results of EAB infestation; photo by Nate Siegert, USFS
A series of studies by Patrick J. Engelken, M. Eric Benbow, Deborah G. McCullough, Nate Siegert, Randall Kolka, Melissa Youngquist and others examine the status of ash (Fraxinus spp.) in the aftermath of the emerald ash borer (EAB) invasion. Initial studies documented the crash of biomass supporting EAB numbers when the large ash trees died (Siegert, Engelken, McCullough. 2021; full citation at end of blog.) More recent studies have focused on bogs and forests in the riparian areas where ash were especially numerous and arguably ecologically most important. I posted a blog about black ash bogs earlier.
I will focus here on the studies in riparian areas of southern and northwest Michigan. Information about impacts in forests of southern Michigan are from Engelken, Benbow and McCullough (2020); information about impacts in northwest Michigan are from Engelken and McCullough (2020). Full citations for both are at the end of the blog.
All study areas had high ash densities before EAB’s arrival. One study (Engelken and McCullough 2020) found ash densities high in the immediate riparian areas (in one case, a strip reaching 100 meters from the streambank) but scattered in surrounding forests.
In all these study areas, populations of mature (reproductive age) ash crashed within 10-15 years after EAB invasion:
In northeast Michigan, EAB carrying capacity was reduced by 94% – 99%; total ash basal area was reduced by 87 – 97.7% (Siegert, Engelken, McCullough. 2021);
In southern Michigan, more than 85% of the basal area of green (F. pennsylvanica) and black ash (F. nigra) had been killed by 2020. An estimated 96% of the overstory ash phloem area had died, thus radically reducing EAB carrying capacity (Engelken, Benbow and McCullough 2020);
In northwest Michigan, more than 95% of the overstory ash have been killed. (Engelken and McCullough 2020).
The worst impact has been on black ash– which plays such an important ecological role in riparian areas and wetlands and has enormous importance in Native American cultures. In all these study areas, there is no stump sprouting by black ash (Siegert, Engelken, McCullough. 2021; Engelken, Benbow and McCullough 2020; Engelken and McCullough 2020). In three watersheds of northwest Michigan where black ash constituted up to a quarter of the overstory species before the EAB invasion, scientists found no black ash recruits, only eight saplings, and a single seedling.
Green ash (F. pennsylvanica) has survived in much higher numbers – so far. However, this species’ ability to grow into reproductive size is still uncertain. In northwest Michigan, green ash saplings are abundant in canopy gaps created by EAB-caused mortality of mature ash. These saplings had established before the EAB invasion so some call them the “orphaned cohort”. However, there are few seedlings of any woody plant species in these gaps because sedges form such dense mats.
Green ash reproduction faces many challenges before persistence of the species can be considered assured. First, populations of EAB – now reduced by the lack of mature ash to support them – might resurge when young ash grow to larger sizes. It is not yet clear the extent to which introduced biocontrol agents and native predators, e.g., woodpeckers, will protect these trees as they grow to reproductive size. Here, again, green ash has an advantage over black ash. While green ash produce seed at a relatively young age, black ash don’t produce seed until they reach 30–40 years. Even then, they produce seeds only sporadically, with intervals of five or more years.
A second challenge is the lack of seed sources – at least until and unless young trees are able to reach reproductive size.
A third challenge is competition for resources from other plants. The canopy gaps eliminate competition for light for the taller plants, i.e., the existing ash saplings. However, the sapling cohort is not supported by a seedling cohort. There are very few seedlings of all woody plant species (including invasive species!). Seed germination is suppressed by the dense mats of wetland-adapted sedges and possibly the higher water tables (which resulted from reduced evapotranspiration following mortality of the mature trees).
Competition for resources is also a factor in the forests outside the immediate riparian zone. There, ash seedlings sprout, but shade created by lateral ingrowth suppresses their growth. In southern Michigan, Engelken, Benbow and McCullough (2020) note that the forests are apparently transitioning from red oak dominated forests to red maple and black cherry dominated forests. This transition is apparently intensified by forest mesophication resulting from reduced fire frequency, decreasing light availability in forest understories and increasing soil moisture content.
