New Tool for Evaluating Insect Pests’ Possible Impacts: One Test Shows Great Potential for Identifying the Greatest Threats to our Forests

red spruce (Picea rubens) — the conifer at greatest risk; This grove is in Great Smoky Mountains National Park; photo by Famartin via Wikimedia Commons

Scientists have incorporated into the widely-used urban tree management tool, i-Tree, a tool to help predict the damage that an insect species little known in North America might cause to trees growing in a specific area if it is introduced. This tool is available to all here.

I rejoice that predictive tools are becoming widely available. The tool is obviously the result of a lot of work by participating scientists – who are listed below. I hope many of you will try it out! Perhaps you and your students can join efforts by the tool-development team, especially in analyzing insect species from Central America and Asia that have not yet arrived in North America? If you are interested in helping, contact Katheryn Thomas, Angela Mech, or Ashley Schulz; you can obtain their contact information by visiting their institution’s website. You might choose which insect species to evaluate by consulting your own or colleagues’ research, reviewing the refereed and grey literature, APHIS and CFIA interception databases, databases maintained by several countries, websites such as CABI, EPPO, etc.

The new tool might help create a more effective “early warning” system. Whether this happens depends on what others do now.  Anyone – perhaps a staffer of a federal or state agency, or a city tree manager, or an academic – can apply the tool to meet his/her own objectives. If a more effective national or continental “early warning” system is to be created, someone needs to set up a process for conveying the findings to responsible federal or state/provincial agencies or even the scientific societies, e.g., Entomological Society (and, in the case of beetles transporting associated fungi, American Phytopathological Society). Perhaps the most challenging issue is to find an entity willing to receive these communications, review their accuracy, and – at a minimum – make the results accessible to phytosanitary agencies, interested public, etc. One possible entity is “PestLens”, a web-based early-warning system maintained by APHIS. The project’s objective is to provide early-warning information and facilitate a prompt, coordinated, and appropriate safeguarding response. PestLens posts alerts once a month. These are visible to anyone who subscribes. However, it remains unclear how often APHIS and state agencies act on the notices. The North American Plant Protection Organization (NAPPO) also hosts an alert system, but it records only official notices, leading to some absurdities. (E.g., NAPPO reported Mexico’s designation of the invasive shot hole borers as quarantine pests – without mentioning that they are well-established in California because neither APHIS nor California Department of Food and Agriculture has designated the insects as officially regulated.)

Those applying the tool need to have some knowledge and access to a range of scientific resources (including, in my view, people who can check the accuracy of the data entered into the system). Users must have appropriate skills to conduct some research into the insect and what it feeds on. Information required for the tool includes the following:

  • taxonomic information for the insect (Order, Family, Genus, Species)
  • the feeding guild of the insect (i.e., foliovore, gall, reproductive, root, sap, wood)
  • climate in the native range of the insect (i.e., Tropical, Dry, Temperate, Continental, Polar)
  • native range of the insect (i.e., Afrotropical, Australasian, Indomalayan, Neotropical, Oceanian, Palearctic Asia, Palearctic Europe)
  • the host trees of the insect in its native range (scientific name [Genus species]). The tool warns participants to include the full range of potential tree hosts – by listing either all or a representative sample. The tool will use this information to estimate the evolutionary distance between known native hosts and potential North American hosts using comprehensive phylogenetic tree of plants.

Clearly, those using the tool have their work cut out for them! The tool does provide definitions, descriptors, and drop-down lists for most of the factors, including insect orders and families, tree genera, geographic origins, and climate types. Users are now anticipated to be employees of federal and presumably state agencies; academics – even students!—and others who have the capacity to research what an insect feeds on in its native range.

This tool is intended to predict the probability that an insect species of concern – either newly detected in the country or thought likely to invade based on port detections or other reasons — will become a high impact invader. I rejoice that they are inclusive – the tool can test the vulnerability of 50+ conifer species and 360+ hardwood species native to North America. Assuming the assessor can enter accurate information for the categories outlined above, the tool can then provide a list of probabilities for each relevant North American host tree.

The tool is based on the findings of two studies, Mech et al. and Schulz et al. (full citations at the end of this blog). I discussed these studies in earlier blogs. They were also incorporated into the broader effort to identify predictive traits carried out by Raffa et al. (full citations at the end of this blog) and discussed in a separate blog.  See the section titled “Potential” to see the exciting results of an application of the Mech et al. findings and methods. 

To develop the tool, project scientists synthesized data on traits and factors representing four types of drivers: (1) insect traits, (2) tree traits (especially those associated with host defenses), (3) the relatedness between the insect’s native and North American tree hosts, and (4) the relatedness between the non-native insect and North American insects on the same tree. They tested key hypotheses, e.g., defense free space and enemy release. The team tested the tool with researchers from USDA APHIS and Canadian Food Inspection Agency (CFIA), Northeast Plant Diagnostic Network, and National Invasive Species Council.

Norway spruce (Picea abies) — host of 30 of the 62 insect species analyzed in Uden et al.; photo by Marzena via Pixabay

The research group hopes this tool will stimulate development of a global database of insects which will utilize the results of basic research on phytophagous insects and what they eat. Basic research on insects native to North America is also important and can benefit other countries that might want to develop a similar tool for their own phytosanitary needs.

The Tool’s Potential

Many of the scientists who developed the i-Tree tool have participated in an analysis of the threat to North American conifer species posed by insects native to Europe that have not yet been introduced to North America (Uden et al.). They applied the methodology from Mech et al., which is comparable to, although not identical to, the i-Tree system. They (1) created a list of 62 European insect species that appear to pose a risk to 47 species of North American conifers; (2) identified and compared the predicted likelihoods of high-impact invasion under each of four phylogenetic systems datasets; and (3) evaluated risk and vulnerability trends among insects & conifer hosts, respectively. In total they evaluated 2,914 insect–novel host pairs.

Fraser fir (Abies fraseri) in Great Smoky Mountains National Park; photo by James St. John via Flickr

Among their findings are the following:

  • Of the 2,914 pairs examined, 302 (10.4%) had a predicted risk of high impact. These pairs included 41 (66%) of the insect species and 20 (41.7%) of the conifer species. The proportion of potential invasions posing a significant risk is higher than those indicated by earlier studies.
  • The insect species posing a risk of high-impact invasion were spread among insect orders, with relatively high levels concentrated in Lepidoptera and Coleoptera, fewer in the Hymenoptera and Hemiptera.
  • Consistent with Mech et al., they found a “Goldilocks” period of evolutionary divergence of hosts exposing the North American tree species to the highest risk. Thus, if a North American conifer shared a common ancestor with the insect’s native European host ~2–10 million years ago, it was predicted to be more vulnerable to a high-impact invasion by a conifer specialist.
  • North American fir (Abies) and spruce (Picea) species are more vulnerable to the introduction of European conifer-specialist insects than are pines (Pinus). [Mech et al. found that trees with high shade tolerance and low drought tolerance are more vulnerable. These traits also fit fir and spruce; but not pine.] The most vulnerable tree species was red spruce (Picea rubens).

Uden et al. also say Fraser fir (Abies fraseri) and Carolina hemlock (Tsuga caroliniana) are highly vulnerable to European insect species. They identified 17 high-risk insect species for Fraser fir. Of course, both are already severely depleted by non-native insect pests (Balsam woolly adelgid and hemlock woolly adelgid, respectively). They have also been identified by the Potter et al. “Project CAPTURE” process as having high priorities for conservation efforts.

I worry that fir and spruce are less important as timber species than pines; I hope this does not result in agencies and important stakeholders assigning this risk finding a lower priority.

Uden et al. assert that their study shows that this system can identify vulnerable tree species in the absence of information about which particular insect might invade. This information helps managers focus biosecurity and management program programs on protecting the most vulnerable tree species. However, 57% of the North American conifers (27 species) were found to be vulnerable under at least one of the insect-host pairs. To further set priorities, they suggest combining predictions from this analysis with USFS Forest Inventory and Analysis (FIA) data to identify vulnerable biogeographic regions and vegetation communities. (Fraser fir and Carolina hemlock rank high under this process.) Scientists could also apply species importance indicators, such as the NatureServe Explorer plant community descriptions. They suggest linking these criteria to the USFS Early Detection Rapid Response surveillance program, link to website which currently targets specific insect species.

red pine (Pinus resinosa) – the pine species at greatest risk; photo by Charles Dawley via Flickr

Uden et al. also warn that their analysis focused on a narrow range of possible introduced species: insects from Europe that feed on conifers exclusively. They caution that no one should assume that tree species that have a low “vulnerability” rank in this study should be considered at low risk for all possible introduced insects. They suggest researchers should identify tree species from the wider Palearctic that are within the high-impact “Goldilocks” zone of divergence times in relation to specific North American tree species, and then identify the insects that feed on those Palearctic trees to determine the species that would have the highest predicted risk of causing a high impact on those North American conifers.

Of course, many North American tree species are not conifers! Applying the methods in Schulz et al. – now integrated into the i-Tree tool – would facilitate similar predictive findings for the angiosperms.

Participants 

The importance of this project is seen in the impressive array of funders supporting it. They include:

  • U.S. Geological Survey John Wesley Powell Center for Analysis and Synthesis for a working group titled “Predicting the nest high-impact insect invasion: Elucidating traits and factors determining the risk of introduced herbivorous insects on North American native plants;”
  • USDA Forest Service National Urban and Community Forestry Advisory Council funded a working group titled “Forecasting high-impact insect invasions by integrating probability models with i-Tree from urban to continental scales”;
  • Nebraska Cooperative Fish and Wildlife Research Unit;
  • University of Washington;
  • USDA Forest Service Eastern Forest Environmental Threat Assessment;
  • National Science Foundation Long-Term Ecological Research program;
  • USDA Forest Service International Programs; and
  • USDA National Institute of Food and Agriculture (Hatch and McIntire-Stennis projects).

Scientists who created this tool:
Kathryn A. Thomas (USGS – Southwest Biological Research Center)
Travis D. Marsico (Arkansas State University)
Daniel A. Herms (The Davey Tree Expert Company)
Patrick C. Tobin (University of Washington)
Andrew Liebhold (U.S. Forest Service)
Nathan Havill (U.S. Forest Service)
Angela Mech (University of Maine)
Ashley Schulz (Mississippi State University)
Matthew Ayres (Dartmouth College)
Kamal Gandhi (University of Georgia)
Ruth A. Hufbauer (Colorado State University)

Kenneth Raffa (University of Wisconsin) Daniel

Uden (University of Nebraska-Lincoln)

Carissa Aoki (Maryland Institute College of Art)

Scott Maco (The Davey Tree Expert Company)

Angela Hoover (University of Arizona)

SOURCES

Mech, A.M., K.A. Thomas, T.D. Marsico, D.A. Herms, C.R. Allen, M.P. Ayres, K.J.K Gandhi, J. Gurevitch, N.P. Havill, R.A. Hufbauer, A.M. Liebhold, K.F. Raffa, A.N. Schulz, D.R. Uden, and P.C. Tobin. 2019. Evolutionary history predicts high-impact invasions by herbivorous insects. Ecol Evol. 2019. Nov; 9(21):12216-12230.

Potter, K.M., Escanferla, M.E., Jetton, R.M., Man, G., Crane, B.S. 2019. Prioritizing the conservation needs of United States tree species: Evaluating vulnerability to forest insect and disease threats. Global Ecology and Conservation (2019), doi: https://doi.org/10.1016/j.gecco.2019.e00622.

Raffa, K.F., E.G. Brockerhoff, J-C Gregoire, R.C. Hamelin, A.M. Liebhold, A. Santini, R.C. Venette, and M.J. Wingfield. 2023. Approaches to Forecasting Damage by Invasive Forest P&P: A Cross-Assessment.  BioScience Vol. 73 No. 2: 85–111    https://doi.org/10.1093/biosci/biac108 

Schulz, A.N.,  A.M. Mech, M.P. Ayres, K. J. K. Gandhi, N.P. Havill, D.A. Herms, A.M. Hoover, R.A. Hufbauer, A.M. Liebhold, T.D. Marsico, K.F. Raffa, P.C. Tobin, D.R. Uden, K.A. Thomas. 2021. Predicting non-native insect impact: focusing on the trees to see the forest. Biological Invasions.

Uden, D.R, A.M. Mech, N.P. Havill, A.N. Schulz, M.P. Ayres, D.A. Herms, A.M. Hoover, K.J. K. Gandhi, R.A. Hufbauer, A.M. Liebhold, T.D. M., K.F. Raffa, K.A. Thomas, P.C. Tobin, C.R. Allen. 2023. Phylogenetic risk assessment is robust for forecasting the impact of European insects on North American conifers. Ecological Applications. 2023; 33:e2761.

