area devastated by EAB; photo by Nathan Siegert, USFS
The emerald ash borer (EAB; Agrilus planipennis) is the most damaging forest insect ever introduced. In late June 2022 it was detected in Forest Grove, Oregon — 26 miles from Portland. This is the first confirmation of EAB on the West Coast – a jump of over 1,000 miles from outbreaks in the Plains states. The infested ash trees were immediately cut down and chipped (see Oregon Department of Agriculture website; full link at end of blog). See my earlier blog on EAB’s threat to ash-dominated riparian wetlands in Oregon.
ash-dominated swamp along the Willamette River in Oregon; photo by William Wyatt, ODF
Oregon has been preparing for the EAB:
The state finalized its response plan in March 2021; see reference at end of blog.
The state sought and received funds from USDA APHIS to initiate a biocontrol program. The funds were not from APHIS’ operational budget, but from the agency’s Plant Pest and Disease Management and Disaster Prevention Program (PPDMDPP) (Farm Bill money).
State and federal agencies have begun collecting seeds for resistance screening and a possible breeding program.
EAB: Why Quarantines Are Essential
As you might remember, in January 2021 APHIS dropped its federal regulations aimed at curtailing EAB’s spread via movement of wood and nursery plants. This shifted the responsibility for quarantines to state authorities. Instead, APHIS reallocated its funding to biological control. I raised objections at the time, saying the latter was no substitute for the former.
A new academic study shows that APHIS’ action was a costly mistake.
Hudgins et al. (2022; full citation at end of this blog) estimate EAB damage to street trees alone – not counting other urban trees – in the United States will be roughly $900 million over the next 30 years. These costs cannot be avoided. Cities cannot allow trees killed by EAB to remain standing, threatening to cause injury or damage when they fall.
ash fallen onto house in Ann Arbor, Michigan; photo courtesy of former mayor John Hieftje
The authors evaluated various control options for minimizing the number of ash street trees exposed to EAB. They assessed the trees’ exposure in the next 40 years, based on management actions taken in the next 30 years.
In their evaluation of management options, Hudgins et al. tried to account for the fact that the effect of management at any specific site depends on the effects of previous management. Additional complexity comes from the facts that the EAB is spread over long distances largely by human actions (i.e., movement of infested wood); and that biocontrol organisms also disperse.
They conclude that efforts to control spread at the invasion’s leading edge alone – as APHIS’ program did – are less useful than accounting for urban centers’ role in long-distance pest dispersal via human movement. Cities with infested trees are hubs for pest transport along roads. Hudgins et al. say that quarantine programs need to incorporate this factor.
Hudgins et al. concluded that the best management strategy always relied on site-specific quarantines aimed at slowing the EAB spread rate. This optimized strategy, compared to conventional approaches, could potentially save $585 million and protect an additional 1 million street trees over the next 40 years. They also found that budgets should be allocated as follows: 74-89% of funds going to quarantine, the remaining 11% to 26% to biocontrol.
In other words, a coherent harmonized quarantine program – either through reinstatement of the federal quarantine or coordination of state quarantines — could save American cities up to $1 billion and protect 1 million trees over several decades. Since street trees make up only a small fraction of all urban trees, up to 100 million urban ash trees could be protected, leading to even greater cost savings.
Unfortunately, such a coordinated approach seems unlikely. States continue to have very different attitudes about the risk. For example, Washington has no plans to adopt EAB regulations, despite it being detected in Oregon. To the north, Canada already has EAB quarantines and Hudgins et al. advise that they be maintained.
The authors recognize that quarantines’ efficacy is a matter of debate. Quarantines require high degrees of compliance from all economic agents in the quarantine area. Also they need significant enforcement effort. Some argue that meeting either requirement, let alone both, is unrealistic. However, under Hudgins et al.’s model, use of quarantines was always part of the optimal management method across a variety of quarantine efficiency scenarios. Again, these models point to allocating about 75% of the total budget to quarantine implementation. In all scenarios, reliance solely on biocontrol led to huge losses of trees compared to a combined strategy.
Hudgins et al. asked their model for optimal application of both quarantines and biocontrol agents. For example, quarantine enforcement could focus on limiting entry of EAB at sites that: 1) have many ash street trees, 2) currently have low EAB propagule pressure, but 3) are vulnerable to receiving high propagule influx from many sites. Seattle is a prime example of such a vulnerable city with many transportation links to distant cities with significant ash populations.
On the other hand, quarantine enforcement could strive to limit outward spread (emigration) of EAB from which high numbers of pests could be transported to multiple other locales, each with many street trees and low propagule pressure. These sites would be along the leading edge of the invasion and where the probability of long-distance pest dispersal is high.
Authorities should be prepared to adjust quarantine actions in response to changing rates and patterns of invasion spread.
Biocontrol agents should be deployed to sites with sufficient EAB density to support the parasitoids, especially sites predicted to be hubs of spread.
Hudgins et al. concede that they did not explicitly account for:
1) The impact of uncertainty regarding EAB spread on the model;
2) Alternative objectives that might point to other approaches, e.g., minimizing extent of invaded range, or reducing the number of urban and forest trees exposed to EAB;
3) Impacts of predators, such as woodpeckers, on EAB populations;
4) Synergistic impacts from climate change, which by exacerbating stress on ash trees will probably increase tree mortality from EAB infestations; and
5) Variation in management efficiency depending on communities’ capacities.
In the future, Hudgins et al. hope to test their model on other species to determine whether there is a predictable spatial pattern for all wood boring pests, that is, should quarantines always be focused on centers of high pest densities as probable sources of spread. Determining any patterns would greatly assist risk assessment and proactive planning.
dead ash near major road in northern Virginia; photo by F.T. Campbell
In an earlier study, Dr. Hudgins and other colleagues projected that by 2050, 1.4 million street trees in urban areas and communities of the United States will be killed by introduced insect pests – primarily EAB. 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. They predict that 6,747 communities not yet affected by the EAB will suffer the highest losses between now and 2060. However, they evaluated risks more broadly: the potential pest threat to 48 tree genera. Their model indicated that if a new woodboring insect pest is introduced, and that pest attacks 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. I have blogged often about the rising rate of shipments coming directly from Asia to the American South
SOURCES
Hudgins, E.J., J.O. Hanson, C.J.K. MacQuarrie, D. Yemshanov, C.M. Baker, I. Chadès, M. Holden, E. McDonald-Madden, J.R. Bennett. 2022. Optimal emerald ash borer (Agrilus planipennis) control across the U.S. preprint available here: https://doi.org/10.21203/rs.3.rs-1998687/v2
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
Members of this team published an article earlier that evaluated the threat from introduced woodborers as a group to U.S. urban areas; see E.J. Hudgins, F.H. Koch, M.J. Ambrose, B. Leung. 2022. Hotspots of pest-induced US urban tree death, 2020–2050. Journal of Applied Ecology
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
TACF back-crossed chestnut growing in southern Fairfax County, Virginia
same chestnut, dying in 2021; probably from Phytopthora root rot
In the first half of the 20th Century, American chestnut (Castanea dentata) was functionally extirpated from US forests east of the Mississippi River by chestnut blight, caused by a fungus from Asia, Cryphonectria parasitica. Today, only 10% of the pre-blight chestnut population remains, most as root sprouts less than 2.5 cm dbh (Dalgleish et al. 2015; full citation at the end of the blog).
Volunteer organizations — with recent help from federal and state agencies – have worked for more than a century to develop chestnut trees resistant to the blight. Their aim is to restore the species to the forest. Their decades of hybridization efforts now appear unlikely to produce a highly blight-resistant chestnut with a genome that is predominantly American, so TACF now plans to incorporate the use of transgenic techniques to enhance resistance to the blight fungus.
However, restoration of chestnut requires addressing a second Asian pathogen: Phytophthora cinnamomi, which causes a fatal root disease. Several studies indicate that up to 80% of seedlings are killed. The pathogen is widespread in soils south of 40o North Latitude, which falls just north of the Maryland-Pennsylvania line. Thus, P. cinnamomi occupies the southern half of American chestnut’s former range. Scientists expect this pathogen to move north in response to the warming climate; indeed, some project that the root disease could reach throughout the entire current chestnut rangeby 2080.
historic range of American chestnut
Gustafson et al. 2022 modelled chestnut’s vulnerability to P. cinnamomi to current and expected environmental conditions in two state forests in the Appalachians of western Maryland to evaluate the probable impact of the root disease on efforts to restore the tree species.
They found that root rot greatly reduced chestnut biomass on the landscape, even when resistance to root rot was at the target level for selection of root rot-resistant chestnut families using traditional breeding methods.
Gustafson et al. 2022 recommend that chestnut restoration apply the following strategies:
Locate restoration plantings at latitudes, elevations, and sites where root rot is not expected to be present well into the future. This probably means sites in the Northeastern US and Canada (Burgess et al. 2017)
Enhance the planting stock’s resistance to P. cinnamomi through breeding.
Identify soil conditions, including soil microbes, that suppress the pathogen or protect tree roots.
Since planting stock – both bareroot and containerized – can transmit P. cinnamomi, either raise seedlings in nurseries located outside the pathogen’s current range or rely on direct seeding. These strategies have their own downsides. Restricting locations of nurseries might complicate efforts to ensure seedlings are adapted to local conditions in the restoration area and seeds would need to be protected from seed predators.
The authors specify these additional important conditions:
Planting locations: while Canada is currently outside the range of American chestnut, the same climatic warming that will facilitate northward spread of P. cinnamomi will probably allow the tree to thrive farther north (Barnes and Delborne 2019). Perhaps the tree’s range will shift farther north than the pathogen’s.
Breeding: some resistance to Phytophthora root rot has been found in families providing blight resistance used in The American Chestnut Foundation (TACF) breeding program. TACF now plans to cross individuals from those families with transgenic blight-resistant chestnut to combine both resistances.
Soils: P. cinnamomi is favored by compacted soils with poor aeration or that tend to remain saturated. These include heavy clay soils and those highly disturbed by agriculture or mining. Restoration sites should be non-disturbed, well-drained sites. (This recommendation contradicts others’ proposals that chestnuts be planted on reclaimed mining sites.) Silvicultural management should also minimize environmental stresses.
that dying chestnut trying to reproduce!
Restoring chestnut will be challenging in any case: successful restoration requires chestnut trees that can compete successfully in the forest and adapt to conditions which are now quite different from those a century ago when the species was dominant. These include abiotic factors, e.g., climate and atmospheric CO2 levels; and biotic factors, e.g., different forest pests and invasive plant species.
In an earlier publication, Gustafson and colleagues (Gustafson et al. 2018) modelled the effects of warmer temperature and elevated atmospheric CO2 levels on chestnut’s growth and competition and the tree’s adaptation to natural and anthropogenic disturbances. They concluded that aggressive restoration programs – involving clearcutting, then planting chestnuts – could restore chestnut as an important component of forested ecosystems in the Appalachian Mountains.
