resistance breeding – Page 3 – Center for Invasive Species Prevention

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

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

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

In summary, this bill will:

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

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

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

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

Posted by Faith Campbell

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

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

Interactions of 2 (of 3) Threats to Beech

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

Threats

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

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

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

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

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

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

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

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

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

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

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

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

Ontario disease assessment

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

U.S. disease assessment

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

Distribution and Effects of Beech Scale and BBD

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

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

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

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

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

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

Precautionary Research and Management

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

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

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

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

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

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

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

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

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

SOURCES

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

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

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

Posted by Faith Campbell

Breeding Pest Resistance in Trees – Thoughtful Perspectives

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

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

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

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

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

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

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

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

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

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

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

TACF American chestnut; photo by F.T. Campbell

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

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

Other Significant Articles

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

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

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

Port-Orford test seedlings; photo courtesy of Richard Sniezko

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

SOURCE

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

Posted by Faith Campbell

Forest Pests: What’s Improved, What’s Still to Do

sassafras – vulnerable to the rapidly spreading laurel wilt disease; photo by F.T. Campbell

In summer 2019 I posted several blogs summarizing my analysis of forest pest issues after 30 years’ engagement. I reported the continuing introductions of tree-killing insects and pathogens; their relentless spread and exacerbated impacts. I noted the continued low priority given these issues in agencies tasked with preventing and solving these problems. Also, Congress provides not only insufficiently protective policies but also way too little funding. I decried the impediments created by several Administrations; anti-regulatory ideology and USDA’s emphasis on “collaborating” with “clients” rather than imposing requirements.

In my blogs, I called for renewed effort to find more effective strategies – as I had earlier advocated in my “Fading Forests” reports (link provided at the end of this blog), previous blogs, and Lovett et al. 2016

Areas of Progress

Now two years have passed. I see five areas of progress – which give me some hope.

1) Important Activities Are Better Funded than I had realized

a) The US Forest Service is putting significant effort into breeding trees resistant to the relevant pests, more than I had realized. Examples include elms and several conifer species in the West – here and here.

b) USDA has provided at least $110 million since FY2009 to fund forest pest research, control, and outreach under the auspices of the Plant Pest and Disease Disaster Prevention Program (§10201 of the Farm Bill). This total does not include additional funding for the spotted lanternfly. Funded projects, inter alia: explored biocontrol agents for Asian longhorned beetle and emerald ash borer; supported research at NORS-DUC on sudden oak death; monitored and managed red palm weevil and coconut rhinoceros beetle; and detected Asian defoliators. Clearly, many of these projects have increased scientific understanding and promoted public compliance and assistance in pest detection and management.

This section of the Farm Bill also provided $3.9 million to counter cactus pests – $2.7 million over 10 years targetting the Cactoblastis moth & here and $1.2 million over four years targetting the Harissia cactus mealybug and here.

flat-padded *Opuntia* cactus – vulnerable to the *Cactoblastis* moth; National Park Service photo

2) Additional publications have documented pests’ impacts – although I remain doubtful that they have increased decision-makers’ willingness to prioritize forest pests. Among these publications are the huge overview of invasive species published last spring (Poland et al.) and the regional overview of pests and invasive plants in the West (Barrett et al.).

3) There have been new efforts to improve prediction of various pests’ probable virulence (see recent blogs and here.

4) Attention is growing to the importance of protecting forest health as a vital tool in combatting climate change — see Fei et al., Quirion et al., and IUCN. We will have to wait to see whether this approach will succeed in raising the priority given to non-native pests by decision-makers and influential stakeholders.

5) Some politicians are responding to forest pest crises – In the US House, Peter Welch (D-VT) is the lead sponsor of H.R. 1389. He has been joined – so far – by eight cosponsors — Rep. Kuster (D-NH), Pappas (D-NH), Stefanik (R-NY), Fitzpatrick (R-PA), Thompson (D-CA), Ross (D-NC), Pingree (D-ME), and Delgado (D-NY). This bill would fund research into, and application of, host resistance! Also, it would make APHIS’ access to emergency funds easier. Furthermore, it calls for a study of ways to raise forest pests’ priority – thus partially responding to the proposal by me and others (Bonello et al. 2020; full reference at end of blog) to create federal Centers for Forest Pest Control and Prevention.

This year the Congress will begin work on the next Farm Bill – might these ideas be incorporated into that legislation?

What Else Must Be Done

My work is guided by three premises:

1) Robust federal leadership is crucial:

The Constitution gives primacy to federal agencies in managing imports and interstate trade.
Only a consistent approach can protect trees (and other plants) from non-native pests that spread across state lines.
Federal agencies have more resources than state agencies individually or in likely collective efforts – even after decades of budget and staffing cuts.

2) Success depends on a continuing, long-term effort founded on institutional and financial commitments commensurate with the scale of the threat. This requires stable funding; guidance by research and expert staff; and engagement by non-governmental players and stakeholders. Unfortunately, as I discuss below, funding has been neither adequate nor stable.

3) Programs’ effectiveness needs to be measured. Measurement must focus on outcomes, not just effort (see National Environmental Coalition on Invasive Species’ vision document).

Preventing New Introductions – Challenges and Solutions

We cannot prevent damaging new introductions without addressing two specific challenges.

1) Wood packaging continues to pose a threat despite past international and national efforts. As documented in my recent blogs, the numbers of shipping containers – presumably with wood packaging – are rising. Since 2010, CBP has detected nearly 33,000 shipments in violation of ISPM#15. The numbers of violations are down in the most recent years. However, a high proportion of pest-infested wood continues to bear the ISPM#15 mark. So, ISPM#15 is not as effective as it needs to be.

We at CISP hope that by mid-2022, a new analysis of the current proportion of wood packaging harboring pests will be available. Plus there are at least two collaborative efforts aimed at increasing industry efforts to find solutions – The Nature Conservancy with the National Wooden Pallet and Container Association; and the Cary Institute with an informal consortium of importers using wooden dunnage.

2) Imports of living plants (“plants for planting”) are less well studied so the situation is difficult to assess. However, we know this is a pathway that has often spread pests into and within the US. There have been significant declines in overall numbers of incoming shipments, but available information doesn’t tell us which types of plants – woody vs. herbaceous, plant vs. tissue culture, etc. – have decreased.

APHIS said, in a report to Congress (reference at end of blog), that introductions have been curbed – but neither that report nor other data shows me that is true.

Scientists are making efforts to improve risk assessments by reducing the number of organisms for which no information is available on their probable impacts (the “unknown unknowns”).

Solving Issues of Prevention

While I have repeatedly proposed radical revisions to the international phytosanitary agreements (WTO SPS & IPPC) that preclude prevention of unknown unknowns (see Fading Forests II and blog), I have also endorsed measures aimed at achieving incremental improvements in preventing introductions, curtailing spread, and promoting recovery of the affected host species.

citrus longhorned beetle exit hole in bonsai tree; USDA APHIS photo

The more radical suggestions focus on: 1) revising the US Plant Protection Act to give higher priority to preventing pests introductions than to facilitating free trade (FF II Chapter 3); 2) APHIS explicitly stating that its goal is to achieve a specific, high level of protection (FF II Chapter 3); 3) APHIS using its authority under the NAPPRA program to prohibit imports of all plants belonging to the 150 genera of “woody” plants that North America shares with Europe or Asia; 4) APHIS prohibiting use of packaging made from solid wood by countries and exporters that have a record of frequent violations of ISPM#15 in the 16 years since its implementation.

