New report: Forest Disturbances in the West and their implications for sustainability

whitebark pine killed by white pine blister rust; Crater Lake National Park; photo by F.T. Campbell

Increasing frequency and severity of forest disturbances pose significant challenges to the sustainable management of forests in the West and to the goods and services they provide. A recent study (Barrett and Robertson 2021; full citation at end of blog) found that natural and human-caused disturbances affected 22.3% of forest land in the West over a 5-year period.  The study analyzed fire, drought, insects, disease, invasive plants, their interactions, and their socioeconomic impacts. Climate change was found to affect most disturbance processes now and is expected to continue to do so in the future.

The impacts of these disturbances varied; most disturbances did not result in stand-replacing mortality.

Overarching Findings on Disturbance Agents in Western Forests

  • Insect and disease outbreaks were the most extensive disturbance types. Each was estimated to affect 6.1 million hectares. Insect and disease outbreaks also caused the highest levels of tree mortality. This finding resulted from what was described as a relatively “low” threshold for “disturbance.” The authors set this threshold at disturbances that cause damage or mortality to 25% of trees in a stand or 50% of an individual tree species.

The overwhelmingly important causal agent was the mountain pine beetle (MPB; Dendroctonus ponderosae). Even after an approximately 50% drop in mortality after its peak years in the 2000s, MPB caused almost half the total area affected by all bark beetles combined  2000-2016.

The great majority of “pest” organisms causing disturbance in the West are native. Some non-native pests are important, though, and they are expected to become more important in the future. The most damaging non-native agent is white pine blister rust (WPBR; Cronaritum ribicola). Despite the largest control effort (in the 1930s), WPBR has caused drastic declines in white pines in the West. Currently attention focuses on high-elevation pines, especially whitebark pine (Pinus albicaulis), which is suffering extensive mortality from a combination of drought, MPB, and WPBR.

tanoak mortality in Big Sur, California; photo by Matteo Garbelotto

Other non-native pests discussed in the report are balsam woolly adelgid, larch casebearer, spruce aphid, and sudden oak death (SOD). The report notes the presence of a second strain of the causal agent of SOD (Phytophthora ramorum). In June 2021, a third strain was detected in Oregon forests (COMTF newsletter). There are mere mentions of goldspotted oak borer and polyphagous shot hole borer. The California fivespined ips (Ips paraconfusus) is reported to vector the fungus Fusarium circinatum which causes pitch canker disease in Monterey pine (Pinus radiata).

  • The second most extensive disturbance agent in the West is human activity – silvicultural management and conversion to non-forest land uses. These activities affected 4.4 million ha.
  • The third most extensive disturbance agent is grazing (primarily livestock). This affected 3.9 million ha.
  • Fire thus ranks fourth as a disturbance agent – as measured by extent. During a five-year period ending in 2017 or 2018, fire affected 3.7 million ha. (I don’t know whether this ranking will change in response to the fire cataclysms of the most recent years; apparently the latest year included in the data was 2017.) The area affected by fire during this period was double that of the period 1960 to 2000. However, fire frequency and extent were still considerably lower than in the 1920s through1940s, before the advent of fire suppression, especially in the drier forests of the interior West.
  • Other disturbance events – including those caused by weather and vegetation (presumably invasive plants) – affected far smaller areas: a total of 2.3 million ha.

Furthermore, drought and invasive plants – while increasing in extent & intensity – are often considered contributing factors rather than as proximate causes.

Data on past disturbance extents are poor for all these causes except fire. Analysis is further complicated by the high variability of disturbance events – year to year and across space. It is also often difficult to determine the ultimate causes. This makes the implications of these recent increases difficult to ascertain.

As the report points out, forest conditions are inherently dynamic, not stable. They note particularly human manipulation of fire – originally setting fires and then, more recently, suppressing them, has shaped the region’s forests for centuries. Fire suppression has significantly altered forest structure throughout the region, resulting in increasing fuel loads, decreasing resilience to fire and other disturbances.

Impacts of Climate Change

Fire suppression has also increased rates of carbon sequestration (see below).

The report notes that while past timber harvest, land clearing, insect outbreaks, and fires have reduced carbon stocks in forests across the United States to about half their maximum storage potential, recent vegetation and forest cover dynamics have resulted in net increases in carbon stocks in the West – despite CO2 emissions from trees killed by fire and insect damage since 2000.

In the future, climate change is expected to increase tree mortality substantially. In drier forests, mortality would result from increased fire incidence facilitated by a combination of longer fire season and decreased snowpack, reduced summer precipitation, and higher temperatures. In high-elevation and mesic forests, mortality would result from reduced snowpack, precipitation, and temperature.

About half of the West is likely to experience unprecedented climates by the end of this century. This change in climate could trigger changes in vegetation types and extent, net primary productivity, wildfire frequency, and expansion of the range of tree-damaging pests. Grasslands, chaparral, and montane forests are expected to expand; subalpine forests, tundra, and Great Basin woodlands are expected to contract. 

Except in Arizona, California, and New Mexico, bark beetles are having a larger impact on forest CO2 emissions than is fire. Future impacts are unclear. Under moderate climate conditions forests would grow faster than under more severe scenarios, but they would thereby generate more fuel for the fires likely to occur during dry years. These fires might ultimately lead to lower carbon stocks.

I have addressed the invasive plant data in a separate blog.

Reducing Impacts via Management

Barrett and Robertson (2021) suggest management actions that could reduce the impact of these disturbances. First, they mention actions aimed at reducing invasions by non-native insects, pathogens, and plants. Also, they name actions to ameliorate climate change, such as reducing greenhouse gas emissions or increasing carbon sequestration and storage to mitigate expected future damage from wildfire, drought, and beetles.

They recommended a series of on-the-ground management actions: fuel reduction treatments; thinning to reduce tree mortality from drought; favoring species that do not host specific pests; and planting genetically resistant varieties. They call for caution to prevent transport of pathogens to new areas during restoration planting of nursery stock or in “assisted migration” projects. Economic impacts of disturbance events on recreation could be mitigated by altering the timing and duration of recreational site visits. The authors also note that the best choices will differ both by site-specific factors and by management goals. They call for community education programs, cooperative stewardship across multiple agencies and landowners, and local and regional planning.