However, this earlier study did not consider the effects of Phytophthora root rot. The 2022 study demonstrates that these recommendations are probably applicable only to the northernmost portion of former chestnut range, outside the areas infested by Phytophthora root rot, unless breeding is successful in substantially increasing resistance to root rot.
Several studies indicate American chestnut is highly susceptible to P. cinnamomi; rates of root rot induce mortality of 80% or higher have been documented. TACF has found that hybrid chestnut families selected for root rot resistance have a mortality rate of about 45%. Even with this level of tolerance, the model shows that chestnut could not regain anything approaching its former abundance on the landscape. Since the threat of P. cinnamomi to chestnut restoration has become evident, TACF is assessing how to integrate increased tolerance to root rot into their larger blight resistance breeding program (Westbrook et al. 2019).
Soil properties – texture, land use, drainage, waterlogging, drought, temperature, and water-holding capacity – influence infection. So does weather: a single heavy rain event might saturate soil sufficiently to facilitate a P. cinnamomi infection. For these reasons, climate change is expected to exacerbate its geographic spread and pathogenicity.
The sites used in both studies are at the center of chestnut’s former range, which is also at the northern edge of the root rot pathogen’s range. However, the two sites differ in important ways, especially in rainfall and soils. The researchers considered one a mesic site and the other, xeric.
Their 2022 model showed that root rot caused a dramatic reduction in chestnut biomass on both the mesic and xeric sites. Apparently temperature and wetness levels offset each other. That is, higher soil temperatures intensified P. cinnamomi virulence at the xeric site sufficiently to overcome its relative soil dryness. At the mesic site, soil temperature sometimes dropped to levels that are lethal to Phytophthora. On the whole, then, climate change is expected to intensify P. cinnamomi infection rates on both sites and reduce the number of sites where the pathogen is absent.
Gustafson et al. (2022) discuss several assumptions and data gaps that require further study.
SOURCES
Barnes, J.C. and Delborne, J.A., 2019. Rethinking restoration targets for American chestnut using species distribution modeling. Biodiversity and Conservation, 28(12), pp.3199-3220.
Burgess, T.I., Scott, J.K., Mcdougall, K.L., Stukely, M.J., Crane, C., Dunstan, W.A., Brigg, F., Andjic, V., White, D., Rudman, T. and Arentz, F., 2017. Current and projected global distribution of Phytophthora cinnamomi, one of the world’s worst plant pathogens. Global Change Biology, 23(4), pp.1661-1674.
Dalgleish, H.J., Nelson, C.D., Scrivani, J.A. and Jacobs, D.F., 2015. Consequences of shifts in abundance and distribution of American chestnut for restoration of a foundation forest tree. Forests, 7(1), p.4.
Gustafson, E.J., B.R. Miranda, T.J. Dreaden, C.C. Pinchot, D.F. Jacobs. 2022. Beyond blight: Phytophthora root rot under climate change limits populations of reintro Am chestnut Ecosphere. 2022;13:e3917.
Gustafson, E.J., A.M.G. De Bruijn, N. Lichti, D.F. Jacobs, B.R. Sturtevant, D.M. Kashian, B.R. Miranda, and P.A. Townsend. 2018. “Forecasting Effects of Tree Species Reintroduction Strategies on Carbon Stocks in a Future without Historical Analog.” Global Change Biology 24: 5500–17. https://doi.org/10.1111/gcb.14397
Westbrook, Jared W., et al. “Resistance to Phytophthora cinnamomi in American chestnut (Castanea dentata) backcross populations that descended from two Chinese chestnut (Castanea mollissima) sources of resistance.” Plant disease 103.7 (2019): 1631-1641.
Posted by Faith Campbell
[An earlier version of this blog has now been corrected, with additional sources added. I think Cornelia Pinchot, USFS, for the corrections.]
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
Phytopthora cinnamomi on manzanita in California; photo courtesy of Ted Swiecki/Phytosphere
While I blog often about wood packaging the fact is that imports of live plant [= “plants for planting” in USDA’s terms] have historically posed a higher risk of introducing tree-killing pests. In 2012, Liebhold et al. found that nearly 70% of 455 damaging pests introduced to the continental U.S. as of 2006 had probably been introduced via plant imports. These included 95% of sap feeding and 89% of foliage feeding insects and about half of the pathogens. Imported plants not only carry a greater variety of pests than wood packaging; they also carry many more.
Introductions on imported plants for planting is not a rare event. An analysis of data in the Agriculture Quarantine Inspection Monitoring (AQIM) during 2009 found that the approach rate of pests on imported plants was apparently 12% (Liebhold et al. 2012) — more than 100 times higher than the 0.1% approach rate found by Haack et al. (2014) for wood packaging. This alarming statistic receives less attention than warranted because APHIS objected to the accuracy of other aspects of the study.
APHIS has adopted changes to its phytosanitary system for plants for planting in the decade since 2009. The question is, have these changes reduced the known risks associate with live plant imports – especially given skyrocketing imports? Are more measures necessary? Current data and analyses cannot provide a scientifically valid answer.
ohia rust on endangered Hawaiian native plant Eugenia koolauensis
First, most studies focus on insects – they even exclude pathogens. Among pathogens introduced in recent decades, probably by the plant trade, are several Phytophthoras, rapid ‘ōhi‘a death, beech leaf disease, boxwood blight. (I am assuming that the Fusarium dieback disease vectored by Euwallacea beetles was introduced via wood packaging.) There have been repeated detections of the Ralstonia solanacearum Race 3 biovar 2, a bacterium that attacks a range of herbaceous plants, despite APHIS requiring specific integrated pest management programs in producing nurseries located in Central America. Examples of recently introduced leaf feeders include the European beech leaf-mining weevil and elm zigzag sawfly.
I concede that it is difficult to study introduced pathogens. It is nearly impossible to compile a complete list of introduced fungi and related organisms since only the most damaging are typically detected and their native ranges are frequently undeterminable. However, European forest pathologists are much more active on these questions. Why? What can we do to focus Americans on the threats these organism pose?
Second, most studies analyzing the pest risk associated with plant imports use port inspection data. However, port inspection data are not reliable indicators of the pest approach rate – as explained by Liebhold et al. 2012 and Haack et al. 2014 (as it pertains to wood packaging). Thus, most of the analyses carried out by Liebhold et al. and MachLachlan et al. (2022) are based on the pests found by APHIS inspectors: actionable pests were detected on only 2.6% of the incoming plants that they inspected.
Here I discuss two recent discussions of the risk associated with imported plant for planting. One is an analysis of establishments of one order of insects in the United States over 200 years (MacLachlan et al. 2022; full citation at the end of the blog). Again, the focus is on insects! The other is a discussion of the pathway during the recent annual meeting of the Continental Dialogue on Non-Native Forest Insects and Diseases. link to posting of presentations This discussion raised some of the key questions, although no answers were provided.
U.S. imports of plants have increased by more than 400% since the 1960s; 35% in just the last 15 years (in 2007 the U.S. imported approximately 3.7 billion plants [Liebhold et al. 2012]; in 2021 it was about 5 billion [MacLachlan et al. 2022]. Yet establishments of new non-native insects associated with this pathway have not risen commensurately. MacLachlan et al. (2022) attempt to answer why this is so. However, pests are often not detected for several years or a decade after their introduction. Furthermore, I doubt that an analysis based on inspection data, not the more reliable AQIM data, can provide an accurate assessment.
To clarify the pest risk associated with plant imports, studies of some insect types, excluding pathogens, is not sufficient. Again, APHIS should update the Liebhold et al. study to determine the approach rate for all types of organisms that threaten North American tree species. Any such study should include trees on Hawai`i, Guam, Puerto Rico, and other U.S possessions and territories. These islands are usually excluded from analyses of imported pests, including Liebhold et al. 2012. I concede that there are probably scientific and data-management challenges but these islands are immensely important from a biodiversity point of view, and they are parts of the United States!
Cycas micronesica endemic to Guam; threatened by cycad scale & cycad blue butterfly; photo courtesy A. Gawel
MacLachlan et al. (2022) focused their analysis on the insect order Hemiptera, including the so-called true bugs, including cicadas, aphids, planthoppers, and leafhoppers. This is the insect order most frequently transported with imported plants. In addition, establishments of Hemiptera can be attributed to plant imports rather than to wood or other vectors. Of the 3,500 species of non-native insects established in North America (including the contiguous U.S. states, Alaska, and Canada), about 27% are Hemiptera. Many are serious pests, e.g., hemlock woolly adelgid and balsam woolly adelgid). Complicating the analysis, however, is the fact that some Hemiptera are inconspicuous so they are difficult to detect. In fact, MacLaughlan et al. 2022 estimate the median delay between introduction and detection to be 80 years! They believe that many introduced species remain undiscovered, ranging from 21% for Eurasian regions to 38% for the Neotropics and 52% for Australasia.
eastern hemlocks killed by hemlock woolly adelgied; Linville Gorge, NC; photo by Steven Norman, USFS
MacLachlan et al. (2022) compare the relationship between plant imports and discoveries of Hemiptera from 1800 to the present in an attempt to answer the puzzle of why new Hemiptera establishments have remained relatively steady despite quadrupled plant imports. Perhaps the pool of novel insect species in the source region has been depleted. Or other factors might have changed, such as
the commodities imported (plant species or types; or geographic source)
phytosanitary measures applied by the U.S.
MacLachlan et al. (2022) tracked plant imports since 1854 from seven ecological regions: Afrotropic, Asian Palearctic, Australasia, European Palearctic, Indomalaya, Nearctic, Neotropic. In the early decades, both imported plants and introduced Hemiptera detected in the U.S., came predominantly from European and Asian Palearctic regions. Now, however, almost no new Hemiptera species are being introduced on plants imported from the European and Asian Palearctic regions. Since the 1950s, estimated establishments from the Indomalaya region have remained relatively stable. Establishments from the Neotropic and Afrotropic regions rose following World War II and have remained relatively high. After also declining in the first half of the 20th century, establishments of new species from Australasia have recently increased.
Generally, the regions associated with declining establishments of new species (Eurasia) are experiencing relatively gradual increases in their exports to the U.S. Those regions which contribute relatively steady or increasing establishments (Neotropics, Indomalaya, Australasia, and Afrotropic) have each undergone rapid increases in exports to the U.S.
Establishment Risk Among Regions
Source regions vary in the type of plants they export (e.g., rootless cuttings v. whole plants) and in the volume of exports. They also differ in the composition of their indigenous and introduced insect populations. Imports from areas with an abundance of species capable of establishing and adapted to environmental conditions in North America pose greater establishment risk, although it is challenging to determine the risk associated with individual species.