Another action leading to stronger programs would be for APHIS to facilitate outside analysis of its programs and policies to ensure the agency is applying the most effective strategies (Lovett et al. 2016). The pending Haack report is an encouraging example.

I have also suggested that APHIS broaden its risk assessments so that they cover wider categories of risk, such as all pests that might be associated with bare-root woody plants from a particular region. Such an approach could speed up analyses of the many pathways of introduction and prompt their regulation.

Also, APHIS could use certain existing programs more aggressively. I have in mind pre-clearance partnerships and Critical Control Point integrated pest management programs. APHIS should also clarify the extent to which these programs are being applied to the shipments most likely to transport pests that threaten our mainland forests, i.e. imports of woody plants belonging to genera from temperate climates. APHIS should promote more sentinel plant programs. Regarding wood packaging, APHIS could follow the lead of CBP by penalizing importers for each shipment containing noncompliant SWPM.

Getting APHIS to prioritize pest prevention over free trade in general, or in current trade agreements, is a heavy lift. At the very least, the agency should ensure that the U.S. prioritize invasive species prevention in negotiations with trading partners and in developing international trade-related agreements. I borrow here from the recent report on Canadian invasive species efforts. (I complained about APHIS’ failure to even raise invasive species issues during negotiation of a recent agricultural trade agreement with China.)

Solving Issues of Spreading Pests

The absence of an effective system to prevent a pest’s spread within the U.S. is the most glaring gap in the so-called federal “safeguarding system”. Yet this gap is rarely discussed by anyone – officials or stakeholders. APHIS quarantines are the best answer – although they are not always as efficacious as needed – witness the spread of EAB and persistence of nursery outbreaks of the SOD pathogen.

APHIS and the states continue to avoid establishing official programs targetting bioinvaders expected to be difficult to control or that don’t affect agricultural interests. Example include laurel wilt, and two boring beetles in southern California – goldspotted oak borer, Kuroshio shot hole borer and polyphagous shot hole borer and their associated fungi.

One step toward limiting pests’ spread would come from strengthening APHIS’ mandate in legislation, as suggested above. A second, complementary action would be for states to adopt quarantines and regulations more aggressively. For this to happen, APHIS would need to revise its policies on the “special needs exemption” [7 U.S.C. 7756]. Then states could adopt more stringent regulations to prevent introduction of APHIS-designated quarantine pests (Fading Forests III Chapt 3).

Finally, APHIS should not drop regulating difficult-to-control species – e.g., EAB. There are repercussions.

APHIS’ dropping EAB has not only reduced efforts to prevent the beetle’s spread to vulnerable parts of the West. It has also left states to come up with a coherent approach to regulating firewood; they are struggling to do so.

Considering interstate movement of pests via the nursery trade, the Systems Approach to Nursery Certification (SANC) program) is voluntary and was never intended to include all nurseries. Twenty-five nurseries were listed on the program’s website as of March 2020. It is not clear how many nurseries are participating now. The program ended its “pilot” phase and “went live” in January 2021. Furthermore, the program has been more than 20 years in development, so it cannot be considered a rapid response to a pressing problem.

Solving Issues of Recovery and Restoration via Resistance Breeding

I endorse the findings of two USFS scientists, Sniezko and Koch citations. They have documented the success of breeding programs when they are supported by expert staff and reliable funding, and have access to appropriate facilities. The principle example of such a facility is the Dorena Genetic Resource Center in Oregon. Regional consortia, e.g., Great Lakes Basin Forest Health Collaborative and Whitebark Pine Ecosystem Foundation are trying to overcome gaps in the system. I applaud the growing engagement of stakeholders, academic experts, and consortia. Questions remain, though, about how to ensure that these programs’ approaches and results are integrated into government programs.

resistant and vulnerable ash seedlings; photo courtesy of Jennifer Koch, USFS

In Bonello et al., I and others call for initiating resistance breeding programs early in an invasion. Often other management approaches, e.g., targetting the damaging pest or manipulating the environment, will not succeed. Therefore the most promising point of intervention is often with by breeding new or better resistance in the host. This proposal differs slightly from my suggestion in the “30 years – solutions” blog, when I suggested that USFS convene a workshop to develop consensus on breeding program’s priorities and structure early after a pest’s introduction.

Funding Shortfalls

I have complained regularly in my publications (Fading Forests reports) and blogs about inadequate funding for APHIS Plant Protection program and USFS Forest Health Protection and Research programs. Clearly the USDA Plant Pest and Disease Management and Disaster Program has supported much useful work. However, its short-term grants cannot substitute for stable, long-term funding. In recent years, APHIS has held back $14 – $15 million each year from this program to respond to plant health emergencies. (See APHIS program reports for FYs 20 and 21.) This decision might be the best solution we are likely to get to resolve APHIS’ need for emergency funds. If we think it is, we might drop §2 of H.R. 1389.

Expanding Engagement of Stakeholders

Americans expect a broad set of actors to protect our forests. However, these groups have not pressed decision-makers to fix the widely acknowledged problems: inadequate resources – especially for long-term solutions — and weak and tardy phytosanitary measures. Employees of federal and state agencies understand these issues but are restricted from outright advocacy. Where are the professional and scientific associations, representatives of the wood products industry, forest landowners, environmental NGOs and their funders, plus urban tree advocates – who could each play an important role? The Entomological Society’s new “Challenge” is a welcome development and one that others could copy.

SOURCES

Bonello, P., Campbell, F.T., Cipollini, D., Conrad, A.O., Farinas, C., Gandhi, K.J.K., Hain, F.P., Parry, D., Showalter, D.N, Villari, C. and Wallin, K.F. (2020) Invasive Tree Pests Devastate Ecosystems—A Proposed New Response Framework. Front. For. Glob. Change 3:2. doi: 10.3389/ffgc.2020.00002

Green, S., D.E.L. Cooke, M. Dunn, L. Barwell, B. Purse, D.S. Chapman, G. Valatin, A. Schlenzig, J. Barbrook, T. Pettitt, C. Price, A. Pérez-Sierra, D. Frederickson-Matika, L. Pritchard, P. Thorpe, P.J.A. Cock, E. Randall, B. Keillor and M. Marzano. 2021. PHYTO-THREATS: Addressing Threats to UK Forests and Woodlands from Phytophthora; Identifying Risks of Spread in Trade and Methods for Mitigation. Forests 2021, 12, 1617 https://doi.org/10.3390/f12121617ý

Krishnankutty, S., H. Nadel, A.M. Taylor, M.C. Wiemann, Y. Wu, S.W. Lingafelter, S.W. Myers, and A.M. Ray. 2020. Identification of Tree Genera Used in the Construction of Solid Wood-Packaging Materials That Arrived at U.S. Ports Infested With Live Wood-Boring Insects. Journal of Economic Entomology 2020, 1 – 12

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. Front. Ecol. Environ. 2012; 10(3):135-143

Lovett, G.M., M. Weiss, A.M. Liebhold, T.P. Holmes, B. Leung, K.F. Lambert, D.A. Orwig, F.T. Campbell, J. Rosenthal, D.G. McCullough, R. Wildova, M.P. Ayres, C.D. Canham, D.R. Foster, SL. Ladeau, and T. Weldy. 2016. NIS forest insects and pathogens in the US: Impacts and policy options. Ecological Applications, 26(5), 2016, pp. 1437–1455

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, & P.C. Tobin. 2019. Evolutionary history predicts high-impact invasions by herbivorous insects. Ecol Evol. 2019 Nov; 9(21): 12216–12230.

