Invasive plants threaten integrity of eastern U.S. forests

garlic mustard (*Alliaria petiolata*); photo by Katja Schulz via Wikimedia

I welcome a recent series of studies documenting the extent of plant invasions in forests of the eastern United States and the socio-economic conditions that contribute to a state of affairs increasingly recognized as a crisis. I wish, however, that the authors had devoted more attention to the role of deliberate planting of non-native species and the resulting propagule pressure.

I summarize here findings of two studies written by largely the same scientists and relying on the same underlying data: surveys of forest plots conducted under the Forest Inventory and Analysis (FIA) program. In this blog, if focus on the extent of invasive plant presence in the forests of the eastern United States. In an accompanying blog I will summarize the status of plant invasions in forests nation-wide.

As I have noted in earlier blogs, link a decade ago one or more invasive plant species had already invaded 46% of FIA plots in the eastern U.S. (Oswald et al. 2015). This situation has worsened. Updated data show that 52.8% of these plots contain invasive plants. In the USFS Southern Region, invasive plants have been documented on 55.3 million ha. In the Northern Region, they are found on 36.9 million ha. (Only ~20% of FIA plots in the Northern Region were surveyed for invasive plants.) In some counties of the 37 states constituting these two USFS regions, 80% of inventoried forest plots contain invasive plants. Areas with lower levels of invasion are found in parts of New England, the Great Lakes states, southern Appalachians, southeastern coastal plain, and western Texas and Oklahoma (Potter et al. 2026). Spread of these bioinvaders is largely unchecked – either throughout the East or “just” in the South. In any case, the extent and intensity of these invasions are so great that their complete removal – or elimination of their impacts – is “practically impossible” (Potter et al., 2024; Potter et al. 2026). [It is not clear whether the scientists mean “nearly” or “in practice”. Or that this difference is important.]

[In comparison, in the West less than 30% of FIA plots are invaded, on average. In Hawai`i, more than70% are (Potter et al. 2026).]

The scientists analyzing the FIA data warn that the extent and impact of plant invasions in eastern forests is undoubtedly worse than these data indicate. The records include only some of the non-native plant species present — those considered to be the worst invaders at the time regional lists were compiled – apparently in the first years of the 21^st Century (Potter et al. 2026).

Japanese honeysuckle (*Lonicera japonica*) photo by Chuck Bargeron

The scientists emphasize the role of disturbance in promoting plant invasions. They cite various studies as well as the FIA data to document that forest edges facilitate non-native plant establishment and spread into forests. They stress various aspects of suburban development, including roads and other transportation corridors. It follows that invasion rates are highest in the “wildland-urban interface (WUI).” [The wildlife-urban interface is the zone of transition between unoccupied land and human development; the zone where structures meet or intermix with undeveloped land and its vegetation.] They worry that the WUI is growing faster than any other land use type in the country – and especially rapidly in the East. As a result, the scientists expect more and worse invasions in the future (Potter et al., 2024 and Potter et al. 2026).

I appreciate that they highlight the uniqueness of WUI ecosystems. Housing development in the WUI has numerous effects on natural ecosystems, including habitat modification and fragmentation followed by diffusion of the direct and indirect effects of anthropogenic activities into neighboring ecosystems at different scales. As regards specifically non-native plants, this transmission occurs through a combination of (1) human-driven disturbances to native ecosystems that promote plant invasion and (2) providing a source of non-native plant propagules in their yards and gardens. These plants can then spread into and establish in nearby ecosystems (in this case, forests). [I note that tree-killing arthropods and pathogens also can be introduced in the WUI.] (Scroll below “Archives” to “Categories”, click on “forest pests” and “wood packaging”.)

They also found that plant invasions are more strongly related to older, than more recent, land-cover changes. Survey plots that have been located in the WUI since 1990 or earlier had on average 2.6% more invasive plant cover and 0.33 more invasive plant species than those that were classified as being in the WUI in 2000 or 2010. Their explanation is that the WUI forests experienced decreased spatial integrity, increased forest-developed area edges, and falling proportions of forest in the surrounding landscapes. In addition, the human population in the vicinity might have grown. All these factors that would increase forest fragmentation and the plots’ susceptibility to invasion.

The other side of the coin is propagule pressure. Both Potter et al (2024) and Potter et al. (2026) note that the flora of residential landscapes – rural as well as suburban – is typically dominated by non-native plant species. Still, I think these studies downplay the impact of this ubiquity of non-native plants in all anthropogenic landscapes.

In discussing the higher invasion rates found in survey plots located in WUIs dating from the 1990s they made no mention of human activities that promote plant invasions. There are several. Plants growing in those older yards had one or two more decades to flower – and for their fruits and seeds to be transported into the forest by birds, wind, or water. Residents might have decided to beautify their neighborhood by planting shrubs or flowers in the woods. Maybe they succumbed to the temptation to dump yard waste in the woods – thinking it would be absorbed by “nature”. Since plant invasions take time to unfold, these additional years of human-mediated exposure are highly relevant. Another factor is that people who choose to live in wooded surroundings probably choose horticultural plants that thrive under such conditions – exactly those best able to establish beyond the property line.

Another opportunity to discuss these factors came from the discovery that plant invasion rates are higher in association with “interface” rather than “intermix” WUI forests. [“WUI interface forests” are those where settled areas abut wildlands. In “WUI intermix forests” the structures are scattered.] They speculate about reasons. Potter, et al. (2024) mention that invasions originating from older housing developments have had more time to establish (or at least to be detected) given the well-known lag associated with plant invasions.

I wish they had focused more on the probable difference in suburban development across time. While I was growing up in expanding suburbs in the 1950s, I observed that the earlier housing developments were either built on land that had been cleared to support agriculture or the builders cleared the forest to make construction easier and cheaper. More recently, wealthier buyers have sought residences on more wooded sites – so creating an “intermix” WUI. Potter et al. (2024) speculate that locations in the “interface” WUI are closer to high-density urbanization so have higher exposure to non-native plants. They do not discuss whether the “interface” WUIs are older, thus giving associated plantings longer years to proceed through the stages of bioinvasion.

burning bush (*Euonymus alatus*) invading a forest in Virginia; photo by F.T. Campbell

The Role of Deliberate Planting?

I recognize that these authors analyzed mountains of data. However, I wish they had incorporated the findings of numerous scientists who have analyzed the role of deliberate planting – especially ornamental horticulture – in facilitating introduction and spread of invasive plants. (Scroll below “Archives” to “Categories” and click on “invasive plants”. Also See Reichard and White 2001 and Mack 2000).

As I hope USFS scientists are aware, recent studies confirm the continuing role of ornamental horticulture in plant invasions. Kinlock et al. (2025) blog 440 found that more than 1,600 plant species sold by nursery and seed catalogs over 200 years had “naturalized” somewhere in the continental 48 states. They do not discuss what proportion of these species are truly damaging invaders. Fertakos and Bradley (2024) found that species were likely to establish if they were introduced to as few as eight locations. Beaury et al. (2024) found that half of 89 plant species recognized as invasive are sold in the same locations where they are invasive. Another 25 species are sold by one or more nurseries located in an area that is currently unsuitable for those species, but that will become more suitable for invasion as temperatures warm.

Potter et al. (2026) acknowledge that the ornamental plant trade is likely to continue introducing new plant species into U.S. forests. However, they recommend only updating the lists of invasive plants to be included in future surveys. Apparently these lists have not been updated since 2004.

Potter et al. (2024) go farther, urging efforts to encourage homeowners to plant more native and environmentally friendly private landscapes. They note that such advocacy is complicated by the fact that non-native – even invasive – species provide valued ecosystem and cultural services.

I add that the nursery industry and their customers enjoy enormous lobbying clout.

Many associations – native plant societies, regional or state invasive plant councils, etc. – are pursuing this approach. To research these efforts, visit the websites for the state native plant societies and the Southeast Exotic Pest Plant Council, Mid-Atlantic Invasive Plant Council, and Midwest Invasive Plant Network. These voluntary efforts have yielded some success. But they have not resulted in adequate protection for our ecosystems. Dr. Douglas Tallamy points out that even non-invasive, non-native plants disrupt food webs.

The insufficient attention to the role of the plant trade in articles intended to be comprehensive has crucially important impacts. As both Potter, et al. (2024) and Potter et al. (2026) affirm, determining which factors are most important in facilitating plant invasions of eastern American forests is the necessary foundation for identifying and implementing the most efficient and effective counter measures.

These scientists are employees of the U.S. Department of Agriculture. If departmental leadership interpret their studies as justifying inaction on regulating plant sales, USDA’s regulatory agencies will not respond. And we will continue failing to curtail introduction and spread of damaging plant invasions.

I agree with the authors on the need for enhanced monitoring and management of WUI zones in the East to detect new species or new locations of invasion and the need to develop better tools for these purposes. However, I ask all stakeholders to follow Evans et al. (2024), who urge prioritizing for state regulation those species in the ornamental trade that are projected to remain or become abundant under evolving climate conditions. Or, more aggressively, follow Beaury et al. (2023)’s call for regulating the nursery trade in a manner consistent with the scope of the horticultural trade at the national level. That would require legislation, since the Federal Noxious Weed Act does not currently address long-established, widespread species. Beaury et al. (2023) also note that existing state restrictions are outdated, tend to include only a few weeds that plague agriculture rather than those that invade natural systems, and are irregularly enforced.

orchids in Everglades National Park; photo by F.T. Campbell

I conclude by agreeing with the scientists that managing the disturbance component of plant invasions points to protecting particularly forests of high conservation value. They suggest adoption of land-use planning rules aimed at this goal. However, as they point out, such action will be extremely unlikely given the magnitude of predicted land-use changes in the country and powerful demographic factors driving them. I would add other barriers: the lobbying clout of the real estate industry and homeowners plus the local nature of zoning decisions.

