forest pests – Page 4 – Center for Invasive Species Prevention

Eastern National Parks: Forest Regeneration Failing in 69%

Gettysburg battlefield; now under attack by emerald ash borer (see below)

Kathryn Miller and colleagues (full citation at end of blog) have published a study that examined the status and trends of forest regeneration in 39 National parks from Virginia to Maine. Four-fifths of the forest plots in the study are classified as mature or late successional – so at first glance the forests look healthy. However, the researchers made an alarming finding: in 27 of 39 parks, forest regeneration is failing – either imminently or probably. Acadia National Park is an exception; it is the only park in the study experiencing healthy regeneration. They warn that without intense, sustained – and expensive! – intervention, these forests are likely to be converted to other types of ecosystems. [I blogged recently about findings regarding regeneration in eastern forests: here and here and here and here.

The forests’ understories have too few seedlings and – especially – saplings to maintain themselves. Worse, in many cases the seedlings and saplings are not the same species as the mature trees that form the canopy. The saplings are shorter species that never reach the canopy. That is, species like pawpaw (Asimina triloba), American holly (Ilex opaca), American hornbeam (Carpinus caroliniana), and eastern redbud (Cercis canadensis) are regenerating, rather than the oaks (Quercus spp.), hickories (Carya spp.), maples (Acer spp.), and pines (Pinus spp.) that constitute the canopies of mature forests in these parks.

Miller and colleagues call these “regeneration mismatches.” In about half of the parks, these native canopy tree species make up less than half of current saplings and seedlings. This situation suggests the forests’ species composition will shift substantially, thereby undermining resilience in the face of other challenges, such as invasive plants and pests and climate change.

In many of these National parks, Miller and colleagues found abundant ash regeneration. For example, ash (Fraxinus spp.) constitute more than half of all seedlings in four parks (Johnstown Flood and Friendship Hill in Pennsylvania; Catoctin Mountain in Maryland; Manassas Battlefield in Virginia). Miller and colleagues consigned ash species to the “subcanopy class” because the emerald ash borer (EAB) has caused such high mortality of mature trees. They think regard it unlikely that current and future seedlings will ever reach full size. The devastating impact is most starkly illustrated in Gettysburg National Battlefield Park. Consistent deer management since 1996 has been rewarded: the Park ranks at the top for regeneration among the 39 parks. However, more than half of the seedlings and a quarter of the saplings are ashes. EAB has shifted the Park’s otherwise secure regeneration status into probable failure.

When regeneration fails: too many deer

Throughout the study region, the overwhelming reason regeneration fails is browsing by overabundant deer. The level of deer browse is considered “acceptable” in only four parks. Deer suppress the number of seedlings and saplings. They also skew species composition of native subcanopy species toward those less palatable. Miller and colleagues found that canopy tree density and cover and past human land use had minimal impacts on seedling and sapling numbers or species composition.

Overabundant deer also promote invasion and spread of non-native plants, which are the second most important factor impeding regeneration. Together, invasive plants and non-native earthworms are ecosystem engineers that negatively impact soil and cause cascades of biotic and abiotic impacts throughout forest ecosystems.

Many of the parks experiencing the most severe impacts of chronic deer browse also have the highest invasions by non-native plants. A natural process of regeneration occurs when the death or collapse of mature trees create gaps in the forest canopy. Where deer and invasive shrubs overlap, this process is often hijacked. Instead of nearby native tree species accelerating their growth toward the canopy, thickets of invasive shrubs crowd the space.

For this reason, Miller and colleagues recommend that park management prioritize treating invasive plants in canopy gaps of disturbed stands to avoid forest loss. They recommend deliberate creation of canopy gaps to promote resilience only for parks, or stands within parks, that have low deer and invasive plant abundance or the capacity to intensively manage invasive plants in gaps.

In most parks, non-native tree species are rare, less than 2% of total regeneration. In seven parks, though, non-native trees exceed ten percent of seedlings and/or saplings. In three parks, saplings of non-native trees are increasing. These are primarily tree-of-heaven (Ailanthus altissima) and Norway maple (Acer platanoides). In Saratoga National Historical Park, seedlings of common buckthorn (Rhamnus cathartica) are increasing.

Beech regeneration in Prince William Forest Park

Role of other pests

Miller and colleagues express fear that beech bark disease and beech leaf disease might have effects similar to those of EAB, leading to a greater “regeneration debt” in parks where American beech (Fagus grandifolia) is the dominant regeneration component. They cite specifically Prince William Forest Park in northern Virginia, [25 mi²] Rock Creek Park in the District of Columbia, [2.7mi²] and Saratoga National Historical Park. [5.3 mi²] The authors also suggest that thickets of beech root sprouts formed in response to BBD can suppress regeneration of other native canopy species and so might need to be managed.

Miller and colleagues mention hemlock woolly adelgid (HWA), but provide very little information. They report that Saint-Gaudens National Historical Park in New Hampshire (the home and studio of sculptor Augustus Saint-Gaudens) is at particular risk because of growth of both beech and eastern hemlock (Tsuga canadensis). I know that Delaware Water Gap National Recreation Area [109m²] has experienced major losses of mature hemlocks. [Shenandoah National Park has also, but it was not included in the study.]

Hemlock Ravine, Delaware Water Gap National Recreation Area; photo by Nicholas T via Flickr

Miller and colleagues report that Acadia National Park is seeing recovery of red spruce (Picea rubens) from a major fire in 1947 and possibly also from acid rain. They do not mention the longer-term threat from the brown spruce longhorned beetle. Their focus is on forest dynamics largely unaffected by deer.

In the same way, the authors make no mention of the absence of dogwood trees, presumably because they had been eliminated by dogwood anthracnose decades ago. Nor do they mention vascular streak dieback of redbud; the causal agent still uncertain. [See Annie Self’s presentation to National Plant Board, August 2023.]

dead ash tree in Shenandoah National Park

One omission is large enough that it might affect the study’s findings. At mi² Shenandoah is the largest National Park in the region. It was not included in the study because the Park’s forest monitoring process is not compatible with those in other NPS units. All the other parks – including Acadia (56² mi) – are much smaller, protecting historic sites like Civil War battlefields.

RECOMMENDATIONS

Miller and colleagues recommend that deer management be initiated in parks classified as at imminent or probable regeneration failure, if such programs are not already under way. They warn that effective deer management requires sustained commitment. Studies of deer exclosures show that full forest recovery from chronic deer overabundance can take as long as 40–70 years.

The authors also recommend actions to open the subcanopy to facilitate growth of saplings belonging to desired species. They caution that deer predation must be controlled. Furthermore, either invasive plant cover must be low, or management must ensure that that the park has sufficient resources to sustain an invasive plant control program – especially if invasive plants are combined with abundant deer.

Parks experiencing compositional mismatches and that are dominated by oak–hickory forest types might also benefit from prescribed burning. Again, deer browse pressure must be minimized. In addition, regeneration of oaks and hickories must already be present.

In park forests dominated by species vulnerable to lethal pests, e.g., beech-, ash-, or hemlock-dominated forest stands, Miller and colleagues recommend considering planting alternative native canopy species and protecting those plantings from deer. Park managers should also consider thinning beech thickets formed after beech bark disease kills canopy trees.

Media coverage

The Washington, D.C., public radio station, WAMU, reported on this research on the air (broadcast December 20) and on its website. It is written by Jacob Fenston, with great photographs by Tyrone Turner. The story emphasized the link between deer and invasive plants – since regeneration in eastern deciduous forest happens by saplings taking advantage of gaps formed when mature trees die. The story quotes DC-area people on their efforts to contain vines. The Natural Resource Manager at Catoctin Mountain Park [8 mi²] describes that park’s longstanding deer control program. The story also mentions impacts of EAB and threat of BLD.

News – Funding for these parks to counter the threats!

Lead author Kathryn Miller has informed me that the Bipartisan Infrastructure Law and Inflation Reduction Act has provided the 39 parks involved in this study over $10 million to improve forest resilience largely through reduction of invasive plants and overabundant deer.

Of course, invasive species threats to National parks are not limited to the Northeast – nor are they new. I have raised this problem from the beginning. To see these blogs, on the “nivemnic” website, scroll down below the archives to the “categories”, then click on “national parks”.

SOURCE

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

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

www.fadingforests.org

Succession: “novel drivers” change the trajectory

hardwood regeneration in northern Virginia forest; photo F.T. Campbell

I have posted several blogs recently about tree species’ regeneration. One blog found poor regeneration of many species throughout forests of the eastern United States. Regeneration is particularly poor in the Great Lakes region, western New York and Pennsylvania, along the Mid-Atlantic and New England coasts, and the coastal plain from southern South Carolina to eastern Texas.

A second blog focused on forest succession in New Hampshire. These findings, by Ducey et al., explicitly recognized the impact of non-native tree-killing insects and pathogens. A third article (Payne and Peet, 2023; full citation at the end of this blog) reports similar findings in North Carolina – and explicitly says that the same conditions are found in forests across the eastern United States.

The locations of neither in-depth study – New Hampshire or North Carolina – include those identified by Potter and Riitters (2022) as suffering particularly poor regeneration.

Payne and Peet find that forest succession in the Piedmont region of North Carolina is not proceeding as expected, based on earlier studies conducted in the same region. The differences are apparent at both the canopy and understory levels. Especially notable is the low recruitment of oaks (Quercus species) and hickories (Carya species) – the genera which previous studies indicated would be the climax taxa. One explanation is the disappearance since early in the 20^th Century of fire as a driver of disturbance.

The understory communities are also novel, due largely to invasive species: dramatic loss of flowering dogwood (Cornus florida) killed by the non-native pathogen dogwood anthracnose (Discula destructiva), plus overcrowding of the shrub level by invasive plant species. Other drivers are probably suppression of growth of woody species caused by excessive deer herbivory, and overall accelerated shifts in successional trajectory due to hurricane damage.

flowering dogwood autumn display; F.T. Campbell

Forests in eastern North America in the 21^st Century face several drivers of change that are either novel or greatly heightened. In addition to the disappearance of chronic fire, these are frequency and timing of hurricanes, feeding by herbivore populations, and introduction of non-native tree-killing pests and plants. Payne and Peet say scientists and managers need to consider these additional drivers – and their interactions! – when anticipating successional change.

Like Ducey et al. in New Hampshire, Payne and Peet used 80 years of data from 33 permanent plots established and 55 years of data from another 3 plots. Twenty-eight of the plots are transitioning from loblolly pine (Pinus taeda) to hardwood dominance; eight plots have been mixed-age hardwood stands since before the study plots were established.

In the North Carolina piedmont, the composition of canopy trees in plots evolving from pine compared to hardwood stands continue to be different 90–120 years after succession began. Canopy trees in upland and bottomland hardwood stands also differ. These differences reflect the relative species in the forest at the initiation of succession dynamics. Hurricanes – especially Hurricane Fran in 1996 – apparently accelerated succession in some plots by toppling the oldest pines. Despite the persistent differences, the species compositions of both canopy and subcanopy layers are trending toward increasing similarity.

deer-damaged red maple; photo by Eli Sagor via Flickr

The impact of deer browsing is complicated. Deer populations in the study area quadrupled after measurement began in 1980. Deer herbivory suppressed growth of all plant species when their stems were thin (3 – 10 cm DBH). However, after 1996 rapid growth of plants in openings caused by Hurricane Fran’s passage began to reverse the effects of deer browsing. Also, while deer browsing decreases regeneration, growth, and abundance of oak and hickory seedlings and saplings, it also decreases the abundance of other tree species that have – nevertheless – increased in abundance, e.g., red maple (A. rubrum) and black cherry(Prunus serotina).