Fourth, while stump sprouting of green ash was noted in southern Michigan, in the northwestern forests all the sprouts died. I have already noted the absence of stump sprouting by black ash at all sites.
Beaver & Green Ash in Northern Virginia
photos of beaver feeding on ash saplings in northern Virginia; photos by F.T. Campbell
In spring 2022 I noticed along one stream in northern Virginia that beavers had cut down green ash saplings; McCullough and Siegert report that this does not appear to be a problem in their study areas.
By December 2022, the beaver-cut trees tried to recover: see the sprouts from a stump [below]. (I think deer or rabbits ate the tips of the sprouts.)
The beavers also continued feeding on the ash — the tree photographed in the spring when it was half-chewed through has now been felled and its branches removed [see below].
Ecosystem Impacts, Especially on Streams
Across much of the upper Midwest, massive ash mortality is causing widespread changes in forest systems.
Riparian forests, i.e., areas adjacent to waterways where periodic inundation occurs, are functionally linked to the aquatic systems. Loss of such a significant proportion of the overstory changes the transfer of energy to adjacent waterways that takes the form of inputs of nutrients from leaf litter and coarse woody debris. Intact forests also stabilize stream banks and maintain channel depth by preventing erosion. Forests moderate temperature of the water. Finally, forests with “coarse woody debris” increase habitat structure. These impacts might be especially important along first order streams, (defined as perennial streams that have no permanently flowing tributaries). These streams are too small to buffer the impacts of major tree loss. The scientists say they are uncertain whether these changes continue to affect larger streams downstream.
Unshaded streams have higher water temperatures that can affect populations of fish, in particular salmonids, by delaying migration, reducing egg viability and increasing egg mortality. Higher temperatures can also alter primary productivity of aquatic algae, potentially increasing eutrophication (Engelken and McCullough 2020).
The scientists expect increasing abundance of coarse woody debris in the forests and streams of northwest Michigan as the 75% of dead ash that are still standing fall. Such debris provides nutrients and habitat for an array of plants and animals, thereby influencing the abundance, activity and species compositions of several ground dwelling insects and seedling establishment. In streams, coarse woody debris provides complex habitat and refuges. It also retains organic matter. Recreationists do find that debris impedes boating.
Loss of ash specifically
As described by Engelken, Benbow and McCullough (2020), and in my earlier blog, ash leaf litter – particularly black ash leaf litter – is highly nutritious. Ash leaf litter has efficient turnover rates and contributes important soil nutrients such as nitrogen, organic carbon and exchangeable cations. Invertebrate communities in headwater streams feed largely on coarse organic material such as leaf litter (Engelken and McCullough 2020). Consequently, loss of the annual influx of ash leaf litter will likely have adverse effects on nutrient availability in riparian forests and adjacent streams.
SOURCES
Engelken, P.J., M.E. Benbow, D.G. McCullough. 2020. Legacy effects of emerald ash borer on riparian forest vegetation and structure. Forest Ecology and Management 457 (2020) 117684
Engelken, P.J. and D.G. McCullough. 2020. Riparian Forest Conditions Along Three Northern Michigan Rivers Following Emerald Ash Borer Invasion. Canadian Journal of Forest Research.
Siegert, N.W., P.J. Engelken, D.G. McCullough. 2021 Changes in demography and carrying capacity of green ash and black ash ten years after emerald ash borer invasion of two ash-dominant forests. Forest Ecology and Management Vol 494, August 2021
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
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.
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
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
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
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.
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
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
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.
One grant program supports research to explore and develop these strategies.
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
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 Lymantriadispar 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
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.
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 Reidet 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
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
I often blog about the forest pest situation in the UK because its scientists provide lots of easily accessible information. This blog has new information on Britain and – I am pleased to add – on Ireland!