Details on Pest Impacts

Disease dominated in the high elevations of interior mountain ranges and in the precipitation-heavy regions of Oregon and Washington. Even in these locations, mortality levels are often low, resulting in multi-aged stands with complex structure. Patterns of disturbance are expected to change as pathogens and their hosts adapt to climate change. The microbes might evolve more rapidly than the host trees.

test planting of rust-resistant seedlings of whitebark pine at Crater Lake NP; photo by Richard Sniezko

Sudden oak death (SOD) is now the leading biotic cause of tree mortality in coastal forests of California [and possibly Oregon?]. In heavily infested areas SOD has caused conversion of previously tanoak-dominated stands. The report provides a summary of Oregon’s attempts to eradicate SOD from 2001 to 2012.

I am surprised by the failure to mention non-native pest impacts on two narrowly endemic species: Port-Orford cedar root disease and pitch canker disease in Monterrey pines – other than to mention the vector (above).

test planting of disease-resistant Port-Orford cedar; photo by Richard Sniezko

Insect outbreaks were most common in pine forests. Decades of fire suppression, and now climate change, have substantially altered forest conditions over millions of hectares, primarily increasing the density of shade-tolerant and fire-intolerant trees (e.g., true firs, Abies spp.). Balsam woolly adelgid (BWA; Adelges piceae) is now threatening subalpine fir stands in British Columbia, Oregon, Washington, Idaho, Montana, and Utah. BWA is ranked as the 10th most damaging forest insect, first among non-native species over the next few decades (2013-2027). The spruce aphid (Elatobium abietinum) is having its most significant impact in coastal Southeast Alaska on Sitka spruce and in Arizona on Engelmann spruce. Projected increases in temperature and the frequency of droughts in the West will likely make spruce aphid a more significant disturbance agent in coming decades.

risk map for goldspotted oak borer

In discussing the goldspotted oak borer (GSOB; Agrilus coxalis sic) in California and emerald ash borer (EAB; Agrilus planipennis) in Colorado, Barrett and Robertson (2021) say that the heterogeneity of western landscapes provides some buffer against invasion.  However, I note that GSOB threatens oaks throughout California (see the map at left). EAB threatens riparian areas of the Pacific states (see map below). These riparian areas are admittedly small in geographic extent but ecologically vital.

Barrett and Robertson (2021) expect seven tree species to suffer substantial levels of tree mortality in the near future.  Six are pines threatened in large part by mountain pine beetle, led by the two high-elevation five-needle pines, whitebark pine (58% of total basal area) and limber pine (44%). These are followed by lodgepole pine (39%), ponderosa pine (28%), pinyon pine (27%), Jeffrey pine (26%). The seventh is grand fir (25% of total basal area); the report does not specify which agents are responsible.

Data Issues

The report notes that insects and pathogens are only partially covered by existing monitoring programs. Pathogens are particularly hard to detect and to make conclusive attributions of causality.

SOURCE

Barrett, T.M. and G.C. Robertson, Editors. 2021. Disturbance and Sustainability in Forests of the Western United States. USDA Forest Service Pacific Northwest Research Station. General Technical Report PNW-GTR-992. March 2021

Posted by Faith Campbell

We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.

For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm

Invasive Plants – an Overview

It’s everywhere! multiflora rose (photo by Famartin)

The United States is overrun with introduced plants. Five years ago, Rod Randall’s database listed more than 9,700 non-native plant species as naturalized in the U.S. Not all of these species were “invasive”.

At that time, regional invasive plant groups listed the following numbers of invasive species in their regions:

  • Southeast Exotic Plant Pest Council – approximately 400 invasive plants
  • Mid-Atlantic Invasive Plants Council – 285 invasive plants
  • Midwest Invasive Plants Network reported that state agencies or state-level invasive plant councils in its region listed more than 270 plant species as invasive, noxious, or pest species
  • California Invasive Plants Council listed 208 species.
  • Texas Invasives reported more than 800 non-native plant species in the state, of which 20 were considered invasive.

Species – Rankings and Extents

We know that these invaders are affecting wide swaths of many ecosystems. A recent study based on Forest Inventory and Analysis (FIA) data (explained here) showed that nation-wide, 39% of forested plots sampled contained at least one invasive species.  Hawai`i was first, at 70%. Eastern forests were second, at 46%. In the West overall, 11% of plots contained at least one invasive species. Plots in both Alaska and the Intermountain states were at 6% of plots invaded.  A different study (Barrett and Robertson 2021; full citation at end of blog) reported the proportion of Western forest covered by invasive plants. This approach resulted in different numbers, but the same general ranking: Hawai`i again “led” at 46%; Pacific Coast states at 3.3%; Rocky Mountain states at 0.75%; coastal Alaska at 0.01%.

In more arid regions, data from the Bureau of Land Management showed that invaded acreage had more than doubled between 2009 and 2015.

buffelgrass removal in Tucson; Photo by Julia Rowe, Arizona Sonora Desert Museum

The situation is expected to get worse: a study of just one small portion of U.S. naturalized plants found that non-native plant species were more widely distributed than native species and that the average invasive plant inhabited only about 50% of its expected range. Furthermore, human actions were more important in facilitating spread than the species’ biological attributes.

Most of the detailed studies have been conducted in the Northeast – by both Forest Service and National Park Service scientists. The USFS’ Northern Region (Region 9) contains 24 states, from Maine to Minnesota, from Delaware to Missouri. A review of forest inventory (FIA) data (Oswalt et al. 2015) provided details on 50 plant species.   (Unfortunately, the Southern Region [Region 8] has chosen to report in different formats, so it is hard to get an overall picture of invasive plants throughout the forests of the entire East. This is especially annoying to those of us who live in Mid-Atlantic states, which are divided between the two regions.)

Oswalt et al. (2015) provided data on the percentage of FIA plots in each state that were reported to have at least one invasive plant species. The northern Midwest ranked highest – e.g., one state (Ohio) at 93%; one state (Iowa) at 81%; two states (Indiana and Illinois) above 70%. Parts of the Mid-Atlantic region were almost as invaded – West Virginia at 79% and Maryland at 65%.  The Northern plains states ranked lowest in invasions – North Dakota at 29% and South Dakota at 15%.