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

or

www.fadingforests.org

Let’s Not Dismiss Conservation of Biodiversity While Seeking Carbon Storage in Forests

red deer on farm in New Zealand; photo by Bernard Spragg via Flickr

Among the non-native species damaging forest systems are mammals – introduced deer, goats and sheep, and swine, … These animals have the greatest impacts on island systems that are sufficiently isolated that they have no native terrestrial mammals, e.g., Hawai`i and New Zealand. Several New Zealanders have published a study of their impacts (Allen et al.; full citation at end of the blog). The focus of their analysis is the native forests’ ability to sequester carbon and thus mitigate climate change. The scientists are well aware, however, that forests provide many other ecosystem values and services, including biodiversity, water supply and quality, etc.

Introduced ungulates can have many direct effects: reduction and damage to understory biomass, depletion of seedling regeneration, exacerbated soil erosion, and local nutrient imbalances. Mammals’ browsing can modify the composition of plant communities by favoring abundance of unpalatable species. Changes also can alter ecosystem functions associated with nutrient cycling, e.g., by reducing nutrient returns to the soil and altering rates of litter decomposition

In these ways, introduced ungulates exert long-term impacts on forests’ capacity to store carbon.

Allen et al. aimed to determine the extent of these effects on forests’ capacity to store carbon, both above- and below-ground, and on forest structure and diversity. The authors compared data from 26 pairs of sites across New Zealand – half with ungulate exclosures and half adjacent unfenced control plots. The ungulate exclosures had all been established for at least 20 years. All the sites were in species-rich communities of conifers and broadleaved evergreen angiosperm trees. These forests (1) cover about one-third of the country’s remaining mature natural forest; (2) contain tree species of a wide range of palatability to ungulate herbivores; and (3) have been named a conservation priority for forest carbon management. The ungulates present on the plots were European red deer (Cervus elaphus), fallow deer (Dama dama), sika deer (Cervus nippon), and feral goats (Capra hircus).

They assert that New Zealand is a good place to do this type of study because ungulate introductions are relatively recent so their impacts are well documented.

Allen et al. found that managing invasive ungulates makes valuable contributions to conserving biodiversity but not to carbon sequestration. They found little difference in total ecosystem carbon between ungulate exclosures and unfenced control plots. Most of the difference they did find was explained by the biomass of the largest tree within each plot. As they point out, these large trees have been unaffected by invasive ungulates introduced during the last 20–50 years. However, they believe ungulate-caused changes in understory biomass, species composition, and functional diversity might result in major shifts in the diversity and composition of regenerating species. Hence, longer term consequences for both ecosystem processes and storage of forest carbon storage can be expected.

Indeed, excluding ungulates did increase the abundance and diversity of saplings and small trees. The basal area of the smallest class of tree size was 70% greater. Species richness of small trees and saplings was 44% and 68% higher, respectively. This difference had little impact on overall carbon storage, however, because the small trees and saplings store only about 5%. In contrast, the largest tree size class (dbh =/>30 cm), with their roots, contributed 44% of total ecosystem carbon in both exclosure and control plots. The largest effects of exclosures on carbon stocks were in early successional stands, e.g., those affected by such major disturbances as windthrow, volcanic activity, or landslides.

Climate change is expected to cause surprising interactions among forest productivity, herbivory, disturbance. Allen et al. suggest that authorities should focus on excluding ungulates on these highly productive regenerating forests rather than old-growth forests. I am disturbed by this suggestion. It exposes the most biologically diverse forests to continuing damage.

Data gaps

New Zealand has many long-lived, slow-growing tree species. Recruitment of understory trees is already low across both main islands. This situation has been attributed to ungulate browsing. Over centuries, this might result in shifts in the canopy composition. Allen et al. call for additional research to increase our understanding of how browsing and other short-and long-term drivers affect the regeneration of large trees. Also, data on soil CO2 emissions needs better integration.

Australian brushtail possum; photo by Peter Firminger via Flickr

The study did not consider the impact of other introduced mammals, such as feral pigs (Sus scrofa), rodents, and Australian brushtail possum (Trichosurus vulpecula). The possum is known to damage New Zealand trees. The scientists did not explain this omission; I assume it might have been the result of either lack of resources to support a broader study or differences in management strategies – or both?

I note that the study also did not address the extent to which non-native pathogens threaten these large trees. In response to my query, Kara Allen said that their plots did not include many kauri (Agathis australis) trees, so the severe dieback disease caused by Phytophthora agathidicida did not affect their results. Naturally regenerating kauri is limited to a small area of warm temperate rainforests located at the top of the North Island. So kauri potentially play a relatively small role in terms of overall carbon stocks in New Zealand’s forests. On the other hand, Allen says that myrtle rust (Austropuccinia psidii) could have a major impact on New Zealand forests’ carbon storage. Trees in the host family, Myrtaceae, are ecologically important across both islands. Also, they comprise a large portion of overall forest carbon stocks (ranked in the top 5 largest families for above- and belowground biomass). An example is southern rata (Mterosideros umbellata), which are preferentially fed on by Australian brush possum.

southern rata; photo by Stan Shebs

Bernd Blossey, (free access!) who has long studied the role of high deer populations in North American forests, praises the study’s attempt to measure data, not just rely on models, and its inclusion of soil. However, he notes other limitations of the New Zealand study:

  • The small exclosures (20 x 20 m) are subject to edge effects. Some of Blossey’s exclosures occupy 2 hectares.
  • Twenty  years  is too short a time for analysis of such long-term processes as carbon sequestration and regeneration of slow-growing trees. Therefore, any results must be considered preliminary. Furthermore, no one recorded any differences in carbon sequestration of the paired plots at the time the exclosures were set up.
  • There’s no mention of possible impacts by introduced earthworms. 

Dr. Blossey recognizes that the current study’s authors cannot re-do actions taken decades in the past. Still, the data gaps reduce the value of the findings.

I conclude that uncertainties continue due to:  the long timelines of species’ regeneration and growth to full sizes; the requirement for large exclosures; the complexity of factors affecting carbon sequestration; and probably other influences.. Managers trying to maximize carbon sequestration are forced to act without truly knowing the best strategy or how their actions will affect the future. 

For more about invasive mammals’ impacts in U.S. forests, also see the study by USFS scientists, Poland et al. (full citation listed in sources). One can enter “mammal” in the search box for the on-line PDF.

SOURCES

Allen, K., P.J. Bellingham, S.J. Richardson, R.B. Allen, L.E. Burrows, F.E. Carswell, S.W.Husheer, M.G. St. John, D.A. Peltzer, M. Whenua. 2023.  Long-term exclusion of invasive ungulates alters tree recruitment and functional traits but not total forest carbon. Ecological Applications. 2023; e2836. https://onlinelibrary.wiley.com/r/eap

Poland, T.M., Patel-Weynand, T., Finch, D., Miniat, C. F., and Lopez, V. (Eds) (2019), Invasive Species in Forests and Grasslands of the United States: A Comprehensive Science Synthesis for the United States Forest Sector.  Springer Verlag. The on-line version as at https://link.springer.com/book/10.1007/978-3-030-45367-1

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

or

www.fadingforests.org

Global Overview of Bioinvasion in Forests

black locust – one of the most widespread invasive tree species on Earth; photo via Flickr

In recent years there has been an encouraging effort to examine bioinvasions writ large see earlier blogs re: costs of invasive species – here and here. One of these products is the Routledge Handbook of Biosecurity and Invasive Species (full citation at end of this blog). I have seen only the chapter on bioinvasion in forest ecosystems written by Sitzia et al. While they describe this situation around the globe, their examples are mostly from Europe.

Similar to other overviews, this article re-states the widely-accepted attribution of rising numbers of species introductions to globalization, especially trade. In so doing, Sitzia et al. assert that the solution is not to curtail trade and movement of people, but to improve scientific knowledge with the goal of strengthening biosecurity and control programs. As readers of this blog know, I have long advocated more aggressive application of stronger restrictions on the most high-risk pathways. Still, I applaud efforts to apply science to risk assessment.

Sitzia et al. attempt to provide a global perspective. They remind readers that all major forest ecosystems of Earth are undergoing significant change as a result of conversion to different land-uses; invasion by a wide range of non-native introduced species—including plants, insects, and mammals; and climate change. These change agents act individually and synergistically. Sitzia et al. give greater emphasis than other writers to managing the tree component of forests. They explain this focus by asserting that forest management could be either the major disturbance favoring spread of non-native species or, conversely, the only way to prevent further invasions. They explore these relationships with the goal of improving conservation of forest habitats.

Japanese stiltgrass invasion; photo by mightyjoepye via Flickr

Sitzia et al. focus first on plant invasions. They contend that – contrary to some expectations – plants can invade even dense forests despite competition for resources. They cite a recent assessment by Rejmánek & Richardson that identified 434 tree species that are invasive around Earth. Many of these species are from Asia, South America, Europe, and Australia. These non-native trees can drive not only changes in composition but also in conservation trajectories in natural forests. However, the example they cite, Japanese stilt grass (Microstegium vimineum) in the United States, is not a tree! Sitzia et al. note that in other cases it is difficult to separate the impacts of management decisions, native competitive species, and non-native species.

Sitzia et al. note that plant invasions might have a wide array of ecological impacts on forests. They attempt to distinguish between

  • “drivers” of environmental change – including those with such powerful effects that they call them “transformers”;  
  • “passengers” whose invasions are facilitated by other changes in ecosystem properties; and
  • “backseat drivers” that benefit from changes to ecosystem processes or properties and cause additional changes to native plant communities.

An example of the last is black locust (Robinia pseudoacacia). This North American tree has naturalized on all continents. It is a good example of the management complexities raised by conflicting views of an invasive species’ value, since it is used for timber, firewood, and honey production.

Sitzia et al. then consider invasions by plant pathogens. They say that these invasions are one of the main causes of decline or extirpations in tree populations. I applaud their explicit recognition that even when a host is not driven to extinction, the strong and sudden reduction in tree numbers produces significant changes in the impacted ecosystems.

American chestnut – not extinct but ecological role gone; photo by F.T. Campbell

Sitzia et al. contend that social and economic factors determine the likelihood of a species’ transportation and introduction. Specifically, global trade in plants for planting is widely recognized as being responsible for the majority of introductions. Introductions via this pathway are difficult to regulate because of the economic importance (and political clout) of the ornamental plants industry, large volumes of plants traded, rapid changes in varieties available, and multiple origins of trade. As noted above, the authors seek to resolve these challenges by improving the scientific knowledge guiding biosecurity and control programs. In the case of plant pathogens, they suggest adopting innovative molecular techniques to improve interception efficiency, esp. in the case of latent fungi in asymptomatic plants.

The likelihood that a pathogen transported to a new region will establish is determined by biogeographic and ecological factors. Like other recent studies, Sitzia et al. attempt to identify important factors. They name a large and confusing combination of pathogen- and host-specific traits and ecosystem conditions. These include the fungus’ virulence, host specificity, and modes of action, reproduction, and dispersal, as well as the host’s abundance, demography, and phytosociology. A key attribute is the non-native fungus’ ability to exploit micro-organism-insect interactions in the introduced range. (A separate study by Raffa et al. listed Dutch elm disease as an example of this phenomenon.)  I find it interesting that they also say that pathogens that attack both ornamental and forest trees spread faster. They do not discuss why this might be so. I suggest a possible explanation: the ornamental hosts are probably shipped over wide areas by the plant trade.

surviving elms in an urban environment; photo by F.T. Campbell

Sitzia et al. devote considerable attention to bioinvasions that involve symbiotic relationships between bark and ambrosia beetles and their associated fungi. These beetles are highly invasive and present high ecological risk in forest ecosystems. Since ambrosia beetle larvae feed on symbiotic fungi carried on and farmed by the adults inside the host trees, they are often polyphagous. Bark beetles feed on the tree host’s tissues directly, so they tend to develop in a more restricted number of hosts. Both can be transported in almost all kinds of wood products, where they are protected from environmental extremes and detection by inspectors. Sitzia et al. specify the usual suspects: wood packaging and plants for planting, as ideal pathways. These invasions threaten indigenous species by shifting the distribution and abundance of certain plants, altering habitats, and changing food supplies. The resulting damage to native forests induces severe alterations of the landscape and causes economic losses in tree plantations and managed forests. The latter losses are primarily in the high costs of eradication efforts – and their frequent failure.

Eucalyptus plantation in Kwa-Zulu-Natal, South Africa; photo by Kwa-Zulu-Natal Department of Transportation

Perhaps their greatest contribution is their warning about probable damage caused by invasive forest pests in tropical forests. (See an earlier blog about invasive pests in Africa.) Sitzia et al. believe that bark and ambrosia beetles introduced to tropical forests threaten to cause damage of the same magnitude as climate change and clear cutting, but there is little information about such introductions. Tropical forests are exposed to invading beetles in several ways:  

1) A long history of plant movement has occurred between tropical regions. Sitzia et al. contend that the same traits sought for commercial production contribute to risk of invasion.

