Category: drivers of extinction

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 81st 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

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

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

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 

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 81st 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.

Another New Pest Detected in California; Possible Threat to Native Shrubs

Rhus integrifolia

The California Department of Food and Agriculture (CDFA) is seeking comments  on the appropriate pest rating for Leptosillia pistaciae, a recently discovered fungus that causes pistachio canker.

The Department’s draft pest ranking assigns the highest Economic Impact score – three. It assigns a medium Environmental Impact – two. This is because the pathogen can kill an important native shrub, with possible follow-on consequences of reduced biodiversity, disrupted natural communities, or changed ecosystem processes.

CDFA states that there is no uncertainty in its evaluation, but I see, and describe here, numerous questions about the possible true extent of the invasion and possible host range.  

Comments are due on April 4, 2020.

The pathogen was detected in June 2019, when a habitat manager from an ecological reserve in San Diego County noticed multiple dead lemonade berry shrubs (Rhus integrifolia) in one of the parks. This is the first known detection of Leptosillia pistaciae in the United States and on this host. USDA APHIS has classified Leptosillia pistaciae as a federal quarantine pest. Rhus and Pistacia are in the same family, Anacardiaceae (cashews and sumacs).

According to the CDFA, Leptosillia pistaciae is the only member of this fungal genus known to be associated with disease symptoms on plants. Other species are endophytes or found in dead plant tissues. [It is not at all unusual for fungal species to be endophytes on some plant hosts but pathogenic on others. A California example is Gibberella circinata (anamorph Fusarium circinatum), which causes pitch canker on Monterey pine (Pinus radiata) but is an endophyte on various grass species (Holcus lanatus and Festuca arundinacea).]

(Reminder: this is the second new pest of native species detected in California state in 2019; I blogged about an ambrosia beetle in Napa County here. )

Rhus integrifolia (lemonade berry or lemonade sumac) is native to California. It grows primarily in the south, along the coast – from San Diego to San Luis Obispo. However, some populations are also found in the San Francisco Bay area. This and other sumacs are also sold in the nursery trade.

On pistachio trees in Italy, symptoms are observed in the winter and late spring. During the winter dormant season, trees had gum exudation and cracking and peeling of bark on trunks and branches. On trunks and large branches, cankers appeared first as light, dead circular areas in the bark; subsequently they became darker and sunken. Under the bark, cankers were discolored with necrotic tissues; in some cases, these extended to the vascular tissues and pith. During the active growing season, the symptomatic plants also showed canopy decline. Inflorescences and shoots, originating from infected branches or twigs, wilted and died. When the trunk was girdled by a canker, a collapse of the entire tree occurred.

range map for Rhus integrifolia

On lemonade berry, large clumps of dead adult shrubs were observed on the edge of hiking trails. Some shrubs that had completely dead foliage were re-sprouting from their bases. Trunks of shrubs that were not completely dead were copiously weeping sap and fluids and showed foliage browning and die back with symptoms of stress.

It is thought that spores could be spread by wind, rain splashing, and the movement of dead or dying trees, greenwaste, and infected nursery stock. Contaminated pruning tools might also transport the spores. The possibility of a latent phase – or perhaps asymptomatic hosts – adds to the probability of anthropomorphically assisted spread.

I question how much effort has been put into detection surveys, especially in natural systems with native Rhus species. California has three other native sumacs: R. ovata, R. aromatica, and Malosma laurina (CNPS; full citation at the end of the blog). In addition, there are numerous other species in the family, including poison oaks (Toxicodendron spp.) and the widespread invasive plant genus Schinus.

Furthermore, some plants in the family (other than pistachios) are grown for fruit or in ornamental horticulture, including two of the native sumacs and two non-native species, Rhus glabra and R. lanceolata, cashew, mango, and smoke trees (Cotinus spp.).

Yet CDFA confidently states that there are only two hosts and that it has been detected in only one population – that in San Diego. This is because CDFA considers only official records identified by a taxonomic expert and supported by voucher specimens.

CDFA states that the pathogen is likely to survive in all parts of the state where pistachios are grown – primarily in the Central Valley. California supplies 98% of the pistachios grown in the United States; the remainder is raised in Arizona and New Mexico. California production occurred on 178,000 acres in 2012. A map is included in a flyer on production available at the url listed at the end of this blog.

In discussing spread potential, no mention is made of possible human-assisted spread.

The CDFA document includes instructions for submitting comments; the deadline is April 4.

Sources:

Rhus and related species native to California: California Native Plant Society

https://calscape.org/loc-california/Rhus(all)/vw-list/np-1?

Rhus species used in horticultural plantings in California: CalFlora https://www.calflora.org//cgi-bin/specieslist.cgi?where-genus=Rhus 

Pistachio production information: https://apps1.cdfa.ca.gov/FertilizerResearch/docs/Pistachio_Production_CA.pdf

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.

Progress – Now Threatened – On Protecting Our Cacti

prickly pear cacti in Big Bend National Park
photo by Blake Trester, National Park Service

The cacti that are such important components of desert ecosystems across nearly 2 million square miles straddling the U.S.-Mexico border are under threat from non-native insects – as I have noted in earlier blogs. Of course, cacti are important in other ecoregions, too – I wrote recently about the columnar cacti in the dry forests of Puerto Rico.

