May 2020 – Center for Invasive Species Prevention

Pest Threats to Eastern Forests – Focus on the Mid-Atlantic

EAB-kiled ash tree in Shenandoah National Park in 2016
photo by F.T. Campbell

As we have known for years, forests of the eastern United States are under severe pressure from non-native forest insects and diseases. Several recent studies have put this fact into perspective.

Fei et al. (2019) found that the 15 most damaging introduced species threaten 41.1% of the total live forest biomass in the 48 conterminous states. Nine of the 15 species included in this calculation are pests of the eastern forest. Indeed, the greatest increase in biomass loss, as measured by USDA Forest Service Forest Inventory and Analysis (FIA) plot data occurred here. Compensatory growth in unaffected trees and the recruitment of new regeneration occurs only later – as much as two or more decades after the pest invasions began. Fei et al. (2019) expect these losses will be exacerbated in the future due in part to the likelihood that additional pests will be introduced.

Randall Morin found that non-native pests had caused approximately 5% increase in total mortality, by tree volume, nation-wide.

Most widespread pest threats in the East

Scientists have used several methods of measuring introduced pests’ impacts. One measure is the number of counties where the pest is present. A second measure is the proportion of the volume of the host that has been affected. Both metrics are used by Morin. A third method, used by the CAPTURE Project (Potter et al. 2019a), is the number of hosts affected by the pest.

Morin and colleagues found that the European gypsy moth has invaded 630 counties – or 29% of the volume of its principal host, oaks. (In both cases, the gypsy moth trailed white pine blister rust in extent of infestation. The latter is nationwide but having its greatest impacts in the West). The CAPTURE Project found that the gypsy moth affected the largest number of hosts – 65.

Using the “counties invaded” metric, Morin and colleagues found that dogwood anthracnose had invaded 609 counties in the East (and additional areas in the West); the emerald ash borer had invaded 479 counties at the time of analysis; the hemlock woolly adelged had invaded 432 counties. Using the number of hosts impacted measure, oak wilt (Bretziella fagacearum) affected the second largest number of hosts – 61 (Potter et al. 2019a). [All these pests are described briefly here.]

Project CAPTURE (Potter and colleagues 2019a) evaluated 339 serious pests threatening one or more of 419 native tree species in the continental US. They included both native and introduced pests. They analyzed 1,378 pest-host combinations. They found that:

54% of the host tree species (228) are infested by an exotic pest – although only 28% of the 1,378 host/agent combinations involved pests are known to be non-native in origin.
Exotic agents have, on average, considerably more severe impacts than native pests.
Non-native pests had greater average severity on angiosperms than on conifers. (As an earlier blog documented, Mech and colleagues have reached a similar – although tentative – conclusion.)
Their estimate of the threat posed by non-native pests to forests – especially for the East – is an underestimate because established pests could spread to additional vulnerable areas and there is a high likelihood that new pests will be introduced. The Southeast was consistently a “coldspot” – despite the near extirpation of one understory tree – redbay.

Potter et al. (2019a) ranked forest threats in two ways. Four host families were at highest risk to alien pests, as measured by both the numbers of tree species affected and by the most host/agent combinations: Fagaceae (oaks, tanoaks, chestnuts, beech); Pinaceae (pines); Sapindaceae (soapberry family; includes maples and buckeye); Salicaceae (willows, poplars, aspens). When host families were ranked by the severity of the host/pest threat, Fagaceae was still at greatest risk, and Sapindaceae was still in the top four; however, Ulmaceae (elms) and Oleaceae (includes Fraxinus) replaced pines and willows.

A very interesting study was published by scientists based in the Blue Ridge Mountains of Virginia (Anderson-Teixeira et al. 2020). They contend that their area is a good example of what is happening more broadly in the Mid-Atlantic region.

Anderson-Teixeira et al. (2020) found that non-native pests have substantially impacted at least 24% of the 33 tree genera (eight genera) recorded as present in their study plots. They estimated that over the century beginning with the appearance of chestnut blight in the region and ending with the expected extirpation of ash trees, net live aboveground biomass (AGB) loss among affected species totaled roughly 6.6–10 kg m -2. Forty to sixty percent of this loss started before the Park initiated quantitative surveys of permanent plots in 1987. The authors estimated that chestnut contributed up to 50% of estimated AGB losses over the century. Consequently, the estimate has very high uncertainty.

Despite these losses, Anderson-Teixeira et al. (2020) found that both total aboveground biomass and diversity within individual study plots had largely recovered through increases in non-vulnerable genera.

Average above ground biomass across the plots established in Shenandoah National Park increased as the forest recovers from logging, farming, and other disturbances before formation of the Park. These increases were due primarily to reproduction and growth of tulip poplar (Liriodendron tulipifera) and growth (but not reproduction) of oaks. Net AGB biomass was lost in oak- and hemlock-dominated plots. At plots established in the neighboring Smithsonian Conservation Biology Institute, pests had caused relatively minor impacts on AGB.

Diversity of tree species also did not change much. In the Park, the average number of genera per plot declined only 3% between 1991 and 2013. Diversity at the landscape scale increased by two genera – from 26 to 28. Many individual plots, though, lost three genera due to non-native pests – chestnut, redbud, and hemlock. A fourth genus was lost due to stochastic change. At the same time, the plots gained six native genera). This finding might be skewed by the short duration of the study period, which missed initial declines in several taxa and captured only the initial stages of decline in ash.