Establishment risk of shipments from a particular region also changes over time. The number of potential new species of invaders might shrink as more and more arrive in North America. (This situation has no effect on the continued introduction of insect species already established in North America. These reintroductions might arrive in new areas – so expanding the area at risk; or their increasing number contributes to propagule pressure at establishment sites.) Another factor might be phytosanitary policies. Strengthening of phytosanitary measures might suppress the number of organisms that travel with the plant shipment, enter North America, and establish. The opposite might happen if phytosanitary measures are relaxed or if the sourcing or type of imports diversifies in ways that connect additional species in source regions with trade pathways.
Considering all regional plant sources, MacLachlan et al. (2022) estimate that establishments per unit of additional imports – of Hemipterans – have shrunk because of a combination of increased imports, accumulated introductions associated with past imports, and the passage of time. These decreases are substantial – between 75.2% and 99.8% for the various regions from 1962 to 2012. For the Asian Palearctic and Neotropic regions, MacLachlan et al. (2022) determined that depletion of species pools is a contributing factor. Other factors are thought to explain the substantial decline in establishment likelihood for the other regions. However, note the caveats above re: lag times in detecting introductions.
However, despite that significant decrease in risk per unit of imports, the number of establishments has remained relatively constant over the past century. MacLachlan et al. (2022) attribute this pattern to the decreases in marginal risk from additional imports being offset by substantial increases in overall import levels and diversification of the origins of imports across regions, which exposed the U.S. to new source species pools.
MacLachlan et al. (2022) suggest that APHIS should target biosecurity resources to the specific commodity-country pairs associated with a demonstrated higher relative risk of introducing additional insect species.
MacLachlan et al. (2022) are unable to evaluate the efficacy of APHIS’ most important policy change: creation of the “Not Authorized for Importation Pending Pest Risk Assessment” (NAPPRA) program because it was adopted in 2011 and they analyzed data only through 2012. A decade later this policy restricts imports of about 250 taxa (Regelbrugge to Continental Dialogue). It is certainly time to evaluate its efficacy through a new study of pest approach rates in the “plants for planting” trade.
I do not think that U.S. phytosanitary policy should be based on an analysis of just one of at least three types of pests that travel via the pathway. We need analysis of the risk from pathogens, nematodes, viruses … and other orders of arthropods.
The Continental Dialogue on Non-Native Forest Insets and Pathogens
The Continental Dialogue on Non-Native Forest Insects and Pathogens hosted a discussion of the risk of pest introduction via the plant trade during its recent annual meeting. Participants asked: How can the international phytosanitary system curtail introductions of unknown organisms when it is based on risk assessments that address only species that are fully known and – usually – have proven to be invasive elsewhere.
Rhodomyrtos psidioides in eastern Australia killed by myrtle rust; photo by Peter Entwistle
In recent decades, tens of species of Phytophthora have been introduced to countries around the world. Myrtle rust (Austropuccinia psidii) has been introduced to 27 countries from the U.S. to Australia and South Africa. The two causal agents of boxwood blight has been introduced to at least 24 countries in three geographic areas: Europe and western Asia; New Zealand; and North America. The ash decline fungus has been introduced across Europe. Most of these species were unknown to science at the time of their introduction. Other species were known – but not believed to pose a threat because, in their native regions, their co-evolved hosts are not harmed.
For more than a decade, scientists have noted that the international phytosanitary system has failed to prevent this rapid worldwide spread of significant pathogens via the international nursery trade. Examples include Brasier 2008; Liebhold el. al. 2012; Santini et al. 2013; Roy et al. 2014; Eschen et al. 2015; Jung et al. 2015; Meurisse et al. 2019; O’Hanlon et al. 2021.
During the Continental Dialogue discussion, Craig Regebrugge, Vice President of AmericanHort (the principal nursery trade association) noted the economic importance of greenhouse and nursery production and the importance of offering novel plants to their customers. Also, he noted that U.S. retail nurseries import primarily unrooted plant cuttings. In so doing, they have a strong incentive to ensure that they are pest-free in order to avoid delays arising during inspections. Those delays would probably kill these highly perishable products. Most U.S. imports of “finished” plants come from Canada. There have been pest problems; one of the most recent examples is a moth that attacks boxwoods (Buxus), which is the top-selling shrub crop in the U.S. Earlier there was confusion over whether plants shipped from British Columbia had been infected by the sudden oak death pathogen.
Regelbrugge noted that the industry’s voluntary integrated pest management program – Systems Approach to Nursery Certification (SANC) – currently has about two dozen participating nurseries. Hoped-for adoption by more of the hundreds of production nurseries in the country has been delayed by COVID-related travel restrictions, but he hopes to restore momentum. The industry is looking for opportunities to strengthen the program through marketing messages.
Regelbrugge and a second speaker, Rebecca Epanchin-Niell of the University of Maryland, warned that prohibitions on imports will stimulate smuggling. Both raised concerns about direct-to-consumer sales by e-commerce vendors and sought ideas on how to change the behavior of both exporters and consumers.
Later Sarah Green of British Forest Research asked the APHIS representativewhether the agency’s import procedures are working to prevent introductions. She pointed to the issues raised by the scientific sources I cited above: pest risk analyses address only known organisms, so this process cannot protect importers from unknown organisms. She noted that the United Kingdom is struggling to contain a number of introductions of previously unknown pathogens. Gary Lovett of the Cary Institute noted that this weakness of pest risk assessments also hampers U.S. attempts to prevent introductions – especially of pathogens. He called on the Dialogue to focus on the resource at risk – native and urban forests – and change our phytosanitary programs on this basis. He has advocated halting imports of plants that are congenerics of important North American tree species, in order to minimize the risk that pests that damage those genera will be introduced.
an American elm that has survived DED – at Longwood Gardens; photo by F.T. Campbell
Jiri Hulcr of the University of Florida tried to reassure Dialogue participants by stating that recent research has substantially reduced the threat from “unknown unkowns”. I applaud Dr. Hulcr’s efforts to reduce scientific uncertainty about the invasive potential of pathogens native to regions other than North America. His study might be the largest attempted by U.S.-based scientists. However, I note that his study assessed the threat posed by 55 insect-vectored fungi to two species of oak and two species of pines. The forests of the southeastern U.S. comprise many other tree genera! He also set a very high bar for defining a threat as serious: the damage to the host must be equivalent to that caused by Dutch elm disease or laurel wilt. Both have devastated their respective hosts. I believe U.S. phytosanitary policy must aim at protecting the full range of native species. Furthermore, levels of damage that affect the host’s role in the ecosystem – not just rapid mortality — should not be acceptable.
Li, Y. C. Bateman, J. Skelton, B. Want, A. Black, Y-T. Huang, A. Gonzalez, M.A. Jusino, Z.J. Nolen, S. Freemen, Z. Mendel, C-Y. Chen, H-F. Li, M. Kolarik, M. Knizek, J-H. Park, W. Sittichaya, P.H. Thai, S-I. 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 pathogen. Phytopathology. https://doi.org/10.1094/PHYTO-01-21-0041-R
Liebhold, A.M., E.G. Brockerhoff, L.J. Garrett, J.L. Parke, and K.O. Britton. 2012. Live Plant Imports: the Major Pathway for Forest Insect and Pathogen Invasions of the US. www.frontiersinecology.org
MacLachlan, M.J., A. M. Liebhold, T. Yamanaka, M. R. Springborn. 2022. Hidden patterns of insect establishment risk revealed from two centuries of alien species discoveries. Sci. Adv. 7, eabj1012 (2021).
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
Oregon-ash dominated swamp in the Ankeny National Wildlife Refuge, Willamette Valley, Oregon; photo by Wyatt Williams, Oregon Department of Forestry
In April 2022 I blogged about efforts on the West Coast to prepare for arrival of the emerald ash borer (EAB).
That blog focused on Oregon ash (Fraxinus latifolia), which is an important component of riparian forests. I alerted you to the availability of ODA/ODF EAB 2018 Response Plan.
I also mentioned Oregon’s active participation in “don’t move firewood” campaigns.
California has long inspected incoming firewood. In 2021 it establishment of a state quarantine in response to APHIS ending the federal quarantine. Washington State operates a statewide trapping program for invasive insects but does not regulate firewood.
Contributions from the Tualatin Soil and Water Conservation District enabled the USDA Forest Service Dorena Genetic Resource Center to begin testing Oregon ash for resistance to EAB and related genetics work. Other funding came from the USFS Forest Health Protection program.
EAB has now been detected in Oregon — in the Willamette Valley! (See photo above, by Wyatt Williams) Concerned stakeholders have established a new newsletter to keep people informed and promote cooperative efforts.
The newsletter is “Ash across the West”.
The first issue of the newsletter provides the following information:
there are eight ash species in the West; all are vulnerable to the emerald ash borer (EAB)
Single-leaf ash (Fraxinus anomala) CA, NV, AZ, UT, NM, CO, WY
Fragrant ash (Fraxinus cuspidata) NV, AZ, NM, UT
Calif ash (Fraxinus dipetala) CA, NV, AZ, UT
Fresnillo (Fraxinus gooddingii) AZ
Gregg’s ash (Fraxinus greggii) AZ
OR ash (Fraxinus latifolia) WA, OR, CA
Chihuahuan ash (Fraxinus papillosa) AZ, NM, TX
Velvet ash (Fraxinus velutina) CA, NV, AZ, UT, NM, TX
EAB Risk Map for OR: based upon known occurrences of ash & corresponding human activities associated with known pathways of EAB introduction and establishment.
2022 status of the two field trials
the Dorena Genetic Resource Center (DGRC): planted 600 seedlings from 27 families; 85% survival in 2022; controlling competing vegetation
Washington State University Puyallup Research Center: planted seedlings from 26 of these families; 95% survival rate. Possible complication from a foliar disease.
Seedlings from 17 Oregon ash families (including 14 of those in the DGRC field trial) sent to Dr. Jennifer Koch (USFS) in Ohio) for EAB resistance/susceptibility testing.
Seed collections began in 2019; interrupted by COVID-19 in 2020 but resumed in 2021 and continue in 2022. Several consortia are involved in Oregon and Washington. In California and the other states, The Huntington Botanical Gardens will lead the collecting effort. Funding is from USFS Forest Health Protection. Seeds are stored for gene conservation; some are used for the field trials in Oregon and Washington and the initial EAB-resistance studies going on in Ohio.
Penn State Ash Genomic Project: Dr. Jill Hamilton is trying to create a ‘genomic passport’ for Oregon ash populations for use in establishing genotype-environment associations to inform seed transfer guidelines. If you would like to help Dr. Hamilton collect leaves for sampling, contact: Dr. Jill Hamilton at jvh6349@psu.edu
To help with seed collection, ash monitoring, documenting the importance of ash to various communities, and other activities; or to get on the mailing list for the newsletter, contact Richard Sniezko at Richard.sniezko@usda.gov
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 have posted nearly 40 blogs about wood packaging (SWPM) since 2015. [You can view these by scrolling below archives to find category “wood packaging”.]