2) Logging and conversion of tropical forests into plantation forestry and agriculture entails movement of potentially invasive plants to new areas. Canopies, understory plant communities, and soils are all disturbed. Seeds, insects, and pathogens can be introduced via contaminated equipment.

3) Less developed nations are often at a disadvantage in managing potential invasion. Resources may be fewer, competing priorities more compelling, or potential threats less obvious.

Sitzia et al. call for development of invasive species management strategies that are relevant to and realistic for less developed countries. These strategies must account for interactions between non-native species and other aspects of global environmental change. Professional foresters have a role here. One clear need is to set out practices for dealing with conflicts between actors driven by contrasting forestry and conservation interests. These approaches should incorporate the goals of shielding protected areas, habitat types and species from bioinvasion risk. Sitzia et al. also discuss how to address the fact that many widely used forestry trees are invasive. (See my earlier blog about pines planted in New Zealand.)

planted forest in Sardinia, Italy; photo by Torvlag via Flickr

In Europe, bark beetle invasions have damaged an estimated ~124 M m2 between 1958 and 2001. Sitzia et al. report that the introduction rate of non-native scolytins has increased sharply. As in the US, many are from Asia. They expect this trend to increase in the future, following rising global trade and climate change. Southern – Mediterranean – Europe is especially vulnerable. The region has great habitat diversity; a large number of potential host trees; and the climate is dry and warm with mild winters. The region has a legacy of widespread planting of non-native trees which are now important components of the region’s economy, history and culture. These include a significant number of tree species that are controversial because they are – or appear to be – invasive. Thus, new problems related to invasive plants are likely to emerge.

Noting that different species and invasion stages require different action, Sitzia et al. point to forest planning as an important tool. Again the discussion centers on Europe. Individual states set forest policies. Two complications are the facts that nearly half of European forests are privately owned; and stakeholders differ in their understanding of the concept of “sustainability”. Does it mean ‘sustainable yield’ of timber? Or providing multiple goods and services? Or sustaining evolution of forest ecosystems with restrictions on the use of non-native species? Resolving these issues requires engagement of all the stakeholders.

Sitzia et al. say there has recently been progress. The Council of Europe issued a voluntary Code of Conduct on Invasive Alien Trees in 2017 that provides guidelines on key pathways. A workshop in 2019 elaborated global guidelines for the sustainable use of non-native tree species, based on the Bern Convention Code of Conduct on Invasive Alien Trees. The workshop issued eight recommendations:

  • Use native trees, or non-invasive non-native trees;
  • Comply with international, national, and regional regulations concerning non-native trees;
  • Be aware of the risk of bioinvasion and consider global change trends;
  • Design and adopt tailored practices for plantation site selection and silvicultural management;
  • Promote and implement early detection and rapid response programs;
  • Design and adopt practices for invasive non-native tree control, habitat restoration, and for dealing with highly modified ecosystems;
  • Engage with stakeholders on the risks posed by invasive NIS trees, the impacts caused, and the options for management; and
  • Develop and support global networks, collaborative research, and information sharing on native and non-native trees.

SOURCE

Sitzia, T., T. Campagnaro, G. Brundu, M. Faccoli, A. Santini and B.L. Webber. 2021 Forest Ecosystems. in Barker, K. and R.A. Francis. Routledge Handbook of Biosecurity and Invasive Species. ISBN 9780367763213

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

or

www.fadingforests.org

Analysis of Methods to Predict a New Pest’s Invasiveness: Which Work Best Under What Conditions?

spotted lanternfly – could we have predicted its arrival? Its Impacts? Photo by Holly Ragusa, Pennsylvania Department of Agriculture

As readers of my blogs know, I wish to prevent introduction and spread of tree-killing insects and pathogens and advocate tighter and more pro-active regulation as the most promising approach. I cannot claim to have had great success.

Of course, international trade agreements have powerful defenders and the benefit of inertia. And in any case, prevention will be enhanced by improving the accuracy of predictions as to which specific pests are likely to cause significant damage, which are likely to have little impact in a naïve ecosystem. This knowledge would allow countries to can then focus their prevention, containment, and eradication efforts on this smaller number of organisms.

I applaud a group of eminent forest entomologists and pathologists’ recent analysis of widely-used predictive methods’ efficacy [see Raffa et al.; full citation at the end of this blog]. I am particularly glad that they have included pathogens, not just insects. See earlier blogs here, here, here, and here.

I review their findings in some detail in order to demonstrate their importance. National and international phytosanitary agencies need to incorporate this information and adopt new strategies to carry out their duty to protect Earth’s forests from devastation by introduced pests.

Raffa et al. note the usual challenges to plant health officials:

  • the high volumes of international trade that can transport tree-killing pests;
  • the high diversity of possible pest taxa, exacerbated by the lack of knowledge about many of them, especially pathogens;
  • the restrictions on precautionary approaches imposed by the World Trade Organization’s Sanitary and Phytosanitary Agreement (the international phytosanitary system) – here, here, and here.   
  • the high cost and frequent failure of  control efforts.
ash trees killed by emerald ash borer; Mattawoman Creek, Maryland; photo by Leslie A. Brice

The Four Approaches to Predicting Damaging Invaders

At present, four approaches are widely used to predict behavior of a species introduced to a naïve environment:

(1) pest status of the organism in its native or previously invaded regions;

(2) statistical patterns of traits and gene sequences associated with high-impact pests;

(3) sentinel plantings to expose trees to novel pests; and

(4) laboratory tests of detached plant parts or seedlings under controlled conditions.

Raffa et al. first identify each method’s underlying assumptions, then discuss the strengths and weaknesses of each approach for addressing three categories of biological factors that they believe explain why some organisms that are relatively benign, sparse, or unknown in their native region become highly damaging in naïve regions:

(1) the lack of effective natural enemies in the new region compared with the community of predators, parasites, pathogens, and competitors in the historical region (i.e., the loss of top-down control);

(2) the lack of evolutionary adaptation by naïve trees in the new region compared with long-term native interactions that select for effective defenses or tolerance (i.e., the loss of bottom-up control); and

(3) novel insect–microbe associations formed in invaded regions in which one or both members of the complex are non-native, resulting in increased vectoring of or infection courts for disease-causing pathogens (i.e., novel symbioses). I summarize these findings in some detail later in this blog.

Most important, Raffa et al. say none of these four predictive approaches can, by itself, provide a sufficiently high level of combined precision and generality to be useful in predictions. Therefore, Raffa et al. outline a framework for applying the strengths of the several approaches (see Figure 4). The framework can also be updated to address the challenges posed by global climate change.

Raffa et al. repeatedly note that lack of information about pests undercuts evaluation efforts. This is especially true for pathogens and the processes determine which microbes that are innocuous symbionts in co-evolved hosts become damaging pathogens when introduced to naïve hosts in new ecosystems.

Findings in Brief

Raffa et al. found that:

  1. Previous pest history in invaded environments provides greater predictive power than population dynamics in the organism’s native regions.
  2. Models comparing pest–host interactions across taxa are more predictive when they incorporate phylogenies of both pest and host. Traits better predict a pest’s likelihood of transport and establishment than its impact.
  3. Sentinel plantings are most applicable for pests that are not primarily limited to older trees. Ex patria sentinel plantations are more likely to detect pest species liberated by loss of bottom-up controls than top-down controls, i.e., most fungi and woodborers but not insect defoliators.
  4. Laboratory tests are most promising for pest species whose performance on seedlings and detached parts (e.g., leaves) accurately reflects their performance on live mature trees. They are thus better at predicting impacts of insect folivores and sap feeders than woodborers or vascular wilt pathogens.

Raffa et al. also ask some fundamental questions:

  • How realistic is it to expect reliable predictions, given the uniqueness of each biotic system?
  • When should negative data – lack of data showing a species is invasive – justify decisions not to act? Especially when there are so many data gaps?  
  • Who should make decisions about whether to act? How should the varying values of different social sectors be incorporated into decisions?

Raffa et al. identify critical areas for improved understanding:

1) Statistical tools and estimates of sample size needed for reliable forecasts by the various approaches.

2) Reliability, breadth, and efficiency of bioassays.

3) Processes by which some microorganisms transition from saprophytic to pathogenic lifestyles.

4) Procedures for scaling up results from bioassays and plantings to ecosystem- and landscape-level dynamics.

5) Targetting and synergizing predictive approaches and methods for more rapid and complete information transfer across jurisdictional boundaries.

I am struck by two generalizations:

  1. While most introduced forest insects are first detected in urban areas, introduced pathogens are more commonly detected in forests. I suggest that more intensive surveys of urban trees and “sentinel gardens” might result in detection of pathogens before they reach the forest.
  2. Enemy release is rarely documented as the primary basis for pathogens that cause little or no impact in their native region but become damaging in an introduced region. Enemy release appears generally more important with folivores and sap feeders than with woodborers.

Detailed Evaluation of Predictive Methodologies

white pine blister rust-killed whitebark pine at Crater Lake National Park; photo by F.T. Campbell
  1. Empirical assessment of pest status in previously occupied habitats

This is the most commonly applied method now, partly because it seems to follow logically from the World Trade Organization’s requirement that national governments provide scientific evidence of risk to justify adopting phytosanitary measures. The underlying assumption is that species that have caused damage in either their native or previously invaded ranges are those most likely to cause damage if introduced elsewhere. The corollary is that species that have not previously caused damage are unlikely to cause significant harm in a new ecosystem.

As noted above, Raffa et al. found that a species’ damaging activity in a previously invaded area can help indicate likely pest status in other regions. However, its status — pest or not — in its native range is not predictive. See Table 1 for numerous examples of both pests and non-pests. For example, Lymantria dispar has proved damaging in both native and introduced ranges. Ips typographus has not invaded new territories despite being damaging in its nature range and frequently being transported in wood. White pine blister rust is not an important mortality source on native species in its native range but is extremely damaging in North America.

Raffa et al. also note the importance of whether effective detection and management strategies exist in determining a pest’s impact ranking. Insects are more easily detected than pathogens; some respond to long distance attractants such as pheromones or plant volatiles. These methods can include insect vectors of damaging pathogens.

Re: the difficulty of assessing insect–microbe associations, they name several examples of symbionts which have caused widespread damage to naïve hosts: laurel wilt in North America; Sirex noctilio and Amylosterum areolatum around the Southern Hemisphere; Monochamus spp. and Bursaphelenchus xylophilus in Asian and European pines. Dutch elm disease illustrates a widespread epidemic caused by replacement of a nonaggressive native microorganism in an existing association with a non-native pathogen. Beech bark disease resulted from independent co-occurrence of an otherwise harmless fungus and harmless insect.

In sum, “watch” lists are disappointingly poor at identifying species that are largely benign in their native region but become pests when transported to naive ecosystems. Many of our most damaging pests are in this group. Raffa et al. note that this is not surprising because naïve systems lack the very powerful top-down, bottom-up, and lateral forces that suppress pests’ populations in co-adapted system. Countries often try to overcome this uncertainty by shifting to pathway mitigation and other “horizontal measures” – as I have often advocated. Raffa et al. emphasize that such approaches are costly to implement and constrain free trade.

  • Predictive models based on traits of pests and hosts

Predictive models provide the most all-encompassing and logistically adaptable of the forecasting approaches. Typically, models consider various components of risk, e.g. probability of transport, probability of establishment, anticipated level of damage.

The overriding assumption is that patterns emerging from either previous invasions or basic biological relationships can provide reliable predictions of impacts that might result from future invasions. However, Raffa et al. note that the models’ reliability and specificity are hampered by small sample sizes and data gaps.

They found that specific life history traits have proved to be more predictive of insect — and to a lesser extent fungal – establishment than of impact. Earlier studies [Mech et al. (2019) and Schulz et al. (2021)] found no association between life history traits and impacts for either conifer-feeding or angiosperm-feeding insects.   

Some traits of pathogens have been linked to invasion success, e.g., dispersal distance, type of reproduction, spore characteristics, and some temperature characteristics for growth and parasitic specialization. Raffa et al. say that root-infecting oomycete pathogens have a broader host range and invasive range than those that attack aboveground parts. Oomycetes that grow faster and produce thick-walled resting structures have broader host ranges. Phenotypic plasticity is also important. Raffa et al. say that those organisms that require alternate hosts can be limited in their ability to establish. However, they don’t mention that – once introduced — they can have huge impacts, as the example of white pine blister rust illustrates.

Raffa et al. say that phylogenetic distance of native and introduced hosts is more predictive for foliar ascomycetes than for basidiomycete and oomycete pathogens with broad host ranges. They suggest predictive ability can be improved by incorporating other factors, e.g., feeding guild. They note that the findings of Mech et al. and Schulz et al. (see links above) show the importance of both host associations with pests and phylogenetic relationships between native and naïve hosts for predicting impacts.

Geography is important: while there is a greater chance of Northern Hemisphere pests invading in the same hemisphere, this is not universal, as shown by Sirex (of course, the woodwasp is attacking hosts native to the Northern Hemisphere – pines).