SOURCES

Beaury, E.M., J.M. Allen, A.E. Evans, M.E. Fertakos, W.G. Pfadenhauer, B.A. Bradley. 2023. Horticulture could facilitate invasive plant range infilling and range expansion with climate change. BioScience 2023 0 1-8 https://doi.org/10.1093/biosci/biad069

Evans, A.E., C.S. Jarnevich, E.M. Beaury, P.S. Engelstad, N.B. Teich, J.M. LaRoe, B.A. Bradley. 2024. Shifting hotspots: Climate change projected to drive contractions and expansions of invasive plant abundance habitats. Diversity and Distributions 2024;30:4154

Fertakos, M.E. and B.A. Bradley. 2024. Propagule pressure from historic U.S. plant sales explains establishment but not invasion. Ecology Letters 2024;27:e14494 doi: 10.1111/ele.14494

Kinlock, N.L., D.W. Adams, W. Dawson, F. Essl, J. Kartesz, H. Kreft, M. Nishino, Jan Pergl, P. Pyšek, P. Weigelt and M. van Kleunen. 2025. Naturalization of ornamental plants in the United States depends on cultivation and historical land cover context. Ecography 2025: e07748 doi: 10.1002/ecog.07748

Oswalt, C.M., S. Fei, Q. Guo, B.V. Iannone III, S.N. Oswalt, B.C. Pijanowski, K.M. Potter. 2016. A subcontinental view of forest plant invasions. NeoBiota. 24:49-54 http://www.srs.fs.usda.gov/pubs/48489

Potter, K.M., K.H. Riitters, B.V. Iannone III, Q. Guo and S. Fei. 2024. Forest plant invasions in the eastern United States: evidence of invasion debt in the wildland‑urban interface. Landsc Ecol (2024) 39:207 https://doi.org/10.1007/s10980-024-01985-y

Potter, K.M., B.V. Iannone III, K.H. Riitters, Q. Guo, K. Pandit, C.M. Oswalt. 2026. US Forests are Increasingly Invaded by Problematic Non-Native Plants. Forest Ecology and Management 599 (2026) 123281

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

https://fadingforests.org

Pest Threats to Plantations: Will At-Risk Countries Demand Improvements to IPPC?

pines in a plantation in Argentina killed by *Sirex noctilio*; photo by J. Villacide

A decade ago, Payn et al. (2015) compiled studies from around the globe to evaluate threats to widespread tree plantations. At that time, they said climate change posed the greatest threat to plantation forestry globally, in the forms of storm and flood damage and simultaneous warming and drying trends with extreme temperatures.

Still, the authors warned that forest health would be an increasingly important constraint to plantation productivity. They were optimistic, however, that modern breeding and other technologies could offset losses.

What is the current situation? The countries that depend on these plantations for fiber production are not demanding that leaders of the international phytosanitary structure build a more effective system to protect their investments. Instead, individual scientists struggle to better understand threats. Mostly, they propose expanded research.

Economic Importance of these Species

Eucalypts

“Eucalypts” comprises three genera in the family Myrtaceae: Angophora, Corymbia and Eucalyptus. These include more than 700 tree species native primarily to Australia. A few species are native to Indonesia, New Guinea and the Philippines (Paine et al. 2011; Crous et al. 2019). Some of these species have been extensively planted outside their native ranges for more than 100 years. These plantations have expanded rapidly in recent decades, especially in Southeast Asia and the Southern Hemisphere (Crous et al. 2019). Eucalypts are now the most widely planted hardwood timber in the world (Paine et al. 2011).

Eucalypt plantation in Brazil; photo by Jonathan Wilken via Wikimedia

Eucalypts’ popularity has been driven chiefly by their rapid growth; short rotation times including through coppicing; and adaptability to a very wide variety of sites and climatic conditions (Paine et al. 2011; Crous et al. 2019). Also, these trees are an important source of the short-fiber pulp required for production of high-quality paper used in modern office copiers and printers (Paine et al. 2011). Plantations are increasing even in Australia, where harvesting of native forests is increasingly being restricted (Paine et al. 2011).

Pines

Pines – a genus restricted naturally to the Northern Hemisphere – is second in global popularity. South America hosts 4.6 million hectares of pine plantations (Lantschner and Villacide 2025). South America is more dependent on forestry plantations for wood production than any other region. In 2012, 88% of its industrial roundwood was produced by non-native plantations. This far exceeded the global proportion of approximately 19%.

These intensively managed plantations have enabled Brazil and Chile to become “planted forest powerhouses.” Uruguay and, more slowly, Argentina are following the same path (Payn et al. 2015).

Documentation of the Damage

Euclaypts

The highly diverse eucalypts host an even greater diversity of fungi. As of 30 years ago, scientists were aware of more than 500 species of just one type, the leaf-infecting fungi. Additional fungi are associated with seeds, capsules, twigs, branches, and stems. Little is known about the vast majority of these fungi. Even species considered causal agents of important diseases have not yet been confirmed using Koch’s Postulates. Areas of origin for most is also unknown (Crous et al. 2019).

Crous et al. (2019) compiled information on 110 genera of fungi found on eucalypt foliage. Some genera include well-recognized primary pathogens. They name Austropuccinia and Calonectria, Coniella, Elsinoe, Pseudocercospora, Quambalaria and Teratosphaeria. Other genera are thought to include species that are opportunists that develop on stressed or dying tissues. Many other leaf fungi are putative pathogens, but unstudied. Additional fungi cause vascular wilts (e.g. Ceratocystidaceae), stem canker diseases (Cryphonectriaceae, Botryosphaeriaceae) and root diseases (e.g. Armillaria, Ganoderma) of eucalypts.

Crous et al. (2019) state that the rust Austropuccinia psidii is one of the most damaging of the foliage fungal pathogens. They consider it to be a greater threat to eucalypt plantations outside the trees’ native ranges. (The Myrtaceous species in Australia most damaged by A. psidii are in other genera.)

Two families of leaf fungi – Mycosphaerellaceae and Teratosphaeriaceae – include species that cause serious diseases. Pérez, et al. report a study in plantation in Uruguay that detected six new species. They also discovered new hosts for some known species. (Such initial detections of new fungal species in out-of-native-range plantations is a usual occurrence.)

Over the 100-year history of planting eucalyptus outside Australasia, dozens of leaf pathogens have been transported to novel regions. Crous et al. 2019 report the wide geographic breadth of many of these introductions. For example, Mycosphaerella heimii is crippling plantation forestry in five global regions – South America (Brazil and Venezuela); Asia (Indonesia and Thailand); Africa (Madagascar), Europe (Portugal); and in its presumably native Australia. A second species, M. marksii, has a similarly wide introduced range: Portugal, China and Indonesia, South Africa, Ethiopia, and Uruguay. Pérez et al. calls Mycosphaerella leaf diseases one of the most important impediments to Eucalyptus plantation forestry in Uruguay.

Although Crous et al. do not provide dates of detection, it appears that many of these leaf pathogens were introduced outside Australasia before the mid-990s, when the World Trade Organization (WTO) and International Plant Protection Convention (IPPC) came into force. Together, these agreements govern what actions phytosanitary officials may take to curtail international movement of plant pests. (To see my critique of the WTO/IPPC system, visit here.) The possible exception might be Kirramyces gauchensis, a well-known pathogen of Eucalyptus grandis in South America (Argentina and Uruguay), Hawai`i, and Africa (Uganda and Ethiopia) (Pérez, et al. 2009). Crous et al. (2019) expect another genus, Quambalaria species, to become a threat to eucalypt plantation forestry globally in the future.

*Phoracantha semipunctata*; photo by Umo Schmidt via Flickr

Arthropod pests have also been spread to many Eucalyptus-growing regions in North and South America, Europe and Africa since the 1980s. Some species have colonized virtually all eucalypt-growing regions, e.g., Phoracantha semipunctata. Some have – so far – appeared on only one continent.

In an effort to determine how many of these introductions have occurred after adoption of the WTO/ IPPC system, I Googled the species named by Paine et al. (2011). I used the year 2000 as the cutoff date, to allow for detection lag. Among the insect species that fit this criterion are a lerp psyllid, a leaf beetle, and two gall wasps detected in North America; a true bug, two galling insects, and a leaf beetle in South Africa; and three psyllids in Europe.

Asia stands out as having very few introduced Australian insects plaguing eucalyptus plantations. Only one insect of Australian origin is causing significant damage in this region, Leptocybe invasa. It was detected after 2000, so it might have been introduced under the WTO/IPPC regime. Many widespread species, e.g., Phoracantha semipunctata, are notably absent. Instead, large numbers of endemic insects use these trees. This contrasts with the situation in the Southern Hemisphere, where few of the numerous native insects have shifted onto eucalypts.

New Zealand has detected only two new species of Australian origin since 1999 — two psyllids. This is despite the two nations’ proximity, the large volume of trade that passes between them, and the likelihood that at least some small sap-suckers might be introduced via aerial dispersal. New Zealand is famous for its strict phytosanitary (and sanitary) policies and programs.

Eucalyptus plantation in Kwa-Zulu, South Africa

Plantations’ vulnerability has been increased by expanding reliance on clonal, artificially-induced hybridization. Developers’ goals – and initial results – are enhanced adaptation to specific environments, desired fiber characteristics, and hybrid vigor. However, these vast areas planted in genetically identical trees are sitting ducks. An insect or pathogen that overcomes the host’s defenses can spread rapidly across the entire planting.

These hybrids also can act as “bridges,” facilitating spread of fungi to formerly resistant host species. Crous et al. (2019) fear that this process will undermine resistance in Eucalyptus pellita to the pathogen Teratosphaeria destructans. Plantations in Southeast Asia and South Africa now comprise hybrids between this resistant species and the highly susceptible Eucalyptus brassiana.

Pines

As with the eucalypts, the intensively managed pine plantations are comprised of fast-growing exotic species, all at the same developmental stage, and with minimal genetic diversity, planted to maximize wood production. These practices again lead to biological homogenization and reduced resilience to pests (Villacide and Fuetealba, 2025)

In the Southern Hemisphere, Sirex noctilio has become the most significant economic pest of Pinus species. These attacks can cause up to 80% mortality. Several other Sirex species have also been introduced, all apparently in the 1980s or earlier (Wilcken et al., 2025) – before adoption of the current international phytosanitary regime. However, in 2023, a new species, Sirex obesus, was discovered causing tree mortality in pine plantations in southeastern Brazil. This species is indigenous to the United States and Mexico.

Stazione et al. (2026) discuss two other non-native pine pests that established recently in South America.

Analysis of mitochondrial DNA of Orthotomicus erosus points to a western Eurasian lineage. The low genetic diversity of the introduced population in Argentina and Uruguay suggests a single or limited introduction event followed by regional spread.

The source region of Cyrtogenius luteus is more difficult to determine but is probably somewhere in China. The higher haplotype diversity might reflect multiple introductions. Again, shared haplotypes between Argentina and Uruguay countries indicates a contiguous regional spread, possibly driven by extensive pine plantations & intra-regional trade (Stazione et al. 2026)

Policy Aspects

Some scientists express concern about the failure of international phytosanitary measures. But are their countries speaking up in regulatory bodies, especially the International Plant Protection Convention?

Studies by Crous et al. (2019) and Pérez et al. (2009) clearly show that pathogens from Australia continue to be transported to regions where eucalypt plantations are grown. This happens despite most of the movement of genetic material being in the form of seeds – which should be less likely to transport pathogens than trade in plants. Pérez et al. (2009) explicitly raise concerns about the effectiveness of current quarantine procedures. Crous et al. (2019) state that quarantines continue to fail in many parts of the world.