Several taxa were lost from the monitoring plots but were not completely extirpated from the region. Even those species not “lost” suffered elevated mortality rates and steep declines in abundance and above-ground biomass. These declines have not been reversed. The exception was some oaks, which regained above ground biomass, but not abundance, following the gypsy moth outbreak in the 1980s and early 1990s.

Taxa-specific findings

(Most of these pests are described briefly here.)

Fei et al. (2019) found that losses in biomass due to non-native pests – as measured by FIA plot data – was greatest for ashes, elms, beech trees, and hemlocks..

Morin and colleagues found annual mortality rates had increased three-fold above background levels for ash, beech, and hemlock. They also calculated the present mortality rates for several species for which the majority of loss occurred before their study (consequently, they could not calculate a pre-invasion “background” rate to which present rates could be compared). These included American chestnut (mortality rate of 7%), butternut (mortality rate of 5.6%), and elm trees (mortality rate of 3.5%).

The CAPTURE Project (Potter et al. 2019a) identified fifteen host-agent combinations with the highest severity. Ten of these species are found in the Mid-Atlantic region:

American chestnut (Castanea dentata)
Allegheny chinquapin (C. pumila)
Carolina ash (Fraxinus caroliniana) ,
pumpkin ash (F. profunda)
Carolina hemlock (Tsuga caroliniana)
butternut (Juglans cinerea)
eastern hemlock (Tsuga canadensis)
white ash (Fraxinus americana)
black ash (F. nigra)
green ash (F. pennsylvanica)

Four of these species are in genera included among the eight genera evaluated in the study conducted in the Blue Ridge (Anderson-Teixeira et al. 2020): American chestnut, butternut, eastern hemlock, green and white ash. The four other genera in the Blue Ridge study were elm (Ulmus), oak (Quercus), redbud Cercis, and dogwood (Cornus). All except redbud are recognized by other sources as heavily affected by non-native pests – confirming Anderson-Teixeira et al. (2020)’s conclusion that findings on the Blue Ridge reflect the wider situation.

Anderson-Teixeira et al. (2020) note that several of these tree species have been declared imperiled by the International Conservation Union (IUCN): American chestnut, butternut, American elm, eastern hemlock, and ash species.

Anderson-Teixeira et al. (2020) report data on three taxa previously important in the canopy of Blue Ridge forests – chestnut, elms, and butternut. Chestnuts larger than 10 cm DBH had disappeared from the future site of Shenandoah National Park by 1910. Short-lived sprouts continue to be present in plots in the low-elevation Smithsonian Conservation Biology Institute. Two elm species were described as ‘‘sparse’’ in the 1939 qualitative survey. Elms have persisted at low densities, low biomass, and increasingly small sizes. Butternut was ‘‘common’’ in 1939, but had disappeared from Shenandoah NP by 1987. On the Smithsonian’s property, butternut declined from four living individuals in 2008 to two in 2018. The near disappearance of butternut reflects the national picture: FIA data show the species has decreased about 58% across its U.S. range since the 1980s – which is decades after butternut canker started having a detectable impact in the Midwest.

In the Park, oak-dominated plots lost on average 24.9% of individuals and 15% of aboveground biomass. After 1995, when the gypsy moth was better controlled by spraying of Bacillus thuringiensis var. curstaki, oak aboveground biomass increased gradually, driven by individual tree growth rather than new recruitment. Continued declines in oak abundance are attributable to oak decline and management actions (or inactions) that do not promote regeneration.

In a separate study, a group of oak experts went through a process of queries to identify the greatest threat to oaks now and in the future (Conrad et al. 2020). They initially identified the following threats as most important currently (descending order): gypsy moth, oak wilt, oak decline, climate change, and drought. The top five future threats were initially identified as climate change, oak wilt, sudden oak death, oak decline, and some unknown new or emerging (non-native) pest or pathogen. By the third round, after the experts thought about their colleagues’ responses, oak decline had replaced gypsy moth as the most critical threat currently. Attack by an unknown new or emerging (non-native) pest or pathogen replaced climate change as the most critical future threat. While there was not a complete consensus, the consensus was stronger on the threat from a new pest.

remnant eastern hemlock at Linderlost, Shenandoah National Park
photo by F.T. Campbell

Anderson-Teixeira et al. (2020) reported that eastern hemlock was initially present in ten of Shenandoah plots, but was no longer recorded in the survey plots after 2007. (More than 20,000 insecticide-treated trees remain alive throughout Shenandoah NP).

Before arrival of the emerald ash borer, ash aboveground biomass was increasing in Shenandoah NP and stable on the Smithsonian Institute. EAB-caused mortality was first detected at the Smithsonian site in 2016 and accelerated steeply thereafter, exceeding 12.5% year by 2018. As of 2019, ash had lost 28% of individuals and 30% of aboveground biomass relative to 2016. Ninety-five percent of remaining live trees were considered “unhealthy’’ (Anderson-Teixeira et al. 2020).

eastern (flowering) dogwood; photo by F.T. Campbell

Unlike many studies, the Shenandoah study included understory species. Flowering dogwood declined by up to 90% from plots on the Smithsonian property; 2008–2019 mortality rates averaged 7.1%. Redbud declined by up to 76% from 1995 to 2018. The 2008–2019 mortality rates averaged 6.2% year.

