacted than hosts that diverged somewhere in the middle – the “Goldilocks” range. The divergence period differs among the three pest-risk categories: 3–4 million years ago for conifer specialists, 5–9 million years ago for hardwood specialists, ~1–2 million years ago for hardwood generalists. Schulz et al. suggest that the reason why the peak probability of high impact differs among these groups is that different feeding guilds cause the most damage to the specific host category, and each feeding guild is challenged by different tree host defenses. Bark and wood boring beetles (the hardwood specialists with the greatest impact) must overcome lethal constitutive and induced tree defenses in order to survive for long periods in the cambial layer. These insects have adapted the ability to locate and select poorly defended individuals in the host population. Folivores (i.e., the generalists with the highest impact) adapt to plant chemistry and trichomes (hair-like or scale-like outgrowths), or can avoid host defenses by moving off the foliage. Sap feeders (which include many high impact conifer specialists) are usually tolerated by trees, unless they stimulate hypersensitive reactions or vector pathogens.
Of course, scientists’ estimates of how long ago tree taxa diverged from common ancestors differ. Fortunately, Uden et al. (2022) found that these differences only rarely affect the predicted impact of a non-native insect – at least in the case of the 62 European insects and 47 North American conifer species they analyzed. In only 1.37% of the 2,914 pairs analyzed did the predicted risk differ depending on which source phylogeny was used. These cases were associated with 27 conifer-specialist insects and 9 conifer hosts. The article does not tell us which pest/host pairs these are but, overall, this paper demonstrates that the estimate differences in the phylogenetic trees does not differ enough to be problematic when forecasting insect impact.
Changes Needed in the Way Agencies Set Priorities
Schulz et al. (2025) urged agencies to stop relying only on insect traits as the basis for developing models & phytosanitary regulations. The only insect trait that predicted impact is the insect’s feeding guild. Considering hardwoods, they found that wood borers pose the greatest risk among specialists to hardwoods; folivores among generalists. While sap feeders do not cause statistically higher damage on hardwood tree species, four of the seven high-impact conifer specialists are sap feeders (hemlock woolly adelgid, balsam woolly adelgid, red pine scale, and spruce aphid). Therefore, the i-Tree Pest Predictor tool incorporates consideration of whether a pest of conifers is a sap feeder.
Schulz et al. (2025) also caution agencies against relying on just the number of hosts an insect might exploit. Assessors must consider the range of underlying plant chemistry / host defenses that the insect encounters. They found that hosts that are shade tolerant are more susceptible to high impact from conifer specialists and hosts that have intermediate to no shade tolerance are more susceptible to high impact from generalists.
Uden et al. (2022) identified a possible weakness in USDA efforts to prioritize pest prevention targets. They found that APHIS’ Prioritized Offshore Pest List included only 12 conifer specialists from Europe among the 150 species listed. They go on to note that while sap feeders constitute 53% of tree pest species established in the U.S., APHIS listed none. The models applied by Uden, Schulz, and Mech do not consider whether the insect is likely to become established. Improving our understanding of the many factors influencing an insect’s likelihood of being transported to North America or becoming established requires additional research. This might eventually lead to a usable tool for predicting this aspect of bioinvasion by forest pests.
There is an urgent need for such a tool. As Uden et al. noted, they found that 66% of the insect species they analyzed fell into the “high impact” category. This is a much higher proportion than estimates based on earlier studies, so identifying which of these insects are likely to establish versus not establish in North America can provide more resolution and help identify which insects are going to be most problematic.