European ash trees alongside the Malham Cove path in Yorkshire; KA via geograph
UK Overview
British woodlands cover just 13% of total land area (just over one-third of the European average of 37%). Their value is increasingly recognized—especially their role in combating climate change through carbon sequestration, flood mitigation and urban cooling. Realization of these benefits is driving new policy to increase woodland cover. In 2019–2020, 13,700 ha of new woodland was created in the United Kingdom (Green et al. 2021). The U.K. government has pledged to plant 30,000 ha of broadleaf and coniferous woodland every year as part of its climate change mitigation strategy (Donald et al. 2021). One example, the ‘northern forest’ scheme, involves planting 50 million native trees over 25 years (Green et al. 2021).
Risks Associated with Conservation Plantings or Translocations
On-going conservation planting efforts plus these ambitious new plans prompted Donald et al. (2021) to assess risk that pathogens might be introduced into the environment as a result of putting native plants in natural habitats. The focus of their study is the planting of common juniper (Juniperus communis)in habitats throughout England, Scotland, and Wales.
Juniper killed by Phytophthora austrocedri; photo from British Forest Research
Juniper is one of three native conifers in the UK. It has been in decline for decades, which conservationists hope to reverse. Now, though, juniper populations are experiencing significant mortality from disease. In 2012, the causal agent was determined to be the non-native pathogen Phytophthora austrocedri. The presence of a single genotype in ~60 geographically separate locations across Scotland and England lends support to idea that the pathogen is being introduced to these sites through some human mechanism. Phytophthora austrocedri has not [yet] been detected in Northern Ireland.
British and European nurseries have contained many Phytophthora pathogens. Hence Donald et al. (2021) sought to determine whether the pathogen is being introduced through use of nursery stock in these well-intentioned plantings. (I have blogged about a similar problem in California restoration plantings.)
The authors found that 19% of P. austrocedri detections are within 2 km of a known planting. The more frequently junipers were planted at a site, and the more cuttings planted during each planting effort, the higher the likelihood that nearby junipers would be infested by P. austrocedri. They conclude that transplanting material is a significant risk pathway for the introduction of disease. The key factor appeared to be the origin of the material. A higher percentage of stock at sites with P. austrocedri outbreaks for which data were available had been raised in a central location by the organization doing the planting or obtained from commercial nurseries. No P. austrocedri was detected in Wales. There, unlike in Scotland or England, the majority of plants were sourced from a commercial nursery that only grew juniper collected from Welsh populations and did not trade with other retailers.
Planting juniper has risen rapidly since the mid-1990s. The highest percentage of planting events co-occurring with disease outbreaks were conducted in 2000–2009. It is likely that there is a time lag between planting and disease detectability. If so, the even larger planting effort since 2010 probably will produce many more P. austrocedri outbreaks that will become visible in the future. And that might not be the end. Planting guidelines have been revised based on pathogen detection. However, the entities doing the planting have not changed their approach, especially regarding site selection.
Donald et al. (2021) also found serious data gaps in these programs beyond the health of propagules. They found:
1) very incomplete knowledge of which organizations are doing the planting;
2) poor attention to traceability of source material; and
3) very little follow-up to check the success of planting projects.
The authors concluded that planting projects have had mixed success in restoring juniper populations. They called for changes in planting strategy to reduce the risks of pathogen introduction.
They also note that efforts to slow the spread of P. austrocedri – for which there is no treatment – are more expensive and less likely to succeed than measures aimed at ensuring that nurseries are free of Phytophthoras. California native plant nurseries have shown that nurseries can maintain Phytophthora-free stock.
Risk of Nursery-spread Pathogens & Willingness of UK Nurseries to Adopt BMPs
Great Britain has experienced an accelerating series of Phytophthora outbreaks and disease epidemics affecting British trees. Introductions detected just since early 2000s include P. ramorum, P. kernoviae, P. lateralis, P. austrocedri and P. pseudosyringae. In all the above cases, imported planting material either is confirmed or strongly implicated as the likely route of intro (Green et al. 2021).
