Category: invasive species policy

What Do Invasive Species Cost?

brown tree snake Boiga irregularis; via Wikimedia; one of the species on which the most money is spent on preventive efforts

In recent years a group of scientists have attempted to determine how much invasive species are costing worldwide. See Daigne et al. 2020 here.

Some of these scientists have now gone further in evaluating these data. Cuthbert et al. (2022) [full citation at end of blog] see management of steadily increasing numbers of invasive, alien species as a major societal challenge for the 21st Century. They undertook their study of invasive species-related costs and expenditures because rising numbers and impacts of bioinvasions are placing growing pressure on the management of ecological and economic systems and they expect this burden to continue to rise (citing Seebens et al., 2021; full citation at end of blog).

They relied on a database of economic costs (InvaCost; see “methods” section of Cuthbert et al.) It is the best there is but Cuthbert et al. note several gaps:

  • Only 83 countries reported management costs; of those, only 24 reported costs specifically associated with pre-invasion (prevention) efforts.
  • Data comparing regional costs do not incorporate consideration of varying purchasing power of the reporting countries’ currencies.  
  • Data available are patchy so global management costs are probably substantially underestimated. For example, forest insects and pathogens account for less than 1% of the records in the InvaCost database, but constitute 25% of total annual costs ($43.4 billion) (Williams et al., in prep.) .

Still, their findings fit widespread expectations.  

These data point to a total cost associated with invasive species – including both realized damage and management costs – of about $1.5 trillion since 1960.  North America and Oceania spent by far the greatest amount of all global money countering bioinvasions. North America spent 54% of the total expenditure of $95.3 billion; Oceania spent 30%. The remaining regions each spent less than $5 billion.

Cuthbert et al. set out to compare management expenditures to losses/damage; to compare management expenditures pre-invasion (prevention) to post-invasion (control); and to determine potential savings if management had been more timely.

Economic Data Show Global Efforts Could Be – But Aren’t — Cost-Effective

The authors conclude that countries are making insufficient investments in invasive species management — particularly preventive management. This failure is demonstrated by the fact thatreported management expenditures ($95.3 billion) are only 8% of total damage costs from invasions ($1.13 trillion). While both cost or losses and management expenditures have risen over time, even in recent decades, losses were more than ten times larger than reported management expenditures. This discrepancy was true across all regions except the Antarctic-Subantarctic. The discrepancy was especially noteworthy in Asia, where damages were 77-times higher than management expenditures.

Furthermore, only a tiny fraction of overall management spending goes to prevention. Of the $95.3 billion in total spending on management, only $2.8 billion – less than 3%  – has been spent on pre-invasion management. Again, this pattern is true for all geographic regions except the Antarctic-Subantarctic. The divergence is greatest in Africa, where post-introduction control is funded at more than 1400 times preventive efforts. It is also significant for Asia and South America.

Even in North America – where preventative actions were most generously funded – post-introduction management is funded at 16 times that of prevention.

Cuthbert et al. worry particularly about the low level of funding for prevention in the Global South. They note that these conservation managers operate under severe budgetary constraints. At least some of the bioinvasion-caused losses suffered by resources under their stewardship could have been avoided if the invaders’ introduction and establishment had been successfully prevented.

While in the body of the article Cuthbert et al. seem uncertain about why funding for preventive actions is so low, in their conclusions they offer a convincing (to me) explanation. They note that people are intrinsically inclined to react when impact becomes apparent. It is therefore difficult to motivate proactive investment when impacts are seemingly absent in the short-term, incurred by other sectors, or in different regions, and when other demands on limited funds may seem more pressing. Plus efficient proactive management will prevent any impact, paradoxically undermining evidence of the value of this action!

Aedes aegypti mosquito; one of the species on which the most money is spent for post-introduction control; photo by James Gathany; via Flickr

Delay Costs Money

The reports contained in the InvaCost database indicate that management is delayed an average of 11 years after damage was first been reported. Cuthbert et al. estimate that these delays have caused an additional cost of about $1.2 trillion worldwide. Each $1 of management was estimated to reduce damage by $53.5 in this study. This finding, they argue, supports the value of timely invasive species management.

They point out that the Supplementary Materials contain many examples of bioinvasions that entail large and sustained late-stage expenditures that would have been avoided had management interventions begun earlier.

Although Cuthbert et al. are not as clear as I would wish, they seem to recognize also that stakeholders’ varying perceptions of whether an introduced species is causing a detrimental “impact” might also complicate reporting – not just whether any management action is taken

Cuthbert et al. are encouraged by two recent trends: growing investments in preventative actions and research, and shrinking delays in initiating management. However, these hopeful trends are unequal among the geographic regions.

Which Taxonomic Groups Get the Most Money?

About 42% of management costs ($39.9 billion) were spent on diverse or unspecified taxonomic groups. Of the costs that were taxonomically defined, 58% ($32.1 billion) was spent on invertebrates [see above re: forest pests]; 27% ($14.8 billion) on plants; 12% ($6.7 billion) on vertebrates; and 3% ($1.8 billion) on “other” taxa, i.e. fungi, chromists, and pathogens. For all of these defined taxonomic groups, post-invasion management dominated over pre-invasion management.  

When considering the invaded habitats, 69% of overall management spending was on terrestrial species ($66.1 billion); 7% on semi-aquatic species ($6.7 billion); 2% on aquatic species ($2.0 billion); the remainder was “diverse/unspecified”. For pre-invasion management (prevention programs), terrestrial species were still highest ($840.4 million). However, a relatively large share of investments was allocated to aquatic invaders ($624.2 million).

Considering costs attributed to individual species, the top 10 targetted for preventive efforts were four insects, three mammals, two reptiles, and one alga. Top expenditures for post-invasion investments went to eight insects [including Asian longhorned beetle], one mammal, and one bird.

Asian longhorned beetle

Just two of the costliest species were in both categories: insects red imported fire ant(Solenopsis invicta) and Mediterranean fruitfly (Ceratitis capitate). None of the species with the highest pre-invasion investment was among the top 10 costliest invaders in terms of damages. However, note the lack of data on fungi, chromists, and pathogens. (I wrote about this problem in an earlier blog.)

Discussion and Recommendations

Cuthbert et al. conclude that damage costs and post-invasion spending are probably growing substantially faster than pre-invasion investment. Therefore, they call for a stronger commitment to enhancing biosecurity and for more reliance on regional efforts rather than ones by individual countries. Their examples of opportunities come from Europe.

Drawing parallels to climate action, the authors also call for greater emphasis on during decision-making to act collectively and proactively to solve a growing global and inter-generational problem.

Cuthbert et al. focus many of their recommendations on improving reporting. One point I found particularly interesting: given the uneven and rapidly changing nature of invasive species data, they think it likely that future invasions could involve a new suite of geographic origins, pathways or vectors, taxonomic groups, and habitats. These could require different management approaches than those in use today.

As regards data and reporting, Cuthbert et al. recommend:

1) reducing bias in cost data by increasing funding for reporting of underreported taxa and regions;

2) addressing ambiguities in data by adopting a harmonized framework for reporting expenditures. For example, agriculture and public health officials refer to “pest species” without differentiating introduced from native species. (An earlier blog also discussed the challenge arising from  these fields’ different purposes and cultures.)

3) urging colleagues to try harder to collect and integrate cost information, especially across sectors;

4) urging countries to report separately costs and expenditures associated with different categories, i.e., prevention separately from post-invasion management; damage separately from management efforts; and.

5) creating a formal repository for information about the efficacy of management expenditures.

While the InvaCost database is incomplete (a result of poor accounting by the countries, not lack of effort by the compilers!), analysis of these data points to some obvious ways to improve global efforts to contain bioinvasion. I hope countries will adjust their efforts based on these findings.

SOURCE

Cuthbert, R.N., C. Diagne, E.J. Hudgins, A. Turbelin, D.A. Ahmed, C. Albert, T.W. Bodey, E. Briski, F. Essl, P. J. Haubrock, R.E. Gozlan, N. Kirichenko, M. Kourantidou, A.M. Kramer, F. Courchamp. 2022. Bioinvasion costs reveal insufficient proactive management worldwide. Science of The Total Environment Volume 819, 1 May 2022, 153404

Seebens, H. S. Bacher, T.M. Blackburn, C. Capinha, W. Dawson, S. Dullinger, P. Genovesi, P.E. Hulme, M.van Kleunen, I. Kühn, J.M. Jeschke, B. Lenzner, A.M. Liebhold, Z. Pattison, J. Perg, P. Pyšek, M. Winter, F. Essl. 2021. Projecting the continental accumulation of alien species through to 2050. Glob Change Biol. 2021;27:970-982.

Williams, G.M., M.D. Ginzel, Z. Ma, D.C. Adams, F.T. Campbell, G.M. Lovett, M. Belén Pildain, K.F. Raffa, K.J.K. Gandhi, A. Santini, R.A. Sniezko, M.J. Wingfield, and P. Bonello 2022. The Global Forest Health Crisis: A Public Good Social Dilemma in Need of International Collective Action. submitted

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

Global Loss of Floristic Uniqueness

Hakalau Forest, Hawai“i; nearly 90% of Hawaiian flora is unique to the Islands

A recent article by Yang et al. 2021 (full citation at the end of this blog) seeks to determine the extent to which introduced plants reduce the uniqueness of regional floras. They analyzed data from 658 regions covering about 65.7% of the Earth’s ice-free land surface and about 62.3% of the planet’s known plant species.

They found strong homogenization of plant species’ taxonomic and phylogenetic diversity results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.

There are two aspects of a region’s floral uniqueness. One is the number of species that it shares with other regions. This is taxonomic uniqueness. The other is the distinctiveness of the evolutionary history of the region. When several species are endemic to a region’s flora, and lack close relatives in other regions, that equals phylogenetic uniqueness.

The effect of a species introduction differs depending on which of these aspects one focuses on. Thus, naturalization of a species closely related to native species (e.g., a congeneric species) will have less impact on the phylogenetic floristic uniqueness of the region than naturalization by a distantly related species. Taxonomic uniqueness, however, will be affected to the same degree, irrespective of the phylogenetic distance between the naturalized and native species.

Yang et al. found strong homogenization of plant diversity. They found that species introductions increased the taxonomic similarity in 90.7% of all regional pairs and phylogenetic similarity in 77.2% of all region pairs. Most homogenization results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.

This loss of regional biotic uniqueness or distinctiveness changes biotic interactions and species assemblages. These, in turn, have ecological and evolutionary consequences at larger scales and higher levels.

The degree of homogenization between regions’ floras depends on three factors:

1) The distance between the donor and recipient regions. Since nearby regions share more species, an introduction from a more distant origin is more likely to be a novel species and so contribute to homogenization of “donor” and “receiving” floras.

2) Climatic similarity, especially temperature. A plant species introduced from a climatically similar but geographically distant place is more likely to establish than a species from a different climatic zone. As a result, the recipient area’s flora is changed to more closely resemble the flora of the donor region with which it shares climatic conditions – regardless of the distance between them.

3) The level of exchange of goods and people between two regions. The higher the rate of exchange between two regions, the greater the chance that a species will be introduced and become established. Yang et al. used the existence of current or past administrative relationships (e.g., colonial relationship) between two regions as a proxy for intensity of trade and transport between donor and recipient regions. They found that floras of regions with current or past administrative links have taxonomically become more similar to each other than the floras of regions with no such links.

flora of the Cape Floral Kingdom – South Africa; photo from Michael Wingfield

Establishment of introduced species can increase floristic similarity of the donor and recipient regions (= floristic homogenization) when the species is native to one of the two regions and naturalizes in the other, or when it is not native to both regions and naturalizes in both. On the other hand, a species introduction can decrease the floristic similarity of the two regions (i.e., enhance floristic differentiation) when the species is not native to both regions but naturalized in only one.  

Homogenization hotspots differed slightly depending on whether one focused on taxonomic or phylogenetic aspects.

The regions with the greatest average increase in taxonomic similarity with other regions due to naturalized alien species were New Zealand, portions of Australia, and many oceanic islands. The Australasian situation probably reflects its long biogeographic isolation from other parts of the globe and its highly unique native flora. As a result, nearly all non-native plants introduced to Australasia strongly increase levels of its floristic similarity to the rest of the world. Oceanic islands have species-poor floras with large proportions of unique endemics. They have also received high numbers of naturalized alien plants.

Hotspots of phylogenetic homogenization on continents are the same as those for taxonomic homogenization, but this is not true for islands. Yang et al. think this is because islands’ native floras were established by natural colonization from nearby continental floras so – despite subsequent speciation – they retain their phylogenetic relationship to the donor areas’ floras.  

Yang et al. concede that they lacked high-quality data on native and naturalized alien species lists for a third of Earth’s ice-free terrestrial surface, especially Africa, Eastern Europe, and tropical Asia. They believe, however, that data from these regions are unlikely to change the overall finding.  (Scientists are beginning to compile lists of forest pests in Africa). link to blog

Yang et al. note that introduction and naturalization of alien species are likely to increase in the future, thusaccelerating floristic homogenization. The ecological, evolutionary and socioeconomic consequences are largely unknown.They call for stronger biosecurity regulations of trade and transport and other measures to protect native vegetation.

SOURCE

Yang, Q., P. Weigelt, T.S. Fristoe, Z. Zhang, H. Kreft, A. Stein, H. Seebens, W. Dawson, F. Essl, C. König, B. Lenzner, J. Pergl, R. Pouteau, P. Pyšek, M. Winter, A.L. Ebel, N. Fuentes, E.L.H. Giehl, J. Kartesz, P. Krestov, T. Kukk, M. Nishino, A. Kupriyanov, J.L. Villaseñor, J.J. Wieringa, A. Zeddam, E. Zykova  and M. van Kleunen. 2021. The global loss of floristic uniqueness. NATURE COMMUNICATIONS (2021) 12:7290. https://doi.org/10.1038/s41467-021-27603-y

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

Urban Forests at Risk: Thousands of Communities, Millions of Trees, & Tens of Millions of Dollars

EAB-killed ash tree lying on highway in Fairfax County, Virginia; photo by F.T. Campbell

A recent study (Hudgins, Koch, Ambrose & Leung 2022; full citation at end of blog) projects that, by 2050, 1.4 million street trees in urban areas and communities will be killed by introduced insect pests. This represents 2.1- 2.5% of all urban street trees. Nearly all of this mortality will occur in a quarter of the 30,000 communities evaluated. Additional urban trees – in parks, other plantings, on homeowners’ properties, and in urban woodlands – are also expected to die.

Loss of these trees will undercut all the ecosystem services provided by urban trees.

The principal cause of mortality will be the emerald ash borer (EAB). Already, an estimated 230,000 ash trees have been killed by EAB. The authors predict that 6,747 communities not yet affected by the EAB will suffer the highest losses between now and 2060. Most of these communities are in a 350,000 square mile area of the northeast and central states. However, the risk is far wider, reaching as far as Seattle.

This ash tree has been standing – dead – since 2016. When will it fall?

In the top ‘hotspot cities’ projected mortality is in the range of 5,000–25,000 street trees. These include Milwaukee; the Chicago area (Chicago / Aurora / Naperville / Arlington Heights); Cleveland; and Indianapolis.  As in previous studies, the highest insect impacts are in the Northeast. Pests impacting this region – in addition to the emerald ash borer – include the spongy moth (formerly called gypsy moth) and hemlock woolly adelgid.

Because insect-killed trees must be treated or removed to minimize the risk to human life and property, the pest risk represents an economic as well as ecological threat. Removing and replacing just the street trees is projected to cost cities $30 million per year. Considering the cities I mentioned above, Milwaukee faces costs estimated at $13 million; Warwick, RI $2.5 million; Baltimore $1.7; Richmond and Virginia Beach $7.3 million and $700,000 respectively; and three New Jersey cities (Jersey City, Elizabeth City, and Patterson) $1.6 million combined.

USDA APHIS ended the federal quarantine for EAB in 2021. Therefore these cities and states are on their own to protect themselves from not only this and other damaging insects but also their extraordinarily high economic costs.

The study evaluated the risk to 48 genera of trees in about 30,000 communities. The most widely planted genera are maples (Acer spp.) and oaks (Quercus spp.). Consequently, they will die in largest numbers. An estimated 26.5 million maples and 5.9 million oaks are at risk, primarily in the East. As noted above, EAB is expected to kill 99% of ash trees in 6,747 communities across the country. In the Southwest, there are 3.4 million pines (Pinus spp.); the threat to them is not woodborers, but scale insects (San Jose scale [Quadraspidiotus perniciosus] and calico scale [Eulecanium cerasorum]).

As we know, urban forests are easily invaded because they are close to ports of entry and are often composed primarily of highly susceptible species. Hudgins, Koch, Ambrose and Leung analyzed the potential risk associated with introduction of a new woodboring insect from Asia – which they point out is the source of most imported goods. They determined that if such an introduced pest were to attack maples or oaks, it 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 via a port in the South.

In an earlier blog I reported that the U.S. is currently importing about 20 million shipping containers filled with goods from Asia per year. I have often blogged about the pest risk associated with wood packaging accompanying these imports. The number of containers from Asia entering Southeastern ports rose by more than 10% from December to January.

Hudgins, Koch, Ambrose & Leung combined four sources of information to produce these estimates:

  • a model of spread for 57 species of introduced insect pests already determined to cause significant damage to trees;
  • the distribution of genera of urban street trees across 30,000 US communities;
  • a model of host mortality in response to each insect-host combination; and
  • the cost of removing and replacing dead trees, linked to tree size (dbh).

They excluded several categories of pests. One of the most damaging, Asian longhorned beetle, was excluded because scientists have already developed control methods to limit its spread. Also excluded were species present in the U.S. for less than five years; species with no known economic impacts; and species for which no hosts in natural North American forests have been identified. Also excluded – although the authors do not mention this – are species that did not qualify for inclusion in the Aukema et al. study (see reference at end of this blog) because they have been introduced from nearby portions of North America, e.g., goldspotted oak borer. Finally, the study does not include pathogens. Some pathogens have caused huge losses of urban trees in the past, e.g., Dutch elm disease; some are causing losses now, e.g., sudden oak death. The authors do mention the Fusarium disease vectored by polyphagous (and Kuroshio) shot hole borers in southern California.

elm-lined street; photo from USFS

Consequently, the study’s estimate of 1.4 million street trees dead and costs of $30 million per year are underestimates.

The study has generated considerable media interest, including in the Washington Post.

SOURCES

Aukema, J.E., D.G. McCullough, B. Von Holle, A.M. Liebhold, K. Britton, & S.J. Frankel. 2010. Historical Accumulation of Nonindigenous Forest Pests in the Continental United States. Bioscience. December 2010 / Vol. 60 No. 11

Hudgins, E.J., F.H. Koch, M.J. Ambrose, B. Leung. 2022. Hotspots of pest-induced US urban tree death, 2020–2050. Journal of Applied Ecology 2022;00:1-11 DOI: 10.1111/1365-2664.14141

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

Integrating Invasion & Phytosanitary Sciences & Practice

vegetation killed by Phytophthora cinnamomi in West Australia

Some invasive species practitioners have been trying to develop a standardized framework for describing bioinvasions. Their goal is to overcome disparities in approaches developed by scientists working with various taxonomic groups in hopes of improving understanding of, and communication about, bioinvasions. Prominent among these efforts is the “Unified Framework for Bioinvasion” published by Blackburn et al. in 2011 (full citation at end of blog).  

Now several forest pathologists (Paap et al; full citation at end of blog) say that this framework does not adequately integrate forest pathogens. This omission is particularly unfortunate given the prominence of forest pathogens as damaging invaders – e.g., chestnut blight in Europe and North America; white pine blister rust in North America; sudden oak death in North America and Great Britain; myrtle rust and Phytophthora cinnamomi in Australia. (See profiles of all these pathogens here; I note additional examples in North America, such as laurel wilt disease.)

Paap et al think that this omission impedes understanding of both forest pests and invasive species in general. Also, they say that integrating microorganisms into the broader Blackburn framework would help forest pathologists better understand how and why invasions occur, where they occur, and how they can be stopped or mitigated. 

Furthermore, they note the importance of integrating the diverging terminologies used by invasive species practitioners and plant pathologists and their separate regulatory bodies – the Convention on the Conservation of Biological Diversity (CBD) and the International Plant Protection Convention (IPPC). I concur, since nations’ programs regulating plant diseases and their vectors operate under the IPPC rubric.

Figure 2 and Table 1 lay out Paap et al.’s proposed modification of Blackburn’s framework, and detail strategies linked to management goals appropriate for the stages of plant disease development.

Tanoak mortality in southern Oregon caused by P. ramorum – a pathogen completely unknown until it was introduced to North America and Europe; photo by Oregon Department of Forestry

However, such integration will be impeded by many difficulties (I have re-ordered these points): 

1) The first – which underlies all others — is the paucity of data on microbial taxa, which undermines the pest risk analyses and other systems developed for assessing and managing other types of invasive species. That is,

  • Many of the vast number of microbial taxa have not yet been described.
  • Even species that have been describe often cannot be ascribed to a specific geographic origin. This information gap undercuts efforts to determine whether a disease outbreak is caused by an “introduced” organism.

2) Microbial species are usually detected only when disease impacts become obvious. However, an outbreak might not signal a new or spreading “introduction”. While invasive species must—by definition—cross a geographic boundary (through the assistance of human actions), pathogens can cause disease outbreaks through breaching a wider range of boundaries, including ecological and evolutionary ones. Thus, the disease outbreak doesn’t always fit the definition of “invasive species”.  

3)  Substantial differences exist in training and goals between fields. Forest pathologists are usually trained in plant pathology (often focused on crops) rather than in forestry or ecology. Their goal is to manage the pathogen. Invasion scientists tend to focus on natural ecosystems, study animal and plant invasions, and seek understanding of the invasion process.

4)  A related issue is that the two fields operate under separate regulatory bodies that have different emphases and aims. Paap et al. note that while the IPPC ostensibly includes impacts on natural environments, its members’ priority is plants of economic importance. The World Trade Organization’s Agreement on the Application of Sanitary and Phytosanitary Measures (WTO SPS) seeks primarily to minimize disruption of trade resulting from plant health regulation. On the other hand, the CBD explicitly considers invasive species’ impact to the natural environment (Aichi Biodiversity Target 9). [To read my critique of the WTO SPS and IPPC, read the Fading Forests reports (link at end of this blog), especially FF II.]

Rome – home to the IPPC

They note that in 2004, the IPPC and CBD secretariats established a Memorandum of Cooperation to promote synergy and to avoid duplication. Paap et al. appear disappointed that despite development of joint work plans, phytosanitary programs are still focused largely on crop pathogens.

Disease development – a complex set of circumstances that makes risk assessment less reliable

Since I am not a pathologist (or even a biologist), I learned a lot about the complexities of plant pathology from Paap et al.

While I am certainly familiar with the “disease triangle” concept, I had not thought about certain implications. For example, pathogens can cause severe disease outbreaks by evading any one of three types of barriers: geographic, environmental, or evolutionary. Transport of the micro-organism to a new ecosystem (leaping the geographic barrier and meeting the definition of an “introduction” in invasive species terminology) certainly can facilitate disease outbreaks. However, evolutionary and environmental barriers might also be overcome in other ways.

The result is that a plant disease can develop under multiple scenarios following the introduction of an alien pathogen. These scenarios are:

  • disease on a coevolved host growing as an alien species in the new environment, for example plantations of trees grown for timber (pathogen reunion);
  • disease on a naïve host that is itself alien to the geographic region in question (host jump);
  • disease on an alien host (naïve or coevolved) which supports disease on a host native to the new geographic area that could not be sustained in the absence of the alien host;
  • disease on alien and native hosts; and
  • disease on a host native to the new geographic area but not on an alien host.

Countries’ efforts to conduct pest risk analyses are unlikely to be straightforward – or even possible – with so many disease scenarios

Paap et al. proceed to compare introductory pathways under the CBD categorization and plant pathology. In doing so they point out several aspects of introduction, establishment, and spread that are specific to pathogens. For example, trees’ long life spans and inability to adapt as rapidly as the micro-organism increase their vulnerability to devastating disease outbreaks following the arrival of a novel pathogen.

Participants in the Montesclaros meeting that drafted an early critique of international phytosanitary procedures

Paap et al. reinforce points made by other critics of current phytosanitary programs. (See my earlier blogs under the category “plants as pest vectors”.) In particular, they point out the weakness of visual inspection and note that new molecular assays can detect only known microorganisms. An additional complication is that DNA can persist in soil and plant tissue after death of the organism, leading to false positives. RNA is cannot yet be used as a viability marker.

Paap et al. provide three case studies to illustrate in greater depth several major challenges encountered when managing invasive forest pathogens. Most of these weaknesses are well known to forest pathologists.

1. The inconspicuous nature of microorganisms

As noted by Paap et al. and other authors, the difficulty detecting microbes is exacerbated by the huge volumes of goods, especially live plants, in international trade; the small proportion of those plants that can be inspected; the weakness of visual examination; application of fungicides and fertilizers before export that suppress symptoms. The chosen example is the oomycete genus Phytophthora, specifically P. ramorum.

2. Cryptic status of many species

Current biosecurity programs rely on naming the organism and its place of origin. This is actually impossible for many microorganisms. The tardy response to ash dieback (Hymenoscyphus fraxineus) in Europe illustrates the delay in determining the causal agent and its geographic origin. During this nearly two-decade period the possibility of preventing spread was lost.

3. Rapid evolution

Rapid evolution of the introduced pathogen can overcome resistance in a host. The example described is Cronartium ribicola (causal agent of white pine blister rust) on Western white pine (Pinus monticola) and sugar pine (P. lambertiana). They also mention the threat from hybridization between previously isolated populations, specifically Phytophthora x alni causing a devastating decline of black alder in Europe.

Sugar pine in Sequoia National Park; photo by S. Rae via Flickr

Paap et al. call for increased research to increase our knowledge of microbial diversity, especially in taxonomically rich and poorly studied ecosystems. They praise sentinel plantings as a powerful tool for early warning of pathogen threats.

SOURCES

Blackburn, T.M., P. Pysek, S. Bacher, J.T. Carlton, R.P. Duncan, V. Jarosik, et al. A proposed unified framework for biological invasions. Trends Ecol Evol. 2011; 26(7):333-9.  

Paap, T., M.J. Wingfield, T.I. Burgess, J.R.U. Wilson, D.M. Richardson, A. Santini. 2022. Invasion Frameworks: a Forest Pathogen Perspective.  FOREST PATHOLOGY https://doi.org/10.1007/s40725-021-00157-4

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

Forest Pest Threat to Africa

Eucalyptus plantation in Kwa-Zulu-Natal, South Africa; Kwa-Zulu-Natal Dept. of Transportation

Graziosi et al. (full citation at the end of the blog) point out that trees are crucial for Africa’s future. Eight hundred of the 4,500–6,000 indigenous tree species provide significant food. As elsewhere, trees provide wood and other extractive resources essential for economic growth. They also support biodiversity and mitigate current and impending climatic variations. Africa– especially the Sub-Saharan countries – is already considered highly vulnerable to climate change.

According to Graziosi et al., the cumulative economic impact of all invasive species in Africa is expected to exceed $1.2 billion per year. The total invasion cost as a proportion of GDP for many African countries is among the highest in the world. This raises the stakes for developing locally appropriate management strategies across the continent.

Responding effectively to this threat is hampered by gaps in data as well as some countries’ limited capacity for biosecurity. Graziosi et al. say that improved knowledge of taxonomy, distribution, and damage caused by these organisms is essential. Such knoledge will be crucial to develop continent-wide strategies to manage this emergency and to enhance capacity for country-level interventions.

Native and alien pests. Indigenous and plantation trees

Africa’s trees and their services are threatened by both native pests and accelerating introductions of pests and diseases from elsewhere. Long-established and new invaders increasingly affect planted forests of exotic eucalypts, pines, and Australian acacias, as well as important indigenous trees. Graziosi et al. note that the U.N. Food and Agriculture Organization (FAO) in an annex to a report issued in 2009 recorded about100 species of forest pests affecting trees in planted and natural forests across Africa. Half are native insects and pathogens, a third are alien; about 15% are of unknown origin. Considering all pests, broadleaf trees (predominantly native) are most affected.

The result is damage from the local – e.g., to rural livelihoods – to the continental – e.g., to economic development and biological diversity across Africa. Moreover, pests exacerbate widespread loss of forest cover. Overall, African forests are shrinking at the rate of almost 0.5% annually. This deforestation is affecting particularly natural forests; planted forests are actually growing 1.3% annually.  

Exotic plantation trees face severe threats. More than 47 native and 19 non-indigenous defoliators, sap-feeders, wood- and shoot-borers attack plantations of Acacia spp., Eucalyptus spp., Pinus spp., and teak (Tectona grandis). About 90% of pathogens of plantation forestry are either non-indigenous or of uncertain origin. Eucalyptus alone are severely damaged by 15 species of pathogens. These organisms are listed in Tables 1 and 2.

Numerous native insect species, known as pests of indigenous trees, have reportedly widened their host range and now damage exotic trees too. Some introduced insects appear to pose significant threats to native tree species. One example is the Cypress aphid Cinara cupressi, which is attacking both exotic cypress plantations and the native African cedar Juniperus procera. Some fungi in the family Botryosphaeriaceae are latent pathogens infecting a wide range of hosts including indigenous Acacia. Dieback of large baobab trees was recently reported from southern Africa. While various microorganisms are associated with these symptoms, the specific cause is still uncertain.

A baobab tree in Limpopo region of South Africa; Wikimedia

The risk currently appears to be particularly high in South Africa. The country’s flora is highly diverse and has a high level of endemism. In fact, South Africa is home to the Earth’s smallest floral kingdom, the Cape Floral Kingdom. It is also the apparent hot spot for pest introductions from overseas (see below). Phytophthora cinnamomi is attacking native Proteaceae in South Africa. According to Graziosi et al., an “incredible diversity” of Phytophthora taxa is present, portending threats to additional plant species. Other pathogens are attacking native conifers in the Podocarpus genus, Ekebergia capensis (Meliaceae), and Syzygium trees.

Protea repens and fynbos vegetation near Table Mountain; photo by Mike Wingfield

There is a clear pattern to further spread: pests first introduced to South Africa often spread. Examples include several insects and pathogens on Eucalyptus and the wood-boring pest of pine Sirex noctilio. This pattern is explained by two main factors. South Africa has a high capacity to detect introduced species. Also, there is an active plantation forestry sector that imports propagules. This offers opportunities for contaminating organisms to be introduced simultaneously.

Furthermore, as Graziosi et al. note, determining the geographic origin of significant proportion of pathogens is extremely difficult – an issue I will discuss in a separate blog based on a publication by primarily South African scientists. Some non-indigenous pathogens have been on the African continent for a long time. The Armillaria root rot pathogen apparently was introduced to South Africa with potted plants from Europe in the 1600s! They note also that many non-indigenous pathogens are probably already established on the continent but not yet detected due to the organisms’ cryptic nature and lagging detection abilities.

The future of African forests

African countries expect economic growth with associated increased trade with countries off-continent. The probable result will be to accelerate the rate of species introductions and spread. However, as climate change worsens, managers will find it increasingly difficult both to predict introduced species’ impact and to implement management programs.

This led Graziosi et al. to call for urgent improvements in plant biosecurity across the continent. They advocate improved coordination at regional and international levels. The list of needed actions is a familiar one: development and application of improved diagnostic tools, updated plant exchange regulations, and revised trade policies.

Graziosi et al. also call for development of effective control and management options. They suggest biocontrol, innovative silviculture practices, and selection of resistant trees. The good news is that African countries have already initiated programs to conserve tree germplasm and domesticate indigenous species, including establishment of field gene banks of high-priority indigenous trees. I have previously praised South African efforts, specifically reports here and here.

Mudada, Mapope, and Ngezimana (2022) describe the risk from introduced species to agriculture and human well-being in southern Africa beyond forestry. The region is already ravaged by food insecurities and hidden hunger. It would be devastated if the global average of crop loss due to plant diseases (10-16%) occurs there. They say these losses can be avoided with improved biosecurity mechanisms focused primarily on pest exclusion and plant quarantine regulations.

SOURCES

Graziosi, I. M. Tembo, J. Kuate, A. Muchugi. 2020 Pests and diseases of trees in Africa: A growing continental emergency. Plants People Planet DOI: 10.1002/ppp3.31 

Mudada, N. Mapope, N., and Ngezimana, W. 2022 – The threat of transboundary plant pathogens to agricultural trade in Southern Africa: a perspective on Zimbabwe’s plant biosecurity – A review. Plant Pathology & Quarantine 12(1), 1–33, Doi 10.5943/ppq/12/1/1

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 the United States and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm

Imports from Asia Continue to Surge; Awaiting Better Analysis

port of Long Beach

The surge in US imports from Asia that began in the second half of 2020 continued through 2021 and into January 2022.  As of September 2021, import volumes from Asia averaged almost 20% higher than the historical monthly average for every month of 2021 (Mongelluzzo, October 13, 2021). The surge continued into 2022. In January 2022, US containerized imports from Asia hit the highest monthly total ever recorded — 1.7 million TEU. This was a 14.6% increase over December 2021 – and a 4.5% increase from a year earlier (January 2021). [Mongelluzzo Feb 23]

The 2022 increase in import volumes was on top of the record-breaking levels seen in 2021. For example, average monthly import volumes during 2021 at the principal ports for receipt of goods from Asia — Los Angeles-Long Beach — were 23% over the 2019 average (Mongeluzzo April 2021). 

Increases in volume from December 2021 occurred at ports across the country. Pacific coast ports saw increases – 25.8% at the LA/LB complex (which handles ~50% of US imports from Asia); 39.1% at Northwest Seaport Alliance (Seattle and Tacoma); 19.7% at Oakland. So did ports in the Southeast – 12.7% in Savannah and 14.1% in Charleston. However, New York/New Jersey saw a decrease of 2.2% and Norfolk saw a decrease of 10.6%. [Mongelluzzo Feb 23] New York had seen a steep increase in mid-2021 (Angell Dec. 22, 2021), but apparently this did not hold up through the year.

The southern California ports report that ships leaving China in early March will – as expected – increase import volumes before the end of the month. Long Beach projected that numbers of arriving shipping containers will rise 34% in the week beginning March 20, compared with the week of March 7; Los Angeles projected an increase of 63% [Mongelluzzo March 10].  

port of Mobile

Volumes Will Probably Continue to Rise Along the Gulf 

Containerized imports from Asia through US Gulf ports had risen 27.2% to 1.14 million TEU in 2021. At the port of Mobile, specifically, imports from Asia last year rose 25% from 2020 to 230,347 TEU in 2021. Imports from Asia through Houston jumped 34 % to 807,376 TEU in 2021 [Mongelluzzo Feb 2 2022]

Increasing manufacturing and distribution industries in the Gulf region are probably an important factor in rising import volumes there. Mongelluzzo Feb 2 2022 notes the presence of a Hyundai factory in Alabama, a Tesla factory and Amazon fulfillment center near Austin, as well as several retail chains’ distribution centers near Houston. Many of these facilities opened in 2021.

Import volumes entering via Gulf and Southeastern ports are expected to continue growing in coming months and years. Several carriers have announced new direct Asia-to-US-east coast transport services. These include South Korea’s HMM (to Houston); CMA CGM; and Maersk (Vietnam and China to Houston and Norfolk; China and Indonesia to Charleston and Newark)

Those who follow shipping expect import volumes to drop in February because many factories in Asia were closed for two weeks or more for the Lunar New Year holidays, which began on Febrary 1. Imports should surge again in March. [Mongelluzzo Feb 23]

The Risk

Remember, Asia is the origin of many of the most damaging forest pests. These include Asian longhorned beetle, emerald ash borer, redbay ambrosia beetle, phytophagous and Kuroshia shot hole borers (for profiles of each visit here). Indeed, 15 of 16 non-native bark beetles in the Xyleborini (a tribe of ambrosia beetles) detected in the United States since 2000 are from Asia (Bob Rabaglia, USFS Forest Health Protection, presentation at IUFRO meeting in Prague, September 2021).

It seems to me that the beetles native to southeast Asia, e.g., the phytophagous and Kuroshio shot hole borers, are likely to find the climate along the Gulf of Mexico to their liking.  Indeed, the redbay ambrosia beetle profile already has!

dead redbay in Georgia killed by laurel wilt disease

Li et al. (2021) assessed fungi associated with Eurasian bark and ambrosia beetles and their potential to impact North American trees. They assessed 111 fungal associates of 55 beetle species. They found that none was “highly virulent” on four important pines or oaks of the Southeast. However, I note two caveats.  First, they tested only four host species – two pines (Pinus taeda and P. elliottii var. elliottii) and two oaks (Quercus shumardii and Q. virginiana). They did not test against the many other tree species that comprise important components of forests of the region. Second, their bar for concern was extremely high: to qualify as “highly virulent,” the pathogens had to be as damaging as laurel wilt disease or Dutch elm disease! Both have had extremely damaging impacts on their hosts across North America.

Updated Haack Analysis

As has been documented repeatedly (e.g., my blogs), the current approach to curtailing pest introductions associated with wood packaging is not sufficiently effective. Customs officials continue to detect live quarantine pests in wood packaging as it enters the country. However, the exact level of this threat is unclear since the only assessment was based on data from 2009 (Haack et al., 2014).  I eagerly await the results of Bob Haack’s updated analysis, which I hope will be published by mid-year.

SOURCES

Angell, M. NY-NJ vessel backlog creeps back up amid bigger ship calls. Journal of Commerce. Dec. 22, 2021 https://www.joc.com/maritime-news/ny-nj-vessel-backlog-creeps-back-amid-bigger-ship-calls_20211222.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%2012/23/21%20SUBSCRIBER%20%28Copy%29_PC00000_e-production_E-122936_KB_1223_0617

Angell, M. Maersk to debut new Houston, Norfolk trans-Pac service in March. Journal of Commerce. Feb. 10, 2022 https://www.joc.com/maritime-news/container-lines/maersk-line/maersk-debut-new-houston-norfolk-trans-pac-service-march_20220210.html?utm_campaign=CL_JOC%20Ports%202%2F16%2F22%20_PC00000_e-production_E-127385_TF_0216_0900&utm_medium=email&utm_source=Eloqua

Haack, R.A., K.O. Britton, E.G. Brockerhoff, J.F. Cavey, L.J. Garrett. 2014. Effectiveness of the International Phytosanitary Standard ISPM No. 15 on Reducing Wood Borer Infestation Rates in Wood Packaging Material Entering the United States. PLoS ONE 9(5): e96611. doi:10.1371/journal.pone.0096611

Li, Y., C. Bateman, J. Skelton, B. Wang, A. Black, Y-T. Huang, A. Gonzalez, M.A. Jusino, Z.J. Nolen, S. Freeman, Z. Mendel, C-Y. Chen, H-F. Li, M. Kolařík, M. Knížek, J-H. Park, W. Sittichaya, P. H. Thai, S. Ito, M. Torii, L. Gao, A.J. Johnson, M. Lu, J. Sun, Z. Zhang, D.C. Adams, J. Hulcr. 2021. Pre-invasion assessment of exotic bark beetle-vectored fungi to detect tree-killing pathogens. Phytopathology. https://doi.org/10.1094/PHYTO-01-21-0041-R

Mongeluzzo, B. Additional port capacity alone can’t solve congestion issues: LA-LB. Journal of Commerce. April 2021 https://www.joc.com/port-news/us-ports/additional-port-capacity-alone-can%E2%80%99t-solve-congestion-issues-la-lb_20210407.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%204%2F8%2F21_PC00000_e-production_E-95420_KB_0408_0837

Mongelluzzo, B. September imports show no relief for stressed US ports. Journal of Commerce. Oct. 12, 2021. https://www.joc.com/port-news/us-ports/september-imports-show-no-relief-stressed-us-ports_20211013.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%2010%2F14%2F21_PC00000_e-production_E-116084_KB_1014_0617

Mongelluzzo, B. Gulf Coast import growth propels regional warehousing boom. Journal of Commerce. Feb. 2, 2022. https://www.joc.com/port-news/us-ports/port-mobile/gulf-coast-import-growth-propels-regional-warehousing-boom_20220202.html?utm_campaign=CL_JOC%20Ports%202%2F9%2F22%20%20_PC00000_e-production_E-126647_TF_0209_0900&utm_medium=email&utm_source=Eloqua

Mongelluzzo, B. Asian imports to US surged to new record in January. Journal of Commerce.  Feb 23, 2022 

https://www.joc.com/maritime-news/container-lines/asian-imports-us-surged-new-record-january_20220223.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%202%2F24%2F22%20NONSUBSCRIBER_PC015255_e-production_E-128466_KB_0224_0617

Mongelluzzo, B. Coming LA-LB cargo surge to rebuild vessel backlog, say terminals. Journal of Commerce. March 10, 2022. https://www.joc.com/port-news/us-ports/coming-la-lb-cargo-surge-rebuild-vessel-backlog-say-terminals_20220310.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%203/11/22%20NONSUBSCRIBER_PC015255_e-production_E-130350_KB_0311_0617

Reminder: ask your MC & Senators to sponsor tree-restoration bill!

The Invasive Species Prevention and Forest Restoration Act (H.R. 1389) is before Congress. It is co-sponsored by Reps. Peter Welch [VT], Ann Kuster and Chris Pappas [NH], Chellie Pingree [ME], Elise Stefanik and Antonio Delgado [NY], Brian Fitzpatrick [PA], Mike Thompson [CA], Deborah Ross [NC].

Ask your Member of Congress/Representative to co-sponsor this bill. Ask your Senators to sponsor a companion bill.

In summary, this bill will:

  • Expand USDA APHIS’ access to emergency funds to eradicate or contain newly detected pest outbreaks.
  • Establish a pair of grant programs to support strategies aimed at restoring tree species decimated by non-native plant pests or noxious weeds. Such strategies include biological control of pests and enhancement of a tree host’s pest resistance.
    1. One grant program supports research to explore and develop these strategies.
    2. The second program support application of resistance breeding and other measures to restore forest tree species. Funded programs must incorporate a majority of the following components: collection and conservation of native tree genetic material; production of sufficient numbers of propagules; preparation of planting sites in the species’ former habitat; planting and post-planting maintenance.
  • Mandate a study to identify actions to overcome the shortfall of mission, leadership, and prioritization; identify agencies’ expertise and resources; improve coordination among agencies and with partners; and develop national strategies for saving tree species.

Organizations eligible for these grants include federal agencies; state cooperative institutions; colleges or universities offering a degree in the study of food, forestry, and agricultural sciences; and nonprofit entities with non-profit status per §501(c)(3) of the Internal Revenue Code.

Endorsements: Vermont Woodlands Association, American Forest Foundation, The Association of Consulting Foresters (ACF), Audubon Vermont, Center for Invasive Species Prevention, Ecological Society of America, Entomological Society of America, Maine Woodland Owners Association, Massachusetts Forest Alliance, National Association of State Foresters (NASF), National Woodland Owners Association (NWOA), The Nature Conservancy (TNC) Vermont, New Hampshire Timberland Owners Association, North American Invasive Species Management Association (NAISMA), Pennsylvania Forestry Association, Reduce Risk from Invasive Species Coalition, The Society of American Foresters (SAF), and a broad group of university professors and scientists.

Legislative Point of Contact: Alex Piper, Legislative Assistant, office of Rep. Welch. Contact me – providing your email! – if you wish me to send you Alex’ contact information.  [The “contact” form does not provide your email and I will not reply in a public way.]

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

Canada Again: Report on Forest Pest Efforts

Asian longhorned beetle

I recently posted a blog based on a new report evaluating Canada’s invasive species efforts across all taxa (see Reid et al. reference at the end of this blog). That report focused on federal measures aimed at preventing introductions, including cross-border introductions from the U.S. After posting that blog I learned about a second report by Allison, Marcotte, Noseworthy and Ramsfield (2021) that focuses on non-native insects and diseases that threaten Canada’s trees and forests.

The first report’s authors lamented that invasive species responsibilities are divided among several agencies, depending on the associated commodity or resource. It noted claims by the Canadian government in its 2018 report to the Convention on Biological Diversity to have identified priority pathways. These included several relevant to forest pests: shipping, horticulture, transport containers, and recreation. The Government claimed that the wood packaging, forestry products, and plant products pathways were at least partially regulated and also that national plans had been developed for several priority species, including the Asian subspecies of Lymantria dispar and the emerald ash borer. Reid et al. (2021) included four case studies, two of which dealt with non-native forest pests: successful eradication (the second time around) of the Asian longhorned beetle (ALB); and the high threat to Canada posed by the spotted lanternfly (SLF).

I was pleased to learn of the second 2021 report (Allison, et al.) because of its focus on pests in trees and forests. This is important because U.S. and Canada share four types of forests; evergreen needle leaf forests, sparse trees/parkland, mixed broadleaf / needle leaf forests, and deciduous broadleaf forests (See Fading Forests III, Chap. 1, Fig. 1, Box 2 [link at end of this blog]). Together, North American forests are comprised of 1,165 different native tree species. They offer many potential hosts to any introduced insect or pathogen. Thus the two countries need to coordinate pest-prevention and responses when prevention fails.

It is more likely that a pest from overseas will be introduced first to the U.S. because the U.S. imports a much higher volume of goods and has greater variety of climates and forests. Data show this: as of 2010, more than 181 exotic insects that feed on woody plants were established in Canada (USDA 2009) compared to at least 475 in the U.S. (Aukema et al. 2010). However, sometimes a pest is first introduced in Canada, then spreads south. One examples is beech bark disease.

areas affected by beech bark disease

Forests occupy 40% of Canada’s land cover. Because the country is so large, there are a wide variety of ecozones and forest regions. Almost all (95%) regenerate naturally; 90% are publicly owned (federal and provincial). These forests, which equate to 9% of the Earth’s total forest area, are important not just to Canadians but also to the world for water regulation, carbon sequestration, habitat for biological diversity and the economy. (See Fig.1 in Allison et al.)

Both Canadian reports emphasize Canada’s international obligations, especially under the Convention on Conservation of Biodiversity (CBD). (This is far more than in similar U.S. reports; of course, the U.S. is not a party to this convention.) Allison et al. also mention the Montréal Process Working Group on Criteria and Indicators for the Conservation and Sustainable Management of Temperate and Boreal Forests; international carbon emission mandates and agreements; and the International Plant Protection Convention (IPPC). Allison et al. also stress the importance of international collaborative research, mentioning the International Forestry Quarantine and Regulations Group (IFQRG), expert groups convened by the IPPC and North American Plant Protection Organization (NAPPO), Plant Health Quadrilateral Working Group and participation in the annual USDA interagency research forum on invasive species.

Both new Canadian reports focus on federal efforts, especially those of the Canadian Food Inspection Agency (CFIA), Canada Border Services Agency (CBSA), and Canadian Forest Service of Natural Resources Canada (CFS).

CFIA (Canada’s national plant protection organization, or NPPO in IPPC parlance) is responsible for analyzing risks, setting policy, and managing responses to forest biosecurity incursions. Its authority comes from Canada’s Plant Protection Act and Regulations (S.C. 1990, c. 22). The CBSA enforces regulations at most, but not all, international ports of entry. The CFS conducts research and analysis to support development and implementation of phytosanitary regulations. CFS is also charged with maintaining market competitiveness for forest products and meeting the country’s global commitments to sustainably develop its natural resources. The 10 provinces and three northern territories have jurisdiction over most of the country’s forests, including promoting forest health. Thus, protection and management of Canada’s forests is a shared responsibility among federal, provincial, territorial, municipal and indigenous governments and other stakeholders including the forest industry and non-governmental organizations.

Allison et al. (2021) discuss the effects of invasions on forest ecosystems, including altering forest ecology and the ability of forest ecosystems to provide services.  For instance, invasions can change: competitive interactions among tree species; forest food webs; microenvironments; nutrient cycles; successional trajectories; understory plant communities; transpiration rates; water dynamics; and nitrogen and carbon flows, including carbon sequestration and storage. They cite the emerald ash borer and Dutch elm disease as examples.

Much of these authors’ discussion of invasion processes, bioinvaders’ impacts and biosecurity procedures is familiar from a U.S. point of view. However, I appreciate that they explicitly concede knowledge gaps in three particular situations. 

First, when discussing the absence of recognized ecological impacts associated with most introduced forest insects and pathogens, they state that this lack of known impacts is often likely due to an incomplete understanding of complex phenomena and delays in perception of effects. They cite — again — the case of the emerald ash borer, when impacts were reported only 10 years or more after its establishment.

Second is their discussion of the drivers of invasion. After saying that the interactions of species’ traits, introductory pathways, and receiving habitats are incompletely understood, they note that it is difficult to determine the relative contribution of reproductive traits and propagule pressure in explaining the invasion success of Hemiptera – which reproduce asexually but are also extremely common in invasion pathways. As I have said in a previous blog, the report due by Haack and colleagues later this year can help clarify the current contribution of the wood packaging pathway to propagule pressure.

Third, they note the limited predictive power of border interception records – and possibly other data resources such as risk assessments, surveillance programs – as a basis for understanding pathways. They note that Ips typographus has been intercepted hundreds of times by North American authorities but has never established.

There are some puzzling gaps in Allison, et al. For example, in discussing the costs associated with forest pest introductions, they do not mention the risk to “leaf-peeper” tourism – as U.S. evaluations of the Asian longhorned beetle do. In discussing the role of propagule pressure they mention pathway volume (i.e., the amount and frequency of trade) and the invasive species’ population levels in the point of origin, but not the invader’s ability to exploit the pathway. (Perhaps that is assumed within the pathway volume measurement?) The example cited is heightened arrivals of Lymantria dispar asiatica during periods of outbreak in its native Asian range.

The report provides helpful clarity on Canadian biosecurity practices.  For example, wood packaging entering the country at thefour main Canadian commercial marine ports (Halifax, Montreal, Vancouver, and Prince Rupert) is inspected by CBSA. However, CFIA enforces compliance at the other marine ports and along the 8,891 km (5,500 mile) land-border with the U.S. Apparently CBSA joins in “border blitzes” at selected strategic land-border crossings.

In discussing the provinces’ efforts to slow the spread of established pests, the report mentions the western efforts to prevent introduction of Dutch elm disease. Also, it covers British Columbia’s attempt to prevent spread of balsam woolly adelgid (BWA) into its interior. (I mourn that this effort was not successful.)

elm zigzag sawfly; photo by Georgy Csoka

The report cites the first record of the elm zigzag sawfly, Aproceros leucopoda in North America as an example of citizen detections.

I note that a periodic reassessment of the Canadian regulations governing emerald ash borer in 2014 resulted in a decision to expand the regulated area to reduce regulatory burden, increase awareness of the regulated areas, and maximize compliance. I regret that USDA APHIS decided to fully de-regulate the pest instead. Canada similarly expanded the regulatory zone for a second pest, the brown spruce longhorned beetle (in 2015).

Canada has deregulated pests judged to have spread to the limits of their potential invaded range, e.g., the Pine shoot beetle, Tomicus piniperda.

The report includes three case studies.

whitebark pine infected by WPBR; photo by FT Campbell

White Pine Blister Rust Case Study

This pathogen was introduced more than 100 years ago and has caused extensive damage to Canadian populations of the commercial species western white pine (Pinus monticola) and eastern white pine (P. strobus). It has also contributed to the decline of whitebark pine (P. albicaulis) and limber pine (P. flexilis). An early warning by Gussow (in 1916) about the pathogen’s probable impact apparently led Canada to prohibit further imports of five-needle pines.

Multiple consequences followed the pathogen’s spread. These included reduced volumes of eastern and western white pine due to the combined effects of disease-caused mortality and foresters shifting to alternative species to avoid future losses. Furthermore, both whitebark and limber pines have been ranked as “Endangered” by the Committee on the Status of Endangered Wildlife in Canada. Whitebark pine is also listed on Schedule 1 of the federal Species at Risk Act; limber pine is currently under consideration for such listing.

The report concludes by stating that resistance breeding is an important strategy and that extensive work has been carried out by both the U.S. and Canada.

Asian Longhorned Beetle Case Study

After noting that this pest of maples is of high concern in Canada, the report lays out the history of the first and second detections in Toronto. Because a risk assessment had been completed beforehand, actions could be taken rapidly. CFIA was encouraged to pursue eradication by the success of several previous eradication efforts, as well as the significant negative impact anticipated to the economy. The Ontario Ministry of Natural Resources, NRCan-CFS, the cities of Toronto and Vaughan, and regional authorities and USDA – which shared information – all contributed. The program removed 5,000 trees in the first six months. In 2018, after five consecutive years of no detections, the ALB was declared eradicated. However, four months later, another ALB was reported – two km from the boundary of the first regulated area. The program was renewed, over a larger area. In total, considering both detections, more than 36,000 trees have been removed. Eradication of the pest following the second detection was declared in June 2020.

The report attributes success to 1) surveillance following IPPC guidelines; 2) reliance on science for evidence-based decision-making, including input on several issued by a science committee chaired by CFS; 3) Early engagement of partners and proactive communication that increased public awareness and reporting.

Oak Wilt Case Study

Oak wilt is established in U.S. states that border Ontario. Also, models suggest that the disease (Bretziella fagacearum) would not be limited by Canada’s climate and that it would cause serious economic harm. Therefore, Canada considers it to pose a high risk. Consequently, CFIA led development of a response framework to guide an incursion response. As usual, this was done in collaboration with representatives from federal, provincial and municipal governments, plus New York and Michigan.

Oak wilt is regulated under the federal Plant Protection Act. In addition to adopting various CFIA directives related to imports, the framework includes a communication strategy; provisions for detection and monitoring; and management activities aimed at its eradication if it is discovered in Canada. The framework also identified research needs and provided funding to address them.  map from photos

In their conclusion, Allison, et al. (2021) state that the close relationship between CFIA and CFS is unusual but do not explain how or why.

They also suggest several ways Canada’s biosecurity efforts targetting non-native forest pests could be improved. The first is to increase information-sharing between agencies and with partners, plus better integration of information into policy development. Also, they suggest adoption of new post-border technologies (e.g., “smart” technologies; in-field chemical analyses, and optimization of surveillance programs) and additional research to build a stronger scientific understanding of pest biology, epidemiology, and trade economics. Generally, their recommendations do not overlap with those of Reid et al. (2021) – which was probably being written at the same time. They do both seem to suggest: 1) strengthening partnerships with the public and Indigenous communities and 2) to be prepared to adapt to future conditions.

SOURCES

Allison JD, Marcotte M, Noseworthy M and Ramsfield T (2021) Forest Biosecurity in Canada – An Integrated Multi-Agency Approach. Front. For. Glob. Change 4:700825. doi: 10.3389/ffgc.2021.700825 Frontiers in Forests and Global Change July 2021 | Volume 4 | Article 700825

Aukema, J.E., D.G. McCullough, B. Von Holle, A.M. Liebhold, . Britton, and S.J. Frankel. 2010. Historical Accumulation of Nonindigenous Forest Pests in the Continental United States. BioScience • December 2010 / Vol. 60 No. 11 www.biosciencemag.org

Reid CH, Hudgins EJ, Guay JD, Patterson S, Medd AM, Cooke SJ, and Bennett JR. 2021. The state of Canada’s biosecurity efforts to protect BD from species invasions. FACETS 6: 1922– 1954. doi:10.1139/facets-2021-0012 Published by: Canadian Science Publishing connorreid@cmail.carleton.ca  

United States Department of Agriculture, Animal and Plant Health Inspection Service. 2009. Risk analysis for the movement of solid wood packaging material (WPM) from Canada into the US.

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

Interactions of 2 (of 3) Threats to Beech

American beech (Fagus grandifolia) is a widespread and beautiful tree of the eastern deciduous forest. Its range reaches from Nova Scotia to eastern Wisconsin, then south to Mississippi and Louisiana and east to mid-Georgia. It is an important food source for 40 wildlife species, particularly in the northern parts of its range where few other species produce hard mast. (See Lovett et al. 2006.)

Threats

Unfortunately American beech is under threat from three non-native organisms or complexes: 1) beech bark disease, 2) beech leaf disease, and 3) beech leaf mining weevil. A fourth pest, a previously unknown – and still unnamed bark beetle in the genus Agrilus – has been detected in New York City on European beech trees. It is not yet known whether it will attack American beech and, if so, whether it will also cause serious damage (Michael Bohne, USFS, pers. comm.)  

symptoms of beech bark disease; photo by Linda Haugen, USFS; via Bugwood

Beech bark disease (BBD) results from the interaction of the introduced European beech scale insect (Cryptococcus fagisuga) and several fungi in the Neonectria genus – some of which are also introduced. The resulting disease has been killing American beech trees since the beginning of the 20th Century. It has spread from Nova Scotia to much of the tree’s range. It has dramatically altered the composition and structure of stands containing beech.

symptoms of beech leaf disease; photo provided by Jennifer Koch, USFS

Beech leaf disease (BLD) was initially detected in 2012, near Cleveland. As of December, 2021, it has spread due east across New York, Pennsylvania, and New Jersey to the Atlantic, then up the coast through Connecticut and eastern Massachusetts, with a separate outbreak in central Maine. The disease is apparently associated with a nematode, Litylenchus crenatae ssp. mccanni, although additional pathogens, like bacteria, might also play a role. The origin of the North American population of the nematode is unknown; it is a related but separate subspecies from a Japanese nematode (Reed et al. 2022).

American beech defoliated by leaf mining weevil; photo courtesy of Jon Sweeney, CFS

Beech leaf mining weevil (Orchestes fagi) is, so far, limited to Nova Scotia. However, it is expected that the weevil will continue spreading throughout the range of American beech through both natural dispersal and human-assisted movement. Repeated defoliation by the weevil might increases mortality rates in forests that are surviving in the “aftermath” stage of BBD (Sweeney et al. 2020).

A new study (Reed et al. 2022) concludes that, despite being detected only 10 years ago, BLD has already become pervasive in forests surrounding Lake Erie in the U.S. and Ontario. While somewhat more prevalent in U.S. states on the eastern side of the Great Lakes (on 54% of trees) than in Ontario (on 46% of trees), BLD is spreading rapidly and affecting every canopy layer. Mortality is highest in seedlings and saplings; understory saplings die within 2 – 5 years. The occasional mortality of overstory trees occurs within seven years of [observed] infection. Defoliation and mortality of saplings allow more light to pass through to the understory; this is expected to alter plant communities on the forest floor.

Beech scale is more widespread in Ontario (found on 60% of trees) than in the U.S. (38% of trees). This is not surprising since the scale was detected in Ontario in 1960, 24 years before it was detected in portions of Ohio, New York and Pennsylvania included in the study (in 1984). Throughout this region, beech scale is disproportionately affecting overstory trees.

Only 4% of trees throughout the study area are infected with Neonectria cankers. In other words, full-scale beech bark disease is not yet widespread and is spreading surprisingly slowly. Scientists do not understand this phenomenon.

These findings are based on a network of monitoring plots a network of monitoring plots set up in 2019 set up in areas surrounding the Great Lakes. They comprise 34 plots at 17 locations in southwest Ontario and 30 plots at 25 locations in Ohio, Pennsylvania, and New York. In total the plots hold 646 live American beech trees — 412 saplings; 85 in the intermediate/suppressed (subcanopy) category; and 149 in the dominant/codominant (canopy) class.

Forest composition is similar throughout the study area. The most common species in association with American beech are sugar and red maples (Acer spp.), and white and green ash (Fraxinus spp.). Other tree species present include eastern hemlock (Tsuga canadensis), white pine (Pinus strobus), oaks (Quercus spp.), and birches (Betula spp.). Study plots had few invasive plants – although the invasive species present are well-documented to invade forests.

Ontario disease assessment

In Ontario, BLD was identified in 25 of the 34 plots.  It was present on 171 saplings, 53 intermediate trees, and 70 dominant trees. Both prevalence and severity were greatest on intermediate trees. Beech scale was present at all 34 plots. While scales were found on trees of all sizes, they were almost two times more prevalent and were more severe on mature trees than saplings. Neonectria cankers were detected at 34 plots. Neonectria was rare but most severe on dominant trees. Fewer than one third of saplings and one-sixth of mature trees were pest free.

U.S. disease assessment  

BLD was present in 17 of the 30 plots. It was found on 75 saplings, 30 intermediate trees, and 38 dominant trees. Saplings and dominant trees had similar levels of disease; intermediate trees had significantly less. However, BLD severity was twice as high on saplings compared to mature trees. BLD was present on more than half of the seedlings assessed – 46 out of 82. Beech scale was present in 20 of the 30 plots. It was significantly less common and severe on saplings than on mature trees. Neonectria cankers were present in only 4 of 30 plots. Canker prevalence and severity did not differ significantly among size classes.

Distribution and Effects of Beech Scale and BBD

While beech scale attack facilitates invasion by the Neonectria fungi, the disease – BBD complex – had the most limited distribution of the three pests in this study. It was found on only ~4% of beech trees throughout the study area. The disease was first reported there in the early 2000s. Although no one knows why, it has spread more slowly there than in areas to the east (Reed et al. 2022).

As is the case with beech scale, BBD disproportionately affects large diameter trees. Typically, BBD kills more than half of mature beech within 10 years of its arrival. Dying trees produce prolific root sprouts resulting in dense beech sapling understories that impede regeneration of less shade-tolerant tree species. The persistence of thickets of disease-vulnerable small beech perpetuates the disease. BBD is the only forest disease in North America that can inadvertently intensify itself by increasing densities of its host while suppressing other species.

Beech Forest Community Change in Response to Combined Impacts of BBD and BLD

It is unclear how forests will change as beech die. Some expect saplings of species already present — red maple, white ash, and, especially sugar maple — to exploit the canopy gaps. Of course, white and green ash are under attack by the emerald ash borer; DMF their ability to reach the canopy will depend on the success of biocontrol agents.

However, if BBD or BLD resistant beech survive or if BLD fails to persist, future forests might instead consist of beech thickets that would prevent all but the most shade tolerant species from establishing. Heavy deer predation on maple seedlings and saplings might also play a role. A third possibility is that morbidity from BBD and BLD might lead to uneven-aged conditions that allow younger trees — perhaps even shade intolerant species e.g., oaks — to establish.

Invasive plants also have the potential to fill gaps left by declining beech. While maple-beech forests often have sparse understories due to low understory light levels, pest-caused canopy gaps are expected to increase the abundance of invaders, especially in small woodlots and forests near urban areas. Several shade-tolerant invasive shrubs are already present in low numbers: Japanese barberry (Berberis thunbergii), tatarian honeysuckle (Lonicera tatarica), multiflora rose (Rosa multiflora), and buckthorn (Frangula sp.). Reed et al. (2022) note that these species, plus privet and autumn olive, can take advantage of small canopy gaps, especially when soils are disturbed, e.g., by active intervention to counteract the loss of beech.

Precautionary Research and Management

Reed et al. (2022) call for enhanced monitoring of beech forests focused on

  • the timing of BLD presence relative to tree age and size – which might affect competitiveness of sprouting beech in the understory; and
  • compositional and structural change in forests with BLD or to which it is likely to spread

They also recommend abandoning the management approach for BBD currently recommended by foresters. It calls for removing scale-susceptible beech so that resistant genotypes increase in prevalence. In forests with both BBD and BLD, they conclude, management of natural regeneration is unlikely to succeed because BLD will kill sprouts and saplings that might be resistant to scale. They recommend instead active management of the forest to promote mast-producing, shade intolerant species, such as oaks and hickories.

They also recommend increased support for resistance-breeding programs. Such programs already target BBD, based on the estimated 1% of American beech that show some resistance. Now those programs need to incorporate BLD resistance. (Reed et al. note that small numbers of beech show few or no BLD symptoms so might possess resistance or tolerance.)

grafted beech for resistance breeding; photo by Rachel Kappler, then USFS (now Great Lakes Basin Initiative & Holden Arboretum)

Unfortunately, the Canadian beech breeding program’s future funding is highly uncertain. To counter this threat, in part, Reed et al. (2022) suggest cryopreserving beech embryos from Canada to develop a beech conservation collection that would be available for a more robust, future Canadian breeding program. The USFS is trying to develop methods to screen trees for resistance to BLD, specifically to the nematode (J. Koch, USFS, pers. comm.)

Another approach would actively manage beech stands in which potentially BLD-resistant beech grow to help these trees reach the canopy and reproduce. In the absence of management, any BLD-resistant beech seedlings might be overtopped by faster growing, shade-intolerant species – especially if the gaps promote soil drying or sun scald.

Finally, breeding programs need to factor in the beech leaf mining weevil, DMF which — as I noted in the beginning — is spreading across Nova Scotia and could spread to the rest of the native range of beech (Sweeney et al., 2020).

The Department of Agriculture has created a website on the Department’s plant-breeding efforts. It includes a subwebsite on USFS efforts. However, I did not find much useful information there.

SOURCES

Lovett, G.M., C.D. Canham, M.A. Arthur, K.C. Weathers, and R.D. Fitzhugh. 2006. Forest Ecosystem Responses to Exotic Pests and Pathogens in Eastern North America. BioScience Vol. 56 No. 5 May 2006)

Reed, S.F., D. Volk, D.K.H. Martin, C.E. Hausman, T. Macy, T. Tomon, S. Cousins. 2022. The distribution of beech leaf disease and the causal agents of beech bark disease (Cryptoccocus fagisuga, Neonectria faginata, N. ditissima) in forests surrounding Lake Erie and future implications Forest Ecology and Management 503 (2022) 119753

Sweeney J.D., Hughes, C., Zhang, H., Hillier, N.K., Morrison, A. and Johns R. (2020) Impact of the Invasive Beech Leaf-Mining Weevil, Orchestes fagi, on American Beech in Nova Scotia, Canada. Frontiers in Forests and Global Change | www.frontiersin.org 1 April 2020 | Volume 3 | Article 46

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

Breeding Pest Resistance in Trees – Thoughtful Perspectives

Scott Schlarbaum collecting butternuts; photo by F.T. Campbell

I have blogged several times about the need to enhance efforts to breed trees resistant to the most damaging of the hundreds of introduced insects and pathogens. Others concur – see reports by the National Academy of Sciences in 2018; several publications by USFS scientists Richard Sniezko and Jennifer Koch; a workshop hosted by Purdue w/ USFS support, the creation and efforts of several consortia – Whitebark Pine Ecosystem Foundation, Great Lakes Basin Forest Health Collaborative, Forest Restoration Alliance …

Also, Richard J. A. Buggs, of the Royal Botanic Gardens, Kew, recently summarized barriers to tree breeding. It was published as an especially thoughtful editorial in Plants People Planet in anticipation of the International Year of Plant Health in 2020 (see reference at the end of this blog). That issue included several related articles, also noted below.

 R.J.A. Buggs’ Perspective on Tree Breeding

Buggs says the need for tree resistance research is greater than ever before. First, damage caused by introduced insects and pathogen is rising. Plus, we now recognize trees’ importance in capturing atmospheric carbon. He sees encouraging signs of growing public awareness of both factors. Also, he thinks citizen science might reduce the cost of some activities … although he doesn’t name which they are.

Dr. Buggs lists six major hindrances to breeding programs, including some aspects that I, at least, have not considered:

1) Trees’ size and long generation times mean research is necessarily slow. One result is it is hard to formulate research proposals that match funding cycles. This in turn means a dependence on long-term institutional commitment from well-funded organizations, and such institutions are rare.

I point out that the U.S. government – especially the USFS – is one such institution. Unfortunately, it has so far been reluctant to take commit major resources to breeding pest-resistant trees. Every year I urge you to lobby Congress on appropriations for the agency. In this context, do you understand that while the USFS Research budget receives approximately $300 million each year, less than $5 million of that total is allocated to researching invasive species (of all taxa)? Some gaps are filled by projects funded by the Forest Health Program. You will have a new opportunity to lobby Congress for Fiscal Year 2023 in the spring!

2) On the other hand, reliance on long-term institutional funding shelters projects from multidisciplinary peer-review that could introduce improved technology or methods. This lack of peer review also contributes to a perception among other scientists that tree resistance research is a scientific backwater.

3) Similarly, studies requiring a long time horizon don’t fit publication schedules. Again, the result is that the findings often appear only in institutional reports or conference proceedings. This means they are hard to find and often lack external peer review at not only the proposal stage but also before publication.

4) The long decades without clear success in dealing with Dutch elm disease (but see recent encouraging developments here) and chestnut blight (see The American Chestnut Foundation here) gave the impression that resistance breeding of forest trees is impossible. Buggs says pest resistance problems are easier to tackle for other trees.

TACF American chestnut; photo by F.T. Campbell

5) Those considering what efforts to fund might demand complete resistance to the pest. This goal is not only unrealistic – it is often unnecessary. Often lower levels of resistance or tolerance can result in trees that can be self-sustaining. Dr. Sneizko concurs; see his article appearing in this issue.

6) Forest stakeholders differ over the goal of developing resistant trees. Some think any human intervention is unwarranted in wilderness areas. Some want a tree as similar as possible to pre-epidemic trees. Others want a tree that produces more timber.

Other Significant Articles

A second article in the same issue of Plants People Planet (Federman and Zankowski) discusses the USDA’s commitment to new approaches in tree resistance research.

I found a third article that discusses British approaches to mitigating tree pests to be more informative than Federman and Zankowski – although somewhat worrying. Spence, Hill and Morris praise the U.K.’s Plant Health Risk Register, which they say has enhanced vigilance on possible new pest introductions. However, the authors describe resistance breeding as a strategy to be considered “when a pest has established such that a tree population is unable to recover, and where a genetic basis for resistance is demonstrable in a proportion of the tree population.” Dr. Sneizko, and others – and I!  – call for initiating exploration of the potential for resistance breeding much earlier in an invasion.

A fourth article – by Richard Sniezko and colleagues —  describes encouraging levels of partial resistance to white pine blister rust in two western white pines and evidence for both qualitative and quantitative resistance to Phytophtohora lateralis in Port-Orford Cedar.

Port-Orford test seedlings; photo courtesy of Richard Sniezko

A fifth – by Showalter et al. — reports encouraging levels of resistance to both emerald ash borer DMF and ash dieback in European ash. The authors conclude that a breeding program might be a viable solution to both pests.

SOURCE

Special issue of Plants People Planet for 2020  — the International Year of Plant Health. https://nph.onlinelibrary.wiley.com/toc/25722611/2020/2/1

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