Burgess and Wingfield (2017) list pathogens that have spread widely since the beginning of the 21st Century: Austropuccinia psidii, Calonectria (= Cylindrocladium) eudonaviculata (=Cylindrocladium buxicola), Ceratocystis lukuohia and C. huliohia introduced to Hawai`i. I add that insect-vectored diseases such as Euwallacea species carryingFusarium fungi have also experienced a burst of introductions around the globe since 2000.

Crous et al. (2019) attribute this failure partially to the enormous difficulty of applying effective quarantine to the huge volumes of planting material traded globally. Another factor is undoubtedly the poor understanding of microbial species, their pathogenicity, hosts, pathways of spread, even taxonomies. Some genera cannot be grown in culture.

Furthermore, pathogens’ impacts vary, possibly due to environmental conditions of the location or differing virulence on different hosts. Finally, with so many fungi and so little knowledge, it is difficult to separate true disease agents from multiple secondary infections.

Crous et al. (2019) express the hope that increased recognition of the importance of pathogens, along with improved detection and identification tools, will clarify patterns of spread. But is that enough? Are there no policy changes needed?

Crous et al. (2019) also warn us about additional pathways for spreading pathogens. Some potential pathogens of eucalypts have been moved on plants of other, related genera. Furthermore, Botryosphaeriaceae have been detected in the skins of mangoes (Mangifera indica) and avocados (Persea americana). Both of these fruits move globally in large volumes.

mangoes; photo by Obsidian Soul via Wikimedia

Regarding insects, Paine et al. (2011) focus on a concern that species native to the plantation countries and generalist herbivores from other parts of world will invade Australia and threaten eualypts in their native ranges. See other blog They also call for research to understand international pathways, develop detection methods, improve understanding of patterns of host suitability, susceptibility, and selection.

Villacide and Fuetealba (2025) note that while the introductory pathway for that new species, Sirex obesus, has not been determined, they suspect it might have been wood packaging materials. Villacide and another colleague (Lantschner and Villacide 2025) suggest an initial step would be for Argentina and other countries in the region to negotiate with Brazil to adopt more protective protocols governing trade in wood products, including wood packaging.

I have repeatedly advocated strengthening regulation of wood packaging. Such measures could improve protection of Earth’s forests from pests that use a well-documented high-risk introductory pathway. To see my arguments and underlying data, scoll down below the “archives” to “Categories” and click on “wood packaging”.

SOURCES

Burgess, T.I. and M.J. Wingfield. 2017. Pathogens on the Move: A 100-Year Global Experiment with Planted Eucalypts. Bioscience. Volume 67, Issue 1, January 2017. https://doi.org/10.1093/biosci/biw146

Crous, P.W., M.J. Wingfield, R. Cheewangkoon, A.J. Carnegie, T.I. Burgess, B.A. Summerell, J. Edwards, P.W.J. Taylor, and J.Z. Groenewald. 2019. Folia pathogens o eucalypts. Studies in Mycology 94:125-298 (2019).

Lantschner, V. and J. Villacide. 2025. Invasion Potential of the Recently Established Woodwasp Sirex obesus. Neotropical Entomology. (2025) 54:117 https://doi.org/10.1007/s13744-025-01347-6

Paine, T.D., M.J. Steinbauer, and S.A. Lawson. 2011. Native and Exotic Pests of Eucalyptus: A Worldwide Perspective. Annu. Rev. Entomol. 2011. 56:181-201

Payn, T., J-M. Carnus, P. Freer-Smith, M. Kimberley, W. Kollert, S. Liu, C. Orazio, L. Rodriguez, L. Neves Silva, M.J. Wingfield. 2015. Changes in planted forests and future global implications. Forest Ecology and Management 352 (2015)

Pérez,, C.A., M.J. Wingfield, N.A. Altier, and R.A. Blanchette. 2009. Mycosphaerellaceae and Teratosphaeriaceae associated with Eucalyptus leaf diseases and stem cankers in Uruguay For. Path. 39 (2009) 349–360 doi: 10.1111/j.1439-0329.2009.00598.x www3.interscience.wiley.com

Stazione, L., Soliani, C., Cognato, A. et al. Reconstructing the invasion history of the bark beetles Orthotomicus erosus & Cyrtogenius luteus (Coleoptera, Curculionidae, Scolytinae) in South America. Biol Invasions 28, 49 (2026). https://doi.org/10.1007/s10530-026-03779-6

Villacide, J. and A. Fuetealba. 2025. Pests in plantations: Challenging traditional productive paradigms in the Southern Cone of America. Forest Ecology and Management 597 (2025) 123127

Wilcken, C.F., T.A. da Mota, C.H. de Oliveir, V.R. de Carvalho, L.A. Benso, J.A. Gabia, S.R.S. Wilcken, E.L. Furtado, N.M. Schiff, M.B. de Camargo, M.F. Ribeiro. 2025. Sirex obesus (Hymenoptera: Siricidae) as invasive pest in pine plantations in Brazil. Scientific Reports. 2025. 15:22522 https://doi.org/10.1038/541598-025-06418-7

Posted by Faith Campbell

https://fadingforests.org

Threat to Native Myrtaceae in South America

*Blepharocalyx salicifolius –* a tree in the Myrtaceae native to South America on which found symptoms similar to those caused by Mycosphaerellaceae or Teratosphaeriaceae; photo by Pablo di Flores via Wikimedia

Pests that have followed their hosts to plantations outside the trees’ native ranges might threaten native plants in their new, introduced ranges. That is, the countries where the plantations are located.

Eucalypts

Eucalypts are now the most widely planted hardwood timber taxon in the world (Paine et al 2011). The 700 – 800 species in the three genera considered “eucalypts” (Angophora, Corymbia, and Eucalyptus) host a highly diverse fungal community — more than 500 species have been identified of just one type, leaf-infecting fungi (Crous et al. 2019).

As I described in a related blog, link dozens of leaf pathogens have been transported to countries hosting eucalypt plantations. Among them, two families – Mycosphaerellaceae and Teratosphaeriaceae – are prominent in both numbers of introductions and potential to cause serious diseases.

Pérez et al. (2009) reported that a relatively large number of Mycosphaerellaceae and Teratosphaeriaceae are found on Eucalyptusin Uruguay. The authors cite one troubling case of host shifting: Mycosphaerella lateralis is causing leaf disease on a Musa cultivar (banana!) which is not in the Myrtaceae.

A follow-up study by the same authors (Pérez et al. 2013) surveyed several native forests, paying special attention to those located close to Eucalyptus plantations. They found five species belonging to the Mycosphaerellaceae and Teratosphaeriaceae clades on native Myrtaceous trees; three of these had previously been reported on Eucalyptus in Uruguay. Those occurring on both Eucalyptus and native Myrtaceae included Pallidocercospora heimii, Pseudocercospora norchiensis, and Teratosphaeria aurantia. A fourth species, Mycosphaerella yunnanensis, not previously recorded in Uruguay, was found on the leaves of two native Myrtaceous hosts. Pérez et al. (2013) believe circumstances indicate that all these fungi have been introduced. They warn that these apparent jumps to new hosts have the potential to result in serious disease problems and they should be carefully monitored. This finding is more than a decade old; I have not found a more recent report.

On the global level, Pérez et al. (2013) report, at least 23 species of Mycosphaerellaceae and Teratosphaeriaceae have been found on non-Eucalyptus species in the Myrtaceae. These hosts are in several plant orders, including Myrtales, Proteales, Fabaes and Apiales. The authors express “considerable concern” about the apparent ease of movement in these fungi between hosts. I have been unable to learn more details about these introductions.

Arthropod pests have also been spread to many Eucalyptus-growing regions in North and South America, Europe, and Africa since the 1980s – but not to Asia or New Zealand (Paine et al. 2011). blog

*Myrrhinium atropurpureum* – another South American plant in the Myrtaceae on which symptoms found; photo by Prof. Atilio L, Botanical Garden of Uruguay

Pines

Pines – a genus restricted naturally to the Northern Hemisphere – is second in popularity for intensively managed plantations. South America has 4.6 million hectares of pine plantations (Lantschner and Villacide 2025). Most are in Brazil, Chile, Uruguay, and Argentina (Payn et al. 2015).

*Cinara cupressi;* photo by LBM via Wikimedia

As I reported in an earlier blog, some of the insect pests that followed pines to South America have entered native forests. The most alarming of which I am aware is the aphid Cinara cupressi. It attacks the native conifer Austrocedrus chilensis, which forms pure and mixed stands with southern hemisphere beech (Nothofagus spp.) across approximately 160,000 hectares (Villacide and Fuetealba 2025). Cordilleran cypress is also under attack by the oomycete Phytophthora austrocedri, an oomycete of unknown origin.

Some scientists express concern about phytosanitary measures … but are their countries speaking up in meetings of the International Plant Protection Convention?

Studies by Crous et al. and Pérez et al. clearly show that pathogens from Australia continue to be transported to regions where eucalypt plantations are grown – despite the fact that most of the movement of tree genetic material is in the form of seeds – which should be less likely to transport pathogens than trade in plants. Pérez et al. (2009) explicitly raise concerns about the effectiveness of current quarantine procedures. Crous et al. (2019) state that the quarantines continue to fail in many parts of the world.

See my critique of the international phytosanitary system under the IPPC by visiting the Fading Forest II report (see link below) and reading other blogs under the categories “invasive species policy” and “plants as vectors of pests”.

SOURCES

Crous, P.W., M.J. Wingfield, R. Cheewangkoon, A.J. Carnegie, T.I. Burgess, B.A. Summerell, J. Edwards, P.W.J. Taylor, and J.Z. Groenewald. 2019. Foliar pathogens of eucalypts. Studies in Mycology 94:125-298 (2019)

Lantschner, V. and J. Villacide. 2025. Invasion Potential of the Recently Established Woodwasp Sirex obesus. Neotropical Entomology. (2025) 54:117 https://doi.org/10.1007/s13744-025-01347-6

Paine, T.D., M.J. Steinbauer, and S.A. Lawson. 2011. Native & Exotic Pests of Eucalyptus: A Worldwide Perspective. Annu. Rev. Entomol. 2011. 56:181-201

Payn, T., J-M. Carnus, P. Freer-Smith, M. Kimberley, W. Kollert, S. Liu, C. Orazio, L. Rodriguez, L. Neves Silva, M.J. Wingfield. 2015. Changes in planted forests & future global implications. Forest Ecology and Management 352 (2015)

Pérez, C.A., M.J. Wingfield, N.A. Altier, and R.A. Blanchette. 2009. Mycosphaerellaceae & Teratosphaeriaceae associated with Eucalyptus leaf diseases & stem cankers in Uruguay For. Path. 39 (2009) 349–360 doi: 10.1111/j.1439-0329.2009.00598.x www3.interscience.wiley.com

Pérez, C.A., M.J. Wingfield, N. Altier, and R.A. Blanchette. 2013. Species of Mycosphaerellaceae and Teratosphaeriaceae on native Myrtaceae in Uruguay: evidence of fungal host jumps. Fungal Biology Volume 117, Issue 2, February 2013.