Genomic analyses have been used more often with pathogens. There are two general approaches:

trees killed by chestnut blight; USDA Forest Service photo

1) Comparing the genomes of different species to identify the determinants associated with certain traits or lifestyles. For example, a post hoc analysis of the genus Cryphonectria could distinguish nonpathogenic species from the chestnut blight fungus C. parasitica.

2) Using genomic variation within a single species to identify markers associated with traits. Genome sequencing of a worldwide collection of the pathogens that cause Dutch elm disease revealed that some genome regions that originated from hybridization between fungal species contained genes involved in host–pathogen interactions and reproduction, such as enhanced pathogenicity and growth rate.

Raffa et al. point out that the growth of databases will facilitate genomic approaches to identify important invasiveness and impact traits, such as sporulation, sexual reproduction, and host specificity.

At present, Raffa et al. believe that models based on traits, phylogeny, and genomics offer potential for a rapid first pass to predicting levels of pest damage. However, assessors must first have a list of candidate pest species and detailed information about each. Plus there is still too much uncertainty to rely exclusively on the models.

  • Sentinel trees

Raffa et al. say that sentinel trees can potentially provide the most direct tests of tree susceptibility and the putative impact of introduced pests. Three types of plantations offer different types of information:

  1. In patria sentinels [= sentinel nurseries] = native trees strategically located in an exporting country and exposed to native pests. The intention is to detect problematic hitchhikers before they are transported to a new region. These plantings are useful for commodity risk assessment.  However, all the taxa associated with the sentinel trees must be identified to ascertain whether they can become a threat to plants in the new ecosystem.
  • Ex patria plantings [= sentinel plantations] = trees from an importing country are planted in an exporting country with the aim of assessing new pest–host associations. These plantings are most useful for identifying threats that arise primarily from lack of coevolved host tree resistance (i.e., loss of bottom-up control). They cannot predict the effects of lack of co-adapted natural enemies in the importing region (i.e., loss of top-down control). Plantings are thus more helpful in predicting impacts by pathogens and woodborers than folivores and sap feeders. However, ex patria plantings cannot predict pest problems that arise from novel microbial associations, or increased susceptibility to native pests.
  • Trees in botanic gardens, arboreta, large-scale plantations, and urban parks and yards can provide information on both existing native-to-native associations and new pest–host associations. Analyzing these plantings can be useful for studying host-shift events and novel pest–host associations. Again, all the taxa associated with the sentinel trees must be identified to ascertain whether they can become a threat to plants in the new ecosystem. Monitoring these planting have detected previously unknown plant–host associations (such as polyphagous shot hole borer and tree species in California and South Africa), and entirely unknown taxa. Pest surveillance in urban areas can also facilitate early detection, thereby strengthening the possibility of eradication.
PSHB attack on Erythrina caffri; photo by Paap

Sentinel tree programs are limited by 1) small sample sizes; 2) immature trees; and 3) the fact that trees planted outside their native range might not be accurate surrogates for the same species in native conditions. Some of these issues can be reduced by establishing reciprocal international agreements among trading partners; the International Plant Sentinel Network helps to coordinate these collaborations.

Botanic gardens and arboreta have the advantage of containing adult trees; this is important because pest impacts can vary between sapling to mature trees. However, they probably contain only a few individuals per plant species, usually composed of narrow genetic base.

Large-scale plantations of exotic tree species, e.g., exotic commercial plantations, comprise large numbers of trees planted over large areas with varied environmental conditions, and they stand for longer times. Still, they commonly have a narrow genetic base that might not be representative of wild native plants. Also, only a few species are represented in commercial plantations.

Raffa et al. report that experience in commercial Eucalyptus plantations in Brazil alerted Australia to the threat from myrtle rust (Austropuccinia psidii). However, in an earlier blog I showed that Australia did not act quickly based on this knowledge.

  • Laboratory assays using plant parts or seedlings

Laboratory tests artificially challenge seedlings, plant parts (e.g., leaves, branches, logs), or other forms of germplasm of potential hosts to determine their vulnerability. These tests are potentially powerful because they are amenable to experimental control, standardized challenge, and replication. They also avoid many of the logistical constraints of sentinel plantings. Finally, they can be performed relatively rapidly.

The key underlying assumption is that results can be extrapolated to predict injury to live, mature trees under natural conditions. The validity of this assumption depends on the degree to which exogenous biotic and abiotic stressors affect the outcomes. Raffa et al. report that environmental stressors tend to more strongly influence tree interactions with woodborers than folivores.

These assumption are more likely to be met by pathogens that infect shoots or young tissues, such as the myrtle rust pathogen Austropuccinia psidii, ash dieback pathogen Hymenoscyphus fraxineus, and the sudden oak death pathogen Phytophthora ramorum. 

The host range of and relative susceptibilities to insects is usually tested on twigs bearing foliage for defoliators and sap suckers; bark disks, logs, or branches for bark beetles, ambrosia beetles, and wood borers. These methods do not work as well for bark beetle species that attack mature trees in which active induced responses and transport of resins through established ducts are critically important.

The major advantages of laboratory tests is that they readily incorporate both positive (known hosts) and negative (known nonhosts) controls, can provide a range of environmental conditions, can be performed relatively rapidly, are statistically replicable at relatively low costs, and can test multiple host species and genotypes simultaneously. The ability to statistically replicate a multiplicity of environmental combinations and species is particularly valuable for evaluating relationships under anticipated future climatic conditions.

However, there are several important limitations. In testing pathogens, environmental conditions required for infection are often unknown. Choice of non-conducive conditions might result in false negatives; choice of too-conducive conditions might result in exaggerating the likelihood of infection. Results of tests of insect pests can vary depending on whether the insects are allowed to choose among potential host plants. Other complications arise when the pest being evaluated requires alternate hosts. In addition, seedlings are not always good surrogates for mature trees – especially as regards pathogens and bark, wood-boring and root collar insects. Folivores are less affected by conditions. Plus, the costs can be significant since they involve maintaining a relatively large number of viable and virulent pathogen cultures, insects, and candidate trees in quarantine.

Finally, although lab assays are well suited for identifying new host associations, results might not be amenable to scaling up to predict a pest’s population-level performance in a new ecosystem. Scaling up is especially problematic for those insect species whose dynamics are strongly affected by trophic interactions.

SOURCE

Raffa, K.F., E.G. Brockerhoff, J-C Gregoire, R.C. Hamelin, A.M. Liebhold, A. Santini, R.C. Venette, and M.J. Wingfield. 2023. Approaches to Forecasting Damage by Invasive Forest P&P: A Cross-Assessment.  BioScience Vol. 73 No. 2: 85–111    https://doi.org/10.1093/biosci/biac108  

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

or

www.fadingforests.org

Sooty Bark and Drippy Blight diseaes: “domestic invaders”?

sooty bark disease; photo by Vincet Gaucet via Bugwood

Recently, plant pathologists have paid more attention to pathogens — native to eastern North America — now killing trees on the West Coast. Years ago, the likelihood of such “domestic invaders” was hotly debated. However, these new detections provide examples of native micro-organisms that are apparently benign in the ecosystems in which they evolved, but that cause disease when moved to relatively nearby — but naïve — environments. I would argue that it is the absence of co-evolution of pathogen and host, not distance, that matters. In both cases, environmental stress on the trees appears to play a significant role. Such stress is expected to increase as the climate changes.

[In the past, these issues arose re: goldspotted oak borer – introduced to California from Arizona; and thousand cankers disease – introduced from Arizona to Western states and now to the East.]

Little is known about these diseases. So far, scientists have examined the pathogens’ impacts more thoroughly on plantings that are introduced to the location, so already outside of the forest biomes in which they evolved. Less attention has been paid to hosts native to the regions newly affected. I find this disturbing because I am most concerned about the possible impact to native ecosystems.

I will describe two such diseases. Both attack native tree species, not just the exotic trees described in research. Both were first detected in the Pacific Coast states in the late 1960s – i.e., more than 50 years ago. Why are they becoming prominent now – is it changes in the climate? Evolution? Just slow adaptation?

Example One – Sooty Bark Disease

Sooty-bark disease (Cryptostroma corticale) of maples (Acer spp.) is native to the Great Lakes region, where it causes no problems. However, it has been introduced to the West Coast, where all sources agree that the disease kills trees stressed by heat and drought. These stresses are expected to increase as the climate changes.  The disease is also a serious threat to human health. The fungus’ spores can cause serious pulmonary disease in humans. 

Sooty bark disease was detected in Pullman, Washington, near the Idaho border, in 1968. Now it is more widespread: it was detected in the Seattle area as of 2020 and the Sacramento area of California in 2019 [Curtis Ewing pers. comm.]. Sooty-bark disease has also spread to at least ten countries in Europe, ranging from the United Kingdom to Italy and Bulgaria.

Sources in Washington say hosts include several non-native species widely planted in the area – sycamore maples (Acer pseudoplatanus) [Chastagner], Norway maples (Acer platanoides), Japanese maple (A. palmatum), and horsechestnut (Aesculus hippocastanum) [Chastagner and Washington State University]. More troubling is the fact that several native tree species are also hosts. Big leaf maple (A. macrophyllum), and Pacific dogwood (Cornus nuttallii) are among the most significant. [Big leaf maple is a large hardwood tree in a region dominated by conifers. Pacific dogwood has already been decimated by the introduced disease dogwood anthracnose.] Sources in California add two other maples, these native to eastern North America, red (A. rubrum) and silver (A. saccharinum) maples.

bigleaf maples in Olympic National Park; NPS photo

The fungus initiates infection in a tree’s small branches, then spreads into the heartwood and both up and down the tree. It might also invade pruning wounds. The fungus grows more rapidly at higher temperatures. It is also facilitated by drought stress. When the fungus grows out to the bark, it causes the bark to blister; it then forms spore-forming structures. The spores are dispersed by wind. (Chastagner)

Washington State University’s Ornamental Plant Pathology division has called for more research on all aspects of the disease in trees:  

  • Distribution and spread of the disease;
  • Plant species susceptibility and host range;
  • Diagnostic methods and molecular approaches to improve diagnostic efficiency and capacity;
  • Pathogen life history and genetic diversity;
  • Factors that affect disease development and vulnerability (site, stress, age of host, etc.);
  • Potential of human mediated dispersal and vectors such as pruning tools;
  • Best management practices and worker protection.

So far, little has been done. Scientists in the Pacific Northwest have received a small amount of funding from the USDA Forest Service Forest Health Protection Emerging Pests program link + blog on appropriations to increase diagnostic services and to surveys trees elsewhere in the Puget Sound region.

Example Two – Bacterial Pathogen on Oaks

drippy blight disease on northern red oak; photo by Rachel Sitz, USDA Forest Service; via Bugwood

There is always concern about threats to oaks (Genus Quercus) because of their ecological, economic, and social importance. As Kozhar et al. point out, this genus is one of the most important groups of trees in many regions of the Northern Hemisphere. In North America specifically, oak forests compose a significant part of many forest ecosystems, especially in the East. California has 20 native species, Colorado has one. In addition, oaks are also often planted as shade trees in urban environments, which has resulted in movement of oak species to new geographic areas. There where they experience different environmental conditions, they might find new pests or alert us to the effects of climate change.

The bacterial oak pathogen Lonsdalea quercina is indigenous to the native range of northern red oak (Quercus rubra) in eastern North America. There, it does not cause disease in its co-evolved host. However, it has recently caused two outbreaks in the West – in California and Colorado. In the latter, trees themselves can die; in the former, acorns are damaged, threatening forest regeneration. Other Lonsdalea species have caused similar tree diseases in Europe.

California

In California, the bacterium has been present since at least 1967. It infects acorns of native oaks, including coast live oak (Q. agrifolia), Q. parvula (presumably the mainland subspecies, also named Q. p. var. shrevei),and interior live oak (Q. wislizeni).  The Morton Arboretum link says Q. parvula (presumably – again – the mainland subspecies) is currently threatened by sudden oak death (SOD). Coast live oak has also been highly affected by sudden oak death (SOD), DMF but it is not considered to be threatened. I expect – but sources don’t say – that the bacterium is affecting these species’ reproduction.

Various genotypes of L. quercina are randomly distributed across trees in both native and human-altered habitats and among all host species. Kohzar et al. say this is not surprising since all the host oak species are native to the region. Coast live oak is a major component of native forests and is also widely planted as a shade tree in residential areas. Furthermore, there has been no attempt to restrict the pathogen’s movement by adopting quarantines or other measures by phytosanitary agencies.

As a result, the inoculum can be moved across large distances by insects, birds, small mammals, and humans.

Bacterial pathogens can be associated with insects, relying on their feeding sites and other wounds to facilitate entry to the host’s tissues or for dissemination among hosts. In California, L. quercina might enter host tissue via wounds made by acorn weevils, filbertworms, and some cynipid wasps Kohzar et al.).