Anderson-Teixeira et al. (2020) concede difficulty in estimating mortality due to less virulent or lethal pathogens, including Neofusicoccum spp. on redbud and Dutch elm disease on slippery elm.

Nevertheless, they believe their analysis probably underestimates the overall pest impacts because they did not analyze several other pest/host combinations known to be present in the Park: balsam woolly adelgid (Adelges piceae) on high-elevation populations of Abies balsamea; white pine blister rust (Cronartium ribicola) on eastern white pine (Pinus strobus); beech bark disease (Neonectria spp.) on American beech (Fagus grandifolia); thousand canker disease on walnut and butternut; and emerald ash borer on the novel host fringetree Chionanthus virginicus.

Another possible threat to oaks, winter moth (Operophtera brumata), is apparently now being controlled by the biocontrol agent Cyzenis albicans.

I am uncertain about the current status of two Diplodia fungi – Diplodia corticola and D. quercivora – link to blog which have been detected in both Florida and California. In Florida, almost all the symptomatic trees grow in cultivated settings where they are exposed to various stresses (Mullerin and Smith 2015).

However, host range studies indicate that 33 species of oaks and one species of chestnut that grow in the Southeast are vulnerable, to varying degrees, to D. corticola. Oaks in the red oak group (Section Lobatae) are more vulnerable than are white oaks (Section Quercus) (Mullerin and Smith 2015). In the test, the most vulnerable appear to be the following species native to the Southeast: Q. laurifolia, Q. virginiana, Q. geminata, Q. chapmanni, Q. laevis (turkey oak), Q. phellos, Q. pumila, and Q. incana (Dreaden et al. 2016).

What should we do?

Fei et al. (2019) noted that the losses to biomass would be exacerbated by the likely introduction of additional pests. They did not recommend any prevention actions.

Conrad et al. (2020) said their findings “lend support to national regulatory and awareness efforts to prevent the introduction and establishment of novel exotic insects and pathogens.”

Anderson-Teixeira et al. (2020) join others in declaring that future survival of the IUCN-listed species probably depends on conservation and restoration actions. They cite several sources, but not the CAPTURE Project – although the two studies reinforce each other. They specifically mention limiting invasive species’ spread through strengthened regulations and “enhanced plant biosecurity cyberinfrastructure”.

This last recommendation reinforces the message of Bonello et al. (2019) link to publication. We called for creation of a federal Center for Forest Pest Control and Prevention to implement end-to-end responses to forest pest invasions. One focus would be correcting the currently-inadequate focus on detection, development and deployment of genetic resistance while using modern techniques that allow for much faster breeding cycles.

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

SOURCES

Anderson-Teixeira, K.J., V. Herrmann, W.B. Cass, A.B. Williams, S.J. Paull, E.B. Gonzalez-Akre, R. Helcoski, A.J. Tepley, N.A. Bourg, C.T. Cosma, A.E. Ferson, C. Kittle, V. Meakem, I.R. McGregor, M. N. Prestipino, M.K. Scott, A.R. Terrell, A. Alonso, F. Dallmeier, and W.J. McShea. Date? Long-Term Impacts of Invasive Insects and Pathogens on Composition, Biomass, and Diversity of Forests in Virginia’s Blue Ridge Mountains. Ecosystems

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. 2019. Invasive tree pests devastate ecosystems – A proposed new response framework. Frontiers

Conrad, A.O., E.V. Crocker, X. Li, W.R. Thomas, T.O. Ochuodho, T.P. Holmes, and C. D. Nelson. 2020. Threats to Oaks in the Eastern US: Perceptions and Expectations of Experts. Journal of Forestry, 2020, 14–27

Dreaden, Black, Mullerin, and Smith. Poster presented at the 2016 USDA Invasive Species Research Forum

Fei, S., R.S. Morin, C.M. Oswalt, and A.M. Liebhold. 2019. Biomass losses resulting from insect and disease invasions in United States forests. Proceedings of the National academy of Sciences.

Guo, Q., S. Feib, K.M. Potter, A.M. Liebhold, and J. Wenf. 2019. Tree diversity regulates forest pest invasion. PNAS. www.pnas.org/cgi/doi/10.1073/pnas.1821039116

Morin, R.S., K.W. Gottschalk, M.E. Ostry, A.M. Liebhold. 2018. Regional patterns of declining butternut (Juglans cinerea L.) suggest site characteristics for restoration. Ecology and Evolution.2018;8:546-559

Morin, R. A. Liebhold, S. Pugh, and S. Fie. 2019. Current Status of Hosts and Future Risk of EAB Across the Range of Ash: Online Tools for Broad-Scale Impact Assessment. Presentation at the 81^st Northeastern Forest Pest Council, West Chester, PA, March 14, 2019

Mullerin, S. & J.A. Smith. 2015. Bot Canker of Oak in FL Caused by Diplodia corticola & D. quercivora. Emergent Pathogens on Oak and Grapevine in North America. FOR318

Potter, K.M., M.E. Escanferla, R.M. Jetton, and G. Man. 2019a. Important Insect and Disease Threats to United States Tree Species and Geographic Patterns of Their Potential Impacts. Forests. 2019 10 304.