I first raised the need for APHIS to authorize Robert Haack to update his study analyzing pest “approach rates” in wood packaging in July 2018.
Why?
SWPM has delivered our worst forest pests.
SWPM has been recognized as a major pathway of introduction of wood-boring insects for 30 years. Examples include the Asian longhorned beetle, emerald ash borer, redbay ambrosia beetle, and, possibly, the invasive shot hole borers.
For decades, pest-infested wood packaging has come primarily from the same countries: Mexico, Italy, China, and, more recently, Turkey. Many of our most damaging invaders have come from Asia so growing import volumes from Vietnam and other Asian countries also raise concern.
2) The U.S. and Canada have required that wood be treated to kill pests for at least 16 years.
The U.S. and Canada fully implemented the international standard on wood packaging (ISPM#15) in early 2006 – nearly 17 years ago. They had earlier (1999) required treatment of SWPM from China – nearly 24 years ago.
3) Even old analyses concluded that more than 11,000 incoming containers harbored wood pests each year.
The U.S., Canada, and Mexico import more than 31 million shipping containers per year (see “Background” below). Applying decade-old estimates to this number, we conclude that 11,600 of these containers are probably transporting a quarantine wood-boring pest. About 80% of the containers – and probably the pests! – come to U.S. ports. This pest risk is not limited to the West Coast; expansion of the Panama Canal and congestion at West Coast ports mean that an increasing number of ships are travelling directly to ports on the East and Gulf coasts. These region have already been demonstrated to be highly vulnerable to pests from Asia (ranging from Dutch elm disease and Asian longhorned beetle to laurel wilt and beech leaf disease.)
dead redbay trees – killed by redbay ambrosia beetle + laurel wilt fungus – introduced from Asia to Savannah, Georgia
4) Efforts to reduce the pest “approach rate” have not worked yet.
Meantime, administrative efforts to reduce the numbers of containers carrying pests have not been successful. The Bureau of Customs and Border Protection (CBP) has tried. CBP began penalizing individual shipments that are not in compliance with ISPM#15 in 2017 — 5 years ago.
As of the first three-quarters of Fiscal Year 2022 (John Sagle pers. comm. and Crenshaw-Nolan of CBD to Continental Dialogue on Non-Native Forest Insects and Diseases, September 2022), CBP has issued 510 Emergency Action Notifications (EAN) for noncompliant SWPM. About 38% (194) were issued because actionable pests had been discovered. The rest were issued because the ISPM#15 stamp (attesting to the wood having been treated) was either missing or fraudulent. The full-year interception rate will probably be comparable to interceptions in recent years: in FY2021, 548 EANs; in FY2020, 509; in FY2019, 746. CBP staff are disappointed that interceptions have not declined.
CBP agents inspecting SWPM
5) APHIS has avoided stricter enforcement.
APHIS has not adopted an enforcement stance. It has not stiffened penalties. The agency did not raise these phytosanitary issues when it negotiated a major agriculture trade agreement with China in 2020. The agency continued to insist that ISPM#15 is working – but agreed to work with Robert Haack to re-evaluate the approach rate only in 2021.
Correction: I became alarmed when the study had not been released four months after the analysis was completed (in May). I have since learned that the findings had not yet been completely written up and that internal reviews were proceeding. I apologize for the criticism in the original version of this blog. I impatiently await the study’s release, which I hope will be in a few weeks or months.
In the meantime, APHIS has also hired the Entomological Society to carry out an extensive study that includes analysis of interception data from five ports over a period of five years and rearing insects extracted from incoming wood packaging. I don’t want to postpone action aimed at curtailing introductions via this pathway for another five years!
APHIS has instead tried to improve foreign suppliers’ and phytosanitary agencies’ compliance with ISPM#15 through education. In partnership with Canada and Mexico, APHIS has supported two regional education workshops sponsored by the North American Plant Protection Organization (NAPPO). APHIS is now expanding its outreach to smaller companies, industry associations, and foreign suppliers. APHIS and CBP are now collaborating with an industry initiative to train inspectors that insure other aspects of foreign purchases. In addition, the International Plant Protection Convention (IPPC) is developing a “guidance document”. These educational efforts are supported by the U.S. pallet trade association, National Wooden Pallet and Container Association.
For all of these reasons we urgently need the updated data on the pest approach rate in the analysis by Haack and colleagues. Until we see these results, we can’t know the current level of risk associated with growing volumes of imports or assess the effectiveness of new policies. For example, CBP incorporated compliance with ISPM#15 into its government-importer partnership aimed at ensuring cleanliness of supply chains (C-TPAT) in February 2021. Only by comparing the results of the “approach rate” study with future data collected using the same techniques will it be possible to know how effective this action has been. I greatly appreciate CBP’s efforts.
There is still the issue of untrustworthy stamps.
Past data indicate a high proportion – 87% – 95% — of the SWPM found to be infested bore the ISPM#15 stamp. The same proportion was found in a narrower study in Europe (Eyre et al. 2018). Nor are all problems associated with Asia – importers in Houston have complained that stamps on dunnage from Europe also cannot be trusted.
While there are questions about whether this breakdown results from treatment inadequacy (i.e., 56oC for 30 minutes does not kill the larvae), failure of application, or of fraud –
What matters is that neither regulators nor importers can rely on the stamp to identify pest-free wood packaging.
infested wood packaging bearing ISPM#15 mark; photo courtesy of Oregon Department of Agriculture
(True: ISPM#15 was never intended to prevent pest introductions, only to “reduce the risk of introduction and spread of quarantine pests associated with the movement in international trade of wood packaging material made from raw wood.” Still, we should be trying to minimize pest introductions which threaten our wildland, rural, and urban forests.)
CPB’s experience indicates that cracking down on individual shipments will not be sufficient.
Immediate actions to hold foreign suppliers responsible
U.S. and Canada refuse to accept wood packaging from foreign suppliers that have a record of repeated violations – whatever the apparent cause of the non-compliance. Institute severe penalties to deter foreign suppliers from taking devious steps to escape being associated with their violation record.
APHIS and CBP and their Canadian counterparts provide guidance to importers on which foreign treatment facilities have a record of poor compliance or suspected fraud – so they can avoid purchasing SWPM from them. I am hopeful that the voluntary industry program described here will help importers avoid using wood packaging from unreliable suppliers in the exporting country.
Encourage rapid switch to materials that won’t transport wood-borers. Plastic is one such material. While no one wants to encourage production of more plastic, the Earth is drowning under discarded plastic. Some firms are recycling plastic waste into pallets.
APHIS and CFIA have the authority to take these actions under the “emergency action” provision (Sec. 5.7) of the World Trade Organization’s Agreement on the Application of Sanitary and Phytosanitary Standards (WTO SPS Agreement). (For a discussion of the SPS Agreement, go to Fading Forests II, here.)
APHIS should also release the findings of the 2021-2022 study of approach rates by Haack and colleagues. Then the agency should invite stakeholders to discuss the implications, then develop and implement protective strategy reflecting its findings.
Longer-term Actions
APHIS and CFIA should cite their need for setting a higher “level of protection” to minimize introductions of pest that threaten our forests (described inter alia here.) They should then prepare a risk assessment to justify adopting more restrictive regulations that would prohibit use of packaging made from solid wood – at least from the countries with records of high levels of non-compliance.
Michigan champion green ash killed by emerald ash borer
APHIS and CFIA should also undertake the studies needed to determine the cause of the continuing issue of the wood treatment mark’s unreliability, then act to resolve it. Preferably, this work should be conducted with other countries and such international entities as the IPPC & International Forest Quarantine Research Group (IFQRG). However, if attempting such collaboration causes delays, they should begin unilaterally. Upcoming opportunities to address this issue include:
FAO International Day of Forests in 2023
FAO global assessment of forests & health – pest & disease outbreaks
Of course, these steps should be based on the findings of Haack and colleagues.
Meanwhile, what can we do?
Urge Congress to conduct oversight on APHIS’ failure to protect America’s natural resources from continuing introductions of nonnative insects and diseases.
These hearings should be in the context of drafting the 2023 Farm Bill.
Raise the issue with local, state, and federal candidates for office;
Urge Congress to include provisions of H.R. 1389 in the 2023 Farm Bill;
Ask any associations of which we are members to join in communicating these concerns to Congressional representatives and senators. These include:
if you work for a federal or state agency – raise to leadership; they can act directly or through National Plant Board, National Association of State Departments of Agriculture, National Association of State Foresters, National Governors Association, National Association of Counties
scientific membership societies – e.g., Society of American Foresters, Entomological Society of America, American Phytopathological Society;
individual conservation organizations, either with state chapters or at the national level;
woodland owners’ organizations, e.g., National Woodland Owners Association, National Alliance of Forest Owners (NAFO) and their state chapters
urban tree advocates
International Forest Quarantine Research Group
Write letters to the editors of your local newspaper or TV news station.
BACKGROUND: Calculation of the Number of Infested Containers Entering U.S.
As of 2020 (when trade was greatly depressed by the COVID-19 pandemic), nearly 31 million TEUs [a standardized measure for containerized shipment; defined as the equivalent of a 20-foot long container] entered North America. Ports in the U.S. received 80% (24.5 million); Canada 11.5% (3.5 million); Mexico ~9% (2.7 million). U.S. imports have grown substantially since 2020; during the first quarter of 2022 U.S. imports from Asia each month were 20 to 30% higher than in 2019 before COVID-19 disrupted supply chains (blog #292). The U.S. is projected to handle ~26 million TEUs in 2022 [sources here and here.
A “TEU” equals a 20-feet container. Most containers now are twice as large – 40-feet. Several steps are involved in applying findings of Haack et al. 2014 and Meissner 2009 estimates:
divide estimated number of containers (26 million) in half = 13 million.
Assume that three-quarters of that number (13 million) contain wood packaging (based on Meissner) = 9.75 million.
If 1 out of each thousand of these containers with wood packaging is transporting a pest = 9,750 containers / year.
I performed the same calculation for North America-wide estimate of 31 million TEUs discussed at the beginning of the blog.
container being offloaded at Savannah harbor; photo by F.T. Campbell
A separate study (Hudgins et al. 2022) projected that introduction of a new woodboring insect pest that attacks 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. I have blogged often about the rising rate of shipments coming directly from Asia to the American South.
An analysis of fungi associated with Eurasian bark and ambrosia beetles reached a conclusion that the authors consider to be more optimistic. Li et al. (2021) found that none of the 111 fungi was sufficiently virulent to trigger tree mortality after a single-point inoculation. This level of lethality was considered analagous to Dutch elm disease DMF or laurel wilt DMF. Thirty-eight percent of the fungi were considered to be weak or localized pathogens that could kill trees under certain conditions. However, they tested the fungi against only two oak and two pine species. They did not evaluate fungi that might be lethal when the vector beetle engages in mass attacks. Finally, I think phytosanitary agencies should act promptly when a pathogen threatens levels of mortality somewhat below Dutch elm disease and laurel wilt!