Poland, T.M., Patel-Weynand, T., Finch, D., Miniat, C. F., and Lopez, V. (Eds) (2019), Invasive Spp in Forests and Grasslands of the US: A Comprehensive Science Synthesis for the US Forest Sector. Springer Verlag. (in press).

Roy, B.A., H.M Alexander, J. Davidson, F.T Campbell, J.J Burdon, R. Sniezko, and C. Brasier. 2014. Increasing forest loss worldwide from invasive pests requires new trade regulations. Front Ecol Environ 2014; 12(8): 457–465

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.

United States Department of Agriculture Animal and Plant Health Inspection Service. Report on the Arrival in the US of Forest Pests Through Restrictions on the Importation of Certain Plants for Planting. https://www.caryinstitute.org/sites/default/files/public/downloads/usda_forest_pest_report_2021.pdf

Posted by Faith Campbell

“Global Tree Assessment”: #s at Risk, Threats, & Carbon Sequestration Planting

Kew Gardens U.K., home to Botanic Gardens Conservation International; Wikipedia

A massive international effort has completed a “Global Tree Assessment: State of Earth’s Trees”. This is the result of five years’ effort; it aims at a comprehensive assessment of the conservation status of all the Earth’s trees. As a result of their work, the authors issue a call to action and include specific recommendations.

The leads were the Botanic Gardens Conservation International (BGCI) and International Union for Conservation of Nature’s (IUCN) Species Survival Commission (SSC) Global Tree Specialist Group. They were assisted by about 60 cooperating institutions and more than 500 individual experts. The Morton Arboretum was a major U.S. contributor. Here, my focus is on the global assessment. An accompanying blog contains my analysis of reports on the Morton Arboretum report for the U.S.

The Global Tree Assessment is the largest initiative in the history of the IUCN Red List process. (This process is described in Box 3 of the report, on p. 12; and on p. 40.) As of the end of 2020, IUCN Red List assessments evaluated 28,463 tree species, representing half of all known tree species. Organizers hope to complete comprehensive conservation assessments of all tree species for inclusion on the IUCN Red List by 2023. Other sources utilized included draft Red List profiles and national-level assessments of those species that are found in only one country.

SUMMARY OF FINDINGS

Using these sources, the Global Tree Assessment evaluated 58,497 tree species worldwide. The study determined that 30% are threatened with extinction. This number could change significantly if a large proportion of the 7,700 species (13.2%) recorded as “Data Deficient” turn out to be at risk. At least 142 species are recorded as already extinct in the wild. Two-fifths (41.5%) are considered to be not at risk. Detailed species’ evaluations are provided at GlobalTreeSearch or GlobalTree Portal.

Brazilian forest converted to cattle pasture

The principal threats to trees globally are forest clearance and other forms of habitat loss (at least 65% of species) and direct exploitation for timber and other products (27% or more). The spread of non-native pests is said to affect 5% of the species. Climate change is having a measurable impact on 4% of the species and is expected to increase. (The situation in the United States differs significantly. Overexploitation plays almost no role and on-going habitat loss is important for only a few of the at-risk species.)

The authors decry the lack of attention, historically, to tree endangerment given trees’ ecological, cultural and economic importance. They hope that increased attention to the biodiversity crisis — an estimated 1 million animal and plant species threatened with extinction — and trees’ importance as carbon sinks will lead to increased conservation of trees and forests. They warn, however, that tree-planting programs must put the right species in the right place, including utilizing species that are under threat. In other words, tree planting practices need to change. They note that a community of botanists and conservationists is ready to assist.

Centers of tree species diversity – and of species under threat – are in Central and South America, followed by the other tropical regions of Southeast Asia and Africa. Fifty-eight percent of tree species are single country endemics. The highest proportion of endemism is found in New Zealand, Madagascar and New Caledonia. The region with the highest proportion of native tree species under threat is tropical Africa, especially Madagascar. The highest numbers of species “Not Evaluated” or “Data Deficient” are in IndoMalaya (tropical Asia) and Oceania. In those regions, about a third of species fall in one of those categories.

The assessment authors fear ecosystem collapse caused by major, large-scale disturbance events. Examples are recent unprecedented fires in California, southern Australia, Indonesia, and the Amazon (although they don’t mention Siberia). They also note mass mortality events over large areas of forest caused by other factors, including drought and heat stress and the increased incidence of pests. These events have led to a worrying decline of dominant tree species currently evaluated as “Least Concern.” Citing a 2010 report, they list as examples spruce in Alaska, lodgepole pine in British Columbia, aspen in Saskatchewan and Alberta, and Colorado pinon pine (Pinus edulis) in the American southwest.

The authors emphasize the importance of preventing extinction of monotypic tree families. Such events would represent a disproportionate loss of unique evolutionary history, biological diversity, and potential for future evolution. Of the 257 plant families that include trees, 12 are monotypic. They are scattered around the tropics and former Gondwanaland; none is found in the Neo- or Paleoarctic regions. While extinctions to date have rarely affected plants above the rank of genus, the global assessment authors worry that the on-going sixth extinction wave might result in extinctions at the genus or family level.

In this context, the assessment made a particular effort to evaluate the status of species representing the survival of Gondwanian Rainforest lineages. They found that 29% of these tree species are threatened with extinction. Two case studies focus on Australia. They mention habitat conversion but not two non-native pathogens widespread in Australia, Phytophthora cinnamomi and Austropuccinia psidii.

formerly common, now endangered, Australian tree *Rhodamnia rubescens*, infected by *Austropuccinia psidii*; photo courtesy of Flickr

The proportion of total tree diversity designated as threatened is highest on island nations, e.g., 69% of the trees on St. Helena, 59% of the trees on Madagascar, 57% of the trees on Mauritius. Hawai`i is not treated separately from the United States as a whole. According to Megan Barstow of BGCI (pers. comm.), the just updated IUCN Red List includes 214 threatened tree species in Hawai`i.

[For the U.S. overall, the IUCN reports 1,424 tree species, of which 342 (24%) are considered threatened. In the companion U.S. assessment, the Morton Arboretum and collaborators found that 11% of 841 continental U.S. tree species are threatened.]

MAIN THREATS TO TREES

Habitat loss

Over the past 300 years, global forest area has decreased by about 40%. Conversion of land for crops and pasture continues to threaten more tree species than any other known threat. Additional losses are caused by conversion for urban and industrial development and transport corridors, and by changes in fire regimes. In total, these factors cumulatively threaten 78% of all tree species, 84% if one includes conversion to wood plantations.

Caribbean mahogany (*Swietenia mahogani*); photo by Miguel Vieria

Forest Exploitation

Exploitation, especially for timber, is the second greatest threat globally, affecting 27% of tree species (more than 7,400 tree species). The report focuses on centuries of harvest of valuable tropical timbers and exploitation for fuelwood, with an emphasis on Madagascar, where nearly half of all tree species (117 out of 244 tree species) are threatened.