A study by the National Park Service of part of the Northeast (from Virginia and West Virginia to Maine) found a situation similar to that found by USFS researchers. In 35 of 39 park units, more than half of the plots had at least one invasive plant species when the 2015-2018 survey began. In 10 parks (a quarter of those surveyed), every plot had at least one. Invasions are worsening: 80% of the park units showed there was a significant increase in at least one trend measuring abundance.

Japanese stiltgrass in Shenandoah National Park; Photo by J. Hughes

The USFS and NPS report different species to be most widespread. In the National Park Service-managed units, Japanese stiltgrass (Microstegium vimineum) was found on 30% of all plots, in more than 75% of all NPS-managed  units in the study. This magnitude comes despite the species not being found north of 41o N latitude. In forest plots inventoried by the USDA Forest Service, Japanese stiltgrass was the 14th most widespread species in the Northern region. I speculate that the species might not be common in the upper Midwest, which was not included in the NPS study. Oswalt et al. (2015) noted that Japanese stiltgrass was the 5th most common invasive plant in the Southern region.

Both studies agreed that garlic mustard (Alliaria petiolata) is widespread. The NPS study found it to be the most frequently detected non-grass herbaceous species, detected in 20% of plots throughout the study area (Virginia and West Virginia to Maine). On forest plots monitored by the USFS, garlic mustard was the 3rd most frequently detected species, on 4.5% of the surveyed plots. The species is reported to be present in 36 states & 5 provinces.

Why do Studies Ignore Deliberate Planting as a Factor?

Both USFS & NPS found shrubs and vines to be highly widespread. NPS specified Japanese barberry (Berberis thunbergii), Japanese honeysuckle (Lonicera japonica), multiflora rose (Rosa multiflora), and wineberry (Rubus phoenicolasius). USFS FIA data showed multiflora rose to be the most frequently recorded invasive plant, present on 16.6% of surveyed plots. It is otherwise recorded in 39 states and 5 provinces. Multiflora rose is almost ubiquitous in some states; in Ohio it is recorded on 85% of the plots. “Roses” were reported to be the 3rd most common invasive plant in the Southern Region. Other shrubs also dominated FIA plot detections: European buckthorn was 4th most frequently detected species, present on 4.4% of survey plots; or in 34 states and 8 provinces. Its presence is highest in New York, at 16.8% of plots. If the plots invaded by the various bush honeysuckle species do not overlap, these shrubs occupy 9.5% of all surveyed plots – second to multiflora rose. The vine Japanese honeysuckle ranked 6th – present on 3.6% of survey plots across the region. Japanese honeysuckle is reported to be the most common invasive plant in the Southern region. Other shrubs ranking 12th or above included Autumn olive and Japanese barberry

Tree-of-heaven (Ailanthus altissima) was the most common invasive tree found in National parks, again, despite not growing north of 41o N latitude. It is found in 9% of plots.

Ailanthus

I will say that I find it extremely annoying that the scientists carrying out these studies never mention that virtually all these shrub species had been deliberately planted in forests or nearby lands! Instead, they focus on such factors as histories of agriculture and other disturbances and fragmentation.  It is well documented (e.g., Lehan et al. 2013) that the vast majority of shrub species introduced to the U.S. were introduced deliberately. Furthermore, more than 500 plant species invasive in some region are being sold on-line globally.

Deliberate planting of species that turn out to be invasive is also rarely recognized in the West, e.g., Pearson et. al. There, the motivation for planting might be livestock forage or erosion control rather than wildlife habitat “enhancement” or ornamental horticulture.

I am pleased that the most recent study (Barrett and Robertson 2021) differs somewhat by noting (sometimes) both invasions by forage grasses and the appearance in the mesic forests of Pacific states such planted species as Armenian blackberry. However, while this report notes the potential that pathogens might be transported to new areas by restoration planting and “assisted migration”, it does not mention the concomitant risk of introducing plant species that might prove invasive in the naïve ecosystems.

English ivy invading forest in Washington State; photo from Washington Noxious Weed Board

[Go to the earlier blogs linked here and the Western forests report for discussions of management strategies.]

Annual reports from the NPS Invasive Plant Management Teams (IPMTs; before FY19, “Exotic”, so EPMTs) provide some information about the agency’s efforts to control invasive Plants. Go to Invasive Plant Management Teams – Biological Resources Division (U.S. National Park Service) (nps.gov) . Scroll down to the short paragraph under the heading “Learn about how the teams are actively working …” This link takes you to reports from FYs 2016 – 2018. Reports from FY19 and FY20 will be added soon. Currently at FY 2019 is at https://irma.nps.gov/DataStore/Reference/Profile/2286813  & FY 2020 is at  https://irma.nps.gov/DataStore/Reference/Profile/2286814

New Information from Study of Forests in the West

Barrett and Robertson (2021) state that although invasive plants are increasing in extent and intensity in Western forests, they are usually considered to be contributing factors rather than as proximate causes. However, they note two caveats: 1) determining the ultimate causes and resulting implications of these recent increases is more difficult; and 2) data are particularly poor on plant species’ presence. Indeed, the FIA survey process link is ineffective for early detection and tactical monitoring [that is, identifying particular species in specific habitats of concern] of plant invasions.

Of the 23.4 M ha of forested lands that have experienced a disturbance over a five-year window (the time frame for FIA), only 600,000 ha was affected by the combined categories of geologic, vegetation, and other disturbances. (This is 10% of the area affected by either insects or pathogens.) Cheatgrass (Bromus tectorum) was by far the most abundant species in Western forests, covering 480,000 ha, or about 0.49%cover of all forested land in the conterminous Western United States. Because of the difficulties of surveying, Barrett and Robertson (2021) conclude that the area covered by IAS plants on the Pacific Coast and Rocky Mountains could be twice recorded values.