Villacide, J. and A. Fuetealba. 2025. Pests in plantations: Challenging traditional productive paradigms in the Southern Cone of America. Forest Ecology and Management 597 (2025) 123127

Posted by Faith Campbell

https://fadingforests.org

Pest Threats to Eucalypts and Australia

*Chilecomadia valdiviana* – one of the South American moths that attack Eucalyptus; photo by Natural History Museum of London via Wikimedia

Fifteen years ago, Paine, Steinbauer, and Lawson (2011) worried that insects in South America, Africa, Asia, and Europe that adapt to attacking Eucayptus trees planted there might be introduced to Australasia and threaten the genus in its native range. Their analysis applies to species in all three genera considered to be “eucalypts” — Angophora, Corymbia and Eucalyptus.

Some insects native to those continents have made this host shift already. Paine, Steinbauer, and Lawson reported that such host switching was especially prevalent among lepidopterans. They name several from Brazil, the Chilean cossid moth, Chilecomadia valdiviana, and southern African Coryphodema tristis. In their view, Brazilian eucalypt plantations’ proximity to native vegetation facilitates host-switching. Still, at that time they thought that there were no established pathways for introduction of the South American moths to Australia.

Host-switching is exceptionally common in Asia. Paine, Steinbauer, and Lawson (2011) thought the risk was greatest from insects on native eucalypts in near-neighbors Papua New Guinea, Timor, and The Philippines. An earlier risk assessment evaluating 10 insect species from the region concluded that most are polyphagous and probably switched to eucalypts. Two woodborers – Agrilus opulentus and A. sexsignatus –seem to have coevolved with Eucalyptus deglupta in New Guinea and The Philippines.

According to the same authors, most of the insects that have switched hosts are either polyphagous or normally feed on other myrtaceous species native to these regions. Thus, the Brazilian moth Thyrinteina arnobia feeds on Psidium guajava and several other Myrtaceae. Sarsina violascens is also a pest of Psidium species, as well as species in the Asteraceae, and Oleaceae. And the foliar rust Austropuccinia psidii was first described from Psidium guajava in Brazil and boasts a wide host range in the Myrtaceae in South America. It has been introduced to many regions with plants in the Myrtaceae, notably Hawai`i, Australia, South Africa, New Caledonia, and New Zealand. At least 15 Myrtaceae species in Australia are threatened with extinction.

Still, few non-native insects were damaging eucalypts in Australia’s native forests or plantations as of 2011. Those few are highly polyphagous. Several, if not most, were introduced in the first half of the 20^th Century.

Why so few? Paine, Steinbauer, and Lawson (2011) suggest three possibilities: (a) Australia’s diverse endemic insects already occupy most niches, so they exclude new, foreign competitors; (b) most introduced insects were not previously exposed to Myrtaceae in their native range; and (c) Australia has strong quarantine procedures aiming to limit introductions of non-native herbivores.

The fact that none of the introduced insects has adapted to feed significantly on mature eucalypts’ above-ground tissues seems to me to point to protection provided by the adult trees’ phytochemicals and leaf structure. Paine, Steinbauer, and Lawson (2011) discuss some aspects of leaf structure and wax coatings.

As to Australia’s quarantine procedures, as I reported before, the country has been much less proactive regarding plant pests and diseases that threaten tree species rather than agricultural crops. Significant new programs were established only after 2000, when Plant Health Australia (PHA) was incorporated. The PHA is supposed to facilitate preparedness and response arrangements between governments and industry for plant pests (once an alien pest has become established, management becomes responsibility of the land manager). In 2005, federal, state, and territorial governments and plant industry bodies signed a legally-binding agreement — the Emergency Plant Pest Response Deed (EPPRD). As of 2022, 38 were engaged. It sets up a process to implement management and funding of agreed responses to the detection of exotic plant pests – including cost-sharing and owner reimbursement.

Still, studies documented significant gaps in post-border forest biosecurity systems and the country’s response to the anticipated introduction of the foliar rust Austropuccinia psidii was disappointing. This prompted yet another initiative: development of the National Forest Biosecurity Surveillance Strategy (NFBSS) in 2018. The strategy was; accompanied by an Implementation Plan and appointment of a National Forest Biosecurity Coordinator. The forest sector fund a significant proportion of the proposed activities for the first five years. Still, Drs. Carnegie and Nahrung thought that in-country forest pest surveillance was still too fragmented.

Paine, Steinbauer, and Lawson (2011) consider the Asian spongy moths Lymantria dispar and Orgyia thyellina to pose serious threats. Five eucalypt species were assessed to be at risk of attack as are two preferred host oaks in Europe, Quercus pubescens and Q. robur. They note high volumes of imports from East Asia of containers, vehicles, and machinery, which are known to transport spongy moth egg-masses. It is not known whether the numerous natural enemies of Australia’s diverse lymantriid fauna [which includes four in the genus Lymantria] might provide some protection. These experts also worried that the highly polyphagous Asian longhorned beetle (Anoplophora glabripennis) might arrive in Australia. Eucalypts are not recognized as hosts.

Australia has adopted an enhanced surveillance program for ships arriving from Asian and European Lymantria ranges during female flight periods. Described here. Nahrung and Carnegie (2021) though that the high priority assigned to Lepidoptera exceeded the actual risk; only two non-native species had established in Australia over 130 years.

Paine, Steinbauer, and Lawson (2011) suggest several research topics aimed at reducing the risk to eucalypts in Australia. These include interactions between these insects and mechanisms by which insects adapt to new hosts; host chemistry and resistance mechanisms), chemical ecology (including host selection), population and community dynamics, including possible biocontrol agents, and pathway and risk analysis.

On the other hand, Carnegie and Nahrung (2019) called for developing more effective methods of detection, especially of Hemiptera and pathogens. They also promoted national standardization of data collection. Finally, they advocated inclusion of technical experts from state governments, research organizations and industry in developing and implementing responses to pest incursions. They noted that surveillance and management programs must expect and be prepared to respond to introductions of unanticipated species. They had found that 85% of the pests detected over the last 20 years—and 75% of subsequently mid-to high-impact species established—were not on high-priority pest list.

SOURCES

Carnegie A.J. and H.F. Nahrung. 2019. Post-Border Forest Biosecurity in AU: Response to Recent Exotic Detections, Current Surveillance and Ongoing Needs. Forests 2019, 10, 336; doi:10.3390/f10040336 www.mdpi.com/journal/forests

Nahrung, H.F. and A.J. Carnegie. 2021. Border interceptions of forest insects established in Australia: intercepted invaders travel early and often. NeoBiota 64: 69–86. https://doi.org/10.3897/neobiota.64.60424

Paine, T.D., M.J. Steinbauer, and S.A. Lawson. 2011. Native & Exotic Pests of Eucalyptus: A Worldwide Perspective. Annu. Rev. Entomol. 2011. 56:181-201

Native & Exotic Pests of Eucalyptus: A Worldwide Perspective

Posted by Faith Campbell

https://fadingforests.org

Bird nesting habitats – why no mention of invasive species or deer?

ovenbird (*Seiurus aurocapilla*); photo by Rhododentrities via Wikimedia

Studies of forest ecosystems in eastern North America that claim to be comprehensive still too often make no reference to invasive species – pests, earthworms, or plants. I try here to bridge these gaps.

Akresh et al. (2023) conducted a meta-analysis of bird species’ use of forests as nesting habitat. They applied the Partners-in-Flight to evaluate the community-wide bird conservation values of unmanaged forests compared to various levels of tree removal by harvest. Because of the decline of many bird species that prefer shrubland or early-successional stands, their process gave highest ranks to management approaches that retained 40%–70% of the canopy trees.

Their study notes that habitats for shrubland birds comprise only about 6% of forests in the eastern U.S. They don’t provide data for southeastern Canada. But hasn’t this scarcity of open upland, non-wetland, habitats in this region been true for thousands of years?

The type of forest that undoubtedly has shrunk significantly in recent centuries is “virgin” (or old-growth or late-seral) forests. As Akresh et al. (2023) report, contemporary closed-canopy forests in eastern North America are predominantly structurally homogeneous, mid-seral, even-aged, stands that have regenerated on land previously cleared for either agriculture or timber. These forests are much younger from a forest developmental perspective than precolonial forests; they lack the latter’s range of tree fall gap sizes and multiple age-classes. The tiny fraction of eastern forests that are in the late-seral stage might have higher species richness and conservation value for birds, but since they are usually not under management, Akresh et al. (2023) did not include that question in their analysis.

Akresh et al. (2023) list the bird species whose density appears to be closely linked to various tree canopy densities. For example, ovenbirds and brown creepers promptly decline in abundance in response to any amount of tree harvesting. Two other species — wood thrush and cerulean warbler — have declined steeply range-wide in recent decades. Nesting densities of three of these four species (excluding the warbler) are significantly higher in areas harvested in ways that retain a greater percentage of trees. Densities of another five bird species (Acadian flycatcher, hermit thrush, black-throated green warbler, and red-breasted nuthatch) are also higher in areas with a greater proportion of trees retained.

Another nine species had a more complex relationship with tree densities but still had lower densities in stands with low tree retention. These were blue-gray gnatcatcher, blue-headed vireo, blackburnian warbler, black-throated blue warbler, eastern wood-pewee, least flycatcher, red-eyed vireo, scarlet tanager, and yellow-bellied sapsucker. They found little relationship between bird density and tree retention for five putative mature-forest species (American redstart, great-crested flycatcher, hooded warbler, veery, and yellow-rumped warbler).

scarlet tanager (*Piranga olivacea*); photographed in scrub at Edwin B. Forsythe (Brigantine) NWR by F.T. Campbell

Akresh et al. (2023) claim that silviculture approaches can be used to restore aspects of the structural and compositional conditions found in old-growth forests to second-growth systems, providing a potential pathway for rapidly increasing the conservation value of these areas for bird species. They advocate reducing canopies moderately via variable retention harvests, shelterwood establishment harvests, and irregular shelterwood systems. This strategy can increase understory vegetation density, which they assert can then increase foraging and nesting opportunities for both many mature-forest bird species and many shrubland birds.