Payne and Peet found that soil attributes (wetness, texture, organic matter and chemical components), as well as topographic position were minor factors in determining succession trajectories. Increased light availability due to the new or exacerbated drivers of change (thinning of understory vegetation by disease and deer herbivory and opening of the canopy by hurricanes) overcame the influence of nutrients. At most, a unique soil condition might constraining the impacts of these disturbances. Furthermore, these soil-related conditions and other environmental variables change through time — and as a result so does the vegetation. Specifically, the conditions that once supported establishment of oaks and hickories apparently differ today. Payne and Peet conclude that other drivers might be continuing to impact these species’ maturation.

A partial exception is soil nitrogen, through its influence on mycorrhizal patterns. I review mycorrhizal patterns in the discussion of individual tree species, below.

How are Individual Tree Species Responding?

Oaks and hickories are not expanding as expected – either as canopy-sized trees or as seedlings / saplings in the understory. Payne and Peet agree that century-long suppression of low-intensity ground fires is probably the most significant factor in this compositional shift. This decline has been exacerbated by selective logging and deer herbivory. Hickories have established more widely, possibly because young stems have greater shade tolerance. Only plots located on sandy and acidic soils and plots with the greatest hurricane damage have moderate recruitment of oaks and hickories. Oaks and hickories on the poor soils might be aided by the types of ectomycorrhizal fungi that survive in acidic soils with relatively low nitrogen levels. In addition, these soils’ lower water retention probably impedes competition by more mesic, faster-growing, shade-tolerant species. However, even oaks and hickories that have established as seedlings or saplings only rarely progress to canopy dominance. Payne and Peet conclude that oaks might have lost competitive advantage in many of the undisturbed stands.

More mesophytic hardwoods, especially red maple (Acer rubrum), are becoming more numerous and larger – a trend seen throughout forests of the eastern United States. Damage from Hurricane Fran apparently accelerated this trend. However, red maple growth is significantly inhibited by competition from thicket-forming shrubs, especially in bottomland plots. The invasive non-native species thorny olive or oleaster Elaeagnus pungens increased dramatically following Hurricane Fran in 1996. The situation is likely to worsen: two other invasive species, Amur honeysuckle Lonicera maackii and privet Ligustrum japonicum were first detected in the Duke Forest plots in the 2013 survey.

[In New Hampshire, Ducey et al. detected an unexpected levelling off of red maple increases and decline in sugar maple (Acer saccharum); they were unable to determine a cause.]

beech-dominated understory in northern Virginia; F.T. Campbell

Another mesophytic hardwood – American beech (Fagus grandifolia) – has become very abundant in bottomland hardwood stands, especially in small-stem size classes in the understory. Beech prefers sandy soils and its ectomycorrhizal associations are apparently more tolerant of more acidic soils.

Payne and Peet mention – briefly and vaguely – uncertainty about the future of beech. The reference cited discusses the impact of beech bark disease (BBD) in the northeast. Range maps indicate that BBD is well established in the southern Appalachians along the North Carolina/Tennessee border; it has apparently not spread as far east as the study area. There is no mention of beech leaf disease (BLD), which is the primary threat to seedlings and saplings. BLD is currently known to be in northern Virginia. It is unknown whether the disease has any climatic or other barrier that would prevent its moving farther south.

Another bottomland indicator taxon that is also increasing in abundance is ash (Fraxinus species). Along with sweetgum (Liquidambar styraciflua), tulip poplar (Liriodendron tulipifera) and black cherry Prunus serotina, ash density and basal area increased dramatically in plots heavily damaged by Hurricane Fran. Payne and Peet expect most ash trees to be killed by emerald ash borer (Agrilus planipennis) by 2022. The beetle was detected in the study area in 2015.

ash killed by EAB on Potomac lowlands; F.T. Campbell

Flowering dogwood (Cornus florida)was one of the most abundant understory species throughout the study area until the late 1980s. The species has declined by more than 80% since then due to the non-native disease dogwood anthracnose (Discula destructiva). No other species has experienced as precipitous a decline. There is now almost no regeneration in most upland sites.

A second species almost eradicated from the study area by a non-native pathogen is American elm (Ulmus americana). Its basal area in 2013 was 5% of peak levels in the 1950s. Most of this loss occurred by the 1960s, shortly after arrived of Dutch elm disease (DED) in North Carolina. A congeneric species, slippery elm U. alata, is reported to beabundant; it is somewhat resistant to DED. There is no mention of the zig-zag sawfly (Aproceros leucopoda) which has been detected in North Carolina, a few counties away from the study area. The foliage-feeding insect’s long-term impact on elm species is not yet understood.

Payne and Peet note that the study area has twice experienced loss of important components due to specialist non-native pathogens: elms and dogwoods. A third similar event looms: ash [The article does not discuss prospects for biological control.] A fourth is less certain: beech. [This numbering assumes that American chestnut and eastern hemlock were not significant components of forests in the study area.] In their view, these events demonstrate the drastic impacts such non-native organisms can have, especially when the host species is highly abundant or otherwise dominant in a specific community. The resulting shifts in community dynamics and modifications to light and water availability due to such losses, can be dramatic and long-lasting, even resulting in novel successional trajectories.

Members of the 23rd Civil Engineer Squadron/23rd Wing chainsaw a tree lying across a street in the NCO housing area- damage to piedmont North Carolina by **Hurricane Fran**. Photo courtesy of U.S. National Archives.

Payne and Peet also emphasize the impact of large, episodic disturbances (in their case, hurricanes). These can have widespread and long-lasting impacts on plant community dynamics. Hurricanes’ frequency, intensity, and timing relative to successional stage are key in determining their impacts on successional trajectories. E.g., strong storms that felled the even-aged pine canopy accelerated succession toward more mixed hardwoods. These changes affect biomass, diversity, competitive dynamics, and invasion by invasive plant species, especially in sites with advantageous soil conditions.

Scientists must also evaluate interactions (both reinforcing and antagonistic) between these drivers. For example, in this study deer herbivory and damage from episodic storms had opposite effects on the density of stems in the understory and therefore the future dynamics of forested stands. Hurricane aftereffects frequently accelerated existing or developing trends resulting from various other drivers (e.g., loss of dogwood to anthracnose disease). [While Ducey et al. also detected lasting impacts from hurricane damage in New Hampshire, these effects did not include changes in tree species composition.] Broader regional and global drivers of change, especially those associated with climate change and nitrogen deposition, interact with these many indicators in novel ways based on their own local loadings.

The Nature Conservancy focuses on fire

The Nature Conservancy magazine for Winter 2023 carries an article describing the organization’s experimental efforts to promote oak succession in the Piedmont forests of North Carolina. Greg Cooper, TNC’s forest ecologist in North Carolina, describes retaining dominance by oaks and hickories – rather than maples and poplars – as vital to protecting the region’s faunal diversity and minimizing impacts from climate change. He says this is because oaks use a quarter of the water of maples and poplars.

Cooper links oaks’ failure to reproduce on fire suppression. TNC kills midstory maples and poplars through hack and squirt methods. This allows more light to penetrate the forest and foster oak seedling recruitment. Then they apply controlled fire. “We currently have 700 acres of [controlled-] burn plots, some of which have been burned twice, some of which have been burned once, [and already] we’re getting more light and an immediate flush of herbaceous diversity. We’re getting a lot more berry species, more wildflowers.” TNC is monitoring plots that have been burned, with and without the pre-burn herbicide treatments, and those that have not been burned. They hope to have results in five to ten years that will indicate whether they are achieving the desired improvement in oak regeneration. If so, they also hope is that in future prescribed burns will be sufficient.

Cooper adds that through the Fire Learning Network and a 23-person fire crew they carry out similar work not just on TNC properties, but also federal and state properties.

SOURCES

Ducey, M.J, O.L., Yamasaki, M. Belair, E.P., Leak, W.B. 2023. Eight decades of compositional change in a managed northern hardwood landscape. Forest Ecosystems 10 (2023) 100121

Payne, C.J. and R.K. Peet. 2023. Revisiting the model system for forest succession: Eighty years of resampling Piedmont forests reveals need for an improved suite of indicators of successional change. Ecological Indicators 154 (2023) 110679

Posted by Faith Campbell

www.fadingforests.org

Imports from Asia rise; perhaps 2,000 or more containers carrying insect pests enter U.S. in one month

Wood packaging – crates, pallets, spools for wire, etc. — has been recognized as a major pathway for introduction of tree-killing pests since the Asian longhorned beetle was detected in New York and Chicago in the late 1990s. As of 2021, 65 new species of non-native wood- or bark-boring Scolytinae had been detected in the United States (Rabaglia; full citation at end of the blog).

As I have often reported [To see my 40+ earlier blogs about wood packaging material, scroll down below archives to “Categories,” click on “wood packaging”.], the international phytosanitary community adopted the International Standard for Phytosanitary Measures (ISPM) #15. The goal of ISPM#15 is to “significantly reduce” [not eliminate] the risk of pests associated with solid wood used for constructing packaging (e.g., crates, pallets), from being introduced to other countries through international trade.

I recently reviewed the first 20+ years of implementation of ISPM#15 including two analyses by Robert Haack and colleagues in a blog in December 2022. I have also provided the broader context of the World Trade Organization (WTO) in my Fading Forests II report.

I last blogged about U.S. import volumes in June. My silence since reflected the significant decline in U.S. imports from Asia. This reduction had reduced the likelihood that a new tree-killing pest would be introduced from that region – or that an already-established pest would be introduced to a U.S. region that had escaped it so far.

However, U.S. imports from Asia have suddenly grown! In October 2023, containerized imports from Asia were 12.4% higher than a year ago – and 6% higher than in September. According to the Journal of Commerce (full citation at end of blog), U.S. retailers anticipate consumers will purchase lots of gifts for the upcoming Christmas season.

The U.S. imported 1.57 million TEU from Asia in October. This volume exceeded even the pre-COVID levels. How great is the associated risk of a pest introduction? To calculate that, I apply the following:

most U.S. imports arrive in 40-foot-long containers, so divide TEU by 2 = 785,000
a decade-old estimate that 75% of containers in maritime shipments contain wood packaging (Meissner et al.) = 588,750 containers with wood packaging (I suspect it is more).
the estimate by Haack et al. 2014 that 0.1% (1/10^th of 1 percent) of consignments (which usually means a single container) harbor tree-killing pests;
the estimate by Haack et al. 2022 that 0.22% of consignments harbor tree-killing pests.

The result of these calculations is an estimate of 648 containers (using the 2009 global estimate), or 1,727 containers (using the 2022 global estimate), or 5,730 containers (using the 2010-2020 estimate for China specifically) entering the country in one month harbored tree-killing pests. Since West Coast ports received 54% of those containers, the estimated number of containers transporting pests that enter California, Washington, or Oregon ranged from 349 to 3,042. The rest are scattered among the dozens of ports on the East and Gulf coasts.