FIA surveys detected the highest number of non-native plant species in the forests of the continental Pacific states — 259 species. Many were grasses (although different species than in the Rockies), but shrubs and other forbs were also present. In the Rocky Mountain states the surveys detected a total of 195 non-native species, primarily grasses. FIA surveys in Hawai`i detected 136 non-native species. The most abundant was strawberry guava, which was detected on 9% of the forested area in the state. Surveys of FIA plots in coastal Alaska detected only 8 non-native plant species; common dandelion was the most abundant. Except in Hawai`i, the plants were expected to have substantially lower impacts than in eastern forests

I note that the US Geological Service (Simpson and Eyler, 2018) reports there are approximately 1,754 non-native plants in Hawai`i and 424 in Alaska. Not all are necessarily invasive. And the USGS study covered all of Alaska, not just the southeastern coastal region.

Barrett and Robertson (2021) found that plant invasions are less extensive in older forest stands, mesic stands in contrast to drier areas and those with sparse or open tree canopies, and farther from roads. Thus, invasive plant cover was higher in hardwood and low-elevation and dry conifer forest types than in high-elevation and moist conifer types. In Hawai`i, mean plant cover was more than 40 % in all forest types except cloud forest, where it was 7.8 %. Again, proximity to roads was mentioned in the context of the likelihood of disturbance but no mention was made of the fact that households and businesses (e.g., tourist facilities, even agency facilities!) might deliberately introduce plants – e.g., horticulture.

Barrett and Robertson (2021) expect the impacts of NIS plants on forest lands to increase in the future, due to both additional introductions (despite efforts to prevent such) and spread of established species. They note that every disturbance creates an opportunity for the many ruderal and graminoid species to establish – facilitated by their abundance nearby. They note the significant challenge presented by secondary invaders, which often respond to space made available by “weed control” projects better than natives.

I welcome their concern about shade-tolerant plants apparently increasing in wetter areas of the Pacific coast states. They note that the presence of non-native plants in a forest is less obvious, and the impacts might be more subtle, perhaps primarily affecting tree regeneration through competition or other effects (e.g., promoting fire). Barrett and Robertson (2021) note that many of the shade-tolerant non-native species abundant in temperate Eastern U.S. forests (e.g., garlic mustard) are present in the West and are likely to become important.

SOURCES

Barrett, T.M. and G.C. Robertson, Editors. 2021. Disturbance and Sustainability in Forests of the Western US. USDA Forest Service Pacific Northwest Research Station. General Technical Report PNW-GTR-992

March 2021

Simpson, A., and Eyler, M.C., 2018, First comprehensive list of non-native species established in three major regions of the United States: U.S. Geological Survey Open-File Report 2018-1156, 15 p., https://doi.org/10.3133/ofr20181156.

ISSN 2331-1258 (online)

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 tree-killing 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  (These reports do not discuss invasive plants.)

Tuning in to the News – Mostly Depressing

In late July I participated in the annual meeting of the National Plant Board (NPB) – the organization representing the states’ phytosanitary agencies. USDA’s APHIS, DHS’ Bureau of Customs and Border Protection (CBP), and various industry associations also participated in the meeting. As usual, I learned lots of depressing developments.

A. Old problems continue to vex:

rhododendron plant infested by P. ramorum; photo by Jennifer Parke, Oregon State University

1) Sudden Oak Death in the Nursery Trade – Again!!!

As you might remember, spring 2019 saw an alarming number of plants infested by the sudden oak death pathogen (Phytophthora ramorum) shipped from west coast nurseries to nurseries in 18 states. Another major incident occurred in 2021. The California Oak Mortality Task Force (COMTF) newsletter for June 2021 reports that one nursery in Oregon shipped plants exposed to P. ramorum to big-box stores in 36 states — twice the number of states that received pathogen-exposed plants in 2019.

The first such incident was in 2004 – 17 years ago! Officials of the states that receive these infested plants are angry that every few years they must divert their resources from other duties to inspect nurseries in their states that have been exposed to the pathogen. They note that these “trace-forward” projects cost state governments money and prevent their carrying out other duties; they also impose significant costs on the in-state nurseries due to holds on sales. When infested plants are found, all these costs rise substantially.

The plant health official from Alabama noted that a single west coast nursery that had repeatedly been found to have infected plants shipped 29 lots of host plants to her state in spring 2021. As is clear from the COMTF article, other states also received thousands of plants that had been exposed to the pathogen. The Alabama official questioned why APHIS tilts so far toward a regulatory system that makes it possible for the “exporting” nurseries to ship. The result – too often – is that an infection at one small business can (repeatedly) impose high costs on hundreds of receiving nurseries and states. [I wonder whether anyone has considered a lawsuit against the source nurseries claiming damages? Would that be successful if the regulatory agencies approved the shipments because – at that time – their inspections had failed to detect the problem?]

Officials from the three west coast states, however, want to support their own nurseries’ efforts to relax regulations and maintain or open markets in the central and eastern states. They point to their own considerable efforts to inspect and certify the pest-free status of nurseries in their states.

Because of the different points of view among the states, the National Plant Board per se has never taken a position on the issue.

However, many states – and even APHIS Deputy Administrator El-Lissy – agree that something is not working. So APHIS is in the midst of reviewing its program, with input from NPB members. Such program reviews have been undertaken several times over the past 18 years. So far, they have never produced a program that effectively stops sales of pathogen-infested plants.

2) Contaminated Wood Packaging

Kevin Harriger of CBP reported that over the nine-month period October 2020 – June 2021, CBP intercepted 1,563 shipments that were in violation of ISPM#15, the international rule that requires that wood packaging be treated to kill pests. Most, or 1,148 shipments (73%), lacked the required mark certifying treatment. Four hundred fifteen (26%) of the total number of shipments had a live pest present. Nearly three quarters of the non-compliant shipments transported miscellaneous cargo. This is not a surprise: all of these characteristics are in keeping with past experience.

Meanwhile, APHIS Deputy Director El-Lissy said APHIS was working with importers, exporting countries’ departments of agriculture, and others to improve compliance. Apparently there were two high-profile incidents when shipments of car components were rejected because of ISPM#15 issues. I am trying to learn more about these incidents.

I recently blogged about the pest risk associated with incoming shipping containers and dunnage.  

3) Asian Gypsy Moths (Tussock moths) Still Infesting Ships

Harriger also said that the period 2019-2020 saw the largest number of ships infested by Asian tussock moth eggs since the program began in 2012.  [I am aware that the Entomological Society is searching for a new name for this group of insects.] On average, 12 of 100 approaching vessels was infested. CBP is using sophisticated models to identify regions within Asian ports where conditions exacerbate the risk of moth contamination. CBP can match individual ships’ loading records to this information to pinpoint which are most likely to be infested.