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

Serious Invasive Species Damage to High-Elevation Sites in the West

Dream Lake, Rocky Mountain National Park, with limber pine
photo by F.T. Campbell

In this blog, I summarize two pest threats to the unique ecosystems on high-elevation mountain ridges in the West. At risk are several keystone tree species: the five-needle pines growing at high elevations (“high-five” pines) and subalpine fir. The invasive species causing this damage – white pine blister rust (WPBR; Cronartium ribicola) and balsam woolly adelgid (BWA; Adelges piceae) – are two of the most widespread non-native species threatening North American trees and affecting the highest proportion of host volumes (Morin).

The pines being killed by white pine blister rust are whitebark pine (Pinus albicaulis), limber pine (P. flexilis), Rocky Mountain bristlecone pine (P. aristata), foxtail pine (P. balfouriana), and southwestern white pine (P. flexilis var. reflexa). As of 2010, infestations had not been reported on Great Basin bristlecone pine (P. longaeva) and the Mexican white pine species. [Unless otherwise indicated, information on white pine blister rust is from a comprehensive review and synthesis published in the August 2010 issue of Forest Pathology (Vol. 40:3-4).]

As noted above, sub-alpine fir (Abies lasiocarpa) is also being affected – although less uniformly than the pines – by the balsam woolly adelgid.

Both of these pests arrived approximately a century ago, but they are still spreading and causing additional damage. White pine blister rust had spread widely throughout the West within 40 years of its introduction. Meanwhile, BWA spread among lowland and subalpine firs along the Pacific coast from California to British Columbia within 30 years of its first detection. Its spread eastward was slower, but relentless. It reached Idaho, Montana, Utah and interior British Columbia within 50 years. Also, BWA reached Alaska within 90 years of its introduction in California. These pests are perfect examples of how invasive species introduced long ago are dreaded “gifts that keep on giving”.

For a detailed discussion of these pests’ impacts, see the descriptions posted here. To summarize, though, WPBR is present in the ranges of eight of the nine vulnerable western white pines and has caused severe mortality to some species (Sniezko et. al. 2011). For example, 88% of the limber pine range in Alberta is affected (Dawe et al. 2020). WPBR is generally causing more damage to its hosts’ northern populations. Impact of the BWA are more subtle than WPBR. Also, impacts’ severity is linked to climatic conditions. For example, measurable decline on the Olympic Peninsula was greater on south-facing slopes. However, the study did not determine whether this reflected heat-loading and tree stress or more abundant subalpine fir on these slopes. An estimated 19-53% (average 37%) of subalpine fir trees had died on sample plots on one ridge over the 19 years since BWA was first detected there. Overall forest growth after 2007 could indicate partial recovery, a momentary pause in BWA invasion, or tree growth after severe weather events (Hutton 2015).

Ranges of Trees at Risk

Many of the host trees of these two pests are widespread; others are more narrowly endemic.

Limber pine reaches from Alberta and British Columbia south to mountain peaks in Arizona and New Mexico. Whitebark pine is found from Alberta and British Columbia to California and Nevada (USDA Plants database. Subalpine fir stretches from southeast Alaska along the Canadian Rockies coast into Washington, Oregon, east into Idaho, Montana, Wyoming, Colorado, Utah, even into scattered mountain ranges of Nevada and New Mexico (Hutton 2015).

Limber pine and subalpine fir are also found in a wide range of ecosystems within these ranges. Limber pine is found at both upper and lower tree lines in grassy, open forests; on exposed rocky slopes; and in dense, mixed-conifer stands. Subalpine fir is a pioneer species on ridges, alpine meadows, avalanche chutes, and lava beds (Ragenovich and Mitchell, 2006).

Before arrival of non-native pests or pathogens, these tree species have persisted for thousands of years under harsh conditions (Hutton 2015). Many of the individual trees were long-lived; some five-needle pines, e.g., bristlecone pines, have famously live for thousands of years. Core studies demonstrated that subalpine firs trees could live 272 years in the forests of Olympic National Park and 240 years in Glacier National Park (Hutton 2015). Surely loss of these trees – or even their conversion from large and old to small and short-lived – will result in significant destruction of these unique biomes.

All these trees play important roles in high altitude, unique ecosystems (Pederson et al. no date; Dawe 2020; Hutton 2015):

They retain ground water, slow the rate of snow melt, and maintain stream flow characteristics and water quality;
They curtail soil erosion and maintain slope stability; and
They provide high-value food and shelter to wildlife.

Whitebark and limber pines are famous for providing critical food for many wildlife species at high elevations —notably bears and nutcrackers (Compendium and Dawe 2020).

More Pest Threats

Other diseases, insects, and disturbances also pose serious threats to these tree species. The threats vary by region and age of the stand. They include – for the pines — mountain pine beetle (Dendroctonus ponderosae), dwarf mistletoe (Arceuthobium spp.), and various shoot, cone or foliage insects and pathogens. For subalpine fir, threats include western balsam bark beetle (Dryocoetes confusus), fir engraver (Scolytus ventralis), and the fir root bark beetle (Pseudohylesinus granulatus) (Hutton 2015). Trees are also damaged by bear and deer, seed predation by squirrels, wildfire, and biotic succession.

On Washington’s Olympic Peninsula, BWA initiates or predisposes subalpine fir for a novel disturbance complex. BWA-caused stress makes the trees more susceptible to moisture stress and endemic bark beetle attack. Surviving trees are subsequently subject to toppling by wind. A tree can die in a few years, survive with insects for up to 20 years, or recover, depending on duration, severity, and location of infestation, and local environmental conditions (Hutton 2015).