With drought limiting container ship transits through the Panama Canal (Szakonyi 2023), the threat to East and Gulf coast ports might not rise commensurately.

Because of the low levels of imports in previous months, U.S. imports from Asia remain significantly below levels in previous years: 16.6% lower for the January – September period compared to 2022.

The 2022 analysis found that the rate of wood packaging from China that is infested has remained relatively steady since 2003: 1.26% during 2003–2004, and ranged from 0.58 to 1.11% during the next three time periods analyzed. Packaging from China made up 4.6% of all shipments inspected, but 22% of the 180 consignments with infested wood packaging. Thus the proportion of Chinese consignments with infested wood is five times greater than would be expected based on their proportion of imports. Note the great impact of this high infestation rate on the number of containers transporting tree-killing pests to the U.S. in the paragraph above: more than 8,000 containers compared to about 2,000.

I remind you that the U.S. and Canada have required treatment of wood packaging from China since December 1998. Why are the responsible agencies in the United States not taking action to correct this problem? [which has persisted for 2 decades]

The fact is – as I have argued numerous times — a pallet or crate bearing the ISPM#15 mark has not proved to be a reliable indicator as to whether the wood is pest-free. (This might be because the wood had not been treated, or if it was, the treatment failed). All the pests detected in the Haack et al. studies (after 2006) were in wood packaging bearing the ISPM#15 mark. As noted in my past blogs [click on the “wood packaging” category to bring up blogs about wood packaging and enforcement], Customs and Border Protection also report that nearly all the wood packaging in which that they detected insect pests bore the ISPM#15 mark.

According to Angell in November (full citation at end of blog), U.S. imports from India to the east coast fell by 15% in the first 10 months of 2023 compared to last year – to a total of 623,356 TEUs. This might change in the future: a shipper has promised to start weekly arrivals from India beginning in May 2024. the company plans calls at New York-New Jersey, Savannah, Jacksonville, Charleston, and Norfolk. The ships will call, en route, at ports in Saudi Arabia, Egypt, and Spain. What pests might be hitching a ride on these shipments?

SOURCES

Haack RA, Britton KO, Brockerhoff EG, Cavey JF, Garrett LJ, et al. 2014. Effectiveness of the International Phytosanitary Standard ISPM No. 15 on reducing wood borer infestation rates in wood packaging material entering the United States. PLoS ONE 9(5): e96611. doi:10.1371/journal.pone.0096611

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

Meissner, H., A. Lemay, C. Bertone, K. Schwartzburg, L. Ferguson, L. Newton. 2009. Evaluation of pathways for exotic plant pest movement into and within the greater Caribbean Region.

Mongelluzzo, B. 2023. U.S. imports from Asia hit 2023 high in October despite muted peak season. Journal of Commerce https://www.joc.com/article/us-imports-asia-hit-2023-high-october-despite-muted-peak-season_20231116.html (access limited to subscribers, unfortunately)

Angell, M. 2023. ONE readies India-US East Coast service as part of 2024 network rollout. Journal of Commerce. November 27, 2023

Rabaglia, R. 2021. The increasing number of non-native bark and ambrosia beetles in North America. International Union of Forest Research Organizations. Prague, Czech Republic. September 2021

Szakonyi, M. 2023. Carriers Weigh Options as Panama Canal restrictions become fact of life. Journal of Commerce. November 21, 2023. (Access limited to subscribers, unfortunately)

Posted by Faith Campbell

www.fadingforests.org

USFS Forest Health Protection program: what it funds

affects of mountan pine beetle on lodgepole pine in Rocky Mountain National Park, Colorado photo from Wikimedia; one of pests addressed by USFS FHP

Several USFS scientists have published an assessment of the agency’s program to enhance forest health across the country: the Forest Health (FHP) program. [see Coleman et al., full citation at end of this blog.] The program assists cooperators (including other federal agencies) to prevent, suppress, and eradicate insect and pathogen outbreaks affecting trees, regardless of land ownership.

Each year, I advocate for adequate funding for the FHP program — which comes from annual Congressional appropriations. Funding has remained static at about $100 million per year. I interpret the article as providing support for my call for increased appropriations. First, it reports that the number of projects and extent of area treated have declined from 2011 to 2020. This is because static funding levels are stretched increasingly thin as costs to implement the same activities rise. Second, the program does not address many damaging forest pests already in the country. The result is growth of established threats to forest health. Finally, new insects and pathogens continue to be introduced. Protecting forest health necessitates tackling these new pests – and that requires money and staff.

Coleman et al. analyzed data from the decade 2011- 2020 to determine the most frequently used project types, integrated pest management (IPM) strategies and tactics, dominant forest pests and associated hosts managed, and most comprehensive forest IPM programs in practice. While there is a wide range of possible projects, most of those funded consist of some form of treatment (more below). The databases relied on do not include funding through the National Forest System aimed at improving forest health through such management activities as stand thinning treatments and prescribed fire. Nor are all pest management activities recorded in the centralized databases. I regret especially the fact that “genetic control” (= resistance breeding) are left out.

Port-Orford cedar seedlings in trial for resistance to *Phytophthora lateralis* at Dorena center; photo courtesy of Richard Sniezko, USFS

Summary of Findings

The data are sorted in various categories, depending on whether one wishes to focus on the type of organism being managed or the management approach. All presentations make evident a dramatic imbalance in the projects funded. Again and again, spongy moth (Lymantria dispar dispar), southern pine beetle (SPB, Dendroctonus frontalis), and several bark beetles attacking conifers in the West (in particular mountain pine beetle, [MPB] Dendroctonus ponderosae) dominate, as measured by both funding and area treated.

oak trees in Shenandoah National Park killed by spongy moth; photo by F.T. Campbell

The bulk of the funding went to the above species, plus hemlock woolly adelgid (HWA; Adelges tsugae); emerald ash borer (EAB, Agrilus planipennis), oak wilt (caused by Bretziella fagacearum), and white pine blister rust (WPBR, Cronartium ribicola).
95% of the projects focused on only four taxa: oaks, Quercus spp. [spongy moth suppression and eradication]; loblolly and ponderosa pines [bark beetle prevention and suppression]; and eastern hemlock [HWA suppression].
Projects seeking to suppress an existing pest outbreak covered 87% of the total treatment area. However, 98% of the treated area was linked to only 20 taxa; again, spongy moth dominated.
Projects seeking to prevent introduction or spread of a pest constituted only 30% of all projects and covered only 11% of the total treatment area.
Eradication and restoration projects each equaled less than 5% of total projects and treatment areas.
Native forest pests were targetted by 79% of projects; non-native pests by 21%. However, non-native pests accounted for 84% of the total treatment area (again, the spongy moth).
While 67% of projects took place on USFS lands (focused on MPB and SPB), 89% of the total treatment area was on lands managed by others (state or other federal agencies, or private landowners). Again, the size of the non-USFS area treated was driven primarily by the spongy moth Slow the Spread program.
Insect pests received nearly all of the funding: 70% of funding targetted phloem-feeding insects, especially SPB and MPB; 10% targetted foliage feeders, especially spongy moth; 6% targetted sap feeders. 4% tackled rusts (e.g., WPBR); just 2% addressed wood borers (e.g., Asian longhorned beetle, emerald ash borer).
The ranking by size of area treated differs. In this case, 82% of areas treated face damage by foliage feeders (e.g., spongy moth); 15% of the treated areas are threatened by phloem feeders (e.g., MPB); only 1.4% of the area is damaged by sap feeders (e.g., HWA); 0.6% is threatened by rust; and 0.2% by wood borers.
Re: control strategies, 32% of projects relied on silvicultural strategies; 22% used semiochemical strategies; 21% exploited other chemical controls; and 18% used physical/mechanical control methods.

Coleman et al. regretted that few programs incorporated microbial/biopesticide control strategies; these were applied on only 10% of total treated area. Again, the vast majority of such projects were aerial applications of spongy moth controls, Bacillus thuringiensis var. kurstaki (Btk) and nucleopolyhedrosis viruses (NPV) (Gypchek). Coleman et al. called for more research to support this approach efforts to overcome other obstacles (see below).

Coleman et al. also called for better record-keeping to enable analysis of genetic control/ resistance breeding projects, treatment efficacy, and survey and technical assistance activities.

History

The article provides a brief summary of the history of the Forest Service’ pest management efforts. Before the 1960s, the USFS relied on labor-intensive physical control tactics, classical biocontrol, and widespread chemical applications. Examples include application of pesticides to suppress or eradicate spongy moth; decades of Ribes removal to curtail spread of white pine blister rust; salvage logging and chemical controls to counter phloem feeders / bark beetles in the South and West. These strategies were increasingly replaced by pest-specific management tactics during the 1970s.

Over the decade studied (2011-2020), tree defoliation attributed to various pests (including pathogens) affected an estimated 0.7% of the 333 million ha of U.S. forest land annually. Mortality attributed to pests impacted an estimated 0.8% of that forest annually. See Table 1. Two-thirds of the area affected by tree mortality is attributed to phloem feeders; a distant second agent is wood borers. These data are incomplete because many insects, diseases, and parasitic higher plants are not tracked by aerial surveys.

As I noted above, these data do not include projects that screen tree species to identify and evaluate genetic resistance to a pest; or efforts to collect cones, seed, and scion. I consider these gene conservation and resistance programs to be some of the most important pest-response efforts. I have blogged about the USFS’ Dorena Genetic Resource Center’ efforts to breed five-needle pines, Port-Orford cedar, and ash. link

41% of silvicultural control treatments targetted phloem feeders; 48% addressed cankers and rusts together. Restoration planting was done in response to invasions by ALB, EAB, and WPBR, as well as native bark beetles and mistletoes.

effort to eradicate SOD in southern Oregon; partially funded by USFS FHP. Photo courtesy of Oregon Department of Forestry

Physical/mechanical control projects were most widely applied in the Rocky Mountains in response particularly to diseases: vascular wilts, rusts, and cankers, including WPBR. This type of project was also used to deal with non-native diseases in other parts of the country, e.g., oak wilt, sudden oak death (SOD), Port-Orford cedar root rot, and rapid ʻōhiʻa death. Sanitation treatments (i.e., removal of infected/infested trees) was used for native mistletoes and root rots, and some non-native insects, e.g., EAB and coconut rhinoceros beetle (Oryctes rhinoceros). Pruning is a control strategy for WPBR. Trenching is applied solely to suppress oak wilt.

Chemical controls were limited to small areas. These projects targetted seed/cone/flower fruit feeders, foliage and shoot diseases, sap feeders [e.g., balsam woolly adelgid (BWA), HWA], wood borers (e.g., EAB) and phloem feeders (e.g., Dutch elm disease; DMF oak wilt vectors). Cover sprays have been used against goldspotted oak borer (GSOB); and many native insects. Fungicides are rarely used; some is applied against the oak wilt pathogen in areas inaccessible by heavy equipment.

treating hemlock trees in Conestee Falls, NC; photo courtesy of North Carolina Hemlock Restoration Initiative

Classical biocontrol projects funded by the program targetted almost exclusively HWA. Some 4.3 million predators have been released since the early 1990s; 820,057 in just the past 10 years.