To address this threat – and with massive planting projects proposed – in 2016 the British forest research entity initiated the multidisciplinary ”Phyto-threats” project. Its goal was to understand the drivers of rising Phytophthora infestations and opportunities for mitigating them. The project:
(i) examined Phytophthora distribution and diversity in different nursery management systems;
(ii) assessed the social and economic feasibility of a nursery accreditation programs to curb the risk; and
(iii) identified Phytophthora risks by modelling introduction, establishment and spread of species in relation to biological characteristics, environmental factors and trade flows.
The assessment of Phytophthora presence in nurseries involved collecting 3,624 water and root samples from 163 host genera growing in plant nurseries across the U.K. over a three-year period. Sampling was not random but targetted to facilities thought to harbor Phytophthora. About half of the samples tested positive. They identified 63 species of Phytophthora. Among the most commonly detected species are several that are considered pathogenic — P. cinnamomi, P. cryptogea/pseudocryptogea, P. syringae, P. cactorum, P. cambivora, P. plurivora and P. nicotianae. P. ramorum was found in 12 samples; P. lateralis and P. austrocedri were each found in 10 samples. Several Phytophthora species are potential new records for the U.K. (i.e., P. castanetorum, P. palmivora, P. pseudotsugae,P. tentaculata,P. terminalis, P. uliginosa).
They also saw evidence for Phytophthora root infections in newly arrived plants imported from the European Union.
Their finding raised question about whether Phytophthora can be transported in peat-free potting media, that is, coconut fiber or coir.
The widespread presence and the diversity of Phytophthora found in nurseries was linked to high-risk management practices. These included: careless disposal of culled plants, the near presence of trees along nursery boundaries, and, especially, open water sources. [These factors are essentially identical to infection-facilitating factors found by researchers in California, Oregon, and Washington State. See advisory issued by Oregon State University Extension.]
The project also assessed the feasibility of nursery accreditation programs. The authors consulted widely with nursery owners and customers and conducted a cost-benefit analysis. Regarding nursery practices, owners claimed they were already addressing issues related to water storage in enclosed tanks, clean/covered storage of growing media, installation of drains or free-draining gravel beds, raised benches, and tool disinfestation stations. Therefore the new analysis focused on seven other topics: water testing for pathogens; water treatment s; quarantine holding areas for imported plants; composting or incineration of culled plants; boot and vehicle washing stations; and purchase from only trusted or accredited UK suppliers.
The study found that nurseries would support an accreditation program. However, their support required that costs not be “prohibitive”, actions required not be “unreasonable”, the scheme provide a safety net; and that measures exist to deter non-compliance. Nursery staff wanted to see evidence of consumer demand – a willingness to drive farther to buy “clean” plants, or to pay higher prices for them. The cost-benefit analysis reached a worrying conclusion: nurseries would benefit financially from introducing best practices only when the program would prevent introduction of a wider range of pests and pathogens, not only Phytophthoras. Green et al. (2021) note that the overall net benefit to society from nurseries adopting best practices would be much more substantial. That is, healthy trees are important in meeting carbon sequestration goals. They did not explore whether society should subsidize nurseries’ participation in BMP accreditation programs.
Ireland and Northern Ireland
The island of Ireland (Ireland and Northern Ireland) is thought to have fewer plant pests than other European countries due to its island status and because of its national and international phytosanitary regulations. O’Hanlon et al. (2022) do not mention another possible factor: the likelihood that import volumes to Ireland were probably much lower until the recent vitalization of the Republic’s economy.
O’Hanlon et al. (2022) sought to establish baseline information so scientists can track changes as trade increases and the climate changes. Their search of the literature and unpublished sources identified 396 forest pests on the island, including 11 bacteria, 20 oomycetes, 150 fungi and 215 arthropods. They believe these figures are all probably underestimates. At least 44 of the pests or pathogens are probably non-native to Ireland. (Determining original ranges is difficult, especially for pathogens.)