Pests and diseases

Tree species are impacted by a wide range of pests and diseases that are spread by natural and artificial causes. Invasive and other problematic species are recorded as threats for 1,356 tree species (5%) recorded on the IUCN Red List. This figure might be low because some of the information is outdated (see my discussion of American beech in the companion blog about the North American report, here.) Also, climate change is altering the survival opportunities for many pests and diseases in new environments. The example given is the ash genus (Fraxinus), under attack by not only the emerald ash borer in North America and now Russia and Eastern Europe but also the disease Ash Dieback across Europe. The report refers readers to the International Plant Sentinel Network for early warning system of new and emerging pest and pathogen risks, as well as help in coordinating responses.

black ash (*Fraxinus nigra*) swamp; Flickr

Climate Change

Climate change is impacting all forest ecosystems and is emerging as a significant recorded threat to individual tree species. In the IUCN Red List assessments, climate change and severe weather is recorded as a threat in 1,080 (4%) cases. Trees of coastal, boreal and montane ecosystems are disproportionately impacted. The authors note that the actual impact of climate change is probably more widespread, as it is also impacting fire regimes and the survival, spread, and virulence of pests.

CURRENT CONSERVATION EFFORTS

In Protected areas

Currently, 15.4% of the global terrestrial surface has formal protection status. The IUCN study authors recognize in situ conservation of trees through protection of existing natural habitats as the best method for conserving tree diversity. It is therefore encouraging that at least 64% of all tree species are included in at least one protected area. However, representation is higher for species that are not threatened – 85% are represented in a conservation area while only 56% of threatened trees species are. Nor does the report assess the effectiveness of protection afforded by the various in situ sites. The authors express hope that the parallel IUCN Red List of Ecosystems will contribute to understanding of the efficacy of conservation efforts targetting forests.

The Global Trees Campaign is a joint initiative of Fauna & Flora International (FFI) and BGCI. Since 1999 the campaign has worked to conserve more than 400 threatened tree species in more than 50 countries. The current focus is on six priority taxa = Acer, Dipterocarps, Magnolia, Nothofagus, Oak, and Rhododendron.

Rhododendron in Cook Forest State Park, PA; photo by F.T. Campbell

In Botanic gardens and seed banks

Especially for species under threat, conservation outside their native habitat – ex situ conservation – is an essential additional component. Currently 30% of tree species are recorded as present in at least one botanic garden or seed bank. Again, representation is higher for species that are not threatened – 45% are represented compared to only 21% of threatened tree species. For 41 species, ex situ conservation provides the only hope of survival, since they are extinct in the wild.

AN URGENT CALL FOR ACTION

The authors and collaborators who prepared the Global Tree Assessment hope that this report will help prompt action and better coordination of priorities and resources to better ensure that all tree species are supported by in situ conservation sites and by appropriate management plans. They state several times the importance of restoration plantings relying on native species. The purpose of plantings needs to include conservation of biological diversity, not just accumulation of carbon credits. The Ecological Restoration Alliance of Botanic Gardens (https://www.erabg.org/) is demonstrating that forest restoration can benefit biodiversity conservation. In many cases, propagation methods need to be developed. Also, projects must include aftercare and monitoring to ensure the survival of planted seedlings.

The IUCN assessment notes that ex situ conservation is an important backup. Education, capacity-building and awareness-raising are needed to equip, support, and empower local communities and other partners with the knowledge and skills to help conserve threatened trees.

Policy

The report say it does not address policy and legislation – a gap that fortunately is not quite true. The report both summarizes pertinent international agreements but also provides specific recommendations.

The international agreements that pertain to tree and forest conservation include:

Convention on Biological Diversity (CBD) and several specific programs: the Forestry Programme, Protected Area Programme and Sustainable Use Programme.
Global Strategy for Plant Conservation (GSPC), which is now developing post-2020 targets.
United Nations Framework Convention on Climate Change (UNFCCC) and countries’ implementing pledges to conserve carbon sinks, e.g., REDD+ (Reducing Emissions from Deforestation and Forest Degradation)
United Nations Strategic Plan for Forests 2017-2030
Global Plan of Action for the Conservation and Sustainable Use of Forest Genetic Resources
Convention on International Trade in Endangered Species, which currently protects 560 tree species, including 308 of the most threatened timbers

The report also mentions the voluntary New York Declaration on Forests, under which more than 200 entities – including governments, businesses, and Indigenous communities — have committed to eliminating deforestation from their supply chains. The supply chains touched on include those for major agricultural commodities, production of which is one of the greatest threat to trees.

SPECIFIC RECOMMENDATIONS

1. Strengthen tree conservation action globally through the formation of a new coalition that brings together existing resources and expertise, and applies lessons from the Global Trees Campaign to radically scale up tree conservation.

2. Use information in the GlobalTree Portal on the conservation status of individual tree species and current conservation action to plan additional action at local, national, and international levels, and for priority taxonomic groups. Build on the Portal by strengthening research on “Data Deficient” tree species, and collating additional information threatened species to avoid duplication of efforts and ensure conservation action is directed where it is needed most.

3. Ensure effective conservation of threatened trees within the protected area network by strengthening local knowledge, monitoring populations of threatened species and, where necessary, increasing enforcement of controls on illegal or non-sustainable harvesting of valuable species. Extend protected area coverage for threatened tree species and species assemblages that are currently not well-represented in protected areas.

4. Ensure that all globally threatened tree species are conserved in well-managed and genetically representative ex situ living and seed bank collections, with associated education and restoration programs.

5. Align work with the UN Decade on Ecosystem Restoration 2021–2030, engaging local communities, government forestry agencies, the business community, and other interested parties to ensure that the most appropriate tree species, including those that are threatened, are used in tree planting and restoration programs.

6. Improve data collection for national inventory and monitoring systems and use this information to reduce deforestation in areas of high tree diversity in association with REDD+ and Nationally Determined Contributions (NDCs).

7. Increase the availability of government, private and corporate funding for threatened tree species, and ensure that funding is directed to species and sites that are in greatest need of conservation.

SOURCE

Global Tree Assessment State of Earth’s Trees September 2021 Botanic Gardens Conservation International available here

Posted by Faith Campbell

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

USFS report: treatment of introduced forest pests

still-healthy hemlocks in Cook Forest State Park, PA; photo by F.T. Campbell

In a separate blog, I evaluated several aspects of the report as they apply to invasive species generally. Here I focus on invasive insects and pathogens that attack North American tree species (that is, forest pests).

As I said in the separate blog, I doubt that the book will stimulate policy-makers to increase Forest Service resources allocated to invasive species research, much less management. Sections 14.5 and 16.5 of the report state that the continued absence of a comprehensive investigation of the impacts of invasive species, especially the full value of ecosystem services lost, is a barrier to policymakers seeking to develop priorities and realistic management strategies.

I think the book’s editors tried to provide as much information about impacts as possible given existing knowledge. But the book’s length, comprehensive inclusion of all bioinvaders, organizational structure, and the detailed discussions of theories and models reduce the contribution the book might make to management decisions. I did not find “lessons learned” that could be applied in the policy realm.