SOURCES
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 59(5): 1302-1312.
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
ISHB-infested California sycamore; photo by Beatriz Nobua-Behrmann, University of California Cooperative Extension
Numerous ambrosia beetles have become introduced species. Their invasions are facilitated by their cryptic habits and ecologies, wide host ranges, and specialized breeding systems – all of which allow extremely low populations to start an infestation. The way they breed often results in low genetic diversity in their introduced ranges, but this has not hampered their success. [Bierman et al. 2022]
Also, ambrosia beetles carry fungi, which provide food needed by their larvae. While most of these fungi don’t harm living trees, some do. The United States has been invaded by three damaging ambrosia beetle-fungal complexes: laurel wilt in the Southeast, and Fusarium dieback disease, carried to southern California with polyphagous and Kuroshio shot hole borers.
These shot hole borers and their fungi represent an especially high risk to our forests because they can be transported in both living and dead wood. So not only massive U.S. imports of live plants but also the global movement of goods enclosed in solid wood packaging offer ready pathways for them to arrive and spread here. Neither pathway is regulated effectively enough to prevent either pest imports or interstate spread.
Invasive ambrosia beetles in California and Hawai’i
The invasive ambrosia beetles introduced to California are in the genus Euwallacea. This genus has undergone several taxonomic revisions. Now, the Euwallacea are divided into four species (Stouthammer 2017), of which three are in the U.S.:
Euwallacea fornicatus s.s. – common name polyphagous shot hole borer; first came to attention in southern California in 2012; formerly known as E. whitfordiodendrus.
E. perbrevis – common name tea shot hole borer; formerly known as E. fornicatus s.l.
E. kuroshio – unchanged nomenclature since detected in California in 2013;
E. fornicatior — apparently has not invaded outside of its native range in Asia.
Those now in the U.S. have been introduced to naïve habitats here and elsewhere, often with dire consequences. E. perbrevis, and possibly other species in the complex, are established on the Hawaiian islands.
For an extensive discussion of their introduction history go here
The Fungi: U.S. and Worldwide
Several fungal associates are vectored by the polyphagous shot hole borer (PSHB) and Kuroshio shot hole borer (KSHB). The most important are Fusariumeuwallacea and Fusarium kuroshium, respectively. These fungi were only described after they appeared in California in the 2010s. They cause Fusarium dieback disease.
Because the two beetle species are difficult to distinguish and the associated diseases cause very similar impacts, Californians studying them and educating stakeholders now speak of the two beetle-fungus complexes as one unit, “invasive shot hole borers”.
Both PSHB and KSHB have numerous genetic strains, or haplotypes. For PSHB, the greatest haplotype diversity is in Asia – Thailand, Vietnam and China. Remember that these same regions are also a center of diversity for the huge genus Phytophthora, blog a genus widely recognized as containing many plant pathogens. https://www.dontmovefirewood.org/pest_pathogen/sudden-oak-death-syndrome-html/ One of the PSHB haplotypes, H33, has invaded many more regions than the others, including Israel, California, and South Africa. It has also been detected in several tropical plant greenhouses in Europe (where it has been eradicated). H33apparently is native to Vietnam – near Hanoi and Ho Chi Minh City – the country’s major ports (Rugman-Jones et al 2020 and pers. comm.). Does this haplotype’s spread to three continents reflect circumstances, such as the proximity of its native range to major ports and a “bridgehead effect” from its multiple introductions (the insects can be introduced to new regions on shipments from invaded regions established earlier)? Or does it point to an unknown genetic superiority (Bierman et al. 2022). This issue seems worth exploring.
I have blogged about the rising volume of imports from Vietnam, including to ports on the Gulf Coast –a region that has climatic similarities to Vietnam and known host species, so it seems quite vulnerable to invasion by either PSHB or KSHB.
A second species in the genus, KSHB, was detected in southern California in 2012; it has now spread to Mexico. So far, only one haplotype of this species has been detected in North America; this haplotype is widespread in Taiwan.
Finally, E. perbrevis (formerly known as E. fornicatus s.l.) has been detected in Florida, Hawai`i (island of Maui), and West Australia (to which it is probably native). This species has also been detected in nurseries in the Netherlands, where authorities report that it has been eradicated (Rugman-Jones et al. 2020).
Akacia koa – native tree in Hawai“i attacked by Euwallaceae; photo by David Eckhoff, via Flickr
Some species or haplotypes have been detected in only one introduced location: E. fornicatus H35 and E. kuroshio (H20) in California; H38 in South Africa; H43 on Oahu and the Big Island of Hawai`i; and an unnamed haplotype in West Australia (Rugman-Jones et al. 2020).
This is a brief guide to worldwide invasions by one or more Euwallacea-fungus complexes (Rugman-Jones et al. 2020):
Southern California — two haplotypes of E. fornicatuss.s. (H33 & H35) and E. kuroshio (one haplotype).
Hawai`i – a unique haplotype of E. fornicatuss.s. (H43) on Oahu, the Big Island, and possibly other islands; E. perbrevis on Maui and possibly other islands.
Israel — E. fornicatuss.s. haplotype H33 only.
South Africa — E. fornicatuss.s. haplotype H33 and a unique haplotype (H38).
Western Australia — a unique haplotype of E. fornicatuss.s. and E. perbrevis (which is probably native in northern Queensland).
Greenhouses in Europe – both E. fornicatuss.s. (haplotype not specified) and – in the Netherlands — E. perbrevis; both reported eradicated.
When a location has been invaded by two or more species or haplotypes, this is probably an indication of separate introductions. Multiple introductions thus are suspected in California (Stouthamer et al. 2017; Bierman et al. 2022); South Africa (Bierman et al. 2022); and Hawai`i (Bierman et al. 2022).
As is true of other pathogens, e.g., Phytophthoras, there appears to have been a spurt of introductions in recent decades, to, e.g., California, South Africa, and the second species in Hawai`i. Bierman et al 2022 note the constantly growing number of locations with introductions.
Indigofera jucuna – reproductive host of PSHB in South Africa; photo by Giardano de Barcelona
Impact and Spread
As is common in the case of forest pests, especially pathogens, detection occurred only years after the initial introduction. In South Africa this delay was five years – from 2012 to 2017 or 2018. In California, identification of the species as PSHB in 2012 was nine years after the organism was first detected in the state (2003).
Over the decade since 2012, PHSB, KSHB, and the pathogens they transmit have spread through large portions of southern California. KSHB has spread through “jumps” to distant locations in Orange, Los Angeles, and as far as Santa Barbara and Ventura counties. There have also been detections in even more distant San Luis Obispo and Santa Clara. These latter apparently have not become established.
A likely explanation for this pattern is the movement of firewood. (Rugman-Jones et al 2020 and pers. comm.) See the map here The two beetles and the plant pathogens they carry are expected to spread throughout much of California wherever their many host plants occur.
On Hawai`i, PSHB is attacking several endemic species including one of the largest forest trees, Acacia koa, as well as Pipturus albidus and Planchonella sandwicensis. Numerous non-native species growing on the Islandsare also attacked, including crops (Macadamia and Mangifera) and invasive species
In South Africa, PSHB has spread faster and farther. It has been present since at least 2012 (Stouthamer et al. 2017), although it was not identified until 2018. In about a decade it has spread to every province except Limpopo – PSHB’s largest geographical outbreak of this beetle [Bierman et al. 2022]
Hosts and Areas at Greatest Risk
Hundreds of plant species in at least 33 plant families support successful reproduction of both beetle and fungus. These include many species widespread in southern California, other parts of the U.S., and South Africa. Some California ecosystems are at particular risk because they are dominated by susceptible tree or shrub species. These vulnerable ecosystems are mixed evergreen forests, oak woodlands, foothill woodlands, and riparian habitats. In San Diego County alone, more than 58,000 acres of riparian woodlands are at risk (California Forest Pest Council).
Experience with the Kuroshio shot hole borer (KSHB) in the Tijuana River valley along the California-Mexico border demonstrates the importance of ecological factors in determining disease outcomes. Following introduction, the KSHB killed a high proportion of the willows near the main river channel. However, beginning in 2016, these trees have regrown to almost pre-infestation sizes. Lead researcher John Boland is not certain why these new, fast-growing trees have not been attacked by the KSHB which remains in the area. See links to the Boland studies below.
riparian forest in Tijuana River Valley after recovery from KSHB attack; photo by John Bolton
Urban forests are at particular risk. For example, in South Africa, conservative estimates were that 25% of urban trees would be lost (Bierman et al. 2022). In California, a model developed by Shannon Lynch found the cities at greatest jeopardy are San Diego, Los Angeles, the San Francisco Bay area, and Sacramento. In other areas in the state that lack data on city tree composition, Lynch applied climate models; this approach extended the list of threatened areas to the eastern half of southern California and other parts of the Central Valley. (Lynch presentation to ISHB webinar April 2022; 2nd day.) In my view, this model should also be applied to cities in Arizona and Nevada with similar climates.
Management
Symptoms of PSHB attack and fungus infection differ among tree species. For illustrations of the symptoms on various species, visit here.
Most important, prevent the beetles’ spread through movement of dead or cut wood, e.g., green waste, firewood, and even large wood chips or mulch. Websites provide information on managing these sources.
Where the beetles have already established, California scientists recommend focusing management on heavily infested “amplifier trees”. On these trees, dead limbs should be pruned; dying trees and those with beetles infesting the main trunk should be removed. The wood must be disposed of properly.
Sources
Bierman, A., F. Roets, J.S. Terblanche. 2022. Population structure of the invasive ambrosia beetle, Euwallacea fornicatus, indicates multiple introductions into South Africa. Biol Invasions (2022) 24:2301–2312 https://doi.org/10.1007/s10530-022-02801-x
Eskalen, A., Stouthamer, R., Lynch, S. C., Twizeyimana, M., Gonzalez, A., and Thibault, T. 2013. Host range of Fusarium dieback and its ambrosia beetle (Coleoptera: Scolytinae) vector in southern California. Plant Dis. 97:938-951.
Stouthamer, R., P. Rugman-Jones, P.Q. Thu, et al. 2017. Tracing the origin of a cryptic invader: phylogeography of the Euwallacea fornicatus (Coleoptera: Curculionidae: Scolytinae) species complex. Agric For Entomol 19:366-375. https://doi.org/10.1111/afe.12215
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
Horton House on Jekyll Island, Georgia before laurel wilt killed the giant redbay trees; photo by F.T. Campbell
Close to four hundred tree species native to the United States are at risk of extinction. The threats come mainly from non-native insects and diseases – a threat we know gets far too little funding, policy attention, and research.