Gene conservation and breeding projects were directed primary at commercially important hosts, e.g., loblolly Pinus taeda and slash pine P. elliottii; and several non-native pests, including chestnut blight, EAB, HWA, and WPBR.

Survey and technical assistance (i.e., indirectly funded activities) conducted by federal, state, and tribal personnel contributed to education/outreach, evaluating effectiveness, identification, monitoring, and record keeping strategies.

As should be evident from the data presented here, suppression treatments dominated by number of projects and treatment area. The poster child project is the national spongy moth Slow the Spread program. The authors say this program is the most advanced forest IPM program in the world. It has successfully slowed spongy moth’s rate of spread by more than 80% for more than 20 years.

A second widely-used subset of suppression programs consists of physical / mechanical control. This is often the principal suppression strategy in high-visitation sites (e.g., administration sites, campgrounds, picnic areas, and recreation areas). Sanitation harvests are one of the few viable management techniques for suppressing or slowing the spread of recently introduced non-native pests. Nevertheless, the largest number of suppression projects and use of sanitation treatments focused on a native pest, mountain pine beetle, at the height of its outbreak in early 2010s.

Silvicultural control, specifically tree thinning, represents the predominant forest pest prevention tactic, especially on lands managed by the USFS. Two programs dominate: the Southern Pine Beetle Prevention Program and the Western Bark Beetle Initiative. Again, Coleman et al. assess these treatments as very successful. Forest thinning treatments also address other management concerns, i.e., reduce threat of catastrophic wildfires and reduce adverse effects of climate change.

Chemical control tactics are applied to suppress most forest insect feeding guilds in high-value sites and seed orchards. Soil or tree injections of systemic pesticides are used to protect ash and hemlock trees. Topical sprays have been applied to protect whitebark pine (Pinus albicaulis) from mountain pine beetle. Whitebark pine was listed as threatened under the Endangered Species Act in December 2022.

dead whitebark pine at Crater Lake NP; photo by F.T. Campbell

Soil or tree injections target two non-native insects, EAB and HWA.

Genetic control via resistance breeding represents the primary strategy to combat several non-native diseases. (More options are typically available for insects than diseases.) Coleman et al. focus on the extensive effort to protect many of the five-needle pines from WPBR. As I have described in earlier blogs, the Dorena Genetic Resource Center in Oregon has engaged on numerous other species, too.

Coleman et al. describe pest-management associated monitoring efforts as consisting largely of coordinated annual aerial detection surveys, detection trapping, stream-baiting of Phytophthora ramorum, and ground surveys to address site-specific issues.

Coleman et al. call for improvement of record-keeping / databases to encompass all pests, management actions, and ownerships. They also advocate for additional decision-making tools, development of microbial/biopesticides, genetic research and breeding, and biocontrol strategies for several pest groups.

They consider the southern pine beetle and spongy moth programs to be models of comprehensive IPM programs that could be adapted to additional forest health threats. They note, however, that development and implementation of these programs require significant time, financial commitments, and collaborations from various supporting agencies. Not all programs enjoy such resources.

SOURCE

Coleman, T.W, A.D. Graves, B.W. Oblinger, R.W. Flowers, J.J. Jacobs, B.D. Moltzan, S.S. Stephens, R.J. Rabaglia. 2023. Evaluating a decade (2011–2020) of integrated forest pest management in the United States

Journal of Integrated Pest Management, (2023) 14(1): 23; 1–17

Posted by Faith Campbell

www.fadingforests.org

International Phytosanitary System: More Evidence of Failure

Rome: home of the International Plant Protection Convention

I often assert that the international phytosanitary system has proven to be a failure in preventing introductions.

Some of the recent publications support my conclusion – although most don’t say so explicitly. For example, the Fenn-Moltu et al. (2023) study of insect transport and establishment around the world found that the number of invasive species-related treaties, regulations and legislation a country has adopted had no significant effect on either the number of insect species detected at that country’s border or the number of insect species that established in that country’s ecosystems..

Weber et al. also found considerable evidence that international and U.S. phytosanitary systems are not curtailing introduction of insects and entomophagic pathogens. In my earlier blog I review their study of unintentional “self-introductions” of natural enemies of arthropod pests and invasive plants. They conclude that these “self-introductions” might exceed the number of species introduced intentionally. These introductions have been facilitated by the usual factors: the general surge in international trade; lack of surveillance for species that are not associated with live plants or animals; inability to detect or intercept microorganisms; huge invasive host populations that allow rapid establishment of their accidentally introduced natural enemies; and lack of aggressive screening for pests already established. Examples cited include species introduced to the United States’ mainland and Hawai`i specifically.

The U.S. Capitol – one of the entities that can reflect **our** priorities in setting phytosanitary policy

As I point out often, altering human activities that facilitate invasion is a political process. So is amending international agreements that are not effective. We need to determine the cause of the failures of the existing institutions and act to rectify them. See my critiques of both the American and international phytosanitary system Fading Forests II and Fading Forests III (see links at the end of this blog) and my earlier blogs, especially this and this.

SOURCES

Fenn-Moltu, G., S. Ollier, O.K. Bates, A.M. Liebhold, H.F. Nahrung, D.S. Pureswaran, T. Yamanaka, C. Bertelsmeier. 2023. Global flows of insect transport and establishment: The role of biogeography, trade and regulations. Diversity and Distributions DOI: 10.1111/ddi.13772

Weber, D.C., A.E. Hajek, K.A. Hoelmer, U. Schaffner, P.G. Mason, R. Stouthamer, E.J. Talamas, M. Buffington, M.S. Hoddle and T. Haye. 2020. Unintentional Biological Control. Chapter for USDA Agriculture ResearchService. Invasive Insect biocontrol and Behavior Laboratory. https://www.ars.usda.gov/research/publications/?seqNo115=362852

Posted by Faith Campbell

www.fadingforests.org

Predicting Impacts – Can We Do It?

Clive Braser and others study *Phytophthora* species in their native habitats of Vietnam; which will become aggressive invaders in North America?

For years, one focus of this blog has been on scientists’ efforts to improve prevention of new introductions of forest pests. In earlier blogs, I summarized and commented on efforts by Mech et al. (2019) and Schultz et al. (2021), who extrapolate from insect-host relationships of pests already established in North America. [Full citations are presented at the end of this blog.] Both limited their analysis to insects; Mech et al. focused on those that attack conifers, Schultz et al. on those that attack single genera of angiosperms (hardwoods).

However, many of the most damaging agents are pathogens; for an indication, review the list under “invasive species” here. Indeed, Beckman et al. (2021) reported that only three non-native organisms pose serious threats to one or more of the 37 species of Pinus native to the U.S. All are pathogens: white pine blister rust (WPBR), pitch canker, and Phytophthora root rot (Phytophthora cinnamomi).

For this reason I welcome a study by Li et al. (2023), who used laboratory tests to evaluate the threat posed by more than 100 fungi associated with bark beetles. Since there are more than 6,000 species of bark and ambrosia beetles and they are commonly intercepted at the U.S. border, determining which should be priorities is important. Li et al. point out that the vast majority of such introductions have had minimal impacts. Two, however, have caused disastrous levels of damage: Dutch elm disease and laurel wilt disease.

Li et al. tested 111 fungi associated with 55 scolytine beetles from areas of Eurasia with latitudes and ecosystems analagous to those in the southeastern U.S. The beetles assessed included beetle species responsible for recent major tree mortality events in Eurasia: Dendroctonus species, Platypus koryoensis (Korean oak wilt), Platypus quercivorus (Japanese oak wilt) and Tomicus species.

The authors tested the fungi’s virulence on four species of trees native to the Southeast – two pines (Pinus taeda and P. elliottii var. elliottii), and two oaks(Quercus shumardii and Q. virginiana).

Li et al. found that none of 111 fungal associates caused a level of damage on these four hosts equal to Dutch elm disease on elms or laurel wilt disease on trees in the Lauraceae. Twenty-two of the fungi were minor pathogens – meaning they might cause damage under certain conditions or when loads of inoculum are large enough.

redbay trees killed in coastal Georgia by laurel wilt; photo by Scott Cameron

I think Li et al. set an extremely high bar for “serious” damage. Surely we wish to prevent introduction of pathogens that cause damage at a lower level than the catastrophes to which these two diseases have exposed a genus (elms) and a family (Lauraceae)! Still, the scientific approach used here is a step toward addressing pathogens. These agents of tree mortality are addressed much less frequently than insects. I hope that scientists will continue to test the virulence of these fungi on some of the thousands of other species that make up the forests of the United States, or at least the dominant species in each ecosystem.

It is discouraging that Raffa et al. (2023) found none of four approaches to predicting a new pest’s impact to be adequate by itself. Instead, they outlined the relative strengths and weaknesses of each approach and the circumstances in which they might offer useful information. I am particularly glad that they have included pathogens, not just insects. The four approaches they review are:

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

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

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

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

They emphasize that too little information exists regarding pathogens to predict which microbes will become damaging pathogens when introduced to naïve hosts in new ecosystems. See the article, especially Figure 4, for their assessment of the strengths each of the several approaches.

Raffa et al. raise important questions about both the science and equity issues surrounding invasive species. As regards scientific issues, they ask, first, whether it will ever be possible to predict how each unique biotic system will respond to introduction of a new species. Second, they ask how assessors should interpret negative data? In the context of equity and political power, they ask who should make decisions about whether to act?

In my blog I expressed concern about finding that most introduced forest insects are first detected in urban areas whereas introduced pathogens are more commonly detected in forests. I hope scientists will redouble efforts to improve methods for earlier detection of pathogens. Enrico Bonello at Ohio State and others report that spectral-based tools can detect pathogen-infected plants, including trees.

Japanese cherry trees burned on the Washington D.C. mall because infested by scale; on order of Charles Marlatt

Identifying Key Pathways

International trade is considered the single most important pathway for unintentional introductions of insects. Updated figures remind us about the stupendous amounts of goods being moved internationally. According to Weber et al., international shipping moves ~133 million TEU containers per year between countries, the majority between continents. Four times this number move within regions via coastal shipping. On top of that, four billion passenger trips take place by air every year. Air freight carries another ~220 million tons of goods; while this is a tiny fraction of the weight shipped by boat, the packages are delivered in less than a day – greatly increasing the likelihood that any unwanted living organisms will survive the trip. The U.S. also imports large numbers of live plants – although getting accurate numbers is a challenge. MacLachlan et al. (2022) report 5 billion plants imported in 2021, but the USDA APHIS annual report for FY22 puts the number at less than half that figure: 2.2 billion plant units.

Given the high volume of incoming goods, Weber et al. advocate improved surveillance (including analysis of corresponding interceptions) of those pathways that are particularly likely to result in non-native species’ invasions, e.g. live plants, raw lumber(including wood packaging), and bulk commodities e.g. quarried rock. Isitt et al. and Fenn-Moltu et al. concur that investigators should focus on the trade volumes of goods that are likely to transport plant pests – in their cases, plant imports.

The importance of the plant trade as a pathway of introduction for has been understood for at least a century – as witnessed by the introductions of chestnut blight DMF and white pine blister rust, DMF and articles by Charles Marlatt. A decade ago, Liebhold et al. (2012) calculated that the approach rate of pests on imported plants was 12% — more than 100 times higher than the 0.1% approach rate found by Haack et al. (2014) for wood packaging.