Colorado

The situation in Colorado is different. Significant dieback of exotic oaks planted in the state came to attention in the early 2000s. The hosts include non-native northern red oak, pin oak (Q. palustris), and Shumard oak (Q. shumardii). In Colorado, the bacterium causes “drippy blight disease” on the trees, not the acorns. The disease causes abundant ooze on symptomatic tissue. There has been a significant increase in tree mortality – with associated removal costs. The bacterium also has been found attacking Colorado’s one native oak, Q. gambelii; in this case, the pathogen attacks the acorns rather than the tree.

Due to small sample sizes, Kohzar et al. were unable to answer three key questions:

  • whether the Colorado L. quercina population comprises a new taxonomic species;
  • whether genetic variation in the bacterial populations are explained by the habitat (native or human-altered) or host; and 
  • whether the L. quercina infections on native Q. gambelii serves as an inoculum reservoir for planted Q. rubra hosts or vice versa.
gambel oak; photo by Dave Powell, USDA Forest Service (retired); via Bugwood

Surprisingly, despite its more recent emergence, the Colorado population of L. quercina has higher genetic diversity. Kohzar et al. suggest this might be due to repeated introductions of the bacterium on nursery stock brought in from the northern red oak’s native range in the East. [see below]

As in California, L. quercina infections are associated with insects, especially kermes scale (Allokermes galliformis). This insect does not travel long distances, which might help explain why the Colorado genotypes are limited to nearby trees, not dispersed randomly as in California.

However, kermes scale has been present in the state for far longer than the disease. The scale’s population spiked at the same time as the drippy blight outbreak was detected. Kohzar et al. could not determine whether the rise in scale populations and associated increase in number of entry points through feeding sites led to the increase of bacterial populations, or vice versa.

Kohzar et al. did determine that the Colorado populations of L. quercina were not introduced from California. They cannot explain the original introduction but think there might be continuing introductions from the native range of both northern red oak and L. quercina – the northeastern United States. They call for further studies to understand evolutionary relationships among L. quercina populations from different areas, including the native habitat of red oak in the East to clarify possible causes and sources of the recent outbreak of drippy blight in Colorado.

Role of Environmental Conditions

Kohzar et al. stress the importance of factors other than species’ introductions to new environments as the cause of emerging forest diseases. They say such other factors as changes in environmental conditions, new host-vector associations, cryptic disease agents (e.g., pathogens with a very long latency period or endophytes changing their behavior to pathogenic), hypervirulent strains of known pathogenic species, and/or newly emerging species of unknown origin as key factors leading to disease emergence in forest ecosystems around the globe.

In the case of L. quercina in California and especially Colorado, Kohzar et al. point to stress on the trees caused by new environmental factors, e.g., rapid climate change. [Of course, oaks from humid regions of eastern North America are already outside their natural habitat in much-dryer Colorado.] They support this conclusion by noting the simultaneous appearance of four new diseases caused by Lonsdalea species in different parts of the world during the 1990s and early 2000s. These were Lonsdalea quercina in Colorado; L. britannica on oaks in Great Britain; L. iberica in Spain; and L. populi bark canker on poplar species in Hungary, China, and Spain. All cause similar symptoms of drippy blight disease.

SOURCES

Chastagner, Gary. Professor, Plant Pathology, Washington State University. https://pnwhandbooks.org/plantdisease/host-disease/maple-acer-spp-sooty-bark-disease

Ewing, Curtis. Entomologist, CalFire. Pers. comm. March 2023.

Kozhar, O., R.A. Sitz, R. Woyda, L. Legg, J.R. Ibarra Caballero, I.S. Pearse, Z. Abdo, J.E. Stewart. 2023. Population genomic analysis of an emerging pathogen Lonsdalea quercina affecting various species of oaks in western North America. BioRxiv  https://www.biorxiv.org/content/10.1101/2023.01.20.524998v1

Washington State University Ornamental Plant Pathology https://ppo.puyallup.wsu.edu/sbd/

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

or

www.fadingforests.org

Eradicating Invasive Species: You need “social license” to succeed

spread of non-native conifers in mountains of New Zealand; photos by Richard Bowman; New Zealand government website

As those of us who want to “do something” to counter bioinvasions struggle to mobilize both the  resources and the political will necessary, I rejoice that more studies are examining what factors affect “social license” [= public approva] for such programs. One such study was recently published in New Zealand — Mason et al. (full citation at the end of the blog). New Zealand enjoys a greater appreciation of the uniqueness of its biology and awareness of invasive species’ impacts than the United States. However, their findings might provide useful guidance in the US and elsewhere.

Mason et al. sought to understand motivations of, and constraints on, those local groups responsible for controlling the spread of non-native conifers into New Zealand’s remnant native ecosystems. Non-forest ecosystems across much of the country are at risk of rapidly transforming into exotic conifer forests. For these reasons, authorities are pressing for timely removal of existing seed sources, that is, mature non-native conifer trees of several species. The blog I posted earlier apparently describes effects of conifer invasions in lowland ecosystems, whereas the Programme described here is focused on high-elevation systems.

The eradication effort in the study is the National Wilding Conifer Control Programme, establishedin 2016. A large increase in funding provided during the COVID-19 lockdown made it practical to try to eradicate seed sources from large swathes of vulnerable land. The Programme coordinates control efforts across the country, working across property and land-tenure boundaries. Landowners are expected to cover 20% of the cost of removing conifers from their land. Since removing all seed sources of high-risk conifer species from the landscape is key to achieving long-term goals, success is unlikely if significant seed sources are allowed to persist.

Mason et al. combined workshops, questionnaires, and site visits to gather data on particular aspects of this Programme. They found that social resistance, rather than lack of scientific knowledge, was often the main barrier to success in managing widespread invasive species. The authors do not address whether the fact that only 30 people provided information for their study might undermine the reliability of their findings.

map of conifer wilding sites; adapted from Wilding conifers – New Zealand history and research background, a presentation by Nick Ledgard at the “Managing wilding conifers in New Zealand – present and future” workshop (2003)

The authors suggest that the main benefit of scientific information might be to increase stakeholders’ support for management interventions — rather than to guide manager’ decisions about which strategies to pursue.  To support social license, invasive species research programs might need to focus not only on cost-effective control technologies and strategies, but – perhaps especially — the benefits (both tangible and intangible) of invasive species control for society.

Mason et al. found that people were motivated to combat conifer invasions by impacts with direct influence on humans or human activities (e.g., reduced water yield, damage to infrastructure from wildfires, reduced tourist activities due to landscape transformation) and also by impacts affect ecosystems (e.g.,  impacts on biodiversity, aquatic ecosystems and landscapes).  

People objected to control or eradication programs primarily because of social concerns. These included the unwillingness of landowners to participate and regulatory frameworks that had perverse incentives.

Mason et al. called for greater efforts by scientists to persuade stakeholders[p1]  to allow removal of “wilding” conifers from private land and development of more appropriate regulations. They found that forecasting models were particularly effective in persuading people to support these efforts. It seems to me that outreach teams might need “translators” to convert scientists’ findings to information that would be more useful by stakeholders.

The authors concede that the “wilding conifer” situation has unique attributes. First, invading conifers present a stark, easily seen difference between native and invaded ecosystems. Second, some – but not all—stakeholders appreciate the uniqueness of New Zealand’s biomes. Third, the impacts of conifer invasion are sufficiently well known that they can be described succinctly and accurately.

Do these unique attributes undercut the relevance of this study to North America? It is still true that ongoing support from local stakeholders (including landowners and community groups) influences the effectiveness or profitability of managing invasive species. .It is also true that groups’ varying values affect willingness to support the activities.

Mason et al. think through the issue of stakeholders’ conflicting perspectives on the value of particular invasive species and the values threatened by that invader. These can include ethical or safety concerns around management methods, particularly regarding toxins and genetic modification. Economoic costs are also a factor – especially if the landowner must pay all or some of them.

I find it interesting that the government simultaneously funded a 5-year research program to study various issues regarding the spread, ecosystem impacts, and control of wilding conifers. The result is the Mason et al. study discussed here. I wish the U.S. funded independent analyses of its invasive species programs!

Pinus contorta – the most rapidly growing Pinus introduced to New Zealand; photo by Walter Siegmund / Wikimedia

More Details, Policy Suggestions

Workshop attendees unanimously identified landscape impacts as a reason for controlling wilding conifers. This primarily concerned the loss of New Zealand’s visual heritage or cultural identity rather than loss of native species’ habitats. When the landowner was raised in Europe, these cultural or heritage values sometimes had the opposite effect, since they see conifer forests as important components of “natural” landscapes.

Currently, landowner funding and permission is required for conifer removal. Some individual landowners want to establish new forestry plantings. Some resist removal of existing forestry plantations (which provide income) and shelter belts (which provide shelter for livestock in high country landscapes). Some landowners were unwilling to pay their 20% of removal costs. Or they objected to certain conifer control methods—particularly helicopter spraying of herbicides. New Zealand’s regulatory process also requires years of negotiating to remove standing trees – further delaying any action. In theory, landowners who resist removal could be prosecuted under the Biosecurity Act. However, this approach has never been tried for removing wilding conifers.

Mason et al. suggested several changes in policy to overcome some of these barriers.

First, forestry consultants can “game” the wilding conifer “risk calculator” to obtain government approval to establish conifer plantations in high-risk environments. The authors suggest that authorities create a “liability calculator.” Under this system, landowners wishing to retain conifers on their land for whatever reason would be liable for any subsequent containment costs. However, developing such a tool requires more finely-scaled models of conifer spread.

Second, given the high costs of combatting invading conifers if seed sources are allowed to persist, they suggested that it might be more cost-effective for the control program to pay for plantation removal under New Zealand’s Emissions Trading Scheme.

Given the overwhelmingly social and regulatory nature of barriers to success, the primary role for scientific information is providing assessments of outcomes in the absence of wilding conifer control. Preferred messages were return-on-investment estimates and forecasts of ecosystem impacts, particularly relating to biodiversity loss, water yield reduction, and wildfire hazard. Forecasts were key to demonstrating that management interventions reduced future control costs and avoided environmental impacts which large sections of the community value (i.e. biodiversity loss, reduction in water yield and agricultural productivity, increased wildfire risks). Practitioners felt that forecasting models might also channel research toward areas of high uncertainty. Mason et al. recognize the difficulties presented by inherent complexity of ecological systems. However, they think “good practice” guidelines on forecasting are emerging.

The authors find that information content and presentation need to be tailored to the various audiences – most of whom lack experience in interpreting data from environmental forecasting models. They suggest that outreach materials focus on clear illustration of the tangible and intangible benefits of wilding conifer management rather than detailed explorations of scenarios. Participants suggested ways to improve the web tool to make it more accessible to a non-expert audience. 

Mason et al. mention aspects that require balancing, but don’t suggest criteria for making these choices. They say it is important to include all relevant stakeholders in invasive species management governance bodies. The absence of stakeholders with positive attitudes to wilding conifer invasions led to unanticipated external social resistance to the Programme. They recognize that including stakeholders with conflicting interests might obstruct the decision-making process.  Also, in areas where there has been success in containing conifers’ spread, people can’t see invading trees, so they don’t recognize the problem. They also note that existing data do not adequately recognize risks of spread from deliberately planted seed sources such as shelter-belts, plantations and amenity plantings. The authors do not discuss how to integrate these data into analyses and public outreach.

Finally, Mason et al. recognize that many other factors strongly influence stakeholders’ willingness to support invasive species control programs, especially the level of trust and strength of relationships between bioinvasion program staff and stakeholders.

Also, they suggest topics for future research: assessing how well forecasting models are integrated with  communications with stakeholders; how qualitative and quantitative research methods in different fields might support one another; and empirical tests to measure the relative effects on social license of a) involving stakeholders in developing models, b) using forecasts to assess the consequences of different management decisions and, c) the usefulness of different methods for incorporating scientific information in stakeholder engagement.

SOURCE

Mason, N.W.H., Kirk, N.A., Price, R.J. et al. Science for social license to arrest an ecosystem-transforming invasion. Biol Invasions 25, 873–888 (2023). https://doi.org/10.1007/s10530-022-02953-w

see also https://www.doc.govt.nz/nature/pests-and-threats/weeds/common-weeds/wilding-conifers/

Posted by Faith Campbell

What do YOU think about the role “social license” plays in US invasive species programs? 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

or

www.fadingforests.org

 

Help Fight for $$ to Protect Forests

Help Fight for Money to Protect Forests

This blog asks YOU!!! to support funding for some of the key USDA programs. This blog focuses on USDA’s Animal and Plant Health Inspection Service (APHIS). APHIS is responsible for preventing introduction of pests that harm agriculture, including forests; and for immediate efforts to eradicate or contain those pests that do enter. While most port inspections are carried out by the Department of Homeland Security Bureau of Customs and Border Protection, APHIS sets the policy guidance. APHIS also inspects imports of living plants.