As Murphy Westwood, Vice President of Science and Conservation at the Morton Arboretum, which led the U.S. portion of a major new study, said to Gabriel Popkin, writing for Science: “We have the technology and resources to shift the needle,” she says. “We can make a difference. We have to try.”
Staggering Numbers
More than 100 tree species native to the “lower 48” states are endangered (Carrero et al. 2022; full citation at the end of this blog). These data come from a global effort to evaluate tree species’ conservation status around the world. I reported on the global project and its U.S. component in September 2021. This month Christina Carrero and colleagues (full citation at the end of this blog) published a summary of the overall picture for the 881 “tree” species (including palms and some cacti and yuccas) native to the contiguous U.S. (the “lower 48”).
This study did not address tree species in Hawai`i or the U.S. Pacific and Caribbean territories. However, we know that another 241 Hawaiian tree species are imperiled (Megan Barstow, cited here).
Assessing Threats: IUCN, NatureServe, and CAPTURE
Carrero and colleagues assessed trees’ status by applying methods developed by IUCN and NatureServe. (See the article for descriptions of these methods.) These two systems consider all types of threats. Meanwhile, three years ago Forest Service scientists assessed the specific impacts of non-native insects and pathogens on tree species in the “lower 48” states and Alaska in “Project CAPTURE” (Conservation Assessment and Prioritization of Forest Trees Under Risk of Extirpation). All three systems propose priorities for conservation efforts. For CAPTURE’s, go here.
Analyses carried out under all three systems (IUCN, NatureServe, and CAPTURE) concur that large numbers of tree species are imperiled. Both IUCN and CAPTURE agree that non-native insects and pathogens are a major cause of that endangerment. While the overall number of threatened species remained about the same for all three systems, NatureServe rated threats much lower for many of the tree species that IUCN and CAPTURE considered most imperiled.
This difference arises from the criteria used to rate a species as at risk. IUCN’s Criterion A is reduction in population size. Under this criterion, even extremely widespread and abundant species can qualify as threatened if the population declines by at least 30% over three generations in the past, present, and/or projected future. NatureServe’s assessment takes into account rapid population decline, but also considers other factors, for example, range size, number of occurrences, and total population size. As a result, widespread taxa are less likely to be placed in “at risk” categories in NatureServe’s system.
In my view, the IUCN criteria better reflect our experience with expanding threats from introduced pests. Chestnut blight, white pine blister rust, dogwood anthracnose, emerald ash borer, laurel wilt disease, beech leaf disease, and other examples all show how rapidly introduced pathogens and insects can spread throughout their hosts’ ranges. (All these pests are profiled here . ) They can change a species’ conservation status within decades whether that host is widespread or not.
Which Species Are at Risk: IUCN
Carrero and colleagues found that under both IUCN and NatureServe criteria, 11% to 16% of the 881 species native to the “lower 48” states are endangered. Another five species are possibly extinct in the wild. Four of the extinct species are hawthorns (Crataegus); the fifth is the Franklin tree (Franklinia alatamaha) from Georgia. A single specimen of a sixth species, an oak native to Texas (Quercus tardifolia),was recently re-discovered in Big Bend National Park.
Franklinia (with Bachman’s warbler); both are extinct in the wild; painting by John Jacob Audubon
The oak and hawthorn genera each has more than 80 species. Relying on the IUCN process, Carrero and colleagues found that a significant number of these are at risk: 17 oaks (20% of all species in the genus); 29 hawthorns (34.5% percent). A similar proportion of species in the fir (Abies), birch (Betula), and walnut (Juglans) genera are also threatened.
Other genera have an even higher proportion of their species under threat, per the IUCN process:
all species in five tree genera, including Persea (redbay, swampbay) and Torreya (yews);
two-thirds of chestnuts and chinkapins (Castanea), and cypress (Cupressus);
almost half (46.7%) of ash trees (Fraxinus).
Pines are less threatened as a group, with 15% of species under threat. However, some of these pines are keystone species in their ecosystems, for example the whitebark pine of high western mountains.
Carrero et al. conclude that the principal threats to these tree species are problematic and invasive species; climate change and severe weather; modifications of natural systems; and overharvest (especially logging). Non-native insects and pathogens threaten about 40 species already ranked by the IUCN criteria as being at risk and another 100 species that are not so ranked. Climate change is threatening about 90 species overall.
range of black ash
Considering the invasive species threat, Carrero and colleagues cite specifically ash trees and the bays (Persea spp.). In only 30 years, the emerald ash borer has put five of 14 ash species at risk. All these species are widespread, so they are unlikely to be threatened by other, more localized, causes. In about 20 years, laurel wilt disease threatens to cause extinction of all U.S. tree species in the Persea genus.
Carrero and colleagues note that conservation and restoration of a country’s trees and native forests are extremely important in achieving other conservation goals, including mitigating climate change, regulating water cycles, removing pollutants from the air, and supporting human well-being. They note also forests’ economic importance.
As I noted above, USFS scientists’ “Project CAPTURE” also identified species that deserve immediate conservation efforts.
Where Risk Assessments Diverge
All three systems for assessing risks agree about the severe threat to narrowly endemic Florida torreya and Carolina hemlock.
With three risk ranking systems, all can agree (as above), all can disagree, or pairs can agree in four different ways. Groups of trees fall into each pair, with various degrees of divergence. Generally, only two of the three systems agree on more widespread species:
black ash: IUCN and Project CAPTURE prioritize this species. NatureServe ranked it as “secure” (G5) as recently as 2016.
whitebark pine: considered endangered by IUCN, “vulnerable” (G3) by NatureServe. The US Fish and Wildlife Service has proposed listing the species as “threatened” under the Endangered Species Act. https://www.fws.gov/species-publication-action/endangered-and-threatened-wildlife-and-plants-threatened-species-18 However, Project CAPTURE does not include it among its highest priorities for conservation. Perhaps this is because there are significant resistance breeding and restoration projects already under way.
tanoak: considered secure by both IUCN and NatureServe, but prioritized by Project CAPTURE for protection.
dead tanoak in Curry County, Oregon; photo by Oregon Department of Forestry
Carrero notes the divergence between IUCN and NatureServe regarding ashes. Four species ranked “apparently secure” (G4) by NatureServe (Carolina, pumpkin, white, and green ash) are all considered vulnerable by IUCN. They are also prioritized by Project CAPTURE. I have described the impact of the emerald ash borer on black ash. Deborah McCullough, noted expert on ash status after invasion by the emerald ash borer, also objects to designating this species as “secure” (pers. comm.).
Port-Orford cedar is currently ranked as at risk by IUCN and Project CAPTURE, but not NatureServe. Growing success of the restoration breeding project has prompted IUCN to change the species’ rank from “vulnerable” to “near threatened”. IUCN is expected to reclassify it as of “least concern” in about a decade if breeding efforts continue to be successful (Sniezko presentation to POC restoration webinar February 2022).
While these differing detailed assessments are puzzling, the main points are clear: several hundred of America’s tree species (including many in Hawai`i, which – after all – is our 50th state!) are endangered and current conservation and restoration efforts are inadequate.
Furthermore, a tree species loses its function in the ecosystem long before it becomes extinct. It might still be quite numerous throughout its range – but if each individual has shrunken in size it cannot provide the same ecosystem services. Think of thickets of beech root sprouts – they cannot provide the bounteous nut crops and nesting cavities so important to wildlife. Extinction is the extreme. We should act to conserve species much earlier.
YOU CAN HELP!
Congress is considering the next Farm Bill – which is due to be adopted in 2023. Despite its title, this legislation has often provided authorization and funding for forest conservation (for example, the US Forest Service’ Landscape Scale Restoration Program).
There is already a bill in the House of Representatives aimed at improving the US Department of Agriculture’s prevention and early detection/rapid response programs for invasive pests. Also, it would greatly enhance efforts to restore decimated tree species via resistance breeding, biocontrol, and other strategies. This bill is H.R. 1389.
The bill was introduced by Rep. Peter Welch of Vermont, who has been a solid ally and led on this issue for several years. As of August 2022, the bill has seven cosponsors, most from the Northeast: Rep. Mike Thompson [CA], Rep. Chellie Pingree [ME], Reps. Ann M. Kuster and Chris Pappas [NH], Rep. Elise Stefanik [NY], Rep. Deborah K. Ross [NC], Rep. Brian Fitzpatrick [PA].
Please write your Representative and Senators. Urge them to seek incorporation of H.R. 1389 in the 2023 Farm Bill. Also, ask them to become co-sponsors for the House or Senate bills. (Members of the key House and Senate Committees are listed below, along with supporting organizations and other details.)
Details of the Proposed Legislation
The Invasive Species Prevention and Forest Restoration Act [H.R. 1389]
Expands USDA APHIS’ access to emergency funding to combat invasive species when existing federal funds are insufficient and broadens the range of actives that these funds can support.
Establishes a grant program to support research on resistance breeding, biocontrol, and other methods to counter tree-killing introduced insects and pathogens.
Establishes a second grant program to support application of promising research findings from the first grant program, that is, entities that will grow large numbers of pest-resistant propagules, plant them in forests – and care for them so they survive and thrive.
[A successful restoration program requires both early-stage research to identify strategies and other scientists and institutions who can apply that learning; see how the fit together here.]
Mandates a study to identify actions needed to overcome the lack of centralization and prioritization of non-native insect and pathogen research and response within the federal government, and develop national strategies for saving tree species.
Incorporating the provisions of H.R. 1389 into the 2023 Farm Bill would boost USDA’s efforts to counter bioinvasion. As Carrera and colleagues and the Morton Arboretum study on which their paper is based demonstrate, our tree species desperately need stronger policies and more generous funding. Federal and state measures to prevent more non-native pathogen and insect pest introductions – and the funding to support this work – have been insufficient for years. New tree-killing pests continue to enter the country and make that deficit larger –see beech leaf disease here. Those here, spread – see emerald ash borer to Oregon.
For example, funding for the USDA Forest Service Forest Health Protection program has been cut by about 50%; funding for USFS Research projects that target 10 high-profile non-native pests has been cut by about 70%.
H.R. 1389 is endorsed by several organizations in the Northeast: Audubon Vermont, the Maine Woodland Owners Association, Massachusetts Forest Alliance, The Nature Conservancy Vermont, the New Hampshire Timberland Owners Association, Vermont Woodlands Association, and the Pennsylvania Forestry Association.
Also, major forest-related national organizations support the bill: The American Chestnut Foundation (TACF), American Forest Foundation, The Association of Consulting Foresters (ACF), Center for Invasive Species Prevention, Ecological Society of America, Entomological Society of America, National Alliance of Forest Owners (NAFO), National Association of State Foresters (NASF), National Woodland Owners Association (NWOA), North American Invasive Species Management Association (NAISMA), Reduce Risk from Invasive Species Coalition, The Society of American Foresters (SAF).