Since plant-insect interactions are the foundation of food webs, changes to a region’s flora will have repercussions throughout ecosystems, including insect fauna. See findings by teams led by Doug Tallamy and Sara Lalk; and a chapter in the new forest entomology text written by Bohlmann, and Krokene (citation at end of blog under Allison, Paine, Slippers, and Wingfield). Sandy Liebhold and Aymeric Bonnamour also addressed explicitly links between introductions of non-native plant and insect species. Weber et al. call this phenomenon the “receptive bridgehead effect”: a non-native plant growing prolifically in a new ecosystem provides a suitable host for an organism that feeds on that host, raising the chance for its establishment.

Recent studies confirm the importance of the “receptive bridgehead effect”. Isitt and colleagues found that the large numbers of introduced European insect species – all taxa, not just phytophagous insects – established in North America and Australia/New Zealand were best explained by the numbers of European plants introduced to these regions – in other words, the most important driver appears to be the diversity of non-native plants.

The presence of European plants in North America and Australia/New Zealand promoted establishment of European insects in two ways. First, these high-volume imports increased the propagule pressure of insects associated with this trade. Live plant imports might have facilitated the establishment of ~70% of damaging non-native forest insects in North America. Second, naturalization of introduced European plants provided a landscape replete with suitable hosts. This is especially obvious in Australia/New Zealand, which have unique floras. In Australia, nearly 90% of non-native pest insects are associated with non-native plants. Those non-native insects that do feed on native plants are more likely to be polyphagous.

Amur honeysuckle – one of the hundreds of Asian plants invading North American ecosystems; via Flickr

I hope U.S. phytosanitary officials apply these lessons. Temperate Asia is the source of more non-native plants established in both North America and Australia/New Zealand than is Europe. Already, many insects from Asia have invaded the U.S. The logicof the “receptive bridgehead effect” points to prioritizing efforts to prevent even more Asian insects from reaching our shores!

Fenn-Moltu et al. sought to elucidate which mechanisms facilitate species’ success during the transport and introduction/establishment stages of bioinvasion. They studied the transport stage by analyzing border interceptions of insects from 227 countries by Canada, mainland U.S., Hawai`i, Japan, New Zealand, Great Britain, and South Africa over the 60 year period 1960 – 2019. They studied establishment by analyzing attributes of 2,076 insect species recorded as established after 1960 in the above areas plus Australia (North America was treated as a single unit comprised of the continental U.S. and Canada).

The number of species transported increased with higher Gross National Income in the source country. The number of species transported decreased with geographic distance. They suggest that fewer insects survive longer journeys, but say additional information is needed to verify this as the cause. The number of species transported was not affected by species richness in the native region.

More species established when introduced to a country in the same biogeographic region. They were not surprised that environmental similarity between source and destination apparently strongly affected establishment success. The number of species established was not affected by species richness in the native region. For example, the greatest number of established species originated from the Western and Eastern Palearctic regions, which together comprise only the fifth-largest pool of native insect species.

Gaps Despite Above Studies

As I noted at the beginning, most of the studies examining current levels of pests transported on imported plants have been limited to insects. This is unfortunate given the impact of introduced pathogens (again, review the list damaging organisms under “invasive species” here).

In addition, most studies analyzing the pest risk associated with plant imports use port inspection data – which are not reliable indicators of the pest approach rate. The unsuitability of port inspection data was explained by Liebhold et al. in 2012 and Fenn-Moltu et al. a decade later – as well as Haack et al. 2014 (as the data pertain to wood packaging). Fenn-Moltu et al. note that inspection agencies often (and rightly!) target high-risk sources/commodities, so the records are biased. Other problems might arise from differences in import volume, production practices, and differences in records that identify organism only to genus level rather than species. Fenn-Moltu et al. call for relying on randomized, statistically sound inspection systems; one such example is USDA’s Agriculture Quarantine Inspection System (AQIM). Under AQIM, incoming shipments are randomly selected and put through more thorough inspections to produce statistically based estimates of approach rates, defined as the percent of inspected shipments found to be infested with potential pests (Liebhold et al. 2012). I ask why scientists who are aware of this issue have not obtained AQIM data for pests associated with plant imports. Plant imports have been included in the AQIM system since 2008. Have they not been able to persuade APHIS to provide these data? Or are these data available for only limited types of imported plants? Too narrow a focus would create a different source of potential bias.

Both Isitt et al. and Fenn-Moltu et al. list factors not addressed and other caveats of which we should be aware when extrapolating from their findings.

SOURCES

Allison, J. T.D. Paine, B. Slippers, and M.J. Wingfield, Editors. 2023. Forest Entomology and Pathology Volume 1: Entomology. Springer available gratis at https://link.springer.com/book/10.1007/978-3-031-11553-0

Beckman, E., Meyer, A., Pivorunas, D., Hoban, S., & Westwood, M. (2021). Conservation Gap Analysis of Native U.S. Pines. Lisle, IL: The Morton Arboretum.

Hoddle. M.S. 2023. A new paradigm: proactive biological control of invasive insect pests. BioControl https://doi.org/10.1007/s10526-023-10206-5

Isitt, R., A.M. Liebhold, R.M. Turner, A. Battisti, C. Bertelsmeier, R. Blake, E.G. Brockerhoff, S.B. Heard, P. Krokene, B. Økland, H. Nahrung, D. Rassati, A. Roques, T. Yamanaka, D.S. Pureswaran. 2023. Drivers of asymmetrical insect invasions between three world regions. bioRxiv preprint doi: https://doi.org/q0.1101/2023.01.13.523858

Li, Y., C. Bateman, J. Skelton, B. Wang, 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, T-H Pham, S. Ito, M. Torii, L. Gao, A.J. Johnson, M. Lu, J. Sun, Z. Zhang, D.C. Adams, J. Hulcr. 2022. Pre-invasion assessment of exotic bark beetle-vectored fungi to detect tree-killing pathogens. Phytopathology Vol 112 No. 2 February 2022

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

Liebhold, A.M., T. Yamanaka, A. Roques, S. August, S.L. Chown, E.G. Brockerhoff and P. Pyšek. 2018. Plant diversity drives global patterns of insect invasions. Sci Rep 8, 12095 (2018). https://doi.org/10.1038/s41598-018-30605-4

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).

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

Raffa, K.F., E.G. Brockerhoff, J-C. Gregoirem R.C. Hamelin, A.M. Liebhold, A. Santini, R.C. Venette, and M.J. Wingfield. 2023. Approaches to Forecasting Damage by Invasive Forest Insects and Pathogens: A Cross-Assessment. Bioscience Vol. 73, No. 2. February 2023.

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

Posted by Faith Campbell

www.fadingforests.org

Ash – Science Support Protection Efforts

As we know, survival of North American species of ash (Fraxinus spp.) is threatened by the emerald ash borer (EAB). DMF Sadof, McCullough, and Ginzel (full citation at end of the blog) hope to prevent demise of another ~ 135 million urban ash trees by 2050 bycountering persistent myths that have hindered adoption of effective protective measures. As they note, USDA APHIS has dropped regulations that had been intended to slow the EAB’s spread – which I concede were not very effective.

Protecting urban ash trees now falls to municipalities, states, their leaders and citizens, non-governmental organizations, and tree care professionals. If they apply knowledge gained since the detection of EAB 20 years ago – and are not paralyzed by myths – they can successfully manage EAB populations and protect their town’s ash trees. [I have also blogged about efforts to breed ash trees resistant to EAB.]

Since some studies have found that “myth-busting” is not effective, perhaps people advocating for EAB control should avoid mentioning the myths per se and instead emphasize the science supporting the proposed actions.

Sadof, McCullough, and Ginzel first review aspects of the biology of ash and EAB that are relevant to arborists and pest management specialists:

Adult EAB beetles feed on tree leaves for a couple of weeks from mid-May through June. This maturation period provides a 2–3 week opportunity to kill the leaf-feeding beetles with systemic insecticides before any eggs are laid.
Once eggs hatch, the first stage larvae immediately move into the phloem (inner bark) and cambium tissue, where they begin feeding. Systemic insecticides rarely enter the phloem, so they kill few larvae during this stage.
Detection of early stages of invasion is hampered by several factors, including beetles’ initial colonization of branches in the upper canopy; initially minimal effect on healthy ash trees; and the frequency of two-year life cycles when beetle densities are low. However, it is important to detect and treat these early infestations because EAB populations increase, tree health declines to eventual death.
Detection efforts should target the ash trees most likely to be infested early in the invasion: stressed trees, preferred species (especially green ash), trees growing in the open in parks, along roadsides or surrounded by impervious surfaces. Authorities can take advantage of the attractiveness of stressed trees by establishing “trap trees” to attract EAB adults. Beetles that feed on the “trap trees” can be killed by systemic insecticides. Or the trees can be removed and chipped to kill eggs and larvae before they can emerge. Sadof, McCullough, and Ginzel say trap trees are effective in slowing spread of new infestations when most ash trees remain healthy. Once EAB densities build and many trees are stressed by larval feeding, volatile (airborne) compounds released by girdled trees no longer attract the beetles.
Woodpecker holes in branches of the upper canopy are often the first evidence of EAB invasion in an area.
Even in late stages of the invasion, when most ash trees that were not protected with systemic insecticides are dead, EAB populations persist and continue to colonize and kill available ash trees, including some as small as >2.5 cm in diameter.

Myth: There Is No Point in Trying to Protect Ash Trees—

EAB Will Eventually Kill Them Anyway

Answer:

When the EAB was first detected in 2002, control measures were limited in number and efficacy. In the 20 intervening years, scientist have learned much about EAB biology and ash physiology. Insecticide chemistry and application methods have improved. Currently recommended strategies are based on long-term field studies. More effective insecticides have been developed. Emamectin benzoate is particularly efficient, including the fact that it needs to be applied only every third year. Managers must pay attention to the application protocols, including appropriate dose (i.e., the amount of insecticide product applied); spacing injection ports around the trunk to ensure that the xylem will transport the chemical to leaves throughout the canopy; and conduct injections in spring after bud break.

Myth: Wounds From Drilling Trees to Inject Systemic Insecticides Injure Trees

Answer:

In the early years, trunk injections sometimes caused substantial injury to trees. Refinement of delivery devices and reductions in the pressure at which insecticides are injected have virtually eliminated these issues. Staff must be properly trained in use of the equipment.

demonstration of injecting pesticide into ash tree; photo by F.T. Campbell

Myth: Using Systemic Insecticides to Protect Ash Trees Harms

Non-target Species and the Environment

Answer:

Sadof, McCullough, and Ginzel point out that continent-wide loss of a tree genus is likely to adversely affect the more than 200 species of native arthropods that are specialists on ash. On the other hand, systemic insecticides are unlikely to harm beneficial natural enemies of EAB, including parasitoid wasps, predatory insects, or woodpeckers. First, the insecticides are contained within the tree’s tissues; they do not kill insects on contact. Second, parasitoids and predators avoid dead beetles. Honeydew excreted by sucking insects might contain sufficient insecticide residue to harm parasitoids — if the tree is heavily infested. However, these insects are rapidly killed by these insecticides if they are applied at the optimal time (early to mid-spring). Proper timing of application greatly reduces the potential for tainted honeydew to accumulate on infested trees. Furthermore, in cities there are few populations of natural enemies of sucking insects.