Chapters address impacts in terrestrial and aquatic systems; impacts on ecosystem processes; impacts on various sectors of the economy and cultural resources; interactions with climate change and other disturbances; management strategies for species and landscapes; tools for inventory and management. Each chapter evaluates the current status of knowledge about the topic and suggests research needs. There are also summaries of the invasive species situation in eight regions.

The choice to organize the book around the chapters listed above means that some information one might expect to find in a book about invasive species is scattered or even absent. This is not a good resource for concise descriptions of individual invasive species and their impacts. That information is scattered among the chapters depending on whether some aspect of the species was chosen to illustrate a scientific challenge or success. The regional summaries partially remedy this problem – but they do not provide perspective on organisms that have invaded more than one region, e.g., emerald ash borer or white pine blister rust. To some extent, information about individual species is provided in the several subchapters on forest insects and pathogens. Or the reader of the PDF version can use the word search function!

Of course, information on several individual high-profile bioinvaders can be found in other publications; see the species write-ups and references posted at www.dontmovefirewood.org. Under these circumstances, a description of invasive species impacts from the ecosystem perspective is a welcome addition. I have long wished for a “crown to root zone” description of invasive species’ impacts.

HWA-killed hemlocks in Linville Gorge, NC; photo by Steven Norman, USFS

In this blog, I will focus on issues that the report raises that I found most interesting.

Information in the Report on Invasive Insects & Pathogens that Attack North American Trees

At several places the report states that non-native pests that have the potential to threaten the survival of an entire tree genus should be a high priority (p. 136) (what actions should be prioritized are not specified). They name the emerald ash borer (EAB) and Dutch elm disease. Elsewhere, EAB and hemlock woolly adelgid (HWA) are described as among the most significant threats to forests in the Eastern U.S. While EAB and HWA have certainly received considerable attention from the Forest Service, threats to elm have not. (I regret that the timing of the report precluded reference to Kevin Potter’s priority-setting publication. Potter is not listed as a co-author of the book.)

Hemlock woolly adelgid, emerald ash borer, chestnut blight, white pine blister rust (WPBR), and laurel wilt are cited as examples of highly virulent, host-specific agents that kill dominant, abundant, and ecologically unique hosts (p.18), resulting in exceptionally severe long-term impacts. WPBR and HWA specifically can have profound and far-reaching negative effects on ecosystem structure and function. These can rise to the level of an irreversible change of ecological state (p. 97). Of this list, no federal agency has allocated many resources to efforts to slow the spread of laurel wilt. The Forest Service is certainly tracking its spread and impacts.

Exaggerations or Errors

I think the report exaggerates the level of resources allocated to host resistance breeding. The report mentions programs targetting Dutch elm disease, beech bark disease, EAB, HWA and laurel wilt. It describes programs for white pines and Port-Orford cedar as examples of success. However, I would say that all the programs, except American chestnut, are starved for funds and other resources. The report’s authors concede this on p. 195.

TACF American chestnut in field trial; photo by F.T. Campbell

I think the report is too optimistic about the efficacy – so far – of biocontrol agents targeting HWA & EAB. On the other hand, I appreciate the report’s recognition that application of augmentative biocontrol of the Sirex woodwasp is more complicated in North America than in Southern Hemisphere countries (p. 162).

I am concerned about the statement that many plant pathogens are transported, but few have major impact. Examples in the U.S. are said to be WPBR, chestnut blight, and Phytophthora ramorum (p. 97). However, the report does not mention laurel wilt – which has a broad host range; nor rapid ‘ōhi‘a death — which threatens the most widespread tree species on the Hawaiian Islands. Nor does it mention several pathogens attacking single tree species, including beech bark disease, Port-Orford cedar root disease, and butternut canker. The report was written before recognition of beech leaf disease. The report notes that the three diseases it did mention have huge impacts. I am greatly disappointed that the report does not address how scientists and managers should deal with this “black swan” problem other than long discussions of data gaps, and ways to improve models of introduction and spread.

In addition, I am concerned that the discussion of economic factors that influence trade flows and accompanying invasive species (p. 308) focusses too narrowly on inspection alone, rather than other strategies for curtailing introduction. This section also shortens a description of the point made in Lovett et al. (2016). The report notes that Lovett et al. (2016) say that rates of introduction of wood-boring species decreased after ISPM#15 was implemented. However, the report leaves out the major caveat in that paper and the studies by Haack et al. (2014) and Leung et al. (2014) on which it is based: the reduction was insufficient to protect America from damaging introductions! [A further error has crept in: the Haack study explicitly excluded imports from China from their calculations. The Lovett paraphrase is not really clear on this matter.]

Curiosities/Concerns Re: Regional Write-Ups

I wish the sections on the Northwest and Southwest region discussed why areas with so many characteristics favoring introduction of plant pests – major ports, extensive transportation networks, major horticultural industry, extensive agriculture, and abundant urban and native forests – have so few damaging forest pests. (Admittedly, those present are highly damaging: white pine blister rust, sudden oak death, Port-Orford cedar root disease, pitch canker, balsam woolly adelgid, larch casebearer, polyphagous shot hole borer (I add Kuroshio shot hole borer), and banded elm bark beetle). The report does mention the constant threat of introduction of the European and Asian gypsy moths. (The Entomological Society of America has decided to coin a new common name for these insects; they currently to be called by the Latin binomial Lymatria dispar). The report notes that 22 species of non-native bark and ambrosia beetles have recently been introduced in the Southwest.

The report cites a decade-old estimate by Aukema et al. (2010) in saying that a small proportion of introduced pest species has killed millions of trees or pushed ecologically foundational species toward functional extinction. The figure was 14% of the more than 450 non-native forest insect species. I greatly regret that overlapping preparation and publication periods precluded inclusion of data from studies by Potter, Guo, and Fei. http://nivemnic.us/what-fia-data-tell-us-about-non-native-pests-of-americas-forests/

Section 7.3 of the report discusses frameworks for setting priorities. It identifies five factors: 1) pest species having the greatest negative impacts; 2) uniqueness of the affected ecosystem or community; 3) state of the invasion in space and time; 4) management goals; 5) availability of effective tools. Examples of species meeting these criteria include EAB and Dutch elm disease (pest threatens entire host genus); white pine blister rust on whitebark pine (key species in a system with low arboreal diversity).

The report notes increasing understanding of critical aspects of several important pests’ biology and host interactions – but it does not sufficiently acknowledge the decades of effort required to achieve this knowledge. The time required for additional scientific advances will probably be equal or greater, given falling number of “ologists” in government and academia.

I appreciate inclusion of a discussion (Sections 8.3.1 and 8.3.2) on breeding trees resistant to introduced pests

dead Port-Orford cedar in Redwoods National Park; photo by Richard Sniezko, USFS

This section states that host resistance, forest genetics, and tree improvement might be the most effective approaches to managing many established pests. The section says such breeding does not require the use of genetically modified organisms, although transgenic or gene editing technologies can provide useful tools. I appreciate the report conceding that necessary infrastructure and expertise has been declining for two decades (p. 195).

In discussing international cooperation to reduce transport of invasive species, the report refers to increasing availability of data allowing identification of potentially damaging species in their regions of origin. Again, since this chapter was written, the Forest Service has increased its engagement on this approach: the USFS International Program is supporting sentinel plantings managed by the International Plant Sentinel Network (http://www.plantsentinel.org) … see my recent blog here.