Mature Fraser fir killed by balsam woolly adelgid; Clingman’s Dome; photo by Ben Ramsey via Flickr
Tree species at risk
The analysis undertaken by Uden et al. determined that three conifer species face a high level of hazard from European insects if they are introduced. They identified particularly high threats to two species, Fraser fir (Abies fraseri) and Carolina hemlock (Tsuga caroliniana). The fir is determined to be vulnerable to 17 insect species which are predicted to have high likelihood of a high impact. The hemlock is highly vulnerable to one of the insect species they sampled. They note that both of these conifers have a limited geographic range and ecological habitat, so they likely have a relatively narrow genetic pool. A third species said to be at elevated risk is red spruce (Picea rubens) – which, although more widespread, is also under attack by a non-native insect. All three species fit earlier finding by Mech et al. that conifer trees with high shade tolerance but low drought tolerance more vulnerable to non-native pests. In none of these cases do Uden et al. mention that the tree species have already been severely diminished by established non-native insects – i.e., balsam woolly adelgid on the fir (above), hemlock woolly adelgid on the Carolina hemlock. The Schulz/Mech team is working to refine methods for identifying tree species and regions at greatest risk.
Meanwhile, Uden et al. have suggested that phytosanitary authorities and forest managers apply their findings to identify the European herbivorous insects that pose the greatest threat to North American conifer species. They should identify Palearctic tree species that fall within the high-impact “Goldilocks” zone of divergence times in relation to specific North American tree species, then identify the insects that feed on those Palearctic trees. These insects would presumably pose the highest predicted hazard to those North American tree species. They suggest that species so identified should be added to the USFS’ list of species targetted by its wood borer early detection program. To address likelihood of introduction, they suggest incorporating data on insect species commonly intercepted at ports – an indication of high propagule pressure. There will always be exceptions though. For example, Ips typographus feeds on spruce and has been frequently detected at the ports, but it has not established in North America.
For those focused on identifying species or ecoregions at greatest risk, Uden et al. suggest scientists use several sources to identify vulnerable vegetation communities. Sources suggested include USFS Forest Inventory and Analysis (FIA) and NatureServe Explorer plant community descriptions) that have relatively high-value tree species predicted to be at risk from introduced species.
SOURCES
Aukema, J.E., D.G. McCullough, B. Von Holle, A.M. Liebhold, K. Britton, and S.J. Frankel. 2010. Historical Accumulation of Nonindigenous Forest Pests in the Continental United States. BioScience 60(11): 886-897. https://doi.org/10.1525/bio.2010.60.11.5
Mech, A.M., K.A. Thomas, T.D. Marsico, D.A. Herms, C.R. Allen, M.P. Ayres, K.J. K. Gandhi, J. Gurevitch, N.P. Havill, R.A. Hufbauer, A.M. Liebhold, K.F. Raffa, A.N. Schulz, D.R. Uden, & P.C. Tobin. 2019. Evolutionary history predicts high-impact invasions by herbivorous insects. Ecology and Evolution 9(21): 12216–12230. https://doi.org/10.1002/ece3.5709
Schulz, A.N., A.M. Mech, M.P. Ayres, K. J. K. Gandhi, N.P. Havill, D.A. Herms, A.M. Hoover, R.A. Hufbauer, A.M. Liebhold, T.D. Marsico, K.F. Raffa, P.C. Tobin, D.R. Uden, K.A. Thomas. 2021. Predicting non-native insect impact: focusing on the trees to see the forest. Biological Invasions 23: 3921-3936. https://doi.org/10.1007/s10530-021-02621-5
Schulz, A.N., N.P. Havill, T.D. Marsico, M.P. Ayres, K.J.K. Gandhi, D.A. Herms, A.M. Hoover, R.A. Hufbauer, A.M. Liebhold, K.F. Raffa, K.A. Thomas, P.C. Tobin, D.R. Uden, A.M. Mech. 2025. What Is a Specialist? Quantifying Host Breadth Enables Impact Prediction for Invasive Herbivores
Uden, D.R., A.M. Mech, N.P. Havill, A.N. Schulz, M.P. Ayres, D.A. Herms, A.M. Hoover, K.J.K. Gandhi, R.A. Hufbauer, A.M. Liebhold, T.D. Marsico, K.F. Raffa, K.A. Thomas, P.C. Tobin, C.R. Allen. 2022. Phylogenetic risk assessment is robust for forecasting the impact of European insects on North American conifers. Ecological Applications 33(2): e2761. https://doi.org/10.1002/eap.2761
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/
Scientists in New Zealand are saying explicitly that a forest’s unique mixture of species matters when considering the future. This mixture is the result of the forest’s evolutionary history. Losing members of the biological community reduces the forest’s ability to respond to current and future stresses – its resilience.