Please help by contacting your members of the House and Senate Appropriations Committees. I provide a list of members – by state – at the end of this blog. APHIS is funded by the House and Senate Appropriations Subcommittees on Agriculture and Related Agencies. These Subcommittees have scheduled hearings on the topic and I’ve drafted written testimony for them. I expect CISP will be joined by additional members of the Sustainable Urban Forest Coalition in signing the testimony. You can add the crucial voice of constituent’s support.

I will blog soon about funding for USDA’s Forest Service (USFS) – I don’t yet have necessary information to suggest specific funding levels.

Your letter or email need be no more than a couple paragraphs. To make the case for greater funding, feel free to pick-and-choose from the information that follows. Your greatest impact comes from speaking specifically about what you know and where you live.

These are the specific dollar amounts we’d like you to ask for. The rationale for each is below.

Appropriations for APHIS programs (in $ millions)

ProgramFY 2022 (millions)FY 2023FY 2024 Pres.’ request Our ask
Tree & Wood Pest$61$63$64$65 M
Specialty Crops$210$216$222$222 M
Pest Detection$28$29$30$30 M
Methods Development$21$23$23$25 M

The Costs of Introduced Pests

Introduced pests threaten many forest products and services benefitting all Americans, including wood products, wildlife habitat, carbon sequestration, clean water and air, storm water management, lower energy costs, improved health, aesthetic enjoyment, and related jobs. Already, the 15 most damaging non-native pests threaten at least 41% of forest biomass in the “lower 48” states. In total, these 15 species have caused an additional annual conversion of live biomass to dead wood at a rate similar in magnitude to that attributed to fire (5.53 TgC per year for pests versus 5.4 to 14.2 TgC per year for fire) [Fei et al.; full citation at end of blog; see also earlier].

tanoaks killed by SOD; Oregon Department of Forestry photo

These pests also impose significant costs that are borne principally by municipal governments and homeowners. As more pests have been accidentally introduced over time, these costs have risen. A study published last year [Hudgins et al.] projected that by 2050 1.4 million street trees in urban areas and communities will be killed by introduced insect pests. Municipalities on the forefront include Milwaukee and Madison Wisconsin; the Chicago area; Cleveland; and Baltimore, Towson, and Salisbury, Maryland. Removing and replacing these trees is projected to cost cities $30 million per year. Additional urban trees – in parks, on homeowners’ properties, and in urban woodlands – are also expected to die and require removal and replacement.

Pathways of Introduction

Tree-killing pests are linked to the international supply chain. Many pests—especially the highly damaging wood-borers like emerald ash borer, Asian longhorned beetle, polyphagous and Kuroshio shot hole borers, and redbay ambrosia beetle—arrive in inadequately treated crates, pallets, and other forms of packaging made of wood. Other pests—especially plant diseases like sudden oak death and sap sucking insects like hemlock woolly adelgid—come on imported plants. Some pests take shelter, or lay their eggs, in or on virtually any exposed hard surface, such as steel, decorative stone, or shipping containers.

infested wood from a crate; Oregon Department of Agriculture photo

Wood Packaging

Imports from Asia have historically transported the most damaging pests, e.g., Asian longhorned beetle, emerald ash borer, redbay ambrosia beetle, and the invasive shot hole borers. For decades goods from Asia have dominated imports. As of February 2022, U.S. imports from Asia were running at a rate of 20 million shipping containers per year. A recent analysis [Haack et al.; see also here] indicates that at least 33,000 of these shipping containers, perhaps twice that number, are carrying a tree-killing pest. These facts have led scientists to project [Leung et al.] that by 2050, the number of non-native wood-boring insects established in the US could triple. Hudgins et al. say the greatest damage would occur if an Asian wood-boring insect that attacks maples or oaks were introduced. Such a pest could kill 6.1 million trees and cost American cities $4.9 billion over 30 years. The risk would be highest if this pest were introduced to the South – and U.S. southern ports are receiving more direct shipments from Asia after the expansion of the Panama Canal in 2016. https://www.nivemnic.us/?m=202207

After introduction of the ALB, APHIS acted to curtail further introductions in wood packaging from China. First – in 1998 – APHIS required China to treat its wood packaging. Second, it worked with foreign governments to develop the International Standard for Phytosanitary Measures (ISPM) #15. The U.S. and Canada began phasing in ISPM#15 in 2005 with full implementation in 2006. Under ISPM#15, all countries shipping goods to North America must treat their wood packaging according to specified protocols with the goal of “significantly reducing” the risk that pests will be present.

However, as I have often blogged [see blogs under “wood packaging” category on this site] ISPM#15 has fallen short. Haack et al. found that as recently as 2020, 0.22% [1/5th of 1 percent] of the shipping containers entering the U.S. were infested by a tree-killing insect. This equates to tens of thousands of containers harboring tree-killing insects.

Worse, the data indicate that our trade partners’ compliance with the rules has deteriorated; the “approach rate” of pest-infested wood packaging fell in 2005-2006, but has since gone back up.

The most troubling offender is China. Although required since 1998 to treat its wood packaging, China consistently has one of the highest pest approach rates: it was 0.73% [or ¾ of 1%] during the 2010- 2020 period. This is three times the global average for the period. Since China supplied 40.7% of U.S. imports in 2022 [Szakonyi], or 5,655,000 containers. Thus China alone might be sending to the U.S. 30,000 containers infested with tree-killing insects. These pests threaten our urban, rural, and wildland forests and reduce forest productivity, carbon sequestration, the rural job base, water supplies and quality, and many other ecosystem services. 

ISPM#15 falls short at the global level. The fact that a pallet or crate bears the mark indicating that it complies with ISPM#15 has not proved to be reliable.

You might ask your Member of Congress or Senators to ask APHIS what steps it will take to correct the problem of Chinese non-compliance. (Remind him or her that that the Asian longhorned beetle, emerald ash borer, and many other insects of so-far lesser impact were introduced in wood packaging from China.

Asian longhorned beetle

Remind them also that the Department of Homeland Security’s Bureau of Customs and Border Protection has twice enhanced its enforcement of wood packaging rules. In 2017 it began penalizing importers of non-compliant wood packaging under Title 19 United States Code (USC) §1595a(b) or under 19 USC §1592. In 2021, it incorporated the wood packaging requirements into its voluntary C-TPAC program.)  

You might also urge them to ask APHIS what steps it is taking at the global level to improve the efficacy of ISPM#15 – or to replace it if necessary to ensure that pests are not being introduced.

spread of beech leaf disease

Imported Plants (“Plants for Planting”)

Some pest types—especially plant diseases like sudden oak death and sap-sucking insects like hemlock woolly adelgid—come on imported plants. The U.S. imported about 5 billion plants in 2021 [MacLachlan]. Recent introductions probably via this pathway include several pathogens — Phytophthoras, rapid ʻōhiʻa death in Hawai`i, beech leaf disease (established from Ohio to Maine), and boxwood blight. Insects have also been introduced on imported plants recently; one example is the elm zigzag sawfly (present in North Carolina, Virginia, and New York and Ontario). https://www.nivemnic.us/?p=4115

An analysis of data from 2009 [Liebhold et al.] found that approximately 12% of plant shipments were infested by a pest. This pest approach rate is more than 50 times higher than the 0.22% approach rate for wood packaging. APHIS has adopted several changes to its phytosanitary system for imported plants in the decade since 2009. A few studies have been published, but they have focussed on insects and excluded pathogens. We have noted that pathogens continue to be introduced via the plant trade. Therefore, please ask your Member or Senators to ask APHIS to facilitate an independent analysis of the efficacy of the agency’s current phytosanitary programs to prevent introductions of pests on important plants, with an emphasis on introductions of plant pathogens.

APHIS is responsible for preventing spread of the SOD pathogen, Phytophthora ramorum, through trade in nursery plants. In recent years California has had few detections in nurseries and little expansion in forests – but the situation suggests that this good news is probably more the result of the drought than of program efficacy. In cooler, wetter conditions in Oregon and Washington, detections in nurseries and alarming detections in the forest or plantings continue.

In 2022, the APHIS SOD Program supported detection and regulatory activities in 25 states. P. ramorum was detected at 18 establishment, 12 of which were first-time detections. The California nursery regulatory program – which is funded by APHIS – saw reduced funding in 2022. We think these cuts are unwise since this year’s very wet winter will probably lead to a new disease outbreaks. Programs in Oregon and Washington continue to detect infestations in additional retailers brought in by plants bought from other nurseries. Washington responded to four separate “trace forward” incidents, one involving more than 160 residential sites. Clearly, the federal-state program is not succeeding in eradicating P. ramorum from nurseries. Please suggest that your Congressperson and Senators ask APHIS what steps it is taking to improve the efficacy of the SOD program.

SOD-infected rhodoendron on plants in Indiana; photo by Indiana Department of Natural Resources

In the East, P. ramorum was found in three of 65 streams sampled in 10 states in 2022 (reaching across the Southeast from Mississippi through North Carolina, plus Texas, Maryland, Pennsylvania, and Illinois). One stream is troubling: a first-time detection in South Carolina, with no obvious nursery source. Since stream sampling began, P. ramorum has been detected from eight streams in four states, Alabama, Mississippi, North Carolina, and now South Carolina. The pathogen has been present in some of these streams for more than 10 years.

Oregon faces particularly high risks. Three of the four known strains of P. ramorum are established in Oregon forests. One of them, the EU1 lineage, is more aggressive than the NA1 clonal lineage already present in forests. In addition, the EU1 strain might facilitate sexual reproduction of the pathogen, thus exacerbating Oregon’s struggle to contain the disease.

As we know, introduced pests do not stay in the cities where they first arrived — they spread! Often that spread is facilitated by our movement of firewood, plants, or outdoor household goods such as patio furniture.

The beech trees so important to wildlife conservation in the Northeast are under attack by two pathogens and at risk to an insect. Most alarming is the spread – in a dozen years! — of beech leaf disease DMF from Ohio to Maine. A leaf-feeding weevil is spreading south in eastern Canada. Please suggest that your Member or Senators to ask APHIS what steps it is taking to prevent the weevil’s introduction to the U.S.

‘Ōhi‘a trees make up 80% of the biomass of forests in both wet and dry areas of the Hawaiian archipelago. It is under attack by two diseases caused by introduced pathogens first detected in 2010. ‘Ōhi‘a forests support more threatened and endangered species than any other forest system in the U.S. They also play a uniquely important role in providing other ecosystem services, including water supplies.

Asking for the Money Pest Problems Deserve


To respond effectively to these pests and to the others that will be introduced in coming years, the key APHIS programs identified above must have adequate funds. The funding levels I request – and hope you will support – are lower than I would wish, but everyone expects the Congress to refuse significant increases in funding (see table at beginning of this blog).

The Tree and Wood Pests account supports eradication and control efforts targeting principally the ALB and spongy (= gypsy) moth. Eradicating the ALB normally receives about two-thirds of the funds. The programs in Massachusetts, New York, Ohio, and South Carolina must continue until eradication succeeds.

Oregon detected the EAB in 2022. Although the state and Portland have been preparing for a decade for this eventuality, there will still be significant impacts. Four percent of Portland’s street trees are ash – more than 9,000 trees. Young ash constitute three percent of young trees in parks. Loss of Oregon’s ash will also have severe ecosystem impacts. In Willamette Valley wetlands, ash constitutes up to 100% of the forest trees. Washington and California are also concerned. Indeed, the Hudgins study identified Seattle and Takoma as likely to lose thousands of ash trees. The numerous ash in riparian forests, windbreaks, and towns of North Dakota are also at risk since the EAB is established in South Dakota, Minnesota, and Manitoba.

APHIS manages damaging pests introduced on imported plants or other items through its Specialty Crops program. The principal example is its efforts to prevent spread of the SOD pathogen through the interstate trade in nursery plants. We noted above that this program is not as successful as it should be. We support the Administration’s request for $222 million; however, you might suggest that your Member or Senator urge APHIS to allot adequate funding under this budget line to management of SOD, rapid ʻōhiʻa death pathogens in Hawai`i, and beech leaf disease and elm zig-zag sawfly in the East.

The Pest Detection program is key to the prompt detection of newly introduced pests that is critical to successful pest eradication or containment. The “Methods Development” program enables APHIS to improve development of essential detection and eradication tools.

The Administration’s request include a $1 million emergency fund. This is far below the level needed to respond when a new pest is discovered. Funding constraints have hampered APHIS’ response to past pest incursions.

Please note that many of the members of the Agriculture Appropriations Subcommittee are from states where non-native pests are probably not top of mind. It is important that everyone that knows about these threats communicate with your Member/Senators!!