HOUSE AND SENATE AGRICULTURE COMMITTEE MEMBERS – BY STATE
STATE
Member, House Committee
Member, Senate Committee
Key members * committee leadership # forestry subcommittee leadership @ cosponsor of H.R. 1389
Alabama
Barry Moore
Arizona
Tom O’Halleran
Arkansas
Rick Crawford
John Boozman*
California
Jim Costa Salud Carbajal Ro Khanna Lou Correa Josh Harder Jimmie Panetta Doug LaMalfa
Colorado
Michael Bennet #
Connecticut
Jahana Hayes
Florida
Al Lawson Kat Cammack
Georgia
David Scott * Sanford Bishop Austin Scott Rick Allen
Raphael Warnock Tommy Tuberville
Illinois
Bobby Rush Cheri Bustos Rodney Davis Mary Miller
Richard Durbin
Note that the report was led by scientists at the Morton Arboretum – in Illinois!
Indiana
Jim Baird
Mike Braun
Iowa
Cindy Axne Randy Feenstra
Joni Ernst Charles Grassley
Kansas
Sharice Davids Tracey Mann
Roger Marshall#
Kentucky
Mitch McConnell
Maine
Chellie Pingree @
Massachusetts
Jim McGovern
Michigan
Debbie Stabenow *
Minnesota
Angie Craig Michelle Fischbach
Amy Klobuchar Tina Smith
Mississippi
Trent Kelly
Cindy Hyde-Smith
Missouri
Vicky Hartzler
Nebraska
Don Bacon
Deb Fischer
New Hampshire
Ann McLane Kuster @
New Jersey
Cory Booker
New Mexico
Ben Ray Lujan
New York
Sean Patrick Maloney Chris Jacobs
Kristen Gillibrand
North Carolina
Alma Adams David Rouzer
North Dakota
John Hoeven
Ohio
Shontel Brown Marcy Kaptur Troy Balderson
Sherrod Brown
Pennsylvania
Glenn Thompson
South Dakota
Dusty Johnson
John Thune
Tennessee
Scott DesJarlais
Texas
Michael Cloud Mayra Flores
Vermont
Patrick Leahy
Virginia
Abigail Spanberger #
Washington
Kim Schreir
SOURCES
Christina Carrero, et al. Data sharing for conservation: A standardized checklist of US native tree species and threat assessments to prioritize and coordinate action. Plants People Planet. 2022;1–17. wileyonlinelibrary.com/journal/ppp3
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
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
The Nature Conservancy (TNC) and Clemson University have analyzed how to persuade people not to move firewood – and the tree pests that can accompany it. (Full citation at the end of this blog) Their study is based on five surveys conducted by TNC between 2005 and 2016. These surveys guided TNC’s “Don’t Move Firewood” campaign and its outreach efforts since the beginning in 2008
As Solano et al. note, wood-boring pests continue to enter the country and spread, causing immense damage. Firewood transport by campers is a significant contributor to that spread. Millions of individuals decide whether to move firewood. Yet the scientific literature is quite limited regarding their behavior and TNC’s survey data has never been published.
The patchwork of state and federal quarantines is largely reactive and has failed to prevent continuing spread. The regulatory regime has been further fragmented by APHIS’ deregulation of the emerald ash borer. As a consequence, limiting the spread of pests depends even more on educating campers to behave responsibly – voluntarily.
The TNC’s surveys each focused on different geographic areas and asked different questions in each. So their compilation cannot show trends in awareness or other measures. Nevertheless, the authors find:
Most people in the United States don’t know firewood can harbor invasive forest insects and diseases, but when targeted by effective education they can learn and are likely to change their behavior.
The two best ways to reach the public is through emails confirming campsite reservations and flyers handed out at parks. Web-based information seemed less effective. However, most of the surveys were done before 2011, the year when 50% of adults reported using internet media.
Forestry-related public agencies (especially state forestry departments) are the most trusted sources of information about forest health issues.
It works better to “push” information, not expect people to seek it on their own.
Messages should focus on encouraging the public to make better choices, including how they, themselves, will benefit. Positive, empowering calls to action, like “Buy it Where You Burn It” or “Buy Local, Burn Local” are better than negative messages, such as “Don’t Move Firewood”.
People respond to messages that emphasize protecting forest resources, e.g., ecosystem services like clean water. They response less to messages about forest threats.
Hungerford Lake Recreation Area at Equestrian Campground. Original public domain image from Flickr
Solano et al. describe the ways that different socioeconomic groups differ in their awareness of forest pests and in how they respond to various statements about forests, pests, and messengers. The focus is on how to overcome four psychological barriers to changing behavior that had been identified in a study of climate change. In the firewood context, those barriers were: 1) lack of awareness; 2) mistrust and negative reactions to the messengers; 3) habit; and 4) social comparison, norms, conformity, and perceived poor quality of purchased firewood.
From this work, the authors suggested further work::
Development of education and outreach programs that target those with lower education levels, since, on average, ~60% of people who camp did not graduate from college. Further research is probably needed to identify the most effective messengers and messages.
While 80% of the survey respondents were over 40, the proportion of campers made up of Gen X and millennials is increasing. Managers need to improve outreach for younger audiences. This includes engaging the messengers they trust: scientists, environmentalist politicians, peer networks, and social media.
While women trust the USDA Forest Service and conservation organizations, 55% of campers in a given year are men. Further research is needed to clarify the most effective messengers and messages for men. The outreach agencies should select the messengers that both sexes trust.
Levels of awareness should be assessed both before and after implementing new educational strategies so that the strategies’ effectiveness can be determined.
Since 80% of the respondents were white, determining the most effective messages and messengers for other ethnic groups also seems necessary, although the authors did not address this.
SOURCE:
Solano, A., Rodriguez, S.L., Greenwood, L., Rosopa, P.J., and Coyle, D.R. 2022. Achieving effective outreach for invasive species: firewood case studies from 2005-2016. Biological Invasions. https://link.springer.com/article/10.1007/s10530-022-02848-w
To request a copy of this study from the author, contact the lead author at Clemson University.
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
spotted knapweed (Centaurea maculosa); photo by Alan Vernon via Wikipedia
Litt and Pearson (full citation at the end of the blog) are trying to improve scientists’ ability to predict the impact of biological invasions. Their goal is to predict which organisms will be winners, which losers, in the face of anthropogenic ecosystem change.
They focus on exotic plant invasions, because they are ubiquitous. Furthermore, plant invasions affect ecosystems by reassembling the plant community in ways that affect the niches used by native animals and hence the animals’ success under the new conditions. After determining the differences between the traits exhibited by invasive plants vs. the native plants they are displacing, scientists can then identify which native animals are most likely to be affected, as well as how and why they might respond to exotic plant invasion. [Note that Doug Tallamy is looking at similar issues.]
Litt and Pearson have developed a framework to assess how plants’ traits might affect associated wildlife. Applying the framework requires certain baseline information about the ecosystem in question.
This knowledge is applied in stepwise fashion:
1) Identify the fauna of interest and their linkage to the native plant community. This association might be food or habitat values such as shelter. Then the researcher determines the relevant plant traits of importance to that animal and approximates the strength of the animal’s dependence on these traits. Note that the focus is on plant traits relevant to the animal users, rather than specific plant species.
2) Determine overall importance of the plant traits for the area under study by (a) averaging dependence of a representative subsample of individuals to obtain a community-level value for each plant species or functional group and (b) quantifying the relative abundance of the plant functional group in the community (e.g., cover or biomass).
3) Plot the way the animal species’ abundance changes with resource abundance.
4) Understand how the invasive plants will alter the distributions of the native plants’ traits and potentially introducing novel traits that might alter the faunal community.
Litt and Pearson reviewed earlier studies to test how well this framework explained the responses of three groups of fauna to plant invasions in different ecosystems.
searching for spotted knapweed; photo by Oregon Department of Agriculture
Spiders in invaded grasslands
Intermountain grasslands of western Montana are heavily invaded; non-native plants already comprise 25–60% of average total plant cover.
One group of native spiders construct their irregular webs entirely within a single plant. A second group – orb weavers – suspend their larger webs from multiple plants. The former depend on the architectural complexity of individual plants; they can build larger webs in plant species possessing greater branching and/or longer branches of the flowering stalks. Orb spiders depend more on the complexity of the overall plant community.
Plant architecture is closely tied to the plant’s functional groups, that is, whether they are grasses or forbs.
These grasslands are generally dominated by perennial grasses. The irregular-web spiders can use grasses, but strongly favor forbs, particularly those with the most complex flowering structures. Orb weavers are generalists, incorporating multiple plant species; but they also tend to favor forbs, presumably because they are more robust.
Invasive plants in the Western Montana grasslands are of two types: an annual grass, cheatgrass (Bromus tectorum), and numerous perennial and annual forbs. Cheatgrass largely replaces the dominant native grasses with a similar architecture – although cheat is shorter. The exotic forbs, which can collectively invade at levels comparable to cheatgrass, tend to be taller and more complex structurally than the native forbs. Thus, invasion by exotic forbs strongly shifts the community-level distribution of the key trait toward greater structural complexity by replacing the dominant, but structurally simplistic, native grasses, and the more diminutive native forbs. These changes increased the abundance of both spider groups, but especially the specialist irregular web weavers. They find the new conditions meet their needs. Both spider groups appeared to expand their realized niches in response to invasion, i.e., they are able to use a broader range of plant architectures than was available in the native system.
Chaetodipus sp. photo by J.N. Stuart
Rodents in semi-desert grasslands invaded by Lehmann lovegrass
In the semi-desert grasslands of the American southwest, native grasses and forbs provide food and habitat for a variety of rodents. This vegetation influences which species of rodents are present in two ways: the size of the plants’ seeds and the density of vegetative cover. Litt and Steidl examined both. They divided the rodents into separate guilds based on diet and preferred vegetative cover. The two sets of guilds did not overlap for all species.
In southern Arizona, the native plant community is dominated by several grass species and herbaceous forbs; most species produce relatively large seeds. Vegetative cover is generally low, but varies in a patchy fashion. The rodent communities in uninvaded native grasslands are dominated by seed-eaters that prefer sparse cover.
Invasion of these grasslands by Lehmann lovegrass (Eragrostis lehmanniana) results in increased vegetative cover but the grass produces very small seeds that probably provide little to no food for rodents. Another result is a decrease in overall abundance of arthropods. The new conditions favor different rodent species from those most common in uninvaded habitat.