SOURCE

Posted by Faith Campbell

USFS invasive species report: Herculean effort that could have had greater impact

In February the USFS published a lengthy analysis of invasive species: Invasive Species in Forests and Rangelands of the United States. A Comprehensive Science Synthesis for the US Forest Sector (Poland et al. 2021; full citation at the end of the blog). More than 100 people contributed to the book; I helped write the chapters on legislation and regulations and international cooperation. The book is available for download at no cost here.

Miconia – one of many invasive plants damaging ecosystems in Hawai`i

I greatly appreciate the effort. Authors first met in 2015, and most chapters were essentially written in 2016. The long delay in its appearance came largely from negotiations with the publisher. The delay means some of the information is out of date. I am particularly aware that several experts – e.g., Potter, Guo, and Fei – have published about forest pests since the Aukema source cited. I wonder whether inclusion of their findings might change some of the conclusions about the proportion of introduced pests that cause noticeable impacts.

Since the report’s publication in February I have struggled with how to describe and evaluate this book. What is its purpose? Who is its audience? The Executive Summary says the report is a sector-wide scientific assessment of the current state of invasive species science and research in the U.S.

However, the Introduction states a somewhat different purpose. It says the report documents invasive species impacts that affect ecosystem processes and a wide range of economic sectors. This would imply an intention to enhance efforts to counter such effects– not just to shape research but also to change management. Indeed, the Conclusion of the Executive Summary (pp. xvi-xvii) is titled “An Imperative for Action”.

I am not the author to evaluate how effectively the book sets out research agendas. Regarding its usefulness in prompting policy-makers to do more, I regretfully conclude that it falls short.

Getting the balance right between an issue’s status and what needs to be done is difficult, perhaps impossible. I appreciate that the report makes clear how complex bioinvasion and ecosystem management and restoration are. Its length and density highlight the difficulty of making progress. This daunting complexity might well discourage agency leadership from prioritizing invasive species management.

On the other hand, summary sections sometimes oversimplify or bury important subtleties and caveats. The question of whether some key questions can ever be resolved by science is hinted at – but in detailed sections that few will read. The same is true regarding the restrictions imposed by funding shortfalls.

The Report Would Have Benefitted from Another Round of Editing

Editing this tome was a Herculean task. I feel like a curmudgeon suggesting that the editors do more! Nevertheless, I think the report would have been improved by the effort. One more round of editing – perhaps involving a wider range of authors – could have pulled together the most vital points to make them more accessible to policymakers. It could also have tightened the ecosystem-based descriptions of impacts, which are currently overwhelmed by too much information.

A precis for policymakers

A precis focused on information pertinent to policymakers (which the current Executive Summary does not) should contain the statement that the continued absence of a comprehensive investigation of invasive species’ impacts hampers research, management, and policy (mentioned only in §16.5, on p. 332). It should note situations in which insufficient funding is blocking recommended action. I note three examples: programs aimed at breeding trees resistant to non-native pests (resource issues discussed only in §§8.3.1 and 8.3.2, p. 195); sustaining “rapid response” programs (§6.4.3, p. 125); costs of ecosystem restoration, especially for landscape-level restoration (§16.4). I am sure there are additional under-funded activities that should be included!

cross-bred ash seedlings being tested for vulnerability to EAB; photo courtesy of Jennifer Koch

Other important information that should be highlighted in such a precis includes the statement that many ecosystems have already reached a point where healthy functions are in a more tenuous balance due to invasive species (p. 51). Effective carbon storage and maintaining sustainable nutrient and water balance are at risk. Second, costs and losses caused by invasive forest pests generally fall disproportionately on a few economic sectors and households. They cannot be equated to governmental expenditures alone (p. 305). Third, even a brief estimate of overall numbers of invasive species appears only in §7.4. Information about individual species is scattered because it is used as example of particular topic (e.g., impacts on forest or grassland ecosystems, or on ecosystem services, or on cultural values).

Ecosystem Impacts Overwhelmed

As noted above, the report laments the absence of a comprehensive investigation of invasive species’ impacts. Perhaps the editors intended this report to partially fill this gap. To be fair, I have long wished for a “crown to root zone” description of invasive species’ impacts at a site or in a biome. Concise descriptions of individual invasive species and their impacts are not provided by this report, but they can be found elsewhere. (The regional summaries partially address the problem of too much information – but they do not provide perspective on organisms that have invaded more than one region, e.g., emerald ash borer or white pine blister rust.) Another round of editing might have resulted in a more focused presentation that would be more easily applied by policymakers.

Welcome Straightforward Discussion of Conceptual Difficulties

I applaud the report’s openness about some important overarching concepts that science cannot yet formulate. If supportable theories could be conceived, they would assist in the development of policies:

Despite decades of effort, scientists have not established a clear paradigm to explain an ecosystem’s susceptibility to invasion (p. 85). Invasibility is complex: it results from a dynamic interplay between ecosystem condition and ecological properties of the potential invader, especially local propagule pressure.
Scientists cannot predict how climate warming will change distributions of invasive species [see Chapter 4] and alter pathways. This inability hampers efforts to develop effective prevention, control, and restoration strategies (p. xi). Climate change and invasive species need to be studied together as interactive drivers of global environmental change with evolutionary consequences.

The Report’s Recommendations

Policy-oriented recommendations are scattered throughout the report. I note here some I find particularly important:

Measures of progress should be based on the degree to which people, cultures, and natural resources are protected from the harmful effects of invasive species.
Managers should assess the efficacy of all prevention, control, and management activities and their effect upon the environment. Such an evaluation should be based on a clear statement of the goals of the policy or action. [I wish the report explicitly recognized that both setting goals and measuring efficacy are difficult when contemplating action against a new invader that is new to science or when the impacts are poorly understood. Early detection / rapid response efforts are already undermined by an insistence on gathering information on possible impacts before acting; that delay can doom prospects for success.]
- Risk assessment should both better incorporate uncertainty and evaluate the interactions among multiple taxa. Risk assessment tools should be used to evaluate and prioritize management efforts and strategies beyond prevention and early detection/rapid response.
- Economic analyses aimed at exploring tradeoffs need better tools for measuring returns on invasive species management investments (§16.5).
Actions that might be understood as “restoration” aim at a range of goals along the gradient between being restored to a known historic state and being rehabilitated to a defined desired state. The report stresses building ecosystem resilience to create resistance to future invasions, but I am skeptical that this will work re: forest insects and disease pathogens.
Propagule pressure is a key determinant of invasion success. Devising methods to reduce propagule pressure is the most promising to approach to prevent future invasions (p. 115). This includes investing in quarantine capacity building in other countries can contribute significantly to preventing new invasions to the US.
Resource managers need additional studies of how invasive species spread through domestic trade, and how policies may differ between foreign and domestic sources of risk.

I appreciate the report’s attention to such often-ignored aspects as non-native earthworms and soil chemistry. I also praise the report’s emphasis on social aspects of bioinvasion and the essential role of engaging the public. However, I think the authors could have made greater use of surveys conducted by the Wisconsin Department of Natural Resources and The Nature Conservancy’s Don’t Move Firewood program.