Flat-padded prickly pear cacti of the genus Opuntia are threatened by the cactus moth, Cactoblastis cactorum.

In 1989, the cactus moth was found in southern Florida, to which it had spread from the Caribbean islands (Simonson 2005). Recently, the moth was found to have spread west as far as the Galveston, Texas, area and near I-10 in Columbus, Texas, about 75 miles west of central Houston (Stephen Hight, pers. com.) Two small outbreaks on islands off Mexico’s Caribbean coast have been eradicated.

In Florida, the cactus moth has caused considerable harm to six native species of prickly pear, three of which are listed by the state as threatened or endangered.

When the cactus moth reaches the more arid regions of Texas, it is likely to spread throughout the desert Southwest and into Mexico. In the American southwest, 31 Opuntia species are at risk; nine of them are endemic, one is endangered. Mexico is the center of endemism for the Opuntia genus. In Mexico, 54 Opuntia species are at risk, 38 of which are endemic (Varone et al. 2019; full citation at end of this blog).

The long-term effects of the cactus moth on these North American Opuntia are unknown because there may be substantial variations in tolerance. The attacks observed in the Caribbean islands have shown great variability in various cactus species’ vulnerability (Varone et al. 2019).

The Opuntia cacti support a diversity of pollinators as well as deer, javalina (peccaries), tortoises, and lizards. Prickly pears also shelter packrats and nesting birds (which in turn are fed on by raptors, coyotes, and snakes), and plant seedlings. Their roots hold highly erodible soils in place (Simonson 2005).

While scientists have been concerned about the possible impacts of the cactus moth since it was detected in Florida 30 years ago, a substantial response began only 15 years later. The U.S. Department of Agriculture began trying to slow the spread of the cactus moth in 2005 (Mengoni Goñalons et al. 2014), with a focus on surveys and monitoring, host (cactus) removal, and release of sterile males. This program was successful at slowing the moth’s spread and eradicating small outbreaks on offshore islands of Alabama, Mississippi, and Mexico.

Cactus moth damage to native cacti in Florida
photo by Christine Miller, UF/IFAS

However, the moth continued to spread west and the program never received an appropriation from Congress. The primary funding source was a US – Mexico Bi-National Invasive Cactus Moth Abatement Program. Both countries contributed funds to support the research and operational program to slow the spread in the U.S. Funds were provided through USDA Animal and Plant Health and Inspection Service (APHIS) and the Mexican Secretariat of Agriculture, Livestock, Rural Development, Fisheries and Food (SEGARPA). Unfortunately, funding was reduced by both entities and became inadequate to maintain the Bi-National Program.

Therefore, in 2012, APHIS abandoned its regional program and shifted the focus to biocontrol. This is now considered the only viable control measure in the desert Southwest where vulnerable cacti are numerous and grow close together. The biocontrol project has been funded since 2012 through the Plant Pest and Disease Management and Disaster Prevention program (which receives funding through the Farm Bill). It has received a total of slightly more than $2 million over seven years. More than half the funds went to the quarantine facility to support efforts to rear non-target hosts and verify the biocontrol agent’s host specificity. About a quarter of the funds supported complementary work of an Argentine team (both the cactus moth and the most promising biocontrol agent are native to Argentina). Much smaller amounts have supported U.S.-based scientists who have studied other aspects of the cactus moth’s behavior and collected and identified the U.S. moths being tested for their possible vulnerability to attack by a biocontrol wasp.

Here are details of what these dedicated scientists achieved in just the past seven years at the relatively low cost of roughly $2 million. Unfortunately, the project now faces a funding crisis and we need to ensure they have the resources to finish their work.

Some Specifics of the BioControl Program

After literature reviews, extensive collections, and studies in the cactus moth’s native habitat in Argentina (Varone et al. 2015), a newly described wasp, Apanteles opuntiarum (Mengoni Goñalons et al. 2014), has been determined to be host specific on Argentine Cactoblastis species and the most promising candidate for biocontrol. Wasps were collected in Argentina and sent to establish a colony in a quarantine facility in Florida to enable host specificity studies on North American Lepidoptera (Varone et al. 2015).

Quarantine host specificity studies and development of rearing technology has not been straightforward. Initially, it was difficult to achieve a balanced male/female ratio in the laboratory-bred generations; this balance is required to maintain stable quarantine laboratory colonies for host range testing. This difficulty was overcome. A second challenge was high mortality of the cactus-feeding insects collected in the Southwest that were to be test for vulnerability to the biocontrol wasp. These desert-dwellers don’t do well in the humid, air-conditioned climate of the quarantine facility! For these difficult-to-rear native insects, scientists developed a molecular genetics method to detect whether eggs or larvae of the cactus moth parasitoid were present inside test caterpillars after they were exposed to the wasps. For easy to rear test insects, caterpillars are exposed to the wasps and reared to adulthood. Host specificity tests have been conducted on at least five species of native U.S. cactus-feeding caterpillars and 11 species of non-cactus-feeding caterpillars (Srivastava  et