Oregon and Washington continue to find both Asian and European tussock moths in traps along the Columbia River. Such detections prompt eradication programs of varying expense and disruption.

[In April, I blogged about a report evaluating the risk posed by several Asian tussock moths; the report was prepared by experts under the auspices of the North American Plant Protection Organization.]

B. In addition to the arrival of new pests, there is an alarming spread of established ones:

1) Beech leaf disease

State phytosanitary officials reported detections of beech leaf disease (BLD) in Maine and Virginia. The devastating impact of BLD on this hard mast tree species is described here. BLD has now spread through much of southern New England (Connecticut, Rhode Island, Massachusetts) and up the coast to Maine. Connecticut reports that trees of all sizes are affected. Maine reports that the disease is widespread in the central coastal region.

beech trees in Prince William Forest Park

Virginia reported that the disease has been detected in Prince William Forest Park, a forested area south of Washington, D.C., managed by the National Park Service. This detection is too recent to say how widespread it is.

2) Laurel wilt

Kentucky’s plant health officer reported that laurel wilt disease has been detected on sassafras trees in Louisville, at the northern tip of the state and across the river from Ohio. He noted that a second host plant, spice bush, is in the nursery trade. While laurel wilt is not regulated, officials are concerned about its impact in natural forests. Neighboring states are concerned.

sassafras in northern Virginia; photo by F.T. Campbell

I learned by looking at the map that laurel wilt has also been detected in Sullivan County, Tennessee, on the Virginia border.

3) Spotted Lanternfly

This pest of grapes, tree fruits, and a wide variety of native trees is spreading in Pennsylvania, Delaware, New Jersey, and Maryland. It has also been found in Ithaca, NY, and in Connecticut. The populations in Virginia and West Virginia also continue to spread; a disjunct outbreak has been detected in Prince William County, VA. (south of D.C.). Most alarming are disjunct populations in Ohio on the West Virginia border and in Indiana on the Ohio River border with northern Kentucky. See map here.

The Indiana population has been present for several years. The affected woodland is close to RV parks and other facilities that make further spread likely.

California has established an external quarantine targetting the spotted lanternfly .

C. Wrestling with Continuing Issues:

1) States try to compensate for APHIS’ end of regulating the emerald ash borer and firewood

The members of the NPB have spent years discussing the pros and cons of continuing to regulate ash wood to contain the emerald ash borer (EAB). As I blogged earlier, APHIS has ended its regulatory program. One state – Minnesota – is seeking to use an APHIS procedure to get APHIS’ continued protection from importation of EAB-infested wood (presumably from Canada). Under the Federally Recognized State Managed Phytosanitary Program (FRSMP), a state petitions APHIS to recognize its program for a specific pest. If APHIS grants that recognition, the agency will support the state by continuing to regulate imports of that pest or commodities that might transport the pest when they are destined for the regulating state.

The states have also tried to formulate a system to maintain regulation of firewood (nearly all states’ firewood regulations were based on the federal regulation of all hardwoods to prevent transport of the EAB). As part of this process, the NPB developed guidelines for adoption of regulations by the individual states (available here).  The NPB members are just beginning to explore whether  states might set up third-party certification system(s). Among the challenges to any harmonization are states’ differing legal authorities and disagreement on what threat levels should be applied, and for how long.

2) New information about the Asian longhorned beetle in South Carolina

ALB in South Carolina; photo by R. Brad Thompson, APHIS

South Carolina authorities reported that dendrological studies indicated Asian longhorned beetle (ALB) had been present near Charleston, S.C. since 2012, and possibly earlier. The population has the same genetic makeup as the outbreak in Ohio. This might be explained by either transport of infested wood from Clermont County, Ohio, or that wood packaging entering Charleston harbor came from the same part of China. (Charleston is an important port.) In South Carolina, ALB attacks primarily red maple – as is true at the other infestation sites. However, maple densities are much lower in the swamps of South Carolina and scientists don’t know whether the ALB will fly farther or intensify attacks on other host species. Other questions raised by differences between South Carolina and other, more northern, outbreak sites include possible changes in the beetle’s life cycle and flight periods.

Authorities noted the extremely difficult conditions, which impede survey and control efforts – which I described in an earlier blog.

One innovation was sharing of resources: staff from the North Carolina and Tennessee departments of agriculture went to South Carolina to help with surveys. The Resource Sharing Initiative was started a few years ago as a collaborative effort of APHIS and the NPB. This was the first time states tried it. There were several issues that had to be worked out. One issue was the long time it takes to train people to recognize ALB symptoms. All three states’ officials said the project was worthwhile.

black walnut in Fairfax County, VA — in an area where thousand cankers disease has been present for more than a decade; photo by FT Campbell

3) Recinding quarantines of thousand cankers disease of walnut

States which adopted quarantines targetting this insect/pathogen complex a decade ago now think that it poses little risk to black walnut (Juglans nigra) growing in its native range (as distinct from trees planted in the West). Several are in the process of rescinding their quarantines. I think these states have considered the science carefully and are taking the appropriate action.

4) Nursery self-certification – System Set Up; Will Nurseries Participate? Will Customers Support the Process?

Craig Regelbrugge of AmericanHort noted that the SANC program has now been officially launched – it has graduated from being a pilot program. [SANC stands for Systems Approach to Nursery Certification] Participants are exploring incentives to recruit wider participation by nurseries that produce plants and how to get support from plant retailers. SANC is conceived as an elite program for the best nurseries and marketplace leaders. It was never intended to be a remedial program to clean up problem issues such as the P. ramorum debacle. To work, it seems to me, SANC will need to find a way to persuade customers to want to pay more for quality plants. Hence the critical importance of getting retailers involved.

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

Resistance Breeding – a Useful Strategy

Port-Orford cedar resistance trials at Dorena Center; photo courtesy of Richard Sniezko, USFS

I have written several times about the importance of the United States adopting a comprehensive program to address all aspects of introduced forest pests including breeding of trees resistant to the introduced pests. See Fading Forests III from seven years ago; an earlier blog; and Bonello et al. 2020 (full citation at end of blog), in which we proposed the creation of a federal Center for Forest Pest Control and Prevention to implement end-to-end integrated responses to forest pest invasions. A similar view is being voiced internationally; see, e.g., Buggs et al. 2020.