Lost Opportunities

I am glad that the report makes reference to the “rule of 25” rather than “rule of 10s”. I would have appreciated a discussion of this topic, which is a current issue in bioinvasion theory. As noted at the beginning of this blog, the long time between when the report was written and when it was published might have hampered such a discussion

Also, I wish the report had explored how scientists and managers should deal with the “black swan” problem of infrequent introductions that have extremely high impacts. The report addresses this issue only through long discussions of data gaps, and ways to improve models of introduction and spread.

I wish the section on the Northwest Region included a discussion of why an area with so many characteristics favoring bioinvasion has so few damaging forest pests. Admittedly, those present are highly damaging: white pine blister rust, sudden oak death, Port-Orford cedar root disease, balsam woolly adelgid, and larch casebearer. The report also notes the constant threat that Asian and European gypsy moths will be introduced. (The Entomological Society of America has decided to coin a new common name for these insects; they currently to be called by the Latin binomial Lymatria dispar).

And I wish the section on the Southeast and Caribbean discussed introduced forest pests on the Caribbean islands. I suspect this reflects a dearth of research effort rather than the biological situation. I indulge my disagreement with the conclusion that introduced tree species have “enriched” the islands’ flora.

SOURCE

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

Posted by Faith Campbell

Federal Funding for Forest Pest Programs — Act Now! to Help Congress Decide

If you have not communicated to your Representative and senators your support for adequate funding of U.S. government programs to address non-native insects and pathogens threatening our forests, please do so now!

If political leaders do not hear from us that expanding these programs is important, these programs will continue to languish. It is easiest – and most direct – to inform your representative and Senators of your support. Please do so! If you do not agree that these programs should be expanded & strengthened, I ask that you send a comment outlining what approach you think would be more effective in curtailing introductions, minimizing impacts, and restoring affected tree species. I can then initiate a discussion to explore these suggestions. [I already have endorsed the suggestion to create a CDC-like body to oversee management of non-native forest pests.] You can find your member of Congress here. Your Senators here .

Last week the Biden Administration sent to Congress its proposed budget for the fiscal year beginning October 1, 2021. I find it falls short in key areas. Next, the House and Senate will pass a package of appropriations bills to set actual funding levels. This is the moment to press for boosted funding. In an earlier blog I explained my reasons for seeking specific funding levels.

Asian longhorned beetles – introduced in wood packaging (USDA photo)

Two USDA agencies lead efforts to protect U.S. wildland, rural, and urban forests from non-native insects and pathogens. Their funding is set by two separate – and critical — appropriations bills:

USDA’s Animal and Plant Health Inspection Service (APHIS) has legal responsibility for preventing introduction of tree-killing pests; detecting newly introduced pests; and initiating eradication and containment programs intended to minimize their damage. Funding for APHIS is contained in the Agriculture Appropriations bill.
USDA Forest Service (USFS)
- The Forest Health Management (FHM) program provides funding and applied science to help partners manage pests. The program has two sides: the Cooperative component helps states and private forest managers, so it can address pests where they are first found – usually near cities – and when they spread. The federal lands component helps the USFS, National Park Service, and other federal agencies counter pests that have spread to the more rural/wildland areas that they manage.
- The Research and Development (R&D) program supports research into pest-host relationships; pathways of introduction and spread;; management strategies (including biocontrol); and host resistance breeding

Forest Service funds are appropriated through the Interior Appropriations bill.

APHIS – the Administration’s official budget proposal, and justification, is here.

The Administration proposes a small increase for three of four APHIS programs that are particularly important for preventing introductions of forest pests or eradicating or containing those that do enter. The Administration proposed significant funding for a fourth program that plays a small but important role in managing two specific forest pests.

APHIS Program	Current (FY 2021)	FY22 Administration proposed	FY 2022 Campbell recommended
Tree & Wood Pest	$60.456 million	$61 million	$70 million
Specialty Crops	$196.553 million	209 million	$200 million
Pest Detection	$27.733 million	No change	$30 million
Methods Development	$20.844 million	No change	$25 million

Tree and Wood Pests: It will be a major challenge for APHIS to eradicate the current outbreak of Asian longhorned beetles (ALB) in the swamps of South Carolina. APHIS should also address other pests. Even after cutting spending on the emerald ash borer (EAB), I think APHIS needs significantly more money in this account.

The Specialty Crops program is supported by such traditional USDA constituencies as the nursery and orchard industries, which probably explains the proposed increase. APHIS’ program to curtail spread of the sudden oak death (SOD) pathogen through interstate nursery trade receives funding from this program – about $5 million. I believe this program also now funds the agency’s efforts to slow spread of the spotted lanternfly.

SOD-infected rhododendrons in Indiana nursery in 2019

I would like the Pest Detection program to receive a small increase so the agency and its cooperators can better deal with rising trade volumes and associated pest risk. Similarly, Methods Development should receive a boost because of the need for improved detection and management tools.

USDA Forest Service – the Administration’s official budget proposal is here.

While the Forest Health Management (FHM) and Research and Development (R&D) programs are the principal USFS programs that address introduced forest pests, neither has non-native pests as the principle focus. Non-native forest pests constitute only a portion of the programs’ activities. In the case of Research, this is a very small portion indeed.

President Biden’s budget proposes to spend $59.2 million on the Forest Health Management program and $313.5 million for Research. Both represent significant increases over spending during the current fiscal year. However, the FHM level is still below spending in recent years, although both the number of introduced pests and the geographic areas affected have been rising for decades.

In my earlier blog I suggested the funding levels:

USFS PROGRAM	Current (FY21)	FY22 Administration	FY22 my recommendation
FHP Coop Lands	$30.747 million	$36.747 million	$51 million (to cover both program work & personnel costs)
FHP Federal lands	$15.485 million	22.485 million	$25 million (ditto)

Research & Develop	$258.7 million; of which about $3.6 million allocated to invasive species	$313.560 million	$320 million; I seek report language instructing the USFS to spend more on invasive species

Under the FHM program, a table on pp. 46-47 of the budget justification lists existing and proposed spending on 14 pest taxa (plus invasive plants and subterranean termites). Spending on these 14 species is proposed to total $30.3 million. Of this amount, less than half – $14.9 million – is allocated to such high-profile invasive species of forests as the emerald ash borer (EAB), hemlock woolly adelgid (HWA), sudden oak death (SOD), and threats to whitebark pine (recently listed as a threatened species under the Endangered Species Act). (The USFS does not engage in efforts to eradicate Asian longhorned beetle (ALB) outbreaks; it leaves that task to APHIS.) And of the nearly $15 million allocated to invasive non-native pests, more than half – $8 million – is allocated to European gypsy moths. While I agree that the gypsy moth program has been highly successful, I decry this imbalance. Other non-native pests cause much higher levels of mortality among hosts than does the gypsy moth.

dead whitebark pine at Crater Lake National Park; photo by FT Campbell

I applaud the modest increases in the Administration’s budget for other non-native forest pests. These range from tens to a few hundred thousand dollars per pest. FHM also supports smaller programs targetting rapid ohia death, beech leaf disease, the invasive shot hole borers in southern California, Mediterranean oak beetle, etc. Budget documents don’t report on these efforts.