Most concern is focused on pollinators. Ash trees flower early, before leaves expand. It is reassuring that protocols instruct that the systemic insecticides be applied after bud break — typically after pollen has been shed. I do find it disturbing that apparently there have been no published studies of insecticide concentration in ash pollen.

Myth: It Costs Too Much to Protect Ash Trees

Answer:

Sadof, McCullough, and Ginzel review the several studies and methods developed to estimate the value of urban ash trees – both individually and over a wider area. The value is based on the individual tree’s location, health, and structural condition. These economic studies have consistently shown that it costs less to protect ash trees from EAB with insecticide treatments than to remove ash trees — either proactively or when they decline and die.

Even delaying tree mortality – short of preventing it completely – is worthwhile because it allows municipalities to incorporate tree removal into the budget, rather than be suddenly confronted by large expense that they had not planned for.

Sadof, McCullough, and Ginzel recommend treating ash within a significant area as being most efficient. This approach reduces overall costs and slows rates of ash mortality locally – even for trees that are not treated. In some cases, treating as few as 11% of ash trees slowed the overall rate of ash decline.

An important in comparing costs of treatment to costs of replacement is the high mortality rate of newly planted urban trees: up to two-thirds die shortly after planting. This means that it takes decades to replace a mature tree canopy and the environmental benefits the canopy provides. Sadof, McCullough, and Ginzel conclude that protecting ash trees from EAB has clear positive effects for both the urban forest canopy – and its environmental services – and municipal forestry budgets.

Sadof, McCullough, and Ginzel then outline a viable Integrated Pest Management (IPM) framework that incorporates use of systemic insecticides to protect ash trees from EAB.

1. Define the problem and identify management objectives

Inventory urban trees before EAB is detected. The inventories should identify priority trees based on size (diameter at breast height), tree condition, and suitability of the site where the tree is growing. Focus detection surveillance on green ash trees, especially those in parks, parking lots, and along roads — sites that are sunlit (open) and likely to cause stress to the trees.

2. Monitor and assess the local EAB population to determine when a treatment program should be initiated. Treatment must wait until there is evidence that EAB is present but should not then be delayed, since it should begin while the trees’ vascular systems are still sufficiently healthy to carry the insecticide to branches and leaves. This requires regular inspections of ash trees for visible signs of EAB infestation. Efficiency is improved by focusing on high-risk trees (see above) and noticing woodpecker holes on upper portions of the trunk. Consider debarking symptomatic trees or establishing “trap tree” networks.

3. Identify and gather resources needed to implement an insecticide treatment program. Web-based calculators guide budget decisions based on the municipality’s tree inventory and local costs of treatments. Treating one-third of trees annually with emamectin benzoate can save money while maximizing the number of trees protected. Training city forestry staff in trunk injection methods is cheaper than hiring contractors and ensures better treatment quality and efficiency.

downy woodpecker; photo by Steven Bellovin, Columbia University

4. Incorporate multiple tactics to protect tree health and control EAB.

Ensure trees are actively transpiring when injecting the systemic insecticides; this might require irrigation. Encourage parasitoids and woodpecker foraging on untreated trees. In areas where ash trees are closely spaced, consider an area-wide urban SLAM program. In this strategy, treating a proportion of ash trees at two-year intervals reduces EAB eggs and overall EAB populations. Non-treated trees with EAB larvae might support parasitoid biocontrol populations whose offspring can attack EAB larvae on previously treated ash trees as the emamectin benzoate concentration wanes.

Sadof, McCullough, and Ginzel also suggest establishing a citizen monitoring program to both reduce costs and build community support for ash management. Community participation has been particularly effective when professionals take appropriate and timely action in response to volunteers’ findings.

SOURCE

Sadof, C.S., D.G. McCullough, and M.D. Ginzel. 2023. Urban ash management and emerald ash borer (Coleoptera: Buprestidae): facts, myths, and an operational synthesis. Journal of Integrated Pest Management, 2023, Vol. 14, No. 1 https://doi.org/10.1093/jipm/pmad012

Posted by Faith Campbell

www.fadingforests.org

What do “Self-Introduced” & “Door-Knocker” Species Tell Us?

*Woldstedtius flavolineatus* – one of at least 13 taxa of non-native ichneumonid wasps established in restoration forests in Hawaiian Forest National wildlife rfefuge; photo by Torgrim Breiehagen for the Norwegian Biodiversity Information Centre; via Wikipedia

As we know, non-native insects and pathogens pose a significant and accelerating threat to biodiversity in forests and other ecosystems. They undermine some conservation programs and reduce ecosystem services and quality of life in urban areas. Nevertheless, damaging introductions continue.

Two recent articles have advocated accelerating biocontrol programs. These articles have reminded us of ongoing failures of international and national biosecurity programs, including that of the US. The articles also make interesting suggestions regarding ways to be more pro-active in preventing introductions.

1. “Self-introductions” of invaders’ enemies

Weber et al. (full citation at end of blog) provide many examples of unintentional “self-introductions” of natural enemies of arthropod pests and invasive plants. In fact, “self-introductions” of natural enemies of arthropod pests might exceed the number of species introduced intentionally. These introductions have been facilitated by the usual factors: the general surge in international trade; lack of surveillance for species that are not associated with live plants or animals; inability to detect or intercept microorganisms; huge invasive host populations that allow rapid establishment of their accidentally introduced natural enemies; and lack of aggressive screening for pests already established.

Among the examples illustrating failures of biosecurity programs:

Across six global regions, nearly two-thirds of parasitoid Hymenoptera species were introduced unintentionally. The proportion varies significantly by region. For example, four-fifths of these insects in New Zealand arrived accidentally.
The unintentional spread of the glassy-winged sharpshooter (Homalodisca vitripennis) and a biocontrol agent Cosmocomoidea ashmeadi has been so rapid among islands in the Pacific Ocean (including Hawai`i) they are considered ‘biomarkers’ of biosecurity failures.
Regarding the United States specifically, an estimated 67% of beneficial insects introduced to Hawai`i and 64% of parasitoid Hymenoptera introduced to the mainland U.S. were accidental “self-introductions.”

Weber et al. consider their figures to be underestimates. The situation is particularly uncertain regarding pathogens that kill arthropods. Many microbial species are not yet described.

spotted lanternfly; photo by Stephen Ausmus, USDA

In some cases, these “self-introduced” arthropods have proved beneficial. Two examples are Entomophaga maimaiga and Lymantria dispar nucleopolyhedrovirus (LdNPV), which help control the spongy moth (Lymantria dispar). In other cases the “self-introduced” creatures are pests themselves. A prominent example is the invasion by the spotted lanternfly (Lycorma delicatula). This was facilitated by the widespread presence of the highly invasive plant Ailanthus altissima. It illustrates what Weber et al. call “receptive bridgehead effects.” That is, once an invasive pest is well-established, the chance that its natural enemies will find a suitable host and also establish in the pest’s invaded range is much higher.

Weber et al. reaffirm that there are many good reasons not to allow such random invasions of diverse non-native species – including their natural enemies. Deliberately introduced biocontrol agents are chosen after determining their efficacy, host-specificity, and climatic suitability. Random introductions, on the other hand, might favor generalist species, which could threaten non-target species. Accidental introductions might also be accompanied by pathogens and hyperparasitoids that could compromise the efficacy of biocontrol agents.

In short, unintentionally introduced natural enemies might have about the same level of success in controlling the target pest’s populations as do intentionally introduced agents. However, unintentional introductions of both pests and pathogens carry additional risks of non-target impacts and contamination with their own natural enemies that would hamper the efficacy of the biocontrol agent. Weber et al. conclude that delays in releasing a deliberately chosen and evaluated biocontrol agent reduce the probability that it will successfully establish instead of an unintentionally introduced organism.

cactus moth larva on *Opuntia*; photo by Doug Beckers via Flickr

It is especially likely that an arthropod – whether or not a biocontrol agent – will spread within a geographic region. Weber et al. say both the U.S. and Canada have received more than a dozen species intentionally introduced into the other country. They also cite spread of the cactus moth, Cactoblastis cactorum, into Florida from several Caribbean countries. The cactus moth has spread and now threatens the center of diversity of flat-padded Opuntia cacti in the American southwest and Mexico.

Another example is California: 44% of invading terrestrial macroinvertebrates that have established in the state came from populations established elsewhere in the US and Canada (Hoddle 2023). This number exceeds the total number of invasive macroinvertebrates in the state that originated anywhere in Eurasia (Weber et al.).

True, it is very difficult to prevent natural spread. But a lot of this spread is facilitated by human activities, e.g., transporting vectors such as living plants, firewood, outdoor furniture or storage “pods.” I have complained often — here and here and here — that interstate movement of invasive plant pests is particularly poorly controlled.

Some scientists and regulators have responded to these situations by improving phytosanitary programs. California officials, in 2019, set up a program to fund projects aimed at developing integrated pest management strategies for species thought to have a high invasion potential before they arrive. I urge other states to do the same. This would probably be most effective in controlling the target species – and in relation to cost — if developed by regional consortia.

Weber et al. suggest that given continuing unintentional introductions of non-native species, phytosanitary agencies need to focus on those invasion pathways that are particularly likely to result in invasions, e.g. live plants, raw lumber (including wood packaging), and bulk commodities e.g. quarried rock.

The authors also suggest research opportunities that arise from biocontrol agents’ “self-introductions”. These include:

Comparing actual host ranges to those predicted by laboratory and other studies;
Quantifying the role of Allee effects, for example by studying the spread of the glassy-winged sharpshooter and its biocontrol agent across the Pacific region;
Using molecular analyses to disentangle multiple routes of entry (e.g., the “invasive bridgehead effect”) and hybridization.

2. Door-knocker species

Hoddle (2023) suggests further that early detection programs should focus on “door-knocker” species — those likely to enter and cause significant negative impacts. In an earlier article (Hoddle, Mace and Steggall 2018) argued that the benefits of a pro-active biocontrol program outweigh the costs. The authors say the information gained would cut the time needed to deploy effective biocontrol. Ultimately, this could reduce the prolonged and even irreversible ecological and economic disruption from invasive pests, associated pesticide applications, and lost ecological services.

Asian citrus psyllid (*Diaphorina citri*); USDA photo by Justin Wendell; Hoddle cites this species as one that a pro-active biocontrol program should have targetted

Hoddle calls funding pro-active biocontrol research programs before they’re needed as analogous to buying insurance. The owners of insurance policies hope not to need them but benefit when catastrophe strikes. Furthermore, the information gained from early research might identify natural enemy species that could “self-introduce” along with the invading host. Finally, proactive research might clarify whether the increasing number of natural enemy species that are “self-introducing” pose a threat to non-target organisms.