I have seen efforts to restore pest-decimated tree species to the forest lagging. I complained in a recent blog link that the recent USDA Forest Service report on invasive species (Poland et al. 2021) gave a misleading impression that significant effort was being made on resistance breeding to address several pests.

The USFS does support breeding trees resistant to pests, but in my view this support has been inadequate – including in the USFS report. Others think so, too — see Sniezko and Koch 2017. This insufficiency only grows, despite USDA claims to recognize that promoting resistance to introduced forest pests is an essential component of achieving its strategic goals of maintaining or enhancing productivity while ensuring responsible stewardship of resilient natural resources (Federman and Zankowski 2019).

Work at the Dorena Genetic Research Center

The principal and most notable and successful resistance breeding effort has been the Dorena Genetic Resource Center. The Center was established by, and is funded through the USFS Region 6 Genetic Resource (part of the National Forest System) and Forest Health Management programs. The Center has a solid foundation in the expertise and facilities needed to carry out breeding efforts. Also, it has a 50-plus-year track record.

Dorena has supported breeding of white (five-needle) pines and Port-Orford cedar. Dorena also now provides expertise and some facilities to partners exploring a) breeding Oregon ash to resist the emerald ash borer and b) two Hawaiian trees (koa and ‘ōhi‘a) to resist introduced pathogens (see below). Dorena staff is assisting low-budget, shoe-leather efforts to explore breeding of other trees at risk to non-native pests. These programs are described briefly in Box 8 of Poland et al. 2021. Despite this valuable effort with proven success funding to continue Dorena’s work is tenuous.

White Pine Blister Rust — Efforts to develop resistance to white pine blister rust (WPBR) DMF in five-needle pine species (nine grow across the country) began more than 50 years ago. Currently Dorena focuses on whitebark pine (Pinus albicaulis), denizen of high elevations in the West, along with western white pine (P. monticola), sugar pine (P. lambertiana) , limber pine (P. flexilis), southwestern white pine (P. strobiformis), and foxtail pine (P. balfouriana). Testing whitebark for resistance to WPBR began in 2002. Seedling families from >1,300 parent trees are in various stages of testing. The discovery that some whitebark populations have much higher levels and frequency of partial resistance has allowed rapid distribution of seed. The first restoration plantings in the Pacific Northwest were in 2009.

3-year old seedlings of whitebark pine at Crater Lake National Park; photo by Richard Sniezko, USFS

There are many collaborators – especially the National Park Service, Washington State’s Department of Natural Resources, several Tribes, the Whitebark Pine Ecosystem Foundation, and American Forests. However, planting has been hampered by the high cost of restoration in these high elevation ecosystems, lack of frequent good seed crops on the resistant parent trees, and lack of approval to plant in designated wilderness areas. In the areas with the highest levels of resistant parents, management activities that encourage natural regeneration might be successful. In late 2020 the U.S. Fish and Wildlife Service proposed to list whitebark pine as a Threatened species

Oregon ash (F. latifolia) has not yet been attacked by the emerald ash borer, but all expect EAB to spread to the West coast. Dorena and cooperators have already collected seed from ash trees in Oregon and obtained funding for additional collections, to include Washington and California. The seeds are being stored at both Dorena and the USDA Agriculture Research Service facility at Ft. Collins, Colorado. Seedlings from two dozen families have also been planted at Dorena and a center operated by Washington State University, plus at a USFS Northern Research Station research center in Ohio, where EAB is established and they can be tested for resistance to the insect’s attack.

Koa and ‘ōhi‘a in Hawaii — Regeneration of the koa tree (Acacia koa) has been undercut by the koa wilt pathogen, Fusarium oxysporum f. sp. koae. Dorena initiated efforts with the Hawaii Agricultural Research Center (HARC) to respond in 2003. There has been rapid progress screening seedlings to identify resistant parent trees establishing seed orchards, delineating seed zones, and releasing seed with confirmed levels of resistance for reforestation and restoration (Sniezko and Koch 2017; see also Dudley et al. 2020).

‘ōhi‘a trees killed by rapid ‘ōhi‘a death; photo courtesy of J.R. Friday

When the threat to Hawaii`s most widespread tree ‘ōhi‘a (Metrosideros polymorpha) from rapid ‘ōhi‘a death (ROD) pathogens became apparent, the Dorena staff provided advice on breeding strategies. Its Center Geneticist is part of an ad hoc resistance team. Scientists have identified surviving trees in stands affected by ROD on the Big Island using a variety of methods. These include aerial surveys by drones and fixed-wing aircraft. They then began collecting seeds and cuttings. As of spring 2021, they have collected cuttings or seeds from more than 300 ‘ōhi‘a trees belonging to five varieties. The effort is low-cost, using Americorps volunteers coordinated by a single full-time person, a USFS employee. The program is still in its infancy. It will have to find funding to expand its scope to an operational resistance program once more information on resistance is has been obtained.

Other Efforts

Most other breeding programs are small and poorly funded. In fact, they have been described by one USFS scientist as “hobby projects” of a few scientists determined to try this strategy. Not only are efforts minimal; but also retirement of those few scientists can bring an end to the individual project.

There were greater efforts in the past. I have a document (of unknown origin) from 2011 that describes breeding efforts funded by both the National Forest System and USFS Research and Development. Table 1 listed 16 projects for western conifers; Table 2 listed 32 projects funded by R&D. During this period, the USFS provided start-up funds for the Healthy Forests Initiative, a consortium that sought to prove the concept that genetic engineering could quickly produce an American chestnut able to live and reproduce in its native range. This support was in addition to support for The American Chestnut Foundation backcross hybridization program link.

Part of the problem is the longstanding decline in funding and staffing of USFS research program. A graph in Chapter 6 of FFIII shows the decline in numbers of forest entomologists and pathologists over the 20-year period 1985–2007. Wheeler et al. 2015 discuss the parallel decline in tree breeders and geneticists (citation at end of this blog).