Members of House or Senate Subcommittees that Fund APHIS

(Names of Senators are italicized)

STATEMEMBERAPHIS APPROPHOUSESENATE
AKLisa Murkowski  X
ALJerry Carl Katie BrittXX  X
CalifBarbara Lee David Valadao Josh Harder Diane FeinsteinX X   XX X X        X
FLDebbie Wasserman Scultz Scott FranklinX XX X 
GASanford BishopXX 
IDMike Simpson X 
ILLauren UnderwoodXX 
KSJerry MoranX X
KYMitch McConnellX X
LAJulia Letlow Ashley HinsonX XX X 
MDAndy Harris Chris Van HollenXX    X
MEChellie Pingree Susan CollinsX XX  X
MIJohn Moolenaar Gary PetersX Xx  X
MNBetty McCollumXX 
MSCindy Hyde-SmithX X
MTJon Tester Ryan ZinkeX    XX
NBDeb Fischer  X
NDJohn HoevenX X
NMMartin HeinrichX X
NVMark Amodei X 
OHMarcy KapturXX 
ORJeff MerkleyXXX
PAGuy ReschenthalerXX 
RIJack Reed  X
TXMichael Cloud Jake EllzeyXX X 
UTChris Stewart X 
VABen ClineXX 
WADan Newhouse Derek KilmerXX X 
WVShelly Moore Capito Joe Manchin  X X X
WIMark Pocan Tammy BaldwinX XX  X

SOURCES

Fei, S., R.S. Morin, C.M. Oswalt, and A.M. 2019. Biomass losses resulting from insect and disease invasions in United States forests. PNAS August 27, 2019. Vol. 116 No. 35  17371–17376

Haack R.A., J.A. Hardin, B.P. Caton and T.R. Petrice .2022. Wood borer detection rates on wood packaging materials entering the United States during different phases of ISPM#15 implementation and regulatory changes. Front. For. Glob. Change 5:1069117. doi: 10.3389/ffgc.2022.1069117

Hudgins, E.J., F.H. Koch, M.J. Ambrose, and B. Leung. 2022.  Hotspots of pest-induced US urban tree death, 2020–2050. Journal of Applied Ecology

Leung, B., M.R. Springborn, J.A. Turner, and E.G. Brockerhoff. 2014. Pathway-level risk analysis: the net present value of an invasive species policy in the US. Front Ecol Environ 2014; doi:10.1890/130311

Liebhold, A.M., E.G. Brockerhoff, L.J. Garrett, J.L. Parke, and K.O. Britton. 2012. Live Plant Imports: the Major Pathway for Forest Insect and Pathogen Invasions of the US. Frontiers in Ecology.

MacLachlan, M.J., A. M. Liebhold, T. Yamanaka, M. R. Springborn. 2022. Hidden patterns of insect establishment risk revealed from two centuries of alien species discoveries. Sci. Adv. 7, eabj1012 (2021).

Szakonyi, M. 2023. Sourcing shift from China pulls US import share to more than a decade low.

One State’s Program Illustrates Importance of Federal Funding

Dead ash along Mattawoman Creek in 2019; Mattawoman Creek is a Maryland tributary of the Potomac River, hence of the Chesapeake Bay. Photo courtesy of Leslie A. Brice

In this blog I describe one state’s forest health efforts – Virginia. The pertinent lesson is the importance of external funding, especially that provided by USFS Forest Health Protection program, in supporting states’ efforts. Is your state’s forest health program as dependent upon federal funding as Virginia’s is? If so, there is a role for everyone: lobby your Congressional representative and senators to increase funding for this program!

I have based most of this blog on the Virginia Department of Forestry’s annual report for 2022.

Forests grow on more than 16 million acres in Virginia, or 62% of the Commonwealth’s land area. Eighty percent of these forests are hardwood or hardwood-pine. They break down as follows: 61% oak-hickory; 11% oak-pine; 5% bottomland hardwood; and 2% maple-beech-birch. A fifth of the forest is pine, composed of pine plantation (14%) and natural pine (7%). The long term trend is growth, especially among hardwoods.

The report devotes much of its attention to the agency’s programs to advise private landowners (individuals own 80% of the Commonwealth’s forestland); fire management (including prescribed burns); and state and federal conservation programs (e.g., easements). A major program shares reforestation costs on harvested pine lands. In 2022, this program assisted reforestation practices on 74,702 acres. Virginia has an impressive tree-raising program. VDOF grows more than 40 species, including longleaf and shortleaf pine, several spruce species, and dozens of hardwoods. The aim is to provide stock suited for the Commonwealth’s soils and climate. Many of the hardwood species are grown from acorns and seeds collected and donated by volunteers.

VDOF also helps to protect and improve the Commonwealth’s water quality through tree planting and sound forest management. VDOF has an unusual responsibility: enforcing the Virginia Silvicultural Water Quality Law.

The report also summarizes several urban and community forestry programs focused on education, community engagement, tree selection, and grants for tree planting to ensure canopy retention & management.

Forest Health – Importance of Federal Funding

Spongy Moth

Slightly over 1 million acres was mapped by aerial surveys in FY22. I believe the funding for these surveys came largely from the USFS. The surveys detected heavy to moderate defoliation by the spongy moth on 24,493 acres (almost twice the area detected in FY21). The spongy moth infestation is primarily in counties on the western side of the state, in the mountainous region, which has the highest densities of oaks and other hardwoods.

Spotted Lanternfly

The spotted lanternfly (SLF) was detected in Virginia early – in 2018 in Winchester at the northern end of the Shenandoah Valley. Winchester is connected to central Pennsylvania by Interstate 81, so rapid movement of SLF to Virginia from outbreaks slightly to the east of I-81 in Pennsylvania doesn’t surprise me. SLF has been spreading south along the mountains and over the Blue Ridge to Loudoun and Fairfax counties (in 2022). Fairfax County has announced a four-year, $200,000 effort to try to slow SLF spread by eradicating high densities of its preferred host, Ailanthus, from two county parks in the far south and north ends of the county. Ailanthus removal requires not just cutting the trees, but applying herbicide to prevent sprouting from the roots. This work is funded by the county, the local park authority and a $20,000 grant from the regional energy company, Dominion Energy Charitable Foundation.

Emerald Ash Borer

Virginia has six species of ash: white and green (both common), and smaller populations of black, blue, pumpkin and Carolina. EAB is now confirmed in 84 counties – most of the Commonwealth except the far southeast. The Department of Forestry treats 130 – 150 trees per year – half or more on state lands. At least in FY21, the funding came from federal sources. The report also notes outreach efforts at two minor league baseball games. Virginia recently adopted a priority of protecting the Chesapeake Bay watershed by promoting tree planting in riparian forest buffers. The EAB threatens this goal; see the photo (at top) of ash mortality along a Maryland tributary of the Bay. In 2021, EAB was detected in Gloucester County – a peninsula east of the York River that has Bay shoreline on the eastern side, tributary on the west (see photo).

Gloucester Point – Virginia Institute of Marine Sciences “living shoreline”; EAB was detected in Gloucester County in 2021, threatening riparian areas. Photo courtesy of the Chesapeake Bay Program

Threats to Beech

Beech bark disease is present in the western mountainous parts of the Commonwealth. One new county – Augusta – was detected in 2022. Three other counties are infested with the scale, but the fungal pathogen has not yet been detected. The alarming new threat, beech leaf disease, was detected in Prince William County in 2021. In 2022, it was confirmed in neighboring Fairfax County. The source of funding is not specified.

beech in a typical northern Virginia second-growth forest; photo by F.T. Campbell

Laurel Wilt Disease

Sassafras; photo by David Moynihan

I am pleased that the Commonwealth is paying attention to laurel wilt disease, which has been spreading north on sassafras. The closest outbreaks are in Tennessee, to the southwest of Virginia. The Commonwealth hosted a detection training program attend by 26 participants from six agencies from three states. The report does not specify the source of the funding.

Southern Pine Beetle

Virginia has also utilized funding from the USFS FHP program to manage the southern pine beetle. Since the program’s inception in 2004, Virginia has thinned pines on more than 70,000 acres, including 4,240 acres in FY22.

Invasive Plants

USFS FHP invasive species grants funded control treatments of invasive plants on somewhat less than 1,300 acres of state lands. Different figures on different pages of the report cause confusion. However, it is clear that nearly all the funds came from the USFS FHP program. Ailanthus was the main target; other species mentioned are privet, mimosa, autumn olive and Miscanthus.

State Funding of Conservation Initiatives; Will They Continue?

While the state’s government was controlled by Democrats, the governor and state legislature launched new programs with broader conservation goals. It is unclear whether they will continue now that Republicans have won the governorship and control of the House of Delegates.

Among the programs enjoying increased funding from the state budget during the current two-year cycle are

  • Efforts to restore depleted populations of two groups of tree taxa, shortleaf and longleaf pines. The emphasis has shifted to longleaf pine: the number of projects and acreages rose from 220 acres in FY21 to 1,212 acres in FY22. Restoration of shortleaf pine forests was limited to slightly over 600 acres in both years.
  • Programs to improve management of hardwood stands. These projects included crop tree release, control of “invasive species” (I think probably targetting invasive plants), prescribed burning and commercial thinning. There were also several demonstration projects on state-owned lands, a small land-owner planning assistance program, and training of state foresters and private consulting foresters in hardwood management. Apparently these aspects had been largely ignored in the past.
  • Creation of a dedicated Watershed program focused on increasing riparian forest buffers. This section of the report does not mention the threat posed by loss of ash to the emerald ash borer (EAB) [see EAB section above]
  • Urban forestry projects, many linked to protecting surface and ground water (including Chesapeake Bay watershed).

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

or

www.fadingforests.org

see also the article about beech leaf disease in the mid-Atlantic region written by Gabe Popkin; posted here

Imports from China down slightly, but high pest risk continues

I have blogged often about the pest risk of wood packaging associated with imports from Asia – especially China – and the shift in that risk arising from import volumes and ports at which they are arriving (increasing volumes entering country at ports along Atlantic and Gulf coasts). [See blogs posted on this site, under the “wood packaging” category (listed below the archives by date).] As noted, U.S. imports from Asia are at all-time highs: in the first three months of 2022, they reached 1.62 million TEU (shipping containers measured as twenty-foot equivalents). This was 31.1% higher than in the same period in pre-pandemic 2019 (Mogelluzzo, B. April 22, 2022).

The most recent information (Szakonyi, M. 2023) confirms that U.S. importers are shifting suppliers to countries other than China, primarily because of lengthy shutdowns in Chinese factories linked to the “0 COVID” policy and some U.S. restrictions and tariffs. Over 2022 (full year), China – including Hong Kong – supplied 40.7% of U.S. imports. This is still a huge proportion, but lower than in 2021, when it was 42.4%. The Journal of Commerce calculates that the number of containers coming from China fell by 435,000. At the current rate of infestation in wood packaging from China calculated by Haack et al. 2022, that might mean about 1,200 fewer containers from China with infested wood packaging entering the U.S.

[Explanation of calculations: I divided 435,000 by 2 to convert 20-ft TEU into 40-ft containers that CBP encounters at the ports; multiplied the result by 0.75 – based on the decade-old Meissner estimate of % of containers that have SWPM; then multiplied the result by .0073 because that is infestation rate for China during 2010-2020 period]

This might be progress. China continues to have a terrible record of non-compliant wood packaging 23 years after U.S. and Canada instituted phytosanitary requirements. According to Haack et al. (2022), packaging from China made up 4.6% of all shipments inspected under the terms of their analysis, but 22% of the 180 consignments with infested wood packaging. Thus the proportion of Chinese consignments with infested wood is five times greater than expected based on their proportion of the dataset. The rate of wood packaging from China that is infested has remained relatively steady = 1.26% during 2003–2004, 0.73% during 2010 – 2020. And the insects present belong to the group that causes the greatest damage: longhorned beetles (Cerambycids). Indeed, 78% of beetles in this family that were detected were from China.

There is some good news: some types of goods likely to be enclosed in crates have decreased notably. The proportion of furniture and other home items imported from China has declined from 71.6% of all U.S. imports in 2010 to 52.6% in 2022. As Haack et al. (2022) found, crates are the type of wood packaging where wood pests are most commonly found. While crates constituted only 7.5% of the wood packaging inspected, they made up 29.4% of the infested packaging – or four times greater than their proportion of the dataset.

The pest risk might not be changing significantly, however, because some of the new suppliers are also in Asia. Vietnam’s share of U.S. imports rose from 8.2% to 8.7%. The types of goods most often imported from Vietnam included electronics, shoes, and apparel. The U.S. has already been invaded by insect-pathogen complexes native to Vietnam, Taiwan, and other parts of southeast Asia – e.g., redbay ambrosia beetle and laurel wilt; invasive shot hole borers and Fusarium disease.

U.S. imports from South Korea, mostly electronics and autoparts, climbed from 3.8% to 4.1%. Imports from India also saw a tiny increase – from 3.8% to 3.9%. These shipments were primarily apparel and iron and steel components. These goods prompt concern because wood packaging associated with heavy materials are often infested by insects (Eyre et al. 2018). The Haack et al. (2022) analysis found two interceptions of wood packaging from Vietnam, one from Korea, and three from India.