The Republic of Ireland is one of the least forested countries in Europe. Forests cover ~ 11% of the land area. In Northern Ireland, it is even less: ~ 8%. These forests are predominantly plantations of exotic species. In the Republic, Picea sitchensis makes up 51% of the forest area, Pinus contorta another 10%. Other exotic species planted are Picea abies (4%) and Larix kaempferi. In Northern Ireland, ~ 62% of the forest area is composed of conifer mixtures. Planting of P. sitchensis has accelerated recently, probably as a result of removal of ash and larch because of their vulnerability to pests already established on the island.
Sitka spruce plantation in U.K. Adam Ward, Geograph.org.uk
O’Hanlon et al. (2022) note the great vulnerability of these monocultures to pests. They found 51 pests native to Ireland that are associated with non-indigenous tree genera. They are also concerned about pests introduced from other parts of Europe. For example, green spruce aphid (Elatobium abietinum, native to Central and Eastern Europe) is already attacking Sitka spruce. A second pest of spruce, Ips typographus, which is native to much of Europe but not the British/Irish isles, is not yet established on the island. Northern Ireland imports bark and wood from Europe for processing. Ips typographus has been associated with at least one such shipment.
Non-native forest pests and pathogens also threaten tree species native to Ireland. These include:
Dutch elm disease caused by fungi from the genus Ophiostoma vectored by bark beetles of the genus Scolytus. The second outbreak, caused by the more aggressive pathogen 0. novo-ulmi, was detected in Britain in 1965 and in Ireland in 1977. It caused considerable mortality of elms in Northern Ireland throughout 1970s.
Phytophthora ramorumwas recognized as a threat to forests in Europe only in 2010, when extensive mortality of Japanese larch was detected in Britain. The Republic of Ireland has only the EU1 lineage of the species. Northern Ireland has both the EU1 and EU2 lineages – the former only in nurseries.
Phytophthora disease of alder (caused by several Phytophthora species) was confirmed in Ireland in 2001. However, symptoms of the disease were noted as far back as 1995. It is likely that there are many other Phytophthora species present but not yet recorded.
Ash dieback disease (causal agent Hymenoscyphus fraxineus) on European ash (Fraxinus excelsior) has spread across Europe from Poland beginning in the 1990s. It was confirmed on the Irish island in 2012. Authorities made significant attempts to eradicate the disease, but were not successful. It is now recorded in every county in both Northern Ireland and Ireland. Damage to the economy, environment, and society are expected to be large. The Irish government had helped plant more than 13,000 ha of ash between 1992 and 2012. An estimated 2.9 million ash trees are in Northern Irish hedgerows. British scientists say more than 1,000 fauna species are associated with ash trees.
A second pest on ash — ash sawfly (Tomostethus nigritus) — was detected in Northern Ireland in 2016; it has defoliated hundreds of trees in Belfast.
In recent years, forest pest incursions have increased at a relatively steady rate, comparable to other countries, including Britain. In the 1970s, 26 species were reported; in the 1980s, 27; in the 1990s, 16; in the 2000s, 37; between 2010 and 2017, 28. See the graph in Fig. 2
There is a strong link between pest and pathogen findings in Britain and Ireland. O’Hanlon et al. (2021) list 16 insects and pathogens detected in Britain after 1960 which were later detected in Ireland. The list includes H. fraxineus, 0. novo-ulmi, Phytophthora ramorum, and Phytophthora lateralis. The average delay was 10 years. The authors note that the two islands share similar ecological conditions and hosts, are nearby, plus there is substantial travel and exchange of goods between them. For example, in 2018 an estimated 30,000 metric tonnes of conifer roundwood was sent from Scotland to Northern Ireland for processing.
There are very limited physical checks on plants or plant products moving between Ireland and Northern Ireland. The exception is conifer wood that is not bark-free. European Union regulations require that such shipments be accompanied by a plant passport that certifies that the wood has been inspected by a professional operator authorized by the NPPO of the exporting country. What rules will apply now, after BREXIT, remains unclear. Because of concerns about re-igniting sectarian conflict, most political figures want the border on the island to be almost invisible.