BWA study plots in the Cascade Range experienced subalpine fir mortality ranging from 7 to 79% (measured as stem counts, not basal area) over a 19 to 38 years study period. Higher mortality occurred at low-elevation, mesic sites. One stand experienced 40% mortality in 19 years, but lost the remaining 60% during a subsequent spruce budworm infestation. Most plots continued to show sporadic signs of adelgid presence and continued tree mortality. However, 41-69% of trees survived stem infestations (Hutton 2015).

How to Protect These Ecosystems

The seeds of both whitebark and limber pines are dispersed to newly disturbed, open areas by Clark’s nutcracker (Nucifraga columbiana). Furthermore, whitebark cones open to release seeds only after fire. This had led to expectations that prescribed fire could promote regeneration of these species. However, studies by Dawe (2020) and other have found that nutcracker seed caching behavior and seedling establishment are complex. Fire management might have to vary among regions, demanding consideration of stand characteristics,like openness and the presence of other tree species. For example, in the Colorado Front Range, limber pine can be replaced by subalpine fir when fire-free intervals are long. On the other hand, in Alberta, fire appeared to boost regeneration of the dominant tree species in the stands pre-fire. In the study areas, these were white spruce (Picea glauca) and lodgepole pine (Pinus contorta) (Dawe 2020). Dawe recommends protecting existing stands of limber pine through fire mitigation efforts, e.g., thinning and other fuel treatments, and supplementary planting of seedlings.

Efforts to find biocontrol agents to target the balsam woolly adelgid began in 1957; the original focus was on the insects’ damage to Fraser fir (Abies fraseri) in the southern Appalachians. More than 25 predatory species have been introduced from Europe and Asia. There was simultaneous research on native predators. None has had an impact on BWA populations in either the East or the West.

Neither white pine blister rust nor balsam woolly adelgid is considered a quarantine pest by federal officials, so there is no attempt to prevent their movement via interstate trade in Christmas trees, timber, or nursery stock. Hutton (2015) hypothesizes that the absence of regulatory measures targetting BWA arises from the pest’s gradual effect and the hosts’ not being commercially important as timber species (although several firs are important in horticulture and as Christmas trees). I think another factor is that the pests were introduced so long ago and are now widespread.

Efforts are under way to detect resistant genotypes to be used in breeding programs. Several of the lower-elevation five-needle pines vulnerable to WPBR have benefitted from extensive breeding efforts Whitebark pine has more recently been added to programs.

The eastern Fraser fir is the target of breeding – primarily for Christmas trees (APS). However, at least small-scale volunteer efforts have been carried forward by the Alliance for Saving Threatened Forests.

Hutton (2015) expresses hope that evolutionary pressure by BWA might enhance survival of more resistant forms of subalpine fir and lead to their gradual takeover. However, I ask, why leave it to chance?

In this context, I remind you of my involvement with a group (see Bonello et al. 2019) proposing creation of a federal Center for Forest Pest Control and Prevention to implement end-to-end responses to forest pest invasions – including overcoming the currently inadequate focus on detection, development and deployment of genetic resistance using modern techniques that allow for much faster breeding cycles.

I am puzzled that the Project CAPTURE places whitebark pine and subalpine fir only in Class A4.2, not among the highest priority species (Potter et al. 2019). As I blogged last spring, Project CAPTURE is part of a multi-partner effort to categorize and prioritize US tree species for conservation actions based on the threats and the trees’ ability to adapt to those threats. I find it puzzling because I am not sure I agree that these two species have a moderately high mean pest severity score – as required by the category. I am less puzzled by the assignment of a low adaptive capacity score.

Limber pine apparently ranks even lower in the Project CAPTURE priority process.

Posted by Faith Campbell

SOURCES

A comprehensive review and synthesis of the history, ecology, and management of white pines threatened by white pine blister rust see the August 2010 issue of Forest Pathology (Vol. 40:3-4).

American Phytopathological Society. Science Daily. December 9, 2019 https://www.sciencedaily.com/releases/2019/12/191209161314.htm?utm_source=feedburner&utm_medium=email&utm_campaign=Feed%3A+sciencedaily%2Fplants_animals%2Finvasive_species+%28Invasive+Species+News+–+ScienceDaily%29

Dawe, D.A., V.S. Peters, M.D. Flannigan. 2020. Post-fire regeneration of endangered limber pine (Pinus flexilis) at the Northern extent of its range. Forest Ecology and Management 457 (2020) 117725

Hutton, K.M. 2015. A dissertation submitted in partial fulfillment of the requirements for the degree of Doctor of Philosophy. University of Washington. Available here

Morin, R. Presentation to the 81^st Northeastern Forest Pest Council Northeastern states forst agencies, Philadelphia, Pennsylvania, March 2019.

Potter, K.M., Escanferla, M.E., Jetton, R.M., Man, G., Crane, B.S. 2019. Prioritizing the conservation needs of US tree spp: Evaluating vulnerability to forest P&P threats, Global Ecology and Conservation (2019), doi: https://doi.org/10.1016/

Ragenovich, I.R. and R.G. Mitchell. 2006. Forest Insect and Disease Leaflet (FIDL) #118. http://www.na.fs.fed.us/pubs/fidls/bwa.pdf

Sniezko, R.A., M.F. Mahalovich, A.W. Schoettle, D.R. Vogler. 2011. Past and Current Investigations of the Genetic Resistance to Cronartium ribicola in High-elevation Five-needle Pines. In Keane, R.F., D.F. Tomback, M.P. Murray, and C.M Smith, eds. 2011. The future of high-elevation, five-needle white pines in Western North America. Proceedings of the High Five Symposium. 28-30 June, 2010. Missoula, MT.

International Year of Plant Health: Time to Admit that the International Phytosanitary System is Failing

As I noted last November, the premise of the international phytosanitary system – the Agreement on the Application of Sanitary and Phytosanitary Standards (SPS Agreement) and the International Plant Protection Convention (IPPC) – is that importing countries should, and can, rely on exporting countries to take the actions necessary to meet the importing countries’ plant health goals. However, the experience with the International Standard on Phytosanitary Measures (ISPM) #15 and wood packaging casts doubt on this premise.

Exporters are not reliably ensuring the cleanliness of their wood packaging, putting American forests at risk. Indeed, some experts have concluded that continuing to implement ISPM#15 at current levels could triple the number of non-native wood-boring insects introduced into the U.S. by 2050 (Leung et al. 2014).

Too many shipments carry wood packaging that bears no ISPM#15 stamp. And too many pieces of wood packaging arrive with the ISPM#15 stamp, yet are not reliably pest-free. If we cannot clean up this pathway – which involves boards or even logs that are, after all, already dead — it bodes poorly for limiting pests imported with other commodities that are pathways for tree-killing pests – especially living plants (plants for planting). Living plants are much more easily damaged or killed by treatments than the dead wood used in packaging – so ensuring pest-free status of a shipment is even more difficult. (A longer discussion of the SPS Agreement and IPPC is found in Chapter III of Fading Forests II, available here.

Here are the problems – and the latest evidence.

ALB larva in piece of wood packaging material

Too Many Shipments with Pest-Infested Wood Packaging Are Reaching the Country

My information on Customs and Border Protection (CBP) interceptions comes primarily from Kevin Harriger (see full reference at end of the blog). I will note when it comes from other sources.

In November 2019, Kevin Harriger reported that over the past three years, CBP detected a regulated pest, on average, in 30% of the wood packaging the agency intercepted because it was not compliant with ISPM#15. Non-compliance is defined as wood packaging that either lacks an official mark or is infested by a quarantine pest, or both.

From this and previous reports, I have 10 years of CBP interception data – from 2020 – 2019. These data thus begin four years after the U.S. began implementing ISPM#15 (in 2006) and 11 years after the U.S. began requiring China to treat wood packaging accompanying its exports (in 1999).

Over the period 2010 – 2018, CBP intercepted an average of 3,183 shipments with non-compliant wood packaging each year. On average, 2,100 (66%) of these shipments lacked the required ISPM#15 mark. A live quarantine pest was found in an average of 794 (25%) shipments. (There was some overlap in the categories).

In 2019, CBP intercepted a total of 2,572 non-compliant shipments (Stephen Brady, CBP, April 2020). Those lacking the ISPM#15 mark number 1,825 (71%). Shipments in which a live pest was found numbered 747 (29%).

The 2019 data show decreases, in absolute numbers, from earlier years in all categories: a 19% decrease below 1010-2018 average of shipments intercepted; a 13% decrease in number of shipments intercepted because the wood packaging lacked the ISPM#15 mark; a decrease of 6% in the number of shipments intercepted that had a quarantine pest. It is too early to say whether CBP’s stronger enforcement approach launched in November 2017 has resulted in a lower number of shipments in violation of ISPM#15 approaching our shores.

There has been a dispute about which categories of packaging are most likely to be infested. The categories are pallets, crates, spools for cable, and dunnage (wood used to brace cargo and prevent it from shifting). The CBP data available to me and the study by Krishnankutty et al. (2020b – see full reference at the end of this blog) shed no light on that issue.

What is the actual number of infested containers approaching our shores? We know that CBP inspects, on average, 2% of incoming containers – so the above interception data reflect a small percentage of probable true approach rate.

The first issue is, how many containers arrive here?

I have been unable to find data for 2019 – much less 2020, when the media report that import volumes have crashed. Until recently, import volumes had been rising. According to a U.S. DOT report to Congress (see reference at the end of this blog), 25 U.S. maritime ports received 24,789,000 loaded shipping containers (measured as TEU – 20-foot equivalent) in 2018. The number of incoming containers had increased at the top three ports – Long Beach, Los Angeles, and New York / New Jersey – between 3% and 7% since 2016.

However, APHIS told me in November 2019 that CBP reports that only about 13 million loaded containers enter the country every year by rail, truck, air, or sea. While I can’t yet explain the discrepancy, one possible explanation is that DoT counts 40-foot containers as two 20-foot containers.

(Of course, pests introduced to Canada also threaten North America’s forests. Canada received fewer than 5 million containers via maritime trade in 2016 (Asbil pers. comm. 2018).

Two decade-old estimates of the proportion of incoming containers that hold wood packaging (Haack et al. 2017, Meissner et al. 2009) allow me to estimate the risk associated with these incoming containers. Meissner et al. found that 75% of maritime containers have wood packaging. Haack et al. estimated that the wood in 0.1% of those containers was infested. Applying these two factors, I conclude that as many as 18,590 of incoming containers in maritime trade could have been transporting a woodborer in the regulated families (Cerambycids, Buprestids, Siricids). I am hesitant to apply the calculation to CBP’s estimate because I don’t know how many of the 13 million containers entered by sea. However, if I assume that the same percentage of wood packaging applied to all the CBP-counted containers, I conclude that 9,750 of those containers held infested wood packaging – still a significant number.