Cuts continue. Funding for research conducted by the USFS Research stations on ten non-native pests decreased from $10 million in Fiscal Year 2010 to just $2.5 million in Fiscal Year 2020 – a cut of more than 70%. I have lobbied for increased appropriations for decades.

The need for new approaches and increased effort is more widely asserted. One example is the group I am working with that promotes a new Center for Forest Pest Control and Prevention. Link A second example is the University of Florida’s recent conference of forest health researchers, representatives of the forest products industry, non-governmental organizations, and leaders of universities with forest-resource programs. This group suggested forming a united organization to increase capacity to improve forest health research. An article outlining the proposal is available here.

The Role of Biotechnology in Breeding Resistant Trees

what happened? same tree a few years apart — a TACF hybrid chestnut

Part of the discussion on forest research explores the proper role of biotechnology in tree species’ restoration. Purdue University hosted a related workshop in April 2021, in which I took part. (“Society and Policy Influences on Biotechnology Risk Assessment for Restoration of Threatened Forest Tree Species”). I hope participants will soon publish a paper based on our discussions.

Meanwhile, Revive & Restore, a wildlife conservation organization promoting the incorporation of biotechnologies into standard conservation practice, sponsored a workshop in June 2020. The 57 conservationists, wildlife biologists, restoration specialists, conservation geneticists, ethicists, and social scientists who participated agreed on an appropriate structure for using biotechnology. These included:

  • A broader definition of risk and application of new risk assessment tools;
  • Consideration of the risks of not taking action, as well as going ahead with a proposal;
  • Transparency about social and cultural values and engaging stakeholders
  • Monitoring results to ensure actions have been successful, manage uncertainty, and codify lessons learned.

In the literature I read, the workshops I participate in (e.g., National Academy of Sciences 2019; Purdue’s), biotechnology is seen as a potentially helpful set of tools that must be integrated into broader programs, all having research, tree improvement, restoration, and reforestation components. Such programs must have sustained management and resources stemming from public support. (For more complete descriptions of components of a resistance breeding program, see Sniezko and Koch 2017 (full reference below); or Chapter 6 of FFIII). Activities that must be incorporated include:

  • Germplasm collection and storage (applying the varied strategies that are appropriate);
  • Research to detect and test potential resistance or tolerance;
  • Research to identify techniques for producing propagules;
  • Planting sites that will be secure for decades;
  • Site preparation & planting;
  • Post-planting maintenance; and
  • Monitoring to determine success or problems

During the Purdue workshop, and in my writing, I have emphasized the principal hindrance to progress is the lack of resources being allocated to resistance breeding. USFS and academic scientists determined to pursue breeding approach must scrounge for funds. I describe some of their efforts below.

Collaborations on Breeding for Specific Species

(still) healthy hemlocks in Cook Forest State Park, Pennsylania; photo by F.T. Campbell

USFS Hemlock Woolly Adelgid (HWA) Initiative [apparently no website]

This initiative was developed under the leadership and direction of FHP staff. The list of cooperators includes dozens of state, federal, university and private organizations. The annual budget has averaged between $2.5 and $3.5 million. Most resources are apparently allocated to biocontrol, but some funding has been provided for breeding activities, including:

  • Seed collection and storage for both Carolina and eastern hemlocks. Two seed orchards have been established in western North Carolina. I believe they are protected from the hemlock woolly adelgid (HWA) by application of pesticides.
  • Research on these tree species’ silviculture and ecology, including manipulation of sunlight levels to protect seedlings from the adelgid and promote growth

The 2021-2025 Program – currently under review – foresees more integrated pest management applying biocontrol, chemical control, and silviculture. It aims to maintain the health of hemlocks being used in breeding programs and “explore” hemlock replacement options, such as hybrids or HWA-tolerant hemlocks (Mayfield et al. 2021). This effort is encouraging, but I have heard complaints from academics that they can’t get funding to pursue what they regard as promising breeding strategies.

Other small programs to breed resistant hemlocks are under way. The Forest Restoration Alliance (formerly the Alliance to Save Threatened Forests) asks citizens to identify surviving hemlocks and balsam firs. The Alliance has collected and propagated both cuttings and seeds and is testing their resistance.

Ash and Other Trees of the Upper Midwest

To date, few resources have been allocated to resistance breeding of ash. Between 2003 and 2017, only about 7% of research funds allocated to ash and emerald ash borer DMF have been devoted to host resistance. Of the host resistance research, 61% applied to identifying mechanisms, 14% to use of transgenics to develop resistance, only 7% (0.5% of the total research) has supported actual breeding for resistance (Sniezko and Koch 2017).

In May 2021 the USFS announced it was seeking funds from the water-focused Great Lakes Restoration Initiative. The USFS expects to receive up to $5.4 million for reforestation, ecosystem restoration. and forest health improvements on non-federal lands in the Great Lakes basin. (This includes parts of the states of Illinois, Indiana, Michigan, Minnesota, New York, Ohio, Pennsylvania and Wisconsin). The announcement doesn’t mention resistance breeding for ash, beech, hemlock, or other trees in the region. I hope some of the new funds will be allocated to this strategy.

“lingering” ash; photos courtesy of Jennifer Koch, USFS

In an encouraging show of entrepreneurship, USFS scientists and others have formed the Great Lakes Basin Forest Health Collaborative – a partnership with Holden Forests and Gardens, American Forests, and USFS (Kappler et al. 2021). The purpose of the Collaborative is to advance resistance breeding for these important tree species. The initial focus is on the five ash species in the region, especially black ash (Fraxinus nigra) link to blog The Collaborative is recruiting a network of partners, and will provide training and technology transfer. The partners will provide volunteers and other resources. Partners do not have to be within the region if their work helps the Great Lakes Basin, but plantings have to be there.

Partners will help identify survivor trees with potential resistance (e.g., “lingering” ash); establish clone banks and/or seed orchards; and manage seed collections. Each partner will operate independently, but in collaboration with the others. The initial focus is on obtaining representative seed collections of ash and hemlock. Then cloning, testing resistance, and crosses can begin. Eventually select lines will be chosen for bulking up and reintroduction.

In future the Collaboration hopes to engage in breeding hemlocks and identifying beech trees that remain healthy in areas heavily impacted by beech leaf disease (BLD).