The Europhyt database for the period February 2006 – November 2016 documented interception of numerous high-risk pests at the British and Irish borders, including Anoplophora chinensis and A. glabripennis; I. typographus; Monochamus alternatus; H. fraxineus; and P. ramorum O’Hanlon et al. (2021). believe many more go undetected. O’Hanlon et al. (2021) report specifically on detections on commodities from China, especially on wood packaging. One detection on imported plants of interest to me is that of Discula destructiva(dogwood anthracnose). The article does not mention the origin of the shipment. The native British dogwood, Cornus sanguinea, would presumably be vulnerable to this Asian fungus, which has already caused widespread mortality of woodland dogwoods in North America.
Cornus sanguinea; photo by Hans Hillewaert
In addition to reviewing the current situation, O’Hanlon et al. (2021) note pertinent facts about current policy and future science. First, while the two political units on the island have a history of plant pathology expertise, there has recently been a reduction in the number of practicing forest pathologists, mycologists and entomologists. (I and others have complained about the same deterioration in expertise in the United States.)
Second, they describe the years of delay before official recognition that the pathogen Gremmeniella abietina was present in Northern Ireland. This delay resulted from officials refused to accept data from molecular detection tools.
O’Hanlon et al. (2021) add their voice to others criticizing the international phytosanitary system (they cite six major publications: Brasier 2008; Liebhold el. al. 2012; Santini et al. 2012; Eschen et al. 2015; Jung et al. 2016; Meurisse et al. 2019). The failures are (i) visual inspections can miss asymptomatic infections, (ii) limited resources mean only a small proportion of commodities can be inspected, (iii) allowing the use of fungicides masks disease symptoms on plants, (iv) list-based regulations don’t address undescribed organisms and (v) countries vary in how aggressively they carry out the required phytosanitary procedures. O’Hanlon et al. (2021) conclude that “Until these issues are addressed it is likely further increases in the numbers of non-native pests and pathogens of trees will increase.”
The authors note that Eschen et al. (2018) suggested that risk analysis should focus on the commodity (commodity risk assessment) rather than on an individual pest. I have made a similar suggestion, although less clearly worded.
Finally, O’Hanlon et al. (2021) note that climate change is expected to increase the island’s vulnerability to tree-killing pests and pathogens due to fewer frost days, more rain in winter, increased chance of drought in summer, increased average annual temperatures, and more frequent weather extremes. These changes are likely to affect the amount of damage caused by both native and introduced pests organisms. Range shifts in both pests and pathogens and their natural enemies; physiological or behavioral responses in the pests; phenological changes in the hosts; and increased stress on the trees will combine to affect damage.
SOURCES
Donald, F.; Purse, B.V.; Green, S. 2021. Investigating the Role of Restoration Plantings in Introducing Disease—A Case Study Using Phytophthora [UK] Forests 2021, 12, 764
Green, S., D.E.L. Cooke, M. Dunn, L. Barwell, B. Purse, D.S. Chapman, G. Valatin, A. Schlenzig, J. Barbrook, T. Pettitt, C. Price, A. Pérez-Sierra, D. Frederickson-Matika, L. Pritchard, P. Thorpe, P.J.A. Cock, E. Randall, B. Keillor and M. Marzano. 2021. PHYTO-THREATS: Addressing Threats to UK Forests and Woodlands from Phytophthora; Identifying Risks of Spread in Trade and Methods for Mitigation. Forests 2021, 12, 1617 https://doi.org/10.3390/f12121617ý
O’Hanlon, R., Ryan, C., Choiseul, J., Murchie, A.K. and Williams, C. D. 2021 Catalogue of P&P of trees on the island of Ireland. Biology and Enviro
Proceedings of the Royal Irish Academy 2021. Vol. 121, No. 1.12-45 DOI: 10.3318/ BIOE.2021.02
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
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.
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