I am skeptical; it is much easier to create openings in the canopy than to “create” large trees supporting cavities and associated fauna and flora utilized by some bird species. The authors do advise managers that late-seral, unharvested stands can provide important habitat for old-growth-dependent taxa and any intensive forestry should also take into account other factors.

old-growth hemlock stand in Cook Forest State Forest, Pennsylvania; photo by F.T. Campbell

In addition, often the understory vegetation that responds to the more open environment will be invasive non-native plants. Already about half of eastern U.S. forests have been invaded by non-native plants (Oswalt et al. 2016; Kurtz 2023). Many of these are shrubs: honeysuckles, privets, roses, buckthorn. Management of these plants is difficult – especially when opening the canopy to allow light to reach the forest floor. (at www.nivemnic.us, scroll down to “categories”, click on “invasive plants”.) So the question arises, do the non-native plant species adequately substitute for native shrubs in providing resources needed by those birds?

Maybe. Gleditsch and Carlo (2014) found that a shrub layer dominated by non-native honeysuckle shrubs (Lonicera species) does support nesting populations of several common species, especially catbird (Dumetella carolinensis), American robin (Turdus migratorius ), and northern cardinal (Cardinalis cardinalis). However, they did not consider the species of concern to Akresh et al. (2023) – the rare species that prefer open-canopy, early-successional communities. So they do not inform us whether these high-priority species can utilize shrublands dominated by non-native species. Gleditsch and Carlo (2014) apparently did not find nests of several species considered to be associated with mature forests. So, again, these forests’ value for conservation remains unclear. Gleditsch and Carlo (2014) do counter earlier fears that these non-native shrubs are “traps” for nesting passerine birds. (The concern was that the plants’ structure facilitated nest raiding by predators.) They say, instead, that these plants’ effects are species-specific, context-dependent, and often a mix of both positive and negative outcomes.

invasive shrub honeysuckle; photo by Kevin Casper via public.domain.pictures.net

Akresh et al. (2023) also do not address the impact of browsing by super-abundant deer. Others (at www.nivemnic.us, scroll down to “categories”, click on “deer”.) have demonstrated that interactions of deer predation with invasive plants is especially damaging to native flora. Considering forests from Virginia to Maine, Miller et al. (2023) advise opening the canopy or subcanopy of forests to promote tree regeneration where deer and invasive shrubs overlap only where deer are controlled.

I have seen no recent analyses of the impact of widespread pest-caused tree mortality beyond some early efforts focused on eastern hemlocks and on high-altitude whitebark pines.

SOURCES

Akresh, M.E., D.I. King, S.L. McInvale, J.L. Larkin, and A.W. D’Amato. 2023. “Effects of Forest Management on the Conservation of Bird Communities in E North America: A Meta-Analysis.” Ecosphere 14(1):e4315. https://doi.org/10.1002/ecs2.4315

Gleditsch, J.M. and T.A. Carlo. 2014. Living with Aliens: Effects of Invasive Honeysuckles on Avian Nesting. PLOS One. September 2014. Volume Nine Issue Nine. E107120

Miller, K.M., S.J. Perles, J.P. Schmit, E.R. Matthews, M.R. Marshall. 2023. Overabundant deer and invasive plants drive widespread regeneration debt in eastern United States national parks. Ecological Applications. 2023;33:e2837. https://onlinelibrary.wiley.com/r/eap

Posted by Faith Campbell

https://fadingforests.org

A New Year …Will there be a new priority on countering invasive species?

Alaska yellow cedar (*Chamaecyparis nootkatensis*); one of the species vulnerable to *Phytophthora austrocedri*; APHIS has determined it is too late to try to slow its spread. Photo by Nucatum amygdalarum via Wikimedia

On 30 December 2025, US Department of Agriculture Secretary Brooke L. Rollins issued a Secretary’s Memorandum setting five new priorities for research and development. One is to protect agriculture from invasive species. Another is to resolve longstanding trade barriers due to sanitary and phytosanitary concerns.

The Secretary’s intention is to strengthen US agriculture to benefit both farmers and consumers. He justifies the action by claiming that President Lincoln’s original purpose in establishing USDA was to acquire and diffuse useful information on subjects connected with agriculture. According to this interpretation, Lincoln recognized that working to improve agriculture and secure the nation’s food supply would benefit everyone. The emphasis on research and development was reiterated by the almost simultaneous adoption of the Morrill Act of 1862, which created the system of land-grant universities and development of the Cooperative Extension System via the Smith-Lever Act of 1914.

The memorandum specifies five priority areas of research to be pursued by all USDA agencies and offices – to the maximum extent permitted by law and in accordance with any applicable regulations and procedural requirements.

Increasing Profitability of Farmers & Ranchers — especially reducing volatility in profitability. Goals include reducing inputs or increasing mechanization and automation.
Expanding Markets for US agricultural products. Two approaches are mentioned: generating science and data to resolve longstanding sanitary and phytosanitary trade barriers; and expanding use of agricultural commodities in novel biobased products and bioenergy.
Protecting the Integrity of American agriculture from Invasive Species. The memorandum lists four examples of current invasive pest and pathogen threats: new world screwworm in Mexico; continued westward expansion of spotted lanternfly; persistence of highly pathogenic avian influenza in poultry flocks; and citrus greening. It notes that invasive species threaten both agriculture and natural resources. The research is to focus on new and effective methods for preventing, detecting, controlling,and eradicating these threats.
Promoting Soil Health to Regenerate Long-Term Productivity of Land. The research is to promote soil health practices, increase water-use efficiency, & reduce the need for inputs.
Improving Human Health through Precision Nutrition and Food Quality. Research on “precision nutrition” is said to improve understanding of how healthy dietary patterns impact individuals. Research will also focus on increasing foods’ nutritional content and quality.

*Vaccinium myrtillus* (photo by Anneli Salo via WikiMedia); one of several species in genera shared with North America that are infected by *Phytophthora* spp in the Italian alps

The memorandum also instructs USDA’s Office of the Chief Scientist (that is, the Under Secretary for Research, Education, & Economics) to coordinate these priorities within USDA and among key partners in other federal agencies.

Does This Policy Mean Substantially Stronger USDA Efforts to Counter Bioinvasions?

Can we expect new energy in USDA’s programs aimed at managing non-native forest pests and invasive plants that damage forests, wetlands, grasslands, and other natural systems? The first paragraph of the memorandum states that it is USDA policy to reaffirm a focus on the Department’s original objectives of maximizing and promoting American agriculture; ensuring a safe, nutritious, and secure food supply; enhancing rural prosperity; and protecting our National Forests & Grasslands. That is promising.

The explicit recognition that invasive species pose severe threats to both agriculture and natural resources is also promising. I welcome the inclusion of two plant pests among the examples. Livestock diseases usually receive far more attention in USDA pronouncements.

I note three caveats:

The prominence of enhancing markets for US agricultural exports (# 2). In the past, this longstanding emphasis has led to undercutting phytosanitary agencies’ ability to counter suspected — but incompletely understood — pest risks. I discussed the impracticality of determining a newly detected species’ probable impacts in Chapter 3 of my report, Fading Forests II.
The memorandum makes no reference to implementing stronger sanitary or phytosanitary policies. In my view, the Animal and Plant Health Inspection Service has sufficient knowledge to support adoption of a more assertive regulatory stance with regard to both new introductions and spread within the country? Does the memorandum signal support for such a stance by high-ranking USDA officials?

These officials have often reminded APHIS that it is not a research agency. However, its staff do “methods development” and it funds considerable research through the Plant Pest and Disease Management and Disaster Prevention Programs – Section 7721 of the Plant Protection Act and a matching program for animal diseases.

The US Forest Service does have a research division – although the Trump Administration proposed its virtual elimination in early 2025. The Congressional appropriators have provided funding for USFS R&D – but those bills have not yet been enacted into law. I have complained for years that USFS R&D allocates too few resources (about 1% of the total budget) to research on introduced pests and disease pathogens. Might this new directive help fix this problem?

I hope the emphasis on protecting National Forests & Grasslands does not result in narrowing the types of invasive pests addressed.

Posted by Faith Campbell

https://fadingforests.org

New thinking on how non-native plants invade forests

It used to be thought that closed-canopy forests are resistant to bioinvasion because of the low light availability and relatively infrequent disturbance. Yet many are badly invaded! (On this site, scroll down past the Archives, choose “invasive plants” category.)

Nor is it just temperate forests in North America. Subtropical and tropical forests have also been invaded, as have the temperate forests of South America and, to a lesser extent, Europe. Temperate forests in Asia are less invaded; boreal forests very little (Fridley et al. 2025; see full citation at the end of this blog).

Fridley et al. (2025) have proposed a conceptual model to explain how this happens: “superinvaders” – a special class of woody plants – that achieve competitive dominance across a wide range of forest conditions. The “superinvaders” pose especially grave threats to native biodiversity because they use life-history strategies unlike those of early successional native species.

The result is that existing invasion and succession theories poorly predict which forests are most invasible and by which species. This failure undermines pest risk analyses and early detection.

Fridley et al. have raised lots of interesting ideas – some of which cannot yet be demonstrated by observations.

Temperate forests of North and South America are increasingly dominated by non-native deciduous and semi-evergreen shrubs and trees that combine fast growth rate in high light and high survivorship in forest interiors. These traits enable them to outcompete the native species. Many invaders also produce many more seeds than co-occurring native species. Similar traits are found in the successful non-native plants in subtropical and tropical forests.

Fridley et al. stress that shade tolerance alone does not endow the invasive plants with sufficiently large advantages in their competition with native species. The forest “superinvader” phenotype must combine this ability to persist in shade with high maximum growth rate and high fecundity when conditions become favorable for reproduction.

They explain the invader’s competitive advantage as the result of their experiencing relatively fewer carbon costs because of enemy release in the novel environment, recent environmental changes that alleviate some stress formerly present in the novel environment, or phylogenetic constraints on the local flora that limit natives’ resource-use efficiency. The non-native plant species enjoy this advantage regardless of whether they also possess other competitive mechanisms, e.g., production of allelopathic compounds, denser growth or shading, greater apparent quantum yield. However, Fridley et al. concede that they lack sufficient evidence to incorporate these other competitive mechanisms into their model.

Since any reduction in carbon costs will enhance both shade tolerance and growth rate when light levels are high, these “superinvaders” can outcompete native species in either situation.

To support these concepts, Fridley et al. note that increased abundance of invaders following disturbance is more pronounced in forests than other habitats. They suggest this is because of the much greater magnitude of change in light levels in forests than in open habitats such as grasslands.

They propose that an analogous situation applies to the presence or absence of mutualist microbial associations, although existing studies are insufficient to reach conclusions about the role of carbon allocation to mycorrhizae in the “superinvader” phenotype. The extent to which these forest invasions alter ecosystem-level carbon dynamics, especially soil processes and litter decomposition is also largely unknown.