The actual approach rate might be less – or more! Haack et al. (2014) did not include imports from China in their calculations. Given the history of interceptions, it appears probable to me that a recalculation of the approach rate that included China would probably raise the overall proportion.

Furthermore, 11 years have passed since Haack and Meissner made their calculations. During that time, ISPM#15 has been amended to make it more effective. The most important change was restricting the size of bark remnants that may remain on the wood. I have asked several times that APHIS commission a new analysis of Agriculture Quarantine Inspection Monitoring data to determine the pest approach rate before and after the CBP action in order to determine whether the more aggressive enforcement has led to reductions in non-compliant shipments at the border.

By comparing Dr. Haack’s estimate (see above) with the CBP data, I estimate that Customs is detecting and halting the importation of 4 – 8% of the shipments that actually contain pest-infested wood. Since CBP inspects only about two percent of incoming shipments, the higher detection rate demonstrates the value of CBP’s program to target likely violators – and deserves praise. But it is obviously too low a “catch” rate to provide an adequate level of protection for our forests.

ISPS#15 Is Not Helping to Target Inspections

So – ISPM#15 still allows too many pests to arrive at our shores. Is ISPM#15 at least helping phytosanitary agencies target inspections? No, because both U.S. and European data demonstrate that a high proportion of shipments containing infested wood pieces bore the ISPM#15 stamp. Phytosanitary agencies cannot rely on the presence or absence of the stamp to indicate the pest risk level.

U.S. data:

During the period 2010-2015, CBP found that an average of 95% of pest-infested shipments bore the ISPM#15 mark (Harriger). Unfortunately, CBP data from more recent years don’t provide this breakdown.
In the past two years, CBP inspectors have repeatedly found pests in dunnage bearing the ISPM#15 mark.
Krishnankutty et al. (2020b) analyzed wood packaging from 42 countries of origin intercepted by CBP over six years (April 2012 – January 2018). They found that 87% of the interceptions bore the ISPM mark.

I blogged earlier about the velvet longhorned beetle (Trichoferus (=Hesperophanes) campestris) This pest, like others, has reached our shores and entered the country both before and after implementation of ISPM#15. The predictable result is that VLB is established in three states and has been detected in 14 others plus Puerto Rico (Krishnankutty, et al. 2020a). Apparently we have been lucky that this one isn’t as damaging as so many are!

European data:

For Europe, see Eyre et al. (2018). They concluded that the ISPM-15 mark was of little value in predicting whether harmful organisms were present.

This is alarming and we need to understand the reason – How much is caused by fraud? How much is caused by failure of treatment – either intrinsic weakness or incorrect application? APHIS researchers have found that larvae from wood subjected to methyl bromide fumigationwere more likely to survive to adulthood than those intercepted in wood that had been heat treated (Nadel et al. 2016).

Krishnankutty et al. (2020b) query whether the 2009 requirement that wood be debarked might be less effective in countering insect species that require bark only in the early stages of larval development. Half of the species intercepted in hardwood shipments (e.g., Anoplophora glabripennis, Phoracantha recurva) might fit this profile. They also appear to pose a higher threat since they are polyphagous and known to infest healthy hosts. While some of the softwood-inhabiting species also require bark, they not known to infest living trees and only a quarter were classified in the high-risk group. The Mech et al. 2020 finding that no wood-borers that specialize in conifers posed a high risk appears to support these different impacts.

Krishnankutty, et al. (2020b) also note the risk from pallet recycling. The wood might occasionally be infested by dry-wood borers. One puzzling example was wood packaging shipped from Brazil and bearing a Brazilian ISPM#15 stamp that was infested with a larva of T. campestris (VLB). This is an Asian species not recorded as being present in South or Central America. The authors speculate that the pallets were recycled in Brazil after inadequate treatment in their original places of manufacture.

Of the 17 wood borer species intercepted in hardwoods, three have reproducing populations in the U.S.: A. glabripennis, Phoracantha recurva and T. campestris. Krishnankutty et al. (2020b) say that they are unaware of any of the non-native buprestids and siricids intercepted in softwood SWPM being established in the US. (One Siricid that is established, Sirex noctillio, was not detected in the wood packaging analyzed in this study.)

What Can Be Done to Slow or Eliminate this Pathway?

CBP strengthened enforcement of ISPM#15 in November 2017. CBP’s enforcement actions increased by 400% from 2017 to 2018 (John Sagle, CBP, pers. comm). CBP has also expanded its outreach to shippers and others involved in international trade with the goal of reducing all types of non-compliance – lack of documentation, pest presence, etc. in both wood packaging and shipping containers. The outreach includes awareness campaigns targetting trade, industry, affiliated associations, CBP employees, and international partners (Harriger).

Certain countries have a long-standing record of non-compliance with ISPM#15 – as seen in interception records.

Haack et al. 2014 – Italy was the country of origin for most wood borers intercepted 1985 – 2000.
Haack et al. 2014 – the top 5 countries in the 2003 – 2009 period were Mexico (33.7%), Italy (14.2%), Canada (13.4%), Netherlands (4.4%), China (4.1%).
APHIS’ interception database for FY2011-2016 (provided to me) showed Mexico, China, Italy, and Costa Rica had the highest numbers of interceptions.
Krishnankutty et al. (2020b) found the highest numbers of interceptions came from Mexico, China, and Turkey.