Besides, the Journal of Commerce notes that shifts in suppliers cannot go far. These countries’ manufacturing capacity and transportation infrastructure are far below those of China (Szakonyi, M. 2023).

In February 2023, U.S. imports from Asia continued to decline from record levels in 2021 and 2022 to 1.09 million TEU. This level still exceeds by 25% the 869,091 TEU recorded in March 2020, at the beginning of the COVID-19 shutdown (Mongelluzzo, March 17, 2023).

[Reminder: higher shares of imports from Asia are going to ports along the Atlantic and Gulf coasts – spreading the risk. See earlier blogs. In early March the Port of Savannah posted an advertisement to the on-line Journal of Commerce, crowing that by July it will complete straightening the river at the Garden City Terminal (the container terminal). This fix will enable Savannah to raise its annual container processing capacity by 1.5 million TEU, to 7.5 million.]

The most hopeful finding is that imports from Mexico jumped 19.2% in the first 11 months of 2022 compared to the same period in 2021. Importers have their reasons: a desire to buy from producers closer to the U.S. market. These motivations have nothing to do with the risk of forest pest introductions. However, we can rejoice because Mexico has greatly improved the pest-infestation rates of its exports since 2009. The rate fell from 0.29% in 2003-2004 to 0.04% in 2010-2020 (Haack et al. (2022).  

larval Asian longhorned beetle; Thomas Denholm, NJ Department of Agriculture; Bugwood

I remain outraged that U.S. agencies have not taken effective steps to deal with the nearly 25-year-long problem of Chinese noncompliance with our phytosanitary requirements. As I noted in my previous blog, link to blog 303 Customs and Border Protection officials are disappointed that their enhanced enforcement in 2017 and 2021 has not yet resulted in improved compliance.

I suggested that the U.S. and Canadian government agencies should penalize trade partners with high records of not complying with ISPM#15. Among steps they should consider are

  • U.S. and Canada should refuse to accept wood packaging from foreign suppliers that have a record of repeated violations – whatever the apparent cause of the non-compliance. Institute severe penalties to deter foreign suppliers from taking devious steps to escape being associated with their violation record.
  • APHIS and CBP and their Canadian counterparts should provide guidance to importers on which foreign treatment facilities have a record of poor compliance or suspected fraud – so they can avoid purchasing SWPM from them. I greatly regret that the death of Gary Lovett might put an end to the voluntary industry program he had been developing, described here.
  • Encourage a rapid switch to materials that don’t transport wood-borers. Plastic is one such material. While no one wants to encourage production of more plastic, the Earth is drowning under discarded plastic. Some firms are recycling plastic waste into pallets.

Haack et al. 2022 fully describes the methodology used, the structure of USDA’s Agriculture Quarantine Inspection Monitoring (AQIM) program, detailed requirements of ISPM#15, the phases of U.S. implementation, etc.  Also see the supplemental data sheet in Haack et al. (2022) that compares the methods used in each analysis.

SOURCES

Eyre, D., Macarthur, R., Haack, R.A., Lu, Y. and Krehan, H., 2018. Variation in inspection efficacy by member states of wood packaging material entering the European Union. Journal of Economic Entomology, 111(2), pp.707-715.

Haack RA, Hardin JA, Caton BP and Petrice TR (2022) Wood borer detection rates on wood packaging materials entering the United States during different phases of ISPM#15 implementation and regulatory changes. Frontiers in Forests and Global Change 5:1069117. doi: 10.3389/ffgc.2022.1069117

Meissner, H., A. Lemay, C. Bertone, K. Schwartzburg, L. Ferguson, L. Newton. 2009. Evaluation of Pathways for Exotic Plant Pest Movement into and within the Greater Caribbean Region. A slightly different version of this report is posted at 45th Annual Meeting of the Caribbean Food Crops Society https://econpapers.repec.org/paper/agscfcs09/256354.htm

Mongelluzzo, B. Q1 US imports from Asia show no slowing in consumer demand. Apr 22, 2022. https://www.joc.com/maritime-news/container-lines/q1-us-imports-asia-show-no-slowing-consumer-demand_20220422.html

Mongelluzzo, B. US imports from Asia hit three-year low in February: data. https://www.joc.com/article/us-imports-asia-hit-three-year-low-february-data_20230317.html

Szakonyi, M. 2023. Sourcing shift from China pulls US import share to more than a decade low. https://www.joc.com/article/sourcing-shift-china-pulls-us-import-share-more-decade-low_20230201.html

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

or

www.fadingforests.org

Elm zigzag sawfly invades Eastern U.S.

The characteristic zigzag pattern Picture: Kelly Oten, NC State University.

Guest blog by Kelly Oten, NC State University

The elm zigzag sawfly [EZS; Aproceros leucopoda Takeuchi (Hymenoptera: Argidae)] is the newest invasive forest insect detected in the eastern US. The colloquially-used common name, currently going through the ESA common name approval process, is not only catchy, but perfectly describes this defoliator’s unique feeding damage. As EZS feeds on elm leaves, it weaves a zigzag pattern from the margin of the leaf towards the mid-vein.

An Expansive — and Quickly Growing – Range

Native to East Asia, the first confirmation of EZS in North America occurred in August 2020 in Québec, Canada when an iNaturalist user posted a photo showing the characteristic zigzag defoliation. The observer realized it was potentially EZS and emailed local entomologists in the province who visited the site, collected specimens, and obtained species confirmation through the Canadian Food Inspection Agency Entomology Lab (Martel et al. 2021). However, this detection was not actually the beginning.

Three months before the Canadian detection, the same defoliation pattern was observed in Frederick County, Virginia, USA. Observers suspected EZS, but no specimens were recovered and therefore identification could not be confirmed. A year later, the site was revisited and this time, bingo—specimens were present and confirmed as EZS. Subsequent surveys that summer led to detections in eight additional Virginia counties. At the same time, the telltale defoliation was observed in Lehigh County, Pennsylvania, USA, but no specimen could be recovered for confirmation. In 2022, EZS popped up more widely; four additional states confirmed EZS: Pennsylvania, North Carolina, Maryland, and New York.

map created by Kelly Oten, NCSU

Though new to North America, this insect has a history of invasiveness. First detected in Europe in 2003, it now occupies areas from the United Kingdom and France in the west, to Russia and Kazakhstan in the east (Ashikbayev et al. 2018, iNaturalist 2022).

The strange and unusual biology of elm zigzag sawfly

Like all Hymenopterans, EZS goes through four life stages: egg, larva, pupa, adult. Eggs are laid along leaf margins; after hatching, larvae feed on leaf foliage in a zigzag pattern towards the mid-vein. Older larvae consume the leaf more entirely, leaving behind the mid-vein and thicker lateral veins only. Before pupating, larvae spin a cocoon within which they pupate. Cocoons are seasonally dimorphic; summer pupae (which emerge as adults in 4-7 days) are net-like and attached to leaves or twigs. Overwintering pupae are solid-walled and found in leaf litter or soil. Interestingly, overwintering pupae are not just produced from the last generation of the year. Even early in the summer, overwintering pupae may develop alongside summer pupae. Adults are able to begin oviposition immediately; not only do they not need to feed, but they don’t need to find a mate either! EZS is parthenogenetic, meaning they reproduce without mating. In fact, no male EZS has ever been recorded and it’s believed the species is entirely female.

As elm zigzag sawfly larvae (bottom left on leaf) grow, they feed more wholly on elm leaves. Picture: Kelly Oten, NCSU

The entirety of this life cycle can last ~20-36 days when not overwintering. However, the voltinism of this pest is highly variable. Papp et al. (2018) recorded up to seven generations in a year on lab-reared colonies, but in nature in Europe, anywhere between two and six generations has been recorded (Blank et al. 2010, Mol and Vonk 2015). In Virginia, two generations were recorded in 2021 and 1 in 2022. It is unknown what factors play into the number of generations per year, but it’s clear that it’s highly variable. The ability of EZS to multiply rapidly and have multiple generations per year suggest large populations can build in a relatively short time. In fact, this was observed in North Carolina in 2022 and in Europe several times before. Large populations are capable of severe defoliation and may cause long-term impacts on tree health.

This collective life cycle description is based on Blank et al. 2010, Martel et al. 2021, Martynov and Nikulina 2017, and Wu 2006.

Spread

EZS has astonished many with how it seems to be popping up all over the place in such a short amount of time. Since 2020, it has been detected in five US states and at several sites along the St. Laurence River in the Canadian province. The adults are strong fliers, capable of spreading 45-90 km (~27-55 mi) per year (Blank et al. 2014). Given the fact they’re parthenogenetic, relatively small numbers can disperse to begin new populations. Of perhaps greater concern is the potential for long-range dispersal. In heavy infestations in North Carolina, cocoons were found not just attached to leaves and stems, but also non-living objects, suggesting a possible mechanism of long-range dispersal should they become attached to vehicles or other objects transported long distances. In addition, EZS damage ranges from minor to severe defoliation. When populations are low and feeding is minor, it’s less likely to be detected unless intentional surveys are conducted. This cryptic nature might suggest it’s in more places than we are currently aware.

An elm zigzag sawfly cocoon attached to a fence post.
Picture: Kelly Oten, NCSU

So, what’s the big deal?

In short, we don’t know yet. Generally speaking, defoliation by insects causes little long-term harm to tree health but severe and/or repeated defoliation can weaken or sometimes kill a host. In Europe, trees severely defoliated by EZS are typically able to re-leaf but may suffer branch dieback and/or reduced growth (Blank et al. 2010, Zandigiacomo et al. 2011). Also of note, EZS is attacking elm, an already-threatened tree due to widespread mortality cause by Dutch elm disease in the 1900s. Defoliation by EZS could further weaken infected trees or, at the very least, present an additional threat for remaining elms (Blank et al. 2010). While it seems aesthetic damage will be the primary concern with EZS, the potential for long-term tree health impactsin the US is uncertain and should be investigated. For now, anyone that finds EZS or its characteristic defoliation pattern are encouraged to report it to their respective state agriculture or forestry agency.

A row of winged elm (Ulmus alata) in NC were severely defoliated by elm zigzag sawfly.
Picture: Kelly Oten, NCSU

[See Faith’s earlier blog about the zigzag sawfly here.]

References

Ashikbayev, N. Z., N. S. Mukhamadiyev, G. Z. Mengdibayeva, M. B. Temirzhanov, and N. K. Kuanyshbaev. 2018. Development of forest entomology in Kazakhstan, pp. 42–47.  In T. I. Espolov, K. M. Tireuov, E. I. Islamov, S. B. Baizakov, K. T. Abayeva, E. Z. Kentbaev, and B. A. Kentbaeva (eds.), Actual problems of sustainable development in forestry complex, vol. 2. Aitumar Publishing, Almaty, Kazakhstan.

Blank, S. M., H. Hara, J. Mikulás, G. Csóka, C. Ciornei, R. Constantineanu, I. Constantineanu, L. Roller, E. Altenhofer, T. Huflejt, and G. Vétek. 2010. Aprocerosleucopoda (Hymenoptera, Argidae): an East Asian pest of elms (Ulmus spp.) invading Europe. Eur. J. Entomol. 107: 357–367.

Blank, S. M., T. Köhler, T. Pfannenstill, N. Neuenfeldt, B. Zimmer, E. Jansen, A. Taeger, A.D. Liston. 2014. Zig-zagging across Central Europe: recent range extension, dispersal seed and larval hosts of Aprocerosleucopoda (Hymenoptera, Argidae) in Germany. J. Hymenopt. Res. 41: 57-74.

iNaturalist. Available from https://www.inaturalist.org. Accessed August 2022.

Martel, V., O. Morin, S. Monckton, C. Eiseman, C. Béliveau, M. Cusson, and S. Blank. 2021. Elm zigzag sawfly, Aproceros leucopoda (Hymenoptera: Argidae), recorded for the first time in North America through community science. Can. Entomol. 154: E1.

Martynoz, V. V., and T. V. Nikulina. 2017. Population surge of zigzag elm sawfly (Aproceros leucopoda (Takeuchi, 1939): Hymenoptera: Argidae) in the Northern Ciz-Azov Region. Russ. J. Biol. Invasions 8: 25-34.

Mol, A. W. M., and D. H. Vonk. 2015. De iepenzigzagbladwesp Aproceros leucopoda (Hymenoptera: Argidae), een invasieve exoot in Nederland. Entomol. Ber. 75: 50-63.

Papp, V., M. Ladányi, and G. Vétek. 2018. Temperature-dependent development of Aproceros leucopoda (Hymenoptera: Argidae), an invasive pest of elms in Europe. J. Appl. Entomol. 142: 589-597.

Wu, X. Y. 2006. Studies on the biology and control of Aproceros leucopoda. Plant Prot. 32: 98-100.

Zandigiacomo, P., E. Cargnus, and A. Villani. 2011. First record of the invasive sawfly Aproceros leucopoda infesting elms in Italy. Bull. Insectology 64: 145-149.