Fridley et al. emphasize the role of carbon costs in driving both growth rate and whole-plant light compensation point. This point is defined as the light level at which carbon gain through photosynthesis balances carbon losses from tissue respiration (maintenance and growth) and turnover (shedding and loss from disturbance and herbivory).

To survive in low-light conditions, plants must minimize tissue respiration and turnover. The traits that enable those behaviors have been thought to prevent rapid growth and competitive dominance in high-light conditions. However, the “superinvaders” defy this trade-off by growing faster than most co-occurring native species when light is abundant. Fridley et al. say this is because the plants’ reduced carbon costs enhance competitive advantage in both shade and adequate light conditions.

Fridley et al. name several reasons why a native plant’s carbon costs might exceed those of an introduced species. First on the list is either herbivory or investment in defensive traits. Native plants might be challenged by rising abundance or consumption rates of native or introduced herbivores, such as deer or seed predators, that avoid the introduced species.

A second factor is that the non-native species expends fewer resources to sustain adaptations that confer resistance to other stresses, such as drought or freezing. If a long-standing stress is weakened by global change processes (e.g., atmospheric CO₂ levels, growing season duration, precipitation levels and seasonality, suppression of fire, atmospheric nitrogen deposition), a non-native plant that lacks defenses against that now-weakened stress will have a lower carbon cost and therefore an advantage. In some cases, the non-native species might benefit directly from these changes, e.g., droughts.

In some regions phylogenetic constraints have limited evolution of adaptive solutions to various biotic and abiotic stresses. This is most obvious on tropical oceanic islands. Fridley et al. report that native trees in Hawaiian montane rain forests are less energy-efficient conducting photosynthesis than are the invaders. However, this phenomenon also occurs on continents. Two continents’ floras might experience different climatic histories even when at they are at similar latitudes. For example, Eurasian woody species leaf out earlier and senesce later than North American trees and shrubs – possibly as a result of more predictable spring and autumnal climate across Eurasia. They name as one example Norwegian maple (Acer platanoides) in North America.

The future is uncertain

Fridley et al. consider enemy release to be a key factor in these shrub invasions of closed-canopy forests. Therefore, if enemy release decays over time because the introduced plant species accumulate pests, or the forest environment shifts to favor more stress-tolerant phenotypes of some native species, then the dominance of superinvaders will decline. If, on the other hand, resource enrichment continues, e.g., nitrogen deposition and elevated CO₂, the impacts of woody invaders – present or newly introduced – might continue to rise. The likelihood that additional introductions of more resource-efficient species will continue to damage floras of oceanic islands.

Implications for risk assessments and management

Fridley et al. warn that habitat-matching criteria might be unreliable predictors of forest invasiveness. Among several examples of species that are invasive in interior forest systems in a novel region that do not exhibit this trait in their native range is red oak (Quercus rubra). It is locally dominant in both natural and managed forests in central Europe while in North America, red oak struggles to regenerate in closed-canopy forests. They suggest that Q. rubra in Europe has escaped seedling pathogens present in its native range in North America.

red oak sapling in swampy forest in Virginia; photo by F.T. Campbell

Fridley et al. call for research on traits they have identified as important but that are rarely measured in invasion studies. These include rates of tissue loss and respiratory processes above- and below- ground, plant carbon allocation to tissues and processes, and the whole-plant light compensation points of native and invasive plant species.

The Fridley et al. hypothesis has been supported explicitly by Kinlock et al. (2025). This article says that consistent findings have been reported by earlier small-scale studies in U.S. forests.

I ask for your input on how well the Fridley et al. hypothesis explains shrub and tree invasions in American forests – including those on tropical islands! Is it helpful? Is APHIS incorporating these ideas into plant risk assessments? –

Fridley et al. take pains to reiterate the long-accepted importance of ornamental horticulture in explaining invasive plants’ entry and establishment. They do so in the context of concurring that ruderal traits are not universally advantageous; traits’ benefits depend on the landscape into which the species was being introduced.

SOURCES

Fridley, J.D., P.J. Bellingham, D. Closset-Kopp, C.C. Daehler, M.S. Dechoum, P.H. Martin, H.T. Murphy, J. Rojas- Sandoval, D. Tng. 2025. A general hypothesis of forest invasions by woody plants based on whole-plant carbon economics.

Posted by Faith Campbell

https://fadingforests.org

US imports continue falling … expected to sink further in 2026

Kevin Saville reports for the Journal of Commerce that containerized imports in 2025 are expected to be only 25.2 million TEUs, a decrease of 1.4% from 2024.

Declines are particularly large in the final months of 2025 since importers frontloaded their purchases to try to get ahead of the Trump Administration’s new tariffs. Imports for the first half of the year were up 3.6% compared with 2024 at 12.53 million TEUs. Thus, Saville’s sources expect November import levels to be 11.6% lower than in November 2024; December’s to be almost 13% lower.

Analysts expect the steeper decline to continue into the new year. Ben Hackett, of Hackett Associates, expects import volumes in the first four months of 2026 to be 10.3%, 8.5%, 16.8% , and 11% lower than the corresponding months a year earlier. The data source covers the ports of Los Angeles/Long Beach, Oakland, & Seattle & Tacoma on the West Coast; New York/New Jersey, the Port of Virginia, Charleston, Savannah, Port Everglades, Miami & Jacksonville on the East Coast; & Houston on the Gulf Coast. These are not all the maritime ports, but they are the major ones.

Another JOC reporter, Michael Angell, quoted several sources as saying they expect import volumes for all of 2026 to be flat or down 2% from 2025. Illustrating the reversal from past trends, The Port Authority of New York and New Jersey expects total container volumes in 2026 to be 8.5 million TEUs, a decline of about 2% from 2024. Since 2016, NY-NJ port container volumes have grown at an annual average of 4.2%.

As I have blogged before — see here and here — these swings in import volumes threaten to undermine programs intended to prevent introductions of wood-boring insects hitching rides in wood packaging material. While the higher volumes arriving from Asia in the first half of 2025 pose the most obvious risk, falling volumes reduce fee-based funding that support port inspectors. Another factor is the shift to suppliers other than China – primarily countries in Southeast Asia. Two beneficiaries of this shift are Vietnam and – at least initially – India. They have much better records of compliance with ISPM#15-mandated treatments for wood packaging link than does China.

A third JOC source reports that while US and European imports are down, trade volumes in Asia, Africa, the Middle East and Latin America are rising. I expect this growing trade to facilitate new pest introductions, although we will have to wait several years to see any data.

Posted by Faith Campbell

https://fadingforests.org

Threats to Spring

*Erythronium americanum* dominating herb layer in woods owned by the Institute for Advanced Studies, Princeton, in the 1970s; photo by F.T. Campbell

I fell in love with spring ephemerals in the woods of the Institute for Advanced Studies in Princeton. While the degree I was pursuing had no relationship to birding in the swamp, I spent a lot of time enjoying the woods. At that time, more than 50 years ago, the herbaceous layer was dominated by spring beauties (Claytonia virginica), trout lilies (Erythronium americanum), and violets (Viola species).

Beyond the beauty that delights us (or at least, me!), spring ephemerals are important ecologically. They support specialist pollinators and reduce nutrient losses at a time of year when vegetation cover is low and leaching and runoff rates high.

In the decades since I left Princeton, scientists and nature lovers have observed declines in native understory plant communities. These are predicted to continue due to invasion by plants and worms, worm blogs herbivore pressure by deer, E NPS blog, Blossey blog land use changes, and climate change.

Where I live, in the suburbs of the District of Columbia, these forces are clear. The formerly glorious riparian forests where I walk are overrun by invasive plants. The herb layer is dominated by Japanese stiltgrass (Microstegium vimineum) and – increasingly — lesser celandine (Ficaria verna = Ranunculus ficaria). (I found it interesting that Ficaria began taking over floodplain forests only in the last decades of the 20^th century, although it was introduced more than 100 years earlier.) Many invasive shrubs (Rosa multiflora, various Lonicera species. …) and vines (Ampelopsis sp, Orbiculatus, Lonicera japonica, Hedera helix …) compound the problem. While I am not sure whether most earthworms here are native or not, high deer populations certainly are a factor.

*Ficaria* invasion in Fairfax County, Virginia in 2023; photo by F.T. Campbell

So I rejoice that scientists are studying how one taxon of spring ephemerals, trout lilies – Erythronium species – are coping with individual and combined threats. Gutiérrez and Hovick (full citation at the end of this blog) investigated how two species of Erythronium performed in the absence of a leaf litter layer – with and without competition by Ficaria. They chose to manipulate leaf litter as a proxy for impacts from invasive earthworms and non-native shrubs, especially those with rapidly decomposing leaves. They refer to others’ studies focused on different spring ephemerals.

Gutiérrez and Hovick found that the absence of leaf litter reduced asexual reproduction (corm biomass) in both Erythronium albidum and E. americanum species by 30%. That is, the absence of leaf litter alone reduced the native plants’ performance. This is alarming because persistent leaf litter has been reduced across much of the deciduous forests of eastern North America as a result of action by invasive earthworms and the rapid decomposition of the leaves of most invasive shrubs.

Trout lilies’ performance declined even more when litter absence was coupled with direct competition from Ficaria. Under those conditions, corm biomass declined by 50%. Impacts by lesser celandine occurred despite these plants’ being smaller than counterparts in nearby woodlands. The reduced size of Erythronium corms was sufficient, in their view, to reduce the likelihood that Erythronium would flower to nearly zero. This has clear implications for the long-term population viability of Erythronium andtheir specialist pollinators.

Gutiérrez and Hovick conclude restoration of these floodplain forests’ herb layer must incorporate management strategies that not only reduce Ficaria’s presence but also restore leaf litter.

*Erythronium albidum* along Accotink Creek in Fairfax County, Virginia; photo by F.T. Campbell

Underlying Factors

Native spring ephemerals in eastern North America evolved to emerge through litter layers in early spring. The litter layers impose both costs and benefits. In response to shading by leaf litter, Erythronium produces larger petioles compared to same-sized leaves, thus reducing the proportion of resources allocated to building photosynthetic tissue. In these cases, the corms that both perpetuate the individual and carry out asexual reproduction are smaller.

On the other hand, leaf litter increases moisture retention and reduces frost damage by buffering soil temperatures. While these results were seen in their experiment, Gutiérrez and Hovick believe the benefits are greater in nature than demonstrated in the study using potted plants. Leaf litter also increases nutrient availability, directly by increasing supply and indirectly by facilitating fine root growth. In this context, they note that their experiment used litter composed of just two tree species — red oak (Quercus rubra) and red maple (Acer rubrum). This narrow sample probably failed to capture the varied properties of other tree species’ litter and associated microbial activity.