New Zealand’s forests are part of the broader legacy of the ancient supercontinent of Gondwanaland – the island nation’s plants have close relatives in South America, the Pacific Ocean islands, and Australia. Still, these forests are unique: 80% of New Zealand’s plant species are endemic. The forests are also species-rich. The warm temperate evergreen rain forests of the North Island are home to at least 66 woody plant species that can reach that reach heights above six meters (Simpkins et al. 2024).
These forests have been severely changed by human activity. In just ~ 750 years people have cut down approximately 80% of the original forest cover! (Simpkins et al. 2024) Of the eight million hectares of surviving native forest, a little over five million hectares is managed for the conservation of biodiversity, heritage, and recreation. Another 2 million hectares are plantations of non-native species.
sites in New Zealand where pine plantations are “wilding”
All these forests are challenged by introduced mammals – from European deer to Australian possums. Climate change is expected to cause further disturbance, both directly (through e.g., drought, extreme weather) and indirectly (e.g., by facilitating weed invasion and shifting fire regimes) (Simpkins et al. 2024).
Pathogen threats are also common threats to the native trees of the Pacific’s biologically unique island systems. For example, Ceratocystis lukuohia and C. huliohia (rapid ‘ōhi‘a death, or ROD). The latter is killing ‘ōhi‘a (Metrosideros polymorpha) on the Hawaiian Islands. More than 40% of native plant species in Western Australia are susceptible to Phytophthora cinnamomi. Here I focus on two pathogens, kauri dieback and myrtle rust, now ravaging New Zealand’s native flora. No landscape-level treatment is available for either pathogen.
When considering this suite of challenges, Simpkins et al. focus on these two pathogens’ probable impact on forest carbon sequestration. They worry in particular about erosion of the forests’ resilience due to loss of “ecological memory” – the life-history traits of the species (e.g., soil seed banks) and the structures left behind after individual disturbances.
one of the largest remaining kauri trees, “Tane Mahuta”, in Waipoua Kauri Forest; photo by F.T. Campbell
Kauri Dieback
The causal agent of Kauri dieback, Phytophthora agathidicida, is a soil-borne pathogen that spreads slowly in the absence of animal or human vectors. The disease affects a single species, Agathis australis (kauri, Araucariaceae). However, kauri is a long-lived, large tree that is a significant carbon sink. It probably modifies local soil conditions, nutrient and water cycles, and associated vegetation. Also, kauri has immense cultural significance.
Simpkins et al. note that kauri dieback threatens stand-level loss of A. australis – that is, local extinctions. In the absence of disturbance Kauri trees can grow to awe-inspiring size. In the 19th Century, before widespread logging, some were measured at 20 meters or more in circumference. Consequently, kauri dieback might cause a decline in aboveground live carbon storage of up to 55%. This loss would occur over a period of hundreds of years, not immediately.
Huge kauri are not likely to be replaced by other long-lived emergent conifers (based on an analysis of one species, Dacrydium cupressinum). Instead, kauri are probably going to be replaced by late-successional angiosperms. The authors discuss the ecological implications for levels of carbon storage and proportions of trees composed of Myrtaceae – exacerbating damage caused by myrtle rust (see below).
The expectation of Simpkins et al. that kauri will suffer at least local extinctions is based on an assumption that no kauri trees are resistant to the pathogen. Fortunately, this might not be true: different Agathis populations show various levels of tolerance to Agathis dieback. Identification and promotion of some levels of resistance could enable A. australis to retain a diminished presence in the landscape.