Recognizing the difficulty of identifying an “emerging invasive species” before its introduction, Hoddle endorses other components of prevention programs:

Collaborating with non-U.S. scientists to identify and mitigate invasion bridgeheads. Such efforts would both lessen bioinvasion threats and possibly aid in determining native ranges and facilitating location of natural enemies.
Sentinel plantings, such as those established under the International Plant Sentinel Network. These plantings can also support research on natural enemies of key pests.
Integrating online platforms, networks, professional meetings, and incursion monitoring programs into “horizon scans” for potential invasive species. He mentions specifically PestLens; online community science platforms, e.g., iNaturalist; international symposia; and official pest surveillance, e.g., U.S. Forest Service’s bark beetles survey and surveys done by the California Department of Food and Agriculture and border protection stations.

date palm mealybug (*Pseudaspidoproctus hyphaeniacus*); threat to native *Washingtonia* palms of California; one of pests tracked by PestLens

Weber et al. also support the concept of sentinel plant nurseries – especially because accidental plant and herbivore invasions often occur at the same points of entry.

Both Weber et al. and Hoddle urge authorities not to strengthen regulations governing biocontrol introductions. Weber et al. say that would be to make perfect the enemy of the good. The need is to balance solving problems with avoiding creation of new problems.

SOURCES

Hoddle, M.S., K. Mace, J. Steggall. 2018. Proactive biological control: A cost-effective management option for invasive pests. California Agriculture. Volume 72, No. 3

Hoddle. M.S. 2023. A new paradigm: proactive biological control of invasive insect pests. BioControl https://doi.org/10.1007/s10526-023-10206-5

Weber, D.C. A.E. Hajek, K.A. Hoelmer, U. Schaffner, P.G. Mason, R. Stouthamer, E.J. Talamas, M. Buffington, M.S. Hoddle, and T. Haye. 2020. Unintentional Biological Control Chapter for USDA Agriculture Research Service. Invasive Insect Biocontrol and Behavior Laboratory. https://www.ars.usda.gov/research/publications/publication/?seqNo115=362852

Posted by Faith Campbell

www.fadingforests.org

The White Mountain forest in New Hampshire: 80 years of change, with more ahead

I have been disappointed that a research symposium focused on the northern hardwood forest workshop gave little attention to non-native pests (see citation at end of this blog). A new study based in the Bartlett Experimental Forest in the White Mountains of New Hampshire is more balanced. Ducey et al. (full citation at the end of this blog) analyzed changes in the forest’s species composition and tree size over the past 80 years.

They found that trees of nearly all species are growing into larger sizes as the forest continues to age since the last widespread clearing at the end of the 19^th Century. The same aging is causing a rapid decline in two shade-intolerant species – paper birch (Betula papyrifera) and aspen (Populus tremuloides and P. grandidentata) – which had grown quickly once the cleared areas were abandoned. The mid-shade -tolerant species yellow birch (Betula alleghaniensis) also is declining. Together, the birch and aspen species have declined from a quarter to a third of basal area in 1931 to 10 – 12% in 2015.

Some developments are unexpected. Red maple (Acer rubrum) increased in abundance until the early 1990s, but that growth then levelled off. Sugar maple (Acer saccharum) has declined in abundance except where the forest is managed to retain it.

There is little evidence of tree species migrating upward on slopes in response to changes in the local climate. Major weather events – a hurricane in 1938 and an ice storm in 1998 — caused significant tree mortality across Bartlett Experimental Forest, but not a dramatic change in forest composition.

Eastern hemlock (Tsuga canadensis) is replacing the disappearing birch and aspen on low elevation sites. Hemlock has increased its proportion of basal area from 8 – 10% to a quarter or more. Despite aggressive management aimed at reducing the tree’s presence, American beech (Fagus grandifolia) is on track to dominate large areas of the Bartlett Experimental Forest. Given the tree-killing pests already present in the region, large increases in eastern hemlock, American beech, and red spruce (Picea rubens) are worrying.

Eastern hemlock creates important wildlife habitat for deer and more than 100 other vertebrate species in New England. However, hemlock woolly adelgid (HWA) has been present in New Hampshire since 2000. It is now within 15-20 km of Bartlett Experimental Forest. There is some hope that the region’s cold temperatures might limit HWA’s spread and impacts, but Ducey et al. expect major change when the adelgid arrives.

Ducey et al. cite a separate study demonstrating that mortality caused by beech bark disease (BBD) can be sufficient to upset carbon storage in old-growth forests. On the Bartlett Forest, nearly 90% of beech trees had become diseased by 1950.

Ducey et al. express concern about the possible impact of beech leaf disease (BLD), as well.

BLD has not yet been detected in the White Mountains or New Hampshire, but is in so New England and coastal Maine. Much remains unknown about the disease, including how it spreads and its long-term impacts.

Ducey et al. do not raise pest concerns about red spruce or balsam fir (Abies balsamea), which co-dominate the Bartlett Forest at higher elevations (above 500 m). This silence is disturbing since red spruce can be killed by the brown spruce longhorned beetle, a European woodborer established in Nova Scotia and threatening to spread south. Balsam firs suffer some mortality from feeding by the balsam woolly adelgid, a Eurasian sap-sucker which has been in New England for more than a century.

White ash (Fraxinus americana) is present as a minor component of the Bartlett Forest. Because it is considered to be a valuable timber species, management has resulted in a modest increase in abundance of ash. Ducey et al. expect dramatic reduction — or even elimination of the species — when the emerald ash borer (EAB) arrives. EAB has been detected within ~ 15 km from Bartlett Experimental Forest.

Ducey et al. conclude that silvicultural management applied at the scope and intensity of that in the Bartlett Experimental Forest has moderated some changes. That is, it is maintaining sugar maple and suppressing the increase of beech. Its effect is secondary, however to overall forest development as the forest ages.

SOURCES

Ducey, M.J, O.L., Yamasaki, M. Belair, E.P., Leak, W.B. 2023. Eight decades of compositional change in a managed northern hardwood landscape. Forest Ecosystems 10 (2023) 100121

Proceedings of the First Biennial Northern Hardwood Conference 2021: Bridging Science and Management for the Future. USDA Forest Service Northern Research Station General Technical Report NRS-P-211, May 2023

Posted by Faith Campbell

www.fadingforests.org

USFS Lays Out Incomplete Picture of the Future

tanoak trees in southern Oregon killed by sudden oak death; photo by Oregon Department of Forestry; this pathogen is not mentioned by USFS RPA report

In August the USDA Forest Service published the agency’s 2020 assessment of the future of America’s forests under the auspices of the Resources Planning Act. [See United States Department of Agriculture Forest Service Future of America’s Forests and Rangelands, full citation at the end of the blog.] To my amazement, this report is the first in the series (which are published every ten years) to address disturbance agents, specifically invasive species. In 2023! Worse, I think its coverage of the threat does not reflect the true state of affairs – as documented by Forest Service scientists among others.

This is most unfortunate because policy-makers presumably rely on this report when considering which threats to focus on.

Here I discuss some of the USFS RPA report and what other authors say about the same topics.

The RPA Report’s Principle Foci: Extent of the Forest and Carbon Sequestration

The USFS RPA report informs us that America’s forested area will probably decrease 1- 2% over the next 50 years (from 635.3 million acres to between 619 and 627 million acres), due largely to conversion to other uses. This decline in extent, plus trees’ aging and increases in disturbance will result in a slow-down in carbon sequestration by forests. In fact, if demand for wood products is high, or land conversion to other uses proceeds apace, U.S. forest ecosystems are projected to become a net source of atmospheric CO₂by 2070.

Eastern forests sequester the majority of U.S. forest carbon stocks. These forests are expected to continue aging – thereby increasing their carbon storage. Yet we know that these forests have suffered the greatest impact from non-native pests.

I don’t understand why the USFS RPA report does not explicitly address the implications of non-native pests. In 2019, Songlin Fei and three USFS research scientists did address this topic. Fei et al. estimated that tree mortality due to the 15 most damaging introduced pest species have resulted in releases of an additional 5.53 terragrams of carbon per year. Fei and colleagues conceded this is probably an underestimate. They say that annual levels of biomass loss are virtually certain to increase because current pests are still spreading to new host ranges (as demonstrated by detection of the emerald ash borer in Oregon). Also, infestations in already-invaded ranges will intensify, and additional pests will be introduced (for example, beech leaf disease).

I see this importance of eastern forests in sequestering carbon as one more reason to expand efforts to protect them from new pest introductions, and the spread of those already in the country, etc.

A second issue is the role of non-native tree species in supporting the structure and ecological functions of forests. Ariel Lugo and colleagues report that 18.8 million acres (7.6 million ha, or 2.8% of the forest area in the continental U.S.) is occupied by non-native tree species. (I know of no overall estimate for all invasive plants.) They found that non-native tree species constitute 12–23% (!) of the basal area of those forest stands in which they occur.

Norway maple (*Acer platanoides*); one of the most widespread invasive species in the East. Photo by Hermann Falkner via Flickr

Lugo and colleagues confine their analysis of ecosystem impacts to carbon sequestration. They found that the contribution of non-native trees to carbon storage is not significant at the national level. In the forests of the continental states (lower 48 states), these trees provide 10% of the total carbon storage in the forest plots where they occur. (While Lugo and colleagues state that the proportion of live tree biomass made up of non-native tree species varies greatly among ecological subregions, they do not provide examples of areas on the continent where their biomass – and contribution to carbon storage — is greater than this average.) In contrast, on Hawai`i, non-native tree species provide an estimated 29% of live tree carbon storage. On Puerto Rico, they provide an even higher proportion: 36%.

Brazilian pepper (Schinus terebinthifolius) – widespread invasive in Hawai`i and Florida; early stage invasive in Puerto Rico. Photo by Javier Alexandro via Flickr

In the future, non-native trees will play an even bigger role. Since tree invasions on the continent are expanding at ~500,000 acres (202,343 ha) per year, it is not surprising that non-native species’ saplings provide 19% of the total carbon storage for that size of trees in the lower 48 states (Lugo et al.).

Forming a More Complete Picture: Biodiversity, Disturbance, and Combining Data.

The USFS RPA report has a chapter on biodiversity. However, the chapter does not discuss historic or future diversity of tree species within biomes, nor the genetic diversity within tree species.

Treatment of Invasive Species

The USFS 2020 RPA report is the first to include a chapter on disturbance, including invasive species. I applaud its inclusion while wondering why they have included it only now? Why is the coverage so minimal? I think these lapses undercut the report’s purpose. The RPA is supposed to inform decision-makers and stakeholders about the status, trends, and projected future of renewable natural resources and related economic sectors for which USFS has management responsibilities. These include: forests, forest products, rangelands, water, biological diversity, and outdoor recreation. The report also has not met its claim to “capitalize on” areas where the USFS has research capacity. One excuse might be that several important publications have appeared after the cut-off date for the assessment (2020). Still, the report’s authors cite some of the evaluations that were in preparation as of 2020, e.g., Poland et al.

I suggest also that it would be helpful to integrate data from other agencies, especially the invasive species database compiled by the U.S. Geological Survey, into the RPA. For example, the USGS lists just over 4,000 non-native plant species in the continental U.S. (defined as the lower 48 plus Alaska). On Hawai`i, the USGS lists 530 non-native plant species as widespread. Caveat: many of the species included in these lists probably coexist with the native plants and make up minor components of the plant community.