*Erythronium americanum* along Pohick Creek; photo by F.T. Campbelle

Plants in the Erythronium genus reproduce primarily asexually through producing runners that form corms. The parent corm and runners disintegrate before summer dormancy; the offspring corms persist. Some individuals do not reproduce asexually; they simply replenish their own corm.

The few previous studies give mixed results regarding lesser celandine’s impacts on co-occurring native herbaceous plants (see the summaries in Gutiérrez and Hovick). The authors do not explicitly say whether lesser celandine is usually associated with low litter levels, but that appears to be the implication. They do say that it is not clear whether lesser celandine drives leaf litter loss by altering soil physiochemistry and microbial activity. Or, rather, that it simply performs well when leaf litter is absent.

Where lesser celandine and Erythronium co-occur at high densities, the former’s biomass per square meter can be more than an order of magnitude higher than Erythronium. Gutiérrez and Hovick suggest that competition between the species is primarily belowground. They cite their finding that by the time Erythronium shoots matured, lesser celandine roots occupied most of the belowground pot volume. They expect belowground competition in forests to be even more pronounced because of accumulated lesser celandine root biomass.

Aboveground, the principal factor appears to be the necessity for trout lilies to grow longer petioles to raise their leaves above lesser celandine rosettes, perhaps starving leaf formation. Since leaves are the plant’s photosynthetic organ, this tradeoff could ultimately result in fewer resources returned to the corm for future growth and reproduction. Although Gutiérrez and Hovick also mention that lesser celandine competition might delay Erythronium emergence and flowering, they do not discuss that.

A factor not mentioned by Gutiérrez and Hovick is the probability that Ficaria verna is allelopathic. See the article by Kendra Cipollini listed as a source at the end if this blog.

one of the few floodplains in Fairfax County still dominated by native herbs – Pohick Creek in the Burke area of Fairfax County, Virginia. Note the prevalence of beech in the canopy and subcanopy! photo by F.T. Campbell

Details of Impaired Performance of Erythronium

At the time of senescence, Erythronium plants grown in pots with leaf litter were nearly twice as large as those grown in bare soil conditions. One-third of their offspring corms grew to be larger than the putative biomass threshold for flowering. Only 9% of corms of plants grown in bare soil and 2% (one individual) of those grown with lesser celandine did. As noted above, corms developed by Erythronium grown in the presence of Ficaria actually lost biomass. This is the basis for their conclusion that there would be almost no sexual reproduction the following year where litter was absent and lesser celandine present.

Gutiérrez and Hovick think the principle mechanisms by which leaf litter affects performance of Erythronium plants is by buffering temperature ranges and increasing moisture retention. Indeed, they found that daily temperature ranges and maxima of soil in pots with bare soil or lesser celandine plants were both higher than temperatures under leaf litter. Reducing temperature maxima could be especially important with the increasing frequency and intensity of late-spring heatwaves associated with climate change. Absence of leaf litter advanced trout lily’s shoot emergence, flower emergence, and petal opening by 14 or more days.

This change might expose the plants to increased risk of frost damage. These dynamics will be system-specific, especially with complications added by climate change. However, Therefore, Gutiérrez and Hovick encourage future research to explore species-specific litter effects on spring ephemerals.

Broader Implications

Their findings regarding these two species of spring ephemerals prompt Gutiérrez and Hovick to assert that negative impacts from invasive plant species might be especially underestimated in spring ephemeral communities due to the combination of their short period of annual aboveground activity and tendency towards long lives. Changes might be very subtle over short timeframes.

They add that it is important to learn the role different conditions might play in the futures of related species. The two species’ ranges largely overlap, but E. americanum extends into the extreme southeast and northeast, E. albidum into the prairie states. Although these species’ respond to loss of leaf litter and lesser celandine invasions in similar ways, the fact that E. albidum occurs in areas of higher soil moisture makes it more vulnerable to negative population-level impacts from lesser celandine invasions.

Note about additional threats

Most of the photos of Erythronium americanum in this blog were taken along a particular creek in Fairfax County, Virginia. Ficaria has just begun to invade this area (see photo above); deer are plentiful. These plants face another bioinvasion: beech leaf disease has arrived. Widespread mortality of the predominantly beech understory will presumably open areas to more light, probably spread of the extant invasive plants.

beech in Fairfax County, Virginia with symptoms of beech leaf disease; photo by F.T. Campbell

SOURCES

Cipollini, K. and K.D. Schradin. 2011. Guilty in the Court of Public Opinion: Testing Presumptive Impacts and Allelopathic Potential of Ranunculus ficaria”

Gutiérrez, R.G. and S.M. Hovick. 2025. Compounding negative effects of leaf litter absence and belowground competition from an invasive spring ephemeral on native spring ephemeral growth and reproduction. Biol Invasions (2025) 27:213 https://doi.org/10.1007/s10530-025-03668-4

Actions USDA Could Take to Better Protect Our Forests

ohia trees killed by ROD near Pahoa, Hawai`i; with JB Friday; photo by F.T Campbell … APHIS has not applied NAPPRA to this pathogen

As I have documented numerous times in these blogs, [see here, here, here, here, here, here, here and here] forests throughout the world are being reshaped by rising numbers of introduced, non-native pathogens. Once established, these diseases are nearly impossible to contain, much less eradicate.

While the worst effect of such bioinvasions is widespread mortality of host species, even “lesser” results produce significant changes in the impacted ecosystems.

I believe that the international phytosanitary “system” adopted by the World Trade Organization (WTO) and amended by the International Plant Protection Convention (IPPC) in the mid-1990s impedes efforts to prevent introductions of pathogens. These rules require unattainable levels of certainty about an organism’s impacts before it can be restricted. Scientists such as Haoran Wu and Kenneth Raffa have called for phytosanitary approaches that will be more effective because they are realistic, reflect the true level of threat, and the limits of current science. I agree and have repeated their calls.

How Well Is This “System” Keeping Pathogens At Bay?

If the world’s phytosanitary system worked well, we should be seeing fewer high-risk forest pathogens being introduced to new countries. Instead, examples abound of pests invading new ecosystems in the post-WTO/IPPC era: Austropuccinia psdii — detected in Hawai`i in 2005, Japan in 2009, Australia in 2010, China in 2011, New Caledonia and South Africa in 2013, Indonesia and Singapore in 2016, and New Zealand in 2017.
Phytophthora ramorum – 8 to 14 additional introductions to California after its initial detection.
Fusarium disease vectored by beetles in the Euwallaceae genus:
- Euwallacea fornicatus s.s.— detected in southern California in 2003, Hawai`i in 2007, Israel in 2009, in South Africa in 2012, in Australia in 2021, and in Argentina and Uruguay in 2023 and 2024 . The haplotype detected in South America and several European greenhouses differs from that established elsewhere.
- E. kuroshio detected in southern California in 2013; has spread to nearby Mexico
- E. interjectus detected in central California in 2024.
Boxwood blight fungus Calonectria pseudonaviculata — first detected in the Caucuses in 2010 and the US in 2011. Now established in at least 24 countries in three geographic areas: Europe and western Asia; New Zealand; and North America. Boxwood blight has caused rapid and intensive defoliation of native stands of Buxus sempervirens. Although disease was detected in United Kingdom in the mid-1990s, the causal agent was not determined until 2002.
Beech leaf disease caused by the nematode Litylenchus crenatae subsp. mccannii — detected near Cleveland, Ohio, in 2012. Has since spread east to the Atlantic Ocean, south to Virginia, north into Ottawa.
Phytophthora austrocedrii — detected in nurseries in Ohio and Oregon in 2024. Previously known from Argentina and in England and Scotland. At the latter location it is causing mortality of native Juniperus and introduced Cupressaceae. See here and here.

Most of these pathogens were unknown at the time they were discovered – because they were causing disease in the invaded ecosystems.

beech leaf disease symptoms in northern Virginia; photo by F.T. Campbell

In the Face of International Failures, How Can USDA’s APHIS Succeed?

When countries choose to prioritize preventing bioinvasions, they can impose more restrictive controls than those implemented by the WTO/IPPC system.

I urge USDA to more proactively use its authority to protect America’s plant resources. In particular, I urge USDA leaders to use the NAPPRA authority more effectively and quickly. This allows the agency to temporarily prohibit importation of plants that host potentially damaging pathogens. ). https://www.aphis.usda.gov/plant-imports/nappra

We Americans can’t protect our forests from pathogens without APHIS responding more promptly to recent detections of pathogens in North America and on Pacific islands. Recent events are not encouraging.

The agency did undertake an analysis of Phytophthora austrocedrii after it was detected in nurseries in two states. Unfortunately, in my view, APHIS and the states decided the pathogen was too widespread so they dropped any idea of regulating it. This was despite the apparent threat to junipers across the country. See here and here. P. austrocedri also attacks cypress trees, including Port-Orford cedar, Chamaecyparis lawsoniana. USFS scientists recently announced success in breeding POC trees resistant to a different pathogen.

There are no indications that APHIS will respond to detection of a new pathogen causing wilt disease in elms (Plenodomus tracheiphilus) recently discovered in Alberta, Canada. The pathogen is spread primarily through movement of infected plant material, including on asymptomatic material. Current U.S. regulations do not prohibit importation of plants or cut greenery in the Ulmus genus from Canada. Beyond the risk associated with elm material, I think it is probable that this pathogen also survives on plants in additional taxa, since it was formerly known for causing disease on citrus trees.

Although APHIS has classified Leptosillia pistaciae as a federal quarantine pest, I have learned of no response to detection of the pathogen on the native California shrub, lemonade berry (Rhus integrifolia), in 2019.

Rhus integrifolia – host of *Leptosillia pistaciae*

Has APHIS Changed its Practices in Response to Recent Detections?

We’ve known about gaps and weaknesses in APHIS’ approach for a long time. Here are specifics.

Has APHIS upgraded its attention to nematodes – as should have been prompted by detection of the beech leaf disease nematode (above) and as recommended by Kantor et al.?

Has APHIS changed any of its practices or policies in response to detection of plant and human pathogens associated with wooden handicrafts from countries other than China? Or wood pieces used for unanticipated purposes, e.g., to decorate aquaria? All 31 fungal taxa detected by one of these studies were viable despite having been subjected to various phytosanitary requirements.

USDA has no authority to regulate organisms that pose a risk to non-plant hosts, like us humans! Has APHIS contacted officials at the relevant agency?

Does APHIS respond to detections abroad when pests attack congeners of North American trees? I have blogged about several — see here, here, here and here — detected in Europe or Asia that attack cypress, magnolia, dogwood, Persea, and oaks. PestLens — an alert system created by APHIS — reported these.

How has APHIS incorporated the findings at various “sentinel garden” projects? And the wider implications of findings by Eliana Torres Bedoya and Enrico Bonello regarding findings on asymptomatic plants?