The imbalance of funding allocated to damaging non-native pests compared to other forest management concerns is even worse in the Research program. Of the $313.5 million proposed in the budget for the full research program, only $9.2 million is allocated to the 14 pest taxa (plus invasive plants and subterranean termites) specified in the table on pp. 46-47. Of this amount, less than half — $4.5 million – is allocated to the high-profile invasive species, e.g., ALB, EAB, HWA, SOD, and threats to whitebark pine. The budget does provide extremely modest increases for several of these species, ranging from $12,000 for ALB to $114,000 for EAB. Again, some smaller programs managed at the USFS regional level might address other pests. Still – the budget proposes that USFS R&D allocate only 1.4% of its total budget to addressing these threats to America’s forests! This despite plenty of documentation – including by USFS scientists – that non-native species “have caused, and will continue to cause, enormous ecological and economic damage.” (Poland et al. 2021; full citation at the end of the blog). Poland et al. go on to say:

Invasive insects and plant pathogens (or complexes involving both) cause tree mortality, resulting in canopy gaps, stand thinning, or overstory removals that, in turn, alter microenvironments and hydrologic or biogeochemical cycling regimes. These changes can shift the overall species composition and structure of the plant community, with associated effects on terrestrial and aquatic fauna. In the short term, invasive insects and diseases can generally reduce productivity of desired species in forests. Tree mortality or defoliation can affect leaf-level transpiration rates, affecting watershed hydrology. Tree mortality … also leads to enormously high costs for tree removal, other management responses, and reduced property values in urban and residential landscapes.

eastern hemlock in Shenandoah National Park; photo by FT Campbell

I seek report language specifying that at least 5% of research funding should be devoted to research in pathways of invasive species’ introduction and spread; their impacts; and management and restoration strategies, including breeding of resistant trees. Several coalitions of which the Center for Invasive Species is a member have agreed to less specific language, not the 5% goal.

Two other USFS programs contribute to invasive species management. The Urban and Community Forest program provided $2.5 million for a competitive grant program to help communities address threats to urban forest health and resilience. Of 23 projects funded in FY2020, 11 are helping communities recover from the loss of ash trees to EAB. (On average, each program received $109,000.)

The Forest Service’ International Program is helping academic and other partners establish “sentinel gardens” in China and Europe. North American trees are planted and monitored so researchers can identify insects or pathogens that attack them. This provides advance notice of organisms that could be damaging pests if introduced to the United States.

REFERENCE:

Invasive Species in Forests and Rangelands of the United States. Editors T.M. Poland, T. Patel-Weynand, D.M. Finch, C.F. Miniat, D.C. Hayes, V.M. Lopez Open access!

Posted by Faith Campbell

Congressional Action to Protect Trees from Non-Native Pests

Rep. Peter Welch of Vermont has reintroduced his bill to improve programs intended to prevent introduction of non-native forest pests and enhance efforts to reduce their impacts. The latter provisions include support for breeding trees resistant (or tolerant) to the pest. I hope H.R. 1389 will be adopted – then spur new efforts to conserve and restore forest trees! Please follow my suggestion below.

The Invasive Species Prevention and Forest Restoration Act H.R. 1389 is co-sponsored by Reps. Brian Fitzpatrick (PA), Annie Kuster & Chris Pappas (NH), and Elise Stefanik (NY).

For updates, visit https://www.congress.gov/search?q={%22congress%22:[%22117%22],%22source%22:%22all%22,%22search%22:%22HR%201389%22}&searchResultViewType=expande

When he introduced the bill, Rep. Welch said

“Invasive species are devastating to forests which are a central part of Vermont’s economy and our way of life. This bill will fund efforts to revitalize damaged forests and highlight the need for making this a priority within the federal government.”

Major provisions of H.R. 1389:

Expands USDA APHIS’ access to emergency funding to combat invasive species when existing federal funds are insufficient and broadens the range of actives that these funds can support.
Establishes a grant program to support institutions focused on researching methods to restore native tree species that have been severely damaged by invasive pests.
Authorizes funding to implement promising research findings on how to protect native tree species.
Mandates a study to identify actions needed to overcome the lack of centralization and prioritization of non-native insect and pathogen research and response within the federal government, and develop national strategies for saving tree species.

As I have described in earlier blogs, the measures adopted by federal and state governments to prevent non-native pathogen and insect pest introductions – and the funding to support this work – have been insufficient to meet the growing challenges. In just the past decade, several new tree-killing pests have been detected: polyphagous and Kuroshio shot hole borers, spotted lanternfly, two rapid ʻōhiʻa death pathogens, Mediterranean oak beetle, velvet longhorned beetle. Over the same period, the Asian longhorned beetle has been detected in Ohio and South Carolina; the emerald ash borer expanded its range from 14 to 35 states; the redbay ambrosia beetle and its associated fungus spread from five states to 11; a second strain of the sudden oak death fungus appeared in Oregon forests; and whitebark pine has been proposed by the US Fish and Wildlife Service for listing as Threatened under the Endangered Species Act.

During this same period, funding for the USDA Forest Service Forest Health Protection program has been cut by about 50%; funding for USFS Research projects targetting 10 high-profile non-native pests has been cut by about 70%.

One reason for this disconnect between need and resources is that the non-native tree pest problem is largely out of sight and therefore does not lend itself to the long-term public attention needed to remediate the threats. It is up to us to raise the political profile of these issues.

On the positive side, the passage of time has brought forth new solutions, a deeper understanding of the genetics of plants and animals, new measures for igniting public awareness and invasive identification, new technologies and strategies for helping trees adapt, and a recognition of what resources and organization it will take to mount a proper solution to the problem.

“Project CAPTURE” (Conservation Assessment and Prioritization of Forest Trees Under Risk of Extirpation) has proposed priority species for enhanced conservation efforts. Top priorities in the continental states are listed below. A separate study is under way for forests in Hawai`i, Puerto Rico, and U.S. Virgin Islands.

Florida torreya (Torreya taxifolia)
American chestnut (Castanea dentata)
Allegheny chinquapin (C. pumila)
Ozark chinquapin (C. pumila var. ozarkensis)
redbay (Persea borbonia)
Carolina ash (Fraxinus caroliniana)
pumpkin ash (F. profunda)
Carolina hemlock (Tsuga caroliniana)
Port-Orford cedar (Chamaecyparis lawsoniana)
tanoak (Notholithocarpus densiflorus)
butternut (Juglans cinerea)
eastern hemlock (Tsuga canadensis)
white ash (Fraxinus americana)
black ash (F. nigra)
green ash (F. pennsylvanica).

For a brief explanation of Project CAPTURE, see my earlier blog here. For an in-depth description of the Project CAPTURE process and criteria for setting priorities, read Potter, K.M., M.E. Escanferla, R.M. Jetton, and G. Man. 2019. Important Insect and Disease Threats to United States Tree Species and Geographic Patterns of Their Potential Impacts. Forests 2019, 10. https://www.fs.usda.gov/treesearch/pubs/58290

Please ask your representative to co-sponsor H.R. 1389. Please ask your senators to sponsor a companion bill. For more information, contact Alex Piper at Alex.Piper@mail.house.gov or 202-306-6569 .

H.R. 1389 is endorsed by Vermont Woodlands Association, American Forest Foundation, Center for Invasive Species Prevention, the Reduce Risk from Invasive Species Coalition,, Entomological Society of America, and North American Invasive Species Management Association.

Posted by Faith Campbell