Specifically: Invading Plants

The USFS RPA report gives much more attention to invasive plants than non-native insects and pathogens. The report relies on the findings of Oswalt et al., who based their data on forested plots sampled by the Forest Inventory and Analysis (FIA) program. (The RPA also reports on invasive plants detected on rangelands, primarily grasslands.) Oswalt et al. found that 39% of FIA plots nationwide contained at least one plant species that the FIA protocol considers to be invasive and monitors. The highest intensity of plant invasions is in Hawai`i – 70% of the plots are invaded. The second-greatest intensity is in the eastern forests: 46%. However, the map showing which plots were inventoried for invasive plants makes clear how incomplete these data are – a situation I had not realized previously.

I appreciate that the USFS RPA report mentions that propagule pressure is an important factor in plant invasions. This aspect has often been left out in past analyses. I also appreciate the statement that international trade in plants for ornamental horticulture will probably lead to additional introductions in the future. Third, I concur with the report’s conclusions that once forest land is invaded, it is unlikely to become un-invaded. Invasive plant management in forests often results in one non-native species being replaced by another. In sum, the report envisions a future in which plant invasion rates are likely to increase on forest land.

If you wish to learn more about invasive plant presence and impacts, see the discussion of invasive plants in Poland et al., my blogs based on the work by Doug Tallamy, and several other of my blogs compiled under the category “invasive plants” on this website.

I believe all sources expect that the area invaded by non-native plant species, and the intensity of existing invasions, will increase in the future.

The USFS RPA links these invasions to expansion of the “wildland-urban interface” (“WUI”). These areas increased rapidly before 2010. At that time, they occupied 14% of forest land. The report published in 2023 did not assess their future expansion over the period 2020 to 2070. However, it did project increased fragmentation in many regions, especially in the RPA Western and Southeastern regions. Since “fragmentation” is very similar to wildland-urban interfaces, the report seems implicitly to project more widespread plant invasions in the future.

plant invasions facilitated by fragmentation; northern Virginia; photo by F.T. Campbell

Specifically: Insects and Pathogens

The USFS RPA report on insects and pathogens is brief and contains puzzling errors and gaps. It says that the tree canopy area affected by both native and non-native mortality-causing agents has been consistently large over the three most recent five-year FIA assessment periods. It notes that individual insects or diseases have extirpated entire tree species or genera and fundamentally altered forests across broad regions. Examples cited are chestnut blight and emerald ash borer.

The USFS RPA report warns that pest-related mortality might be underreported in the South, masked by more intense management cycles and higher rates of tree growth and decay. On the other hand, the report asserts that pest-related mortality is probably overrepresented in the Northern Region in the 2002 – 2006 period because surveyors drew polygons to encompass large areas affected by EAB and balsam woolly adelgid (Adelges piceae) infestations. The latter puzzles me; I think it is probably an error, and should have referred to hemlock woolly adegid, A. tsugae. Documented mortality has generally been much more widespread from insects than diseases, e.g., bark beetles, including several native ones, across all regions and over time, especially in the West – where the most significant morality agents are several native beetles. The USFS RPA report mentions that the Northern Region has been particularly affected by non-native pests, including EAB, HWA, BWA, beech bark disease, and oak wilt. It mentions that Hawai`i has also suffered substantial impacts from rapid ʻōhiʻa death.

Defoliating insects have affected relatively consistent area over time. This area usually equaled or exceeded the area affected by the mortality agents. Principal non-native defoliators in the Northern Region have been the spongy moth (Lymantria dispar); larch casebearer (Coleophora laricella); and winter moth (Operophtera brumata). In the South they list the spongy moth.

More disturbing to me is the USFS RPA report’s conclusion that the future impact of forest insects is highly uncertain. The authorsblame the complexity of interactions among changing climate, those changes’ effects on insect and tree species’ distributions, and overall forest health. Also, they name uncertainty about which new non-native species will be introduced to the United States. I appreciate the report’s avoidance of blanket statements regarding the effects of climate change. However, other studies – e.g., Poland et al. – have incorporated these complexities while still offering conclusions about a number of currently established non-native pests. Finally, I am particularly dismayed that the USFS RPA does not provide analysis of any forest pathogens beyond the single mention of a few.

I am confused as to why the USFS RPA report makes no mention of Project CAPTURE (Conservation Assessment and Prioritization of Forest Trees Under Risk of Extirpation). This is a multi-partner effort to prioritize U.S. tree species for conservation actions based on invasive pests’ threats and the trees’ ability to adapt to them. Several USFS units participated, including the Southern Research Station, the Eastern Forest Environmental Threat Assessment Center, and the Forest Health Protection program. The findings were published in 2019. See here. Lead scientist Kevin Potter was one of the authors of the RPA’s chapter on disturbance.

redbay (*Persea borbonia*) trees in Georgia killed by laurel wilt; photo by Scott Cameron. Redbay is ranked by Project CAPTURE as 5th most severely at risk due to a non-native pest

“Project CAPTURE” provided useful summaries of non-native pests’ impacts, including the facts that

54% of the tree species on the continent are infested by one or more non-native insect or pathogen;
nearly 70% of the host/agent combinations involve angiosperm (broadleaf) species, 30% gymnosperms (e.g., conifers). When considering only non-native pests, pests attacking angiosperms had greater average severity.
Disease impacts are more severe, on average, than insect pests. Wood-borers are more damaging than other types of insect pests.
Non-native agents have, on average, considerably more severe impacts than native pests.

Project CAPTURE also ranked priority tree species based on the threat from non-native pests (Potter et al., 2019). Tree families at the highest risk to non-native pests are: a) Fagaceae (oaks, tanoaks, chestnuts, beech), b) Sapindaceae (soapberry family; includes maples, Aesculus (buckeye, horsechestnut); c) in some cases, Pinaceae (pines); d) Salicaceae (willows, poplars, aspens); e) Ulmaceae (elms) and f) Oleaceae (includes Fraxinus). I believe this information should have been included in the Resources Planning Act report in order to insure that decision-makers consider these threats in guiding USFS programs.

I also wish the USFS RPA had at least prominently referred readers to Poland et al. Among that study’s key points are:

Invasive (non-native) insects and diseases can reduce productivity of desired species, interactions at other trophic levels, and watershed hydrology. They also impose enormously high management costs.
Some non-native pests potentially threaten the survival of entire tree genera, not just individual species, e.g., emerald ash borer and Dutch elm disease. I add white pine blister rust and laurel wilt.
Emerald ash borer and hemlock woolly adelgid are listed as among the most significant threats to forests in the Eastern US.
White pine blister rust and hemlock woolly adelgid are described as so profoundly affecting ecosystem structure and function as to cause an irreversible change of ecological state.
Restoration of severely impacted forests requires first, controlling the non-native pest, then identifying and enriching – through selection and breeding – levels of genetic resistance in native populations of the impacted host tree. Programs of varying length and success target five-needle pines killed by Cronartium ribicola; Port-Orford cedar killed by the oomycete Phytophthora lateralis; chestnut blight; Dutch elm disease; butternut canker (causal agent Ophiognomonia clavigignenti juglandacearum), emerald ash borer; and hemlock woolly adelgid.
Climate change will almost certainly lead to changes in the distribution of invasive species, as their populations respond to increased variability and longer-term changes in temperature, moisture, and biotic interactions. Predicting how particular species will respond is difficult but essential to developing effective prevention, control, and restoration strategies.

Poland et al. summarizes major bioinvaders in several regions. Each region except Hawai`i (!!) includes tree-killing insects or pathogens.

It is easier to understand the RPA report’s not mentioning priority-setting efforts by two other entities, the Morton Arboretum and International Union for the Conservation of Nature (IUCN). These studies were published in 2021 and their lead entities were not the Forest Service – although the USFS helped to fund the U.S. portion of the studies.

The Morton Arboretum led in the analysis of U.S. tree species. It published studies evaluating the status of tree species belonging to nine genera, considering all threats. The Morton study ranked as of conservation concern one third of native pine species; 31% of native oak species; significant proportion of species in the Lauraceae. The report on American beech — the only North American species in the genus Fagus – made no mention of beech leaf disease – despite it being a major concern in Ohio – only two states away from the location of the Morton Arboretum near Chicago.

valley oak (*Quercus lobata*) in Alameda Co, California; photo by Belinda Lo via Flickr

Most of the species listed by the Morton Arboretum are of conservation concern because of their small populations and restricted ranges. The report’s coverage of native pests is inconsistent, spotty, and sometimes focuses on odd examples.

Tree Species’ Regeneration

Too late for consideration by the authors of the USFS RPA report come new studies by Potter and Riitters that evaluate species at risk due to poor regeneration. This effort evaluated 280 forest tree species native to the continental United States – two-thirds of the species evaluated in the Kevin Potter’s earlier analysis of pest impacts.

The results of Potter and Riitters 2023 only partially matched those of the IUCN/Morton studies. The Morton study did not mention three genera with the highest proportions of poorly reproducing species according to Potter and Riitters: Platanus, Nyssa, and Juniperus. Potter, Morton, and the IUCN largely agree on the proportion of Pinus species at risk. Potter et al. 2023 found about 11% of oak species to be reproducing poorly, while Morton designated a third of 91 oak species to be of conservation concern.

I believe Potter and Riitters and the Morton study agree that the Southeast and California are geographic hot spots of tree species at risk.

Potter and Riiters found that several species with wide distributions might be at risk because they are reproducing at inadequate rates. Three of these exhibit poor reproduction across their full range: Populus deltoids (eastern cottonwood), Platanus occidentalis (American sycamore), and ponderosa pine(Pinus ponderosa). Four more species are reported to exhibit poor reproduction rates in all seed zones in which they grow (the difference from the former group is not explained). These are two Juniperus, Pinus pungens, and Quercus lobata. As I point out in my earlier blog, valley oak is also under attack by the Mediterranean oak borer.

SOURCES

Fei, S., R.S. Morin, C.M. Oswalt, and A.M. 2019. Biomass losses resulting from insect and disease invasions in United States forests. Proceedings of the National Academy of Sciences. Vol. 116, No. 35. August 27, 2019.

Lugo, A.E., J.E. Smith, K.M. Potter, H. Marcano Vega, and C.M. Kurtz. 2022. The Contribution of Nonnative Tree Species to the Structure and Composition of Forests in the Conterminous United States in Comparison with Tropical Islands in the Pacific and Caribbean. USDA USFS General Technical Report IITF-54

Poland, T.M., T. Patel-Weynand, D.M. Finch, C.F. Miniat, D.C. Hayes, V.M. Lopez, eds. 2021. Invasive Species in Forests and Rangelands of the United States: A Comprehensive Science Synthesis for the United States Forest Sector. Springer Verlag. Available gratis at https://link.springer.com/book/10.1007/978-3-030-45367-1

Potter, K.M., M.E. Escanferla, R.M. Jetton, G. Man, and B.S. Crane. 2019. Prioritizing the conservation needs of United States tree species: Evaluating vulnerability to forest insect and disease threats. Global Ecology and Conservation.

Potter, K.M. and Riitters, K. 2023. A National Multi-Scale Assessment of Regeneration Deficit as an Indicator of Potential Risk of Forest Genetic Variation Loss. Forests 2022, 13, 19. https://doi.org/10.3390/f13010019

United States Department of Agriculture Forest Service. 2023. Future of America’s Forests and Rangelands: The Forest Service 2020 Resource Planning Act Assessment. GTR-WO-102 July 2023

Posted by Faith Campbell