Two more specialized seed-eating rodent species, which seek both lower cover and larger seeds, decreased in abundance. A rodent species which favors lower vegetative cover and feeds on larger invertebrates also declined. In contrast, abundance increased for two other rodent species that prefer more dense cover and are more opportunistic in their feeding. One species surprised the scientists: Dipodomys merriami increased in abundance, despite the fact that this species favors more open environments. Perhaps other functional traits or biotic interactions are important to this species? There was no apparent change in abundance for three other species, suggesting either a lack of statistical power (2 were less abundant) or that these rodents were able to persist through a balance of positive and negative changes in food and habitat characteristics.
Lucy’s warbler [nest in saguaro, not cottonwood); photo by Dominic Sherony
Warblers in Riparian Habitats in the Southwest
Riparian habitats in the same desert region have been aggressively invaded by the exotic shrub saltcedar (Tamarix spp.). Litt and Pearson consider the findings of Mahoney et al. of this invasion’s impact on two ecologically similar warbler species. One, the yellow warbler (Setophaga petechia), is very widely distributed across North America; it is considered a generalist. The other, Lucy’s warbler (Oreothlypis luciae), is endemic to a small region of the southwest United States and northern Mexico.
The two species have similar feeding behaviors but differ in their nesting requirements. The yellow warbler constructs open cup nests in the branches of shrubs and trees. Lucy’s warbler nests in cavities in larger trees excavated by others. Hence, these species were expected to respond similarly to changes in food resources and foraging habitat, but differ in their responses to changes in nesting substrate.
Native vegetation in the region consists primarily of willows and cottonwoods in the riparian corridors, with oak and mesquite woodlands in the adjacent uplands. Saltcedar invasion rapidly displaces the willows; it takes much longer to displace cottonwoods since are large and long-lived. Upland vegetation is uninvaded and unaffected. While saltcedar is structurally similar to native willows, its leaf architecture allows more light to penetrate in saltcedar stands. This can exacerbate heat stress on nestlings in these hot, arid environments, as well as expose the nestlings to nest predation. These effects are exacerbated by the presence of a biocontrol leaf beetle (Diorhabda spp.), which cause widespread defoliation of saltcedar during nesting season. Meantime, the cavity nests used by Lucy’s warbler are barely affected.
The study by Mahoney et al. showed that in low-invasion riparian sites, the two warblers occur at comparable abundances. When saltcedar invasion replaces willows, yellow warblers decline by ~50% while there is no apparent change in abundance of Lucy’s warblers.
Litt and Pearson point out that their framework is based on two key assumptions that establish the context for its efficacy.
The first is that bottom-up forces fuel ecological processes. Plants are key to making the sun’s energy available to consumer animals and – thence to predators. Consumers’ and predators’ top-down effects are secondary. The authors’ framework thus provides better predictions of community outcomes when systems are predominantly structured by bottom-up forces. As top-down forces increase or when invasive plants differentially affect multiple dimensions of the consumer niche space, it will be more challenging to track and predict outcomes, as our rodent example demonstrates.
The second assumption is that exotic plant invasions will most strongly influence bottom-up processes. Invasive plants displace native plants and their plant traits, thus directly affecting consumers by altering the quality and quantity of food and habitat resources. However, plant community changes caused by plant invasions can also affect predators directly and indirectly via several interactions. These changes in predators’ abundance and/or their per capita effects on prey might create feedbacks that can complicate interpreting and predicting invasion outcomes.
Litt and Pearson concluded that their approach is promising but has inherent limitations linked to the dynamic nature of ecological systems.
[Ecologists continue to evaluate the impacts of saltcedar eradication efforts on another bird species, the federally endangered southwestern willow flycatcher (Empidonax extimus trailii). See, for example, Goetz, A., I. Moffit and A.A. Sher. 2022. Recovery of a native tree following removal of an invasive competitor with implications for endangered bird habitat. Biological Invasions Vol. 24, pp. 2769-2793.]
SOURCE
Litt, A.R. and D.E. Pearson. 2022. A functional ecology framework for understanding and predicting animal responses to plant invasion. Biol Invasions https://doi.org/10.1007/s10530-022-02813-7
& Supporting Information [warblers in riparian ecosystems invaded by tamarisk]
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
Pinus radiata plantation in New Zealand; photo by Jon Sullivan
As countries and conservation organizations ramp up tree planting as one solution to climate change, I worry that many of the plantings will use species not native to the region – with the risk of promoting more bioinvasions. My second fear is that inadequate attention will be paid to ensuring that the propagules thrive.
Warning from New Zealand
New Zealand has adopted a major afforestation initiative (“One Billion Trees”). This program is ostensibly governed by a policy of “right tree, right place, right purpose”. However, Bellingham et al. (2022) [full citation at end of blog] say the program will probably increase the already extensive area of radiata pine plantations and thus the likelihood of exacerbated invasion. They say the species’ potential invasiveness and its effects in natural ecosystems have not been considered.
Bellingham et al. set out to raise the alarm by evaluating the current status of radiata, or Monterrey, pine (Pinus radiata) in the country. They note that the species already occupies ~1.6 M ha; the species makes up 90% of the country’s planted forests. Despite the species having been detected as spreading outside plantations in 1904, it is generally thought not to have invaded widely.
The authors contend that, to the contrary, radiata pine has already invaded several grasslands and shrublands, including three classes of ecosystems that are naturally uncommon. These are geothermal ecosystems, gumlands (infertile soils that formerly supported forests dominated by the endemic and threatened kauri tree Agathis australis), and inland cliffs. Invasions by pines – including radiata pine – are also affecting primary succession on volcanic substrates, landslides on New Zealand’s steep, erosion-prone terrain, and coastal sand dunes. Finally, pine invasions are overtopping native Myrtaceae shrubs during secondary succession. Bellingham et al. describe the situation as a pervasive and ongoing invasion resulting primarily from spread from plantations to relatively nearby areas.
kauri; photo by Natalia Volna, iTravelNZ
The New Zealanders cite data from South America and South Africa on the damaging effects of invasions by various pine species, especially with respect to fire regimes.
Furthermore, their modelling indicates that up to 76% of New Zealand’s land area is climatically capable of supporting radiata pine — most of the country except areas above 1000 m in elevation or receiving more than 2000 mm of rainfall per year. That is, all but the center and west of the South Island. This model is based on current climate; a warmer/drier climate would probably increase the area suitable to radiata pine.
These invasions by radiata pine have probably been overlooked because the focus has been on montane grasslands (which are invaded by other species of North American conifers). [See below — surveys of knowledge of invasive plants’ impacts.]
Bellingham et al. recognize the economic importance of radiata pine. They believe that early detection of spread from plantations and rapid deployment of containment programs would be the most effective management strategy. They therefore recommend
1) taxing new plantations of non-indigenous conifers to offset the costs of managing invasions, and
2) regulating these plantations more strictly to protect vulnerable ecosystems.
They also note several areas where additional research on the species’ invasiveness, dispersal, and impacts is needed.
Survey of Awareness of Invasive Plants
A few months later a separate group of New Zealand scientists published a study examining tourists’ understanding of invasive plant impacts and willingness to support eradication programs (Lovelock et al.; full citation at end of the blog). One of the invasive plant groups included in the study are conifers introduced from North America and Europe. These conifers are invading montane grasslands, so they are not the specific topic of the earlier article. The other is a beautiful flowering plant, Russell lupine. These authors say that both plant groups have profound ecological, economic, and environmental impacts. However, the conifers and lupines are also highly visible at places valued by tourists. Lovelock et al. explored whether the plants’ familiarity – and beauty – might affect how people reacted to descriptions of their ecosystem impacts.
Visitors from elsewhere in New Zealand were more aware of invasive plants’ impacts and more willing to support eradication programs for these species specifically. Asian visitors had lower awareness and willingness to support eradication of the invasives than tourists from the United Kingdom, Europe, or North America. This pattern remained after the tourists were informed about the plants’ ecological impacts. All groups were less willing to support eradication of the attractive Russell lupine than the conifers.
Conifers invading montane grasslands are perhaps the most publicized invasive plants in New Zealand [as noted above]. Lovelock et al. report that New Zealand authorities have spent an estimated $NZ166 million to eradicate non-native conifers over large tracts of land on the South Island. Still, only about half the New Zealand visitors surveyed were aware of the ecological problems caused by wild conifers.
invasive lupines in New Zealand; photo by Michael Button via Flickr
Russell lupine (Lupinus × russellii) is invading braided river systems, modifying river flows, reducing nesting site availability for several endangered birds, and provides cover for invasive predators. While initially planted in gardens, the lupines were soon being deliberately spread along the roads to ‘beautify’ the landscape. Foreign tourists often specifically seek river valley invaded by the lupine because pictures of the floral display appear in both official tourism promotional material & tourist-related social media. It is not surprising, then, that even among New Zealanders, only a third were aware of the lupines’ environmental impacts.
The oldest participants (those over 60) had the lowest acceptance of wild conifers. Participants 50–59 years old were most aware of ecological problems caused by wild conifers. Participants 30–39 years old showed the highest acceptance of wild conifers and lowest awareness of ecological issues.
Female participants showed a higher preference for the landscape with wild conifers (45.90%) than males (36.89%). Female participants were also half as aware of ecological problems (25.62% v. 46.12% among male participants).
Nearly all survey participants (96.1%) preferred the landscape with flowering lupine; only 19.4% were aware of associated ecological problems. New Zealand domestic visitors were more aware. After the impacts of lupines were explained, half decided to support eradication. However, the same proportion of all survey participants (42.5%) still wanted to see lupines in the landscape.
Once again, participants older than 50 were more aware of ecological problems arising from lupine invasions. Both men and women greatly preferred the landscape with Russell lupins.
While the authors do not explore the ramifications of the finding that younger people are less aware of invasive species impacts, I think they bode ill for future protection of the country’s unique flora and fauna. They did note that respondents had a high level of acceptance overall for these species on the New Zealand landscapes.
While the study supported use of simple environmental messaging to influence attitudes about invasive species, also showed that need to consider such social attributes as nationality and ethnicity. So Lovelock et al. call for investigation of how and why place of origin and ethnicity are important in shaping attitudes towards invasives. Conveying conservation messages will be more difficult because tourist materials often contain photographs of the lupines. Much of this information comes from informal media such as social media, which are beyond the control of invasive species managers.
SOURCES
Bellingham, P.J., E.A. Arnst, B.D. Clarkson, T.R. Etherington, L.J. Forester, W.B. Shaw, R. Sprague, S.K. Wiser, and D.A. Peltzer. 2022. The right tree in the right place? A major economic tree species poses major ecological threats. Biol Invasions Vol.: (0123456789) https://doi.org/10.1007/s10530-022-02892-6
Lovelock B., Y. Ji, A. Carr, and C-J. Blye. 2022. Should tourists care more about invasive species? International and domestic visitors’ perceptions of invasive plants and their control in New Zealand. Biological Invasions (2022) 24:3905–3918 https://doi.org/10.1007/s10530-022-02890-8
Posted by Faith Campbell
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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