These numbers reflect in part the huge volumes of goods imported from both Mexico and China. But China and Italy stand out for their poor performance. (The U.S. does not regulate – or inspect! – wood packaging from our third-largest trade partner Canada.)

Officials know which individual companies within these countries have a history of non-compliance. For example, 21 of the interceptions on wood packaging made from Populus trees in China (53%) were associated with stone, ceramic, and terracotta commodities. Anoplophora glabripennis was intercepted six times in Populus originating from a single wood-treatment facility in China (Krishnankutty et al. 2020b).

How reduce risk to U.S. forests?

Over the past year or two, I have suggested the following actions:

USDA APHIS join Bureau of Customs and Border Protection in penalizing violators.
Citing the need for setting a higher “level of protection”, APHIS & the Canadian Food Inspection Agency (CFIA) should prepare a risk assessment to justify adopting more restrictive regulations. The new regulations should prohibit use of packaging made from solid wood – at least from the countries with records of high levels of non-compliance (listed above).
USDA Foreign Agriculture Service (FAS) should assist U.S. importers to determine which suppliers reliably provide compliant wood packaging.
USDA FAS and APHIS should help importers convey their complaints about specific shipments to the exporting countries’ National Plant Protection Organizations (NPPOs; departments of agriculture).
APHIS should increase pressure on foreign NPPOs and the International Plant Protection Convention more generally to ascertain the reasons ISPM#15 is failing and to fix the problems.
APHIS should fund more studies and audits of wood packaging to document the current efficacy of the standard, including an urgent update of the Haack study of pest approach rate.

The international standard has demonstrably failed to provide a secure method to evaluate the pest risk associated with wood packaging accompanying any particular shipment. The presence of the stamp on pieces of wood packaging does not reliably show that the wood is pest-free.

The situation is even worse re: movement of plants for planting.

SOURCES

Asbil, W. Canadian Food Inspection Agency, pers. comm. August 2018.

Eyre, D., R. Macarthur, R.A. Haack, Y. Lu, and H. Krehan. 2018. Variation in Inspection Efficacy by Member States of SWPM Entering EU. Journal of Economic Entomology, 111(2), 2018, 707–715)

Haack, R. A., K. O. Britton, E. G. Brockerhoff, J. F. Cavey, L. J. Garrett, M. Kimberley, F. Lowenstein, A. Nuding, L. J. Olson, J. Turner, and K. N. Vasilaky. 2014. Effectiveness of the international phytosanitary standard ISPM no. 15 on reducing wood borer infestation rates in wood packaging material entering the United States. Plos One 9:e96611.

Harriger, K. Executive Director for the Agriculture Programs and Trade Liaison office, Department of Homeland Security Bureau of Customs and Border Protection (CBP), presentations to the Continental Dialogue on Non-Native Forest Insects and Diseases, over appropriate years. https://continentalforestdialogue.org/events/

Krishnankutty, S.M., K. Bigsby, J. Hastings, Y. Takeuchi, Y. Wu, S.W. Lingafelter, H. Nadel, S.W. Myers, and A.M. Ray. 2020a. Predicting Establishment Potential of an Invasive Wood-Boring Beetle, Trichoferus campestris (Coleoptera:) in the United States. Annals of the Entomological Society of America, XX(X), 2020, 1–12

Krishnankutty, S., H. Nadel, A.M. Taylor, M.C. Wiemann, Y. Wu, S.W. Lingafelter, S.W. Myers, and A.M. Ray. 2020b. 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. Commodity Treatment and Quarantine Entomology

Leung, B., M.R. Springborn, J.A. Turner, E.G. Brockerhoff. 2014. Pathway-level risk analysis: the net present value of an invasive species policy in the US. The Ecological Society of America. Frontiers of Ecology.org

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.

Meissner, H., A. Lemay, C. Bertone, K. Schwartzburg, L. Ferguson, L. Newton. 2009. Evaluation of Pathways for Exotic Plant Pest Movement into and within the Greater Caribbean Region. Caribbean Invasive Species Working Group (CISWG) and USDA APHIS Plant Epidemiology and Risk Analysis Laboratory

Nadel, H. S. Meyers, J. Molongoski, Y. Wu, S. Lingafelter, A. Ray, S. Krishnankutty, A. Taylor. 2017. Identification of Port Interceptions in Wood Packing Material Cumulative Progress Report, April 2012 – June 2017

USDA APHIS interception database – pers. comm. January 2017.

USDA APHIS press release dated September 12, 2018

U.S. Department of Agriculture, Press Release No. 0133.20, January 27, 2020

US Department of Transportation. Port Performance Freight Statistics in 2018 Annual Report to Congress 2019 https://rosap.ntl.bts.gov/view/dot/43525

Wu, Y., S.M. Krishnankutty, K.A. Vieira, B. Wang. 2020. Invasion of Trichoferus campestris (Coleoptera: Cerambycidae) into the United States characterized by high levels of genetic diversity and recurrent intros. Biological Invasions Volume 22, pages1309–1323(2020)

Yemshanov, D., F.H. Koch, M. Ducey, K. Koehler. 2012. Trade-associated pathways of alien forest insect entries in Canada. Biol Invasions (2012) 14:797–812

Posted by Faith Campbell