How is APHIS applying the impact assessment tools developed (for insects) by Ashley Schulz and Angela Mech? Has APHIS incorporated Kenneth Raffa’s advice about the strengths and weaknesses of various prediction tools?

I wonder whether APHIS has responded in any way to the rash of woodborer introductions on the west coast, including three species in the invasive shot hole borer complex and the Mediterranean oak borer. Has the agency explored the threat that the spotted poplar borer (Agrilus fleischeri) – another wood-boring beetle native to northern Asia – might pose to North American Populus species? Canada has twice intercepted the species on solid wood packaging material .

USDA APHIS is explicitly not a research agency. However, it claims that its decisions are science-based. In my view, this means APHIS has a responsibility to respond to scientific findings (such as those above) and to bring about research aimed at answering pertinent questions, e.g., those related to risks of pest introduction and establishment, effective detection and management technologies, etc.

APHIS has occasionally done this:

It established the NORS-DUC research facility to study what aspects of nursery management facilitate establishment of Phytophthora ramorum.
It enabled and participated in several studies of wood-borer introduction via wood packaging, including those by Robert Haack and colleagues (see blogs on this website under the category “wood packaging”).
It enabled and participated in a study of introduction pathways that included plants-for-planting – relying on 2009 data. (Liebhold et al. 2012)
Did APHIS support the study by Li et al. to evaluate the vulnerability of two oak and two pine species to 111 fungi associated with Old World bark and ambrosia beetles?

APHIS could do much more to determine whether North American trees are vulnerable to pathogens and arthropods detected on the congeners in trade partner countries. Opportunities include:

studying which North American species might be vulnerable to the growing number of the 38 new Phytophthora species detected overseas. This would be a monumental task: 216 species have been recognized in the genus. I have focused specifically on the 38 species detected by Jung, Brasier, and others in Vietnam and now the 18 Phytophthora species detected in the Alps. (I have already noted that APHIS and the states dropped any idea of regulating one of those species, P. austrocedrii).
Regarding P. ramorum specifically, scientists now recognize 12 genetic strains; 8 are in Southeast Asia, a ninth (EU2) in Europe. How likely is it that some of these will be introduced to the U.S.? Three strains are known to be established in western North American forests – NA1, NA2, and EU1.

In addition, new hosts continue to be identified. APHIS has pledged to update the host list annually. In the past I have criticized APHIS for not accepting hosts identified in the United Kingdom.

While APHIS is not well-funded, it has largely escaped budget slashing by “DOGE,” other Trump Administration cuts, and congressional decreases. Scientific expertise at the USDA Forest Service has been shrinking for decades (see Chapter 6). Now, loss of expertise has reached crisis levels. The result will be less capacity to assist APHIS in evaluating pest risks and research needs.

Earlier, I noted the importance of APHIS using its full NAPPRA authority. Unfortunately, the record is not encouraging here, either.

Since the agency gained this authority in 2011, it has adopted lists of species temporarily prohibited for importation only three times – in 2013, 2017, and 2021. I complained that the last action was tardy and provided insufficient protection to Hawai’i’s unique flora arising from multiple strains of the ‘ōhi‘a rust pathogen Austropuccinia psidii and here. Even worse, four years after promising to close the loophole that allowed continued imports of cut flowers and foliage – the most likely pathway by which the rust was introduced to Hawai`i, APHIS has not proposed the necessary rule.

Pathogens are more difficult to detect and manage than invasive insects. The “disease triangle” is complex! Numerous pathways are involved! But they also get less attention – and this reflects unwise decisions by agency leaders. I suggest that they should respond to this complexity by adding resources. Voglmayr et al. (full reference at the end of this blog) also called for more attention to pathogens. Kantor et al. noted that nematodes are also neglected.

Of course, I have repeatedly urged APHIS leadership to enhance enforcement of regulations governing imports of wood packaging. One suggestion is that it prohibit importation of Chinese wood packaging because of its 25-year record of not complying with – first – U.S. and Canadian regulations and – later – the international regulation known as ISPM#15.

Information Gaps Impede APHIS’ Domestic Program

I have criticized APHIS’ failure to find answers to several questions important to managing the sudden oak death pathogen, Phytophthora ramorum. Like the many questions listed earlier, these also need priority attention.

APHIS has regulated interstate movement of nursery stock to contain P. ramorum for over 20 years. I appreciate its creation of NORS-DUC. But it is also responsible for protecting natural systems in regions not yet invaded, e.g, in the East. APHIS should have studied these issues years ago, given the frequency with which pests spread nationwide via the nursery trade.

Other pathogen systems also have genetic variation that might be important in determining pest-host relationships. As of 2022, scientists had identified 43 haplotypes (genetic variants) of E. fornicatus s.s. worldwide, with the greatest diversity in several Asian countries (P. Rugman-Jones, pers. comm). Other species of plant pathogens also have several haplotypes.

Forests At Risk Outside of North America

North American forests are not alone in being besieged by non-native pathogens. Their numbers have been rising also in Europe and Oceania. The record is less clear in Africa, South America, and Asia.

Reports of tree pathogens in Europe began rising suddenly after the 1980s – admittedly 15 years before the WTO took effect. By 2012, more than half of infectious plant diseases in Europe were caused by introduction of previously unknown pathogens https://www.nivemnic.us/?p=5164

Antonelli et al. (full citation at the end of this blog) report that three previously undetected species of Phytophthora have been detected in European nurseries since 2016. Voglmayr et al. reported that the number of alien fungi in Austria increased 4.6-fold over 20 years. Eighty percent were plant pathogens. The introductory pathway was unclear for the vast majority. They note that differences in research efforts probably explain some discrepancies.

The ash decline pathogen, Hymenoscyphus fraxineus, has apparently been present in eastern Europe since the 1980s, so its spread has probably not been facilitated by the downsides of the WTO/IPPC system.

Other sources report recent introductions of insects to Europe. Musolin et al. reported that 192 species of phytophagous non-native insects had been documented in European Russia as of 2011. They included the emerald ash borer detected in Moscow in 2003. Some of these insects were probably introduced to Europe (outside Russia) before the WTO/IPPC system came into effect. Examples are two insects from North America that were detected in 1999 and 2000, respectively – the western coniferous seed bug, Leptoglossus occidentalis, which vectors a pathogenic fungus Sphaeropsis sapinea (=Diplodia pinea); and the oak lace bug, Corythucha arcuata.

Australia was slow to respond to detection of myrtle rust, Austropuccina psidii. Few federal resources were made available to study its impacts – although the Australian flora includes at least 1,500 species in the vulnerable plant family. Carnegie and Pegg said this experience demonstrated the need to integrate the work of agencies responsible for conservation of natural ecosystems with those determining and implementing phytosanitary policy. New Zealand initially responded more assertively, but also found little funding to support resistance breeding or even to track the rust’s spread.

The record is less clear regarding Africa, South America, and Asia.

Africa

Sitzia et al. expressed concern that bark and ambrosia beetles threaten to cause significant damage to tropical forests. Several factors contribute to these threats: the long history of plant movement between tropical regions; conversion of tropical forests that disturbs canopies, understory plant communities, and soils; and, generally, regions with fewer resources to prevent or respond to invasions.

In Africa, Graziosi et al. reported on the cumulative economic impact of invasive species and the continent’s limited capacity to prevent or respond to introductions. They don’t discuss whether pests attacking plantations of non-native trees followed those trees from their point of origin. They found that some introduced insects pose significant threats to native tree species. They mentioned the Cypress aphid, Cinara cupressi, which was attacking both native African cedar, Juniperus procera, and exotic cypress plantations. All the examples appear to have been introduced before the WTO/IPPC system took effect. All the examples appear to have been introduced before the WTO/IPPC system took effect.

*Cinara cupressi*; photo by Blackman & Eastop via Wikimedia

Graziosi et al. point out that South Africa plays a central role because it imports significant volumes of goods that can transport pests. At most immediate risk is South Africa’s highly diverse and endemic flora. For example Phytophthora cinnamomi is attacking native Proteaceae, which are important components of the unique Cape Floral Kingdom. Other pathogens are attacking native conifers in the Podocarpus genus, Ekebergia capensis (Meliaceae), and Syzygium trees. However, pests first introduced to South Africa often spread. Graziosi et al. name several insects and pathogens of Eucalyptus and the wood-boring pest of pine Sirex noctilio.

Pests in Asia

Available information about China is not definitive. The FAO reports that half of the most damaging forest pests are non-indigenous. They were estimated to occur over an area of 1.3 million ha and to kill over 10 million trees per year. However, the three tree-killing pests which receive the most attention are the pinewood nematode (Bursaphelenchus xylophilus), red turpentine beetle (Dendroctonus valens), and fall webworm (Hyphantria cunea). These were all introduced before the World Trade Organization was founded.

The FAO notes several non-native insects that attack native trees in India, but all were introduced decades before the World Trade Organization began. There is no discussion of tree pathogens.

Thu et al. report a growing number of pest outbreaks damaging plantations of non-indigenous trees in Vietnam. In most cases the pests are indigenous to the country. They report that almost nothing is known about pests that attack species in the highly diverse native forests.

The September 2025 meeting of the International Forest Quarantine Research Group (IFQRG) had a session devoted to the topic “Risk of international trade in plants for planting”. The specific presentations are titled

“Using molecular tools to elucidate the pathways of cryptic pests on plants for planting”
“Risk-based approach to the movement of germplasm into Australia: the luxury afforded to an affluent continent” (note my earlier blog criticizing Australian efforts re forest pests)
“Challenges in the validation of methods for detection of quarantine pathogen – P. ramorum”
“Challenges in surveillance and detection of quarantine fungal tree pathogens in European Union”
“Pathogens in trade and the risk of establishment – update”

I hope that some of these discussions begin to tackle the crucial questions I raised in this blog and earlier. Also, I hope IFQRG continues to explore these important questions.

As Wu and Raffa et al. have said, Earth’s forests cannot afford delay in finding solutions to the challenges posed by introductions of novel pathogens to naïve systems.

SOURCES

Antonelli, C.; Biscontri, M.; Tabet, D.; Vettraino, A.M. 2023. The Never-Ending Presence of

Phytoph Spp in Italian Nurseries. Pathogens 2023, 12, 15. https://doi.org/10.3390/pathogens12010015

Voglmayr, H., A. Schertler, F. Essl, I. Krisai-Greilhuber. 2023. Alien and cryptogenic fungi and oomycetes in Austria: an annotated checklist (2nd edition). Biol Invasions (2023) 25:27–38 https://doi.org/10.1007/s10530-022-02896-2

Posted by Faith Campbell

https://fadingforests.org