However, Lantham, et al. make clear that containing kauri dieback remains “challenging,” despite its discovery nearly 20 years ago (in 2006). Scientists and land managers have little information on the distribution of symptomatic trees, much less of the pathogen itself. This means they don’t know where infection foci are or how fast the disease is spreading.
As is often true, the pathogen is probably present in a stand for years, possibly a decade or more, before symptoms are noticed. This means that the current reliance on public reports of diseased trees, or targetting surveillance on easy-to-access sites (e.g., park entrances and along existing track networks), or at highly impacted areas readily identified through aerial methods, fails to detect early stages of infection. Indeed, it seems probable that P. agathidicida had been present in New Zealand’s ecosystems for decades before its formal identification.
The Waipoua forest is one of the largest areas of forest with old kauri stands in the country. A new analysis of aerial surveys done between 1950 and 2019, shows how the forest is changing. The number of dead trees increased more than four-fold and the number of unhealthy-looking trees increased 16-fold over these 70 years. Kauri dieback is now widespread in this forest, especially in areas near human activities like clearing for pasture or planting commercial pine plantations).
Lantham et al. have developed a model which they believe will help identify areas of higher risk so as to prioritize surveillance and inform responses. These could delimit the disease front and help implement quarantines or other measures aimed at limiting the spread of P. agathidicida to uninfected neighboring sites.
I hope New Zealand devotes sufficient resources to expand surveillance and management to levels commensurate with the threat to this ecologically and culturally important tree species.
Leptospermum scoparia; photo by Brian Gatwicke via Flickr
Myrtle Rust
Myrtle rust is a wind-borne disease that affecting numerous species in the Myrtaceae, including some of the dominant early successional species (e.g., Leptospermum spp.). Simpkins et al. expect that myrtle rust might hasten the decline of two such tree species (L. scoparium and Kunzea ericoides). However, these trees’ small size and rapid replacement by other species during succession minimizes the effect of their demise on carbon storage.
Because I am concerned about the irreplaceable loss to biodiversity, I note that Simpkins et al. also feared immediate threats to some trees in the host Myrtaceae family, specifically highly susceptible species such as Leptospermumbullata.
As I reported in a recent blog, a second group of scientists (McCarthy et al.) explored the threat from myrtle rust more broadly. Austropuccinia psidii has spread through Myrtaceae-dominated forests of the Pacific islands for about 20 years.
Trees in the vulnerable plant family, Myrtaceae, are second in importance (based on density and cover) in New Zealand’s forests. Successional shrub communities dominated by the two species named above, Kunzea ericoides and Leptospermum scoparium, are widespread in the northern and central regions of the North Island and in northeastern and interior parts of the South Island. These regions’ vulnerability is exacerbated by the area’s climate, which is highly suitable for A. psidii infection (Simpkins et al. 2024).
McCarthy et al. concluded that ifLeptospermumscoparium and Kunzea ericoides prove to be vulnerable to myrtle rust, their loss would cause considerable change in stand-level functional composition across these large areas. They probably would be replaced by non-native shrubs, which are already common on the islands. Any resulting forest will differ from that formed via Leptospermeae succession.
These authors also worry that the risk to native ecosystems would increase if more virulent strains of the myrtle rust pathogen were introduced or evolved. They note that A. psidii is known to have many strains and that these strains attack different host species.
SOURCES
Latham, M.C., A. Lustig, N.M. Williams, A. McDonald, T. Patuawa, J. Chetham, S. Johnson, A. Carrington, W. Wood, and D.P. Anderson. 2025. Design of risk-based surveillance to demonstrate absence of Phytophthora agathidicida in New Zealand kauri forests. Biol. Invasions (2025) 27, no. 26
McCarthy, J.K., S.J. Richardson, I. Jo, S.K. Wiser, T.A. Easdale, J.D. Shepherd, P.J. Bellingham. 2024. A Functional Assessment of Community Vulnerability to the Loss of Myrtaceae from Myrtle Rust. Diversity and Distributions, https://doi.org/10.1111/ddi.13928
Simpkins, C.E., P.J. Bellingham, K. Reihana, J.M.R. Brock, G.L.W. Perry. 2024. Evaluating the effects of two newly emerging plant pathogens on North Aotearoa-New Zealand forests using an individual-based model. Ecological Modelling, www.elsevier.com/locate/ecolmodel
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/
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/
discarded pallets next to developed area in Glacier National Park (!); photo by F.T. Campbell
Since July 2015 I have posted nearly 50 blogs about non-native insects introduced via movement of solid wood packaging material (SWPM). Why? Because SWPM is one of two most important pathways by numbers introduced & by impact of the species introduced. (The other pathway is P4P.) To read those earlier blogs, scroll below “archives” to “categories”, choose “wood packaging”.
dead redbay trees in Everglades National Park; killed by laurel wilt vectored by redbay ambrosia beetle
As I have reported in the earlier blogs and in my “Fading Forests” reports (links at the end of this blog), in 2002, the parties to the International Plant Protection Convention (IPPC) adopted an international “standard” to guide countries’ programs intended to reduce the presence of damaging insects in the wood packaging: International Standard for Phytosanitary Measures (ISPM) #15). The U.S. and Canada adopted the standard through a phase-in process culminating in 2006. [For a discussion of the phase-in periods and process, read either of the studies by Haack et al. cited at the end of this blog.] In other words, the U.S. and Canada have implemented ISPM#15 for almost 20 years. China specifically has been subject to requirements that it treat its SWPM even longer – since December, 1998, i.e., more than 25 years.
Unfortunately, ISPM#15 is not intended to prevent pest introductions. As stated in Greenwood et al 2023, “Prior to 2009, the goal of compliance with ISPM 15 was to render the risk of wood-borne pests “practically eliminated,” in 2009 the standard was amended to “significantly reduced”.
Despite almost universal adoption of the standard by countries engaged in international trade, insects have continued to be present in wood packaging. A very high proportion of these infested shipments — 87% – 95% — of the SWPM found by U.S. officials bears the ISPM#15 stamp – that is, is apparently compliant. (See my blogs by clicking on the “Category” “wood packaging” listed below the “Archives”.) The same proportion was found in a narrower study in Europe (Eyre et al. 2018). All the post-2006 examples of infested wood analyzed by Haack et al. (2022) (see below) carry the stamp. I conclude that the ISPM#15 mark has failed in its purpose: to reliably indicate that SWPM accompanying imports has been treated so as to minimize the likelihood that an insect pest will be present.
Dr. Robert Haack, retired USFS entomologist, has twice tried to estimate the “approach rate” of insects in SWPM entering the United States (both studies are cited at the end of this blog). A study published in 2014 that relied on data from 2009 found that U.S. implementation of ISPM#15 was associated with a reduction in the SWPM infestation rate reported of 36–52%. The authors estimated the infestation rate to be 0.1% (1/10th of 1%, or 1 consignment out of a thousand). (See Haack et al. 2014; citation at the end of this blog.)
In their second study, published in 2022, Haack and colleagues found a 61% decrease in rates of borer detection in wood packaging when comparing numbers of wood borer detections in 2003 – before the U.S. implemented ISPM#15 – to those in 2020. Specifically, detections dropped from 0.34% in 2003 to 0.21% in 2020. This decrease occurred despite the volume of U.S. imports rising 68% between 2003 and 2020. (My blogs document a further increase in import volumes over the years since 2020.) In addition, the number of countries from which the SWPM originated more than doubled from 2003–2004 to 2010–2020. This expansion exposes North America to a wider range of insect species that might be introduced, as well as a wider range of individual countries’ effectiveness in enforcing the standard’s requirements (Haack et al. 2022).
These decreases are encouraging. However, Haack et al. (2022) note some caveats:
The reduction in pest presence was greatest during the initial implementation of the program the first phase, 2005-2006 (61%); in subsequent periods pest approach rate inched back up. In the 2010-2020 period, the pest detection rate was only 36% below the pre-ISPM#15 level. Detection rates have been relatively constant since 2005. Does this stasis mean that exporters learned that they could ignore or circumvent the requirements without suffering significant penalties? Or is some of this rise related to increased trade volumes, increasing variety of country of origin for trade, or other global trade patterns unrecognized in the data? (However, see the next bullet point.)
Certain types of commercial goods and exporting countries have consistently fallen short. Specifically, the rate of wood packaging from China that is infested remained relatively steady over the 17 years since 2003. The proportion of consignments with infested wood packaging coming from China was more than five times the proportion of all inspected shipments for this period. In other words, China has had a consistent record of poor compliance with phytosanitary regulations since they were imposed in December 1998. Why is USDA not taking action to correct this problem? (As I note below, DHS CBP has ramped up enforcement efforts.) Some other countries, e.g., Italy and Mexico, have reduced the rate at which wood packaging accompanying their consignments is infested. In fact, Mexico’s improved performance largely explains the overall infestation rate estimate of 0.22% during the period 2010-2010. Mexico’s successes affect the overall statistics in a way that makes other countries’ failure to reduce the presence of pests in wood packaging they ship to the United States far less obvious.
Haack et al. (2022) discuss ten possible explanations for their finding that pest approach rates – as determined by their study — have not decreased more. See the article or my blog about the study.
Although USDA APHIS has not taken steps to strengthen its enforcement, U.S. Customs and Border Protection [an agency in the Department of Homeland Security] has done so twice — see here and here. CBP staff have expressed disappointment that these actions reduced the numbers of shipments in violation of ISPM#15 by only 33% between Fiscal Year 2017 and FY2022. True, more than 60% of these violations consisted of a missing or fraudulent ISPM#15 stamp. However, 194 consignments still harbored live pests prohibited under the standard.
APHIS did agree in 2021 to enable the study by Robert Haack and colleagues, via an interoffice data sharing agreement between USDA APHIS and the Forest Service- this resulted in Haack et al. 2022.
APHIS and CBP also collaborated with an industry initiative to train inspectors that insure other aspects of foreign purchases. The ideas was that CBP or APHIS and their Canadian counterparts would inform importers about which foreign treatment facilities have a record of poor compliance or suspected fraud. The importers could then avoid purchasing SWPM from them. I have heard nothing about this initiative for three years, so I fear it has collapsed.
We lack data on which to base a rigorous analysis
While the two studies by Robert Haack and colleagues are the best available, and they relied on the best data available, the fact is that those available data do not provide a full picture of the risk of pest introduction associated with wood packaging. As pointed out by Leigh Greenwood of The Nature Conservancy in her presentation to 2025 USDA Invasive Species Research Forum, available data have been collected for different purposes than to answer this question. Leigh’s powerpoint is posted here.
Leigh has identified the following data gaps:
In their studies, Haack and colleagues rely on data from the Agriculture Quarantine Inspection Monitoring (AQIM) system. This dataset is based on random sampling of very distinct segments of incoming trade. It is therefore a better measure of insect approach rates than reports of interceptions by either APHIS or CBP.
However, AQIM includes data from only those very distinct segments of trade: perishable goods, SWPM associated with maritime containerized imports, Italian tiles, and “other” goods, AQIM does not contain a segment of trade that includes wood packaging associated with maritime breakbulk or roll-on, roll-off (RORO) cargo. These exclusions have prevented scientists and enforcement officials from determining, inter alia, how great a risk of pest introduction is associated with various types of wood packaging, especially dunnage, as the randomized sample does not include entire pathways for the entrance of dunnage.
Greenwood states that she has not found another country that operates a similar analysis of randomly collected data at ports of entry.
2) USDA does not collect data on consignment size, piece-specific infestation density, nor consignment-wide infestation density. As Haack et al. (2022) point out, reporting detections by consignment doesn’t reveal the number of insects present. If implementation of ISPM#15 resulted in fewer live insects being present in an “infested” consignment, this would reduce the establishment risk because there is lower propagule pressure. However, we cannot know whether this is true.
3) Neither USDA nor CBP reports the inspection effort. Nor do they conduct a “leakage survey” to see how often target pests are missed. This means, inter alia, that we cannot estimate inspectors’ efficiency in detecting infested wood packaging. If their proficiency has improved as a result of improvements in training, inspection techniques, or technology, the apparent impact of ISPM#15 would be under-reported in recent years.
4) USDA does not require port inspectors to report the type of SWPM in which the pest was detected. Leigh participated in an effort that included industry representatives, DHS CBP and USDA APHIS to define the types of wood packaging in legal terminology so that they could be incorporated in the drop-down menu on inspectors’ reporting system. This was first successfully included in the legal glossary within USDA APHIS system of record, ACIR Glossary. Last fall the team was working to integrate the requirement for using these definitions into the inspection data collection system used by DHS CBP, which would then make this data available in Agricultural Risk Management, ARM (see Abstract here for adequate primer on ARM). However, it is unclear now whether the new administration will do so. One potential barrier is that asking the port of entry inspection staff to record these data will add to the time and training required for reporting inspection results.
In summary, Leigh reports that current data systems do not support
estimating probabilities of pest infestation of via volume or type of SWPM (e.g. pallet vs dunnage)
measuring the risk of arrival associated with a specific hazard (in this case, a hazard being a live pest or pathogen associated with SWPM)
extrapolating or supporting findings for some types of wood packaging to other types of wood packaging
Scientists from Canada, Mexico, and the United States have formed a working group under the auspices of the North American Plant Protection Organization (NAPPO). The group is trying to determine whether various types of wood packaging are more likely to harbor pests. This study is currently hampered by the many data gaps, including those Leigh outlined above. The best data available, cited by Haack et al. (2022), found that in maritime containerized shipping, crates were more likely to harbor pests than pallets- however, other forms of SWPM (dunnage, bracing, etc.) had such low sample size that no analysis of those is possible. One of the main objectives of the NAPPO study is to evaluate if dunnage poses the same or higher risk, so this is a major impediment.
Two issues need to be resolved.
One is scientific: why are insects continuing to be detected in wood packaging marked as having been treated? What is the relative importance of insects surviving the treatment versus treatment facilities applying the treatments incorrectly or inadequately?
The second issue is legal and political: what proportion of the detections is due to treatment facilities committing outright fraud – claiming to treat the wood, stamping it with an IPPC stamp, while not actually performing any treatments at all?
Knowing which measures will most effectively solve these quandaries / reduce pest presence in wood packaging depends on knowing what the relative importance of these factors are in causing the problem. The lack of basic data on which to base any analysis certainly hampers efforts to improve protection.
Leigh calls for researchers to recognize these data needs and work to fill them.
•Understand, account for, and communicate data realities
•Work collectively to increase useable data quality
•Use additional research to validate, or to demonstrate disparities
Why Wait for the Science?
In the meantime, however, I assert that more vigorous enforcement efforts by responsible agencies should help reduce the occurrence of fraud. I have suggested the following actions:
U.S. and Canada refuse to accept wood packaging from foreign suppliers that have a record of repeated violations – whatever the apparent cause of the non-compliance. Institute severe penalties to deter foreign suppliers from taking devious steps to escape being associated with their violation record.
APHIS and CBP and their Canadian counterparts follow through on the industry-initiated program described above and here aimed at helping importers avoid using wood packaging from unreliable suppliers in the exporting country.
Encourage a rapid switch to materials that won’t transport wood-borers. Plastic is one such material. While no one wants to encourage production of more plastic, the Earth is drowning under discarded plastic. Some firms are recycling plastic waste into pallets.
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/