Other efforts under way include the Monitoring and Managing Ash (MaMA) Program, based in the Ecological Research Institute in New York State.

Beech trees with resistance to beech bark disease (BBD) were identified as early as the 1980s, but a breeding program was begun only in 2002. A collaborative, multi-agency effort has resulted in the establishment of five regional American beech seed orchards with four others in progress as of 2017. Partners provide a cost-effective process for identifying resistant parent trees. State and National Forest personnel surveyed natural forests for candidate trees and then tested each tree and identified markers associated with resistance (Sniezko and Koch 2017).

Challenges Beyond Breeding

Large-scale restoration of tree species across much of their ranges will require significant inputs of funds, over long time periods, as well as resolving daunting logistical issues.

Some think the most likely scenario will be to plant focal areas, or islands, that can aid future natural regeneration (Sniezko and Koch 2017). The American Chestnut Foundation (TACF) anticipates it will take 1,000 years to re-establish American chestnut DMF across its range through a process of three phases: long-term research and demonstration plantings; a relatively small-scale public horticultural program using trees and/or pollen made available by TACF; and a larger-scale public restoration program using progeny from years of outcrossing and production. (This assumes APHIS approves release of the transgenic “Darling 58” tree, plus – I believe – progress in developing resistance to root disease caused by Phytophthora cinnamomi). Already good progress using focal areas has started with several white pine species, and a national plan is in the works for whitebark pine.

Such efforts will require access to land that can be protected from other uses, e.g., development for decades or centuries. Also it will require management of sites to protect propagules from browsing wildlife (deer, rabbits!), provide adequate water and light, and probably give plantings a competitive advantage in relation to other plants growing there …

non-resistant elms will grow anywhere! photo by F.T. Campbell

And there is the issue of how a relatively small number of resistant propagules will succeed in spreading their improved genetics in areas where non-improved elm, ash, beech and hemlock are reproducing naturally. Is reproduction of unimproved trees likely to continue in the face of new and old pests’ spread? If biocontrol agents succeed in reducing a pest’s impact on a host tree species, will that enhance the competitive ability of unimproved trees to the disadvantage of genetically improved conspecifics? What are realistic expectation for programs, and for their success?

Criteria for Success

Woodcock, Marzano, and Quine (2019) analyzed five breeding programs to identify aspects that contribute to success. Four of the programs were in North America; they targetted chestnut, western white pines, and Sitka spruce & white pine weevil. They concluded that 

  • Success is influenced by the level of resistance present in individual trees, the frequency of resistance in the population, and the heritability of resistance.
  • It is important to consider current and potential future risks to the species in addition to the target pest or pathogen— the benefits of trees resistant to a specific threat are negated if it is susceptible to other threats.
  •  Demand [for a resistant tree to plant] should be evaluated, and the priorities of potential supporters and end users should inform the methods used to produce resistant trees.
  •  Operational deployment should balance the urgency of the threat with the consequences if resistant material does not perform as hoped. Urgency might differ for an emerging pest or pathogen.
  • Deployment strategies should be informed by the risks of imposing a strong selection pressure on the pest or pathogen to evolve to overcome host resistance, and by potential impacts on partially resistant trees.
  • Continued monitoring of field performance is important for evaluation, and can help to identify and mitigate emerging threats (e.g. new pathogen strains).

SOURCES

Bonello, P., F.T. Campbell, D. Cipollini, A.O. Conrad, C. Farinas, K.J.K. Gandhi, F.P. Hain, D. Parry, D.N. Showalter, C. Villari, and K.F. Wallin. 2020. Invasive tree pests devastate ecosystems – A proposed new response framework. Frontiers in Forests and Global Change. January 2020. Volume 3, Article 2 

Buggs, R.J.A. 2020 Changing perceptions of tree resistance research. Plants, People, Planet. 2020; 2: 2– 4. https://doi.org/10.1002/ppp3.10089

Dudley, N.; Jones, T.; Gerber, K.; Ross-Davis, A.L.; Sniezko, R.A.; Cannon, P.; Dobbs, J. 2020. Establishment of a Genetically Diverse, Disease-Resistant Acacia koa A. Gray Seed Orchard in Kokee, Kauai: Early Growth, Form, and Survival. Forests 2020, 11, 1276

Federman, S. and P. Zankowski. 2019. Strategic science planning for responsible stewardship and plant protection at the U.S. Department of Agriculture. Plants, People, Planet © New Phytologist Trust 2019;00:1–4. https://doi.org/10.1002/ ppp3.10075

Kappler, R., C. Blashka, D. burke, E. Hall, C. Pike, J. Koch. 2021. Great Lakes Basin Forest Health Collaborative: What it’s all about. North American Forest Insect Work Conference 28 May 2021

Mayfield, A.E. III, Salom, S., Jetton, R., Havill, N., Rhea, R., and Mausel, D. 2021. North American Forest Insect Work Conference 28 May 2021. Spread, impact and management of HWA in eastern North America

National Academies of Sciences, Engineering, and Medicine. 2019. Forest Health and Biotechnology: Possibilities and Considerations. Washington, DC: The National Academies Press. https://doi.org/10.17226/25221.

Poland, T.M., P. Patel-Weynand, D.M Finch, C.F. Miniat, D.C. Hayes, V.M Lopez, editors. 2021. Invasive Species in Forests and Rangelands of the United States. A Comprehensive Science Synthesis for the US Forest Sector. Springer

Sniezko, R.A. and J. Koch. 2017. Breeding trees resistant to insects & diseases: putting theory into application. Biol Invasions. 2017. 19:3377-3400. DOI 10.1007/s10530-017-1482-5

Wheeler, N.C., K.C. Steiner, S.E. Schlarbaum, D.B. Neale. 2015. The Evolution of Forest Genetics and Tree Improvement Research in the United States, Journal of Forestry, Volume 113, Issue 5, September 2015, Pages 500–510, https://doi.org/10.5849/jof.14-120

Woodcock, P., M. Marzano, C.P. Quine. 2019. Key lessons from resistant tree breeding programmes in the Northern Hemisphere. Annals of Forest Science (2019)76:51 https://doi.org/10.1007/s13595-019-0826-y

Posted by Faith Campbell

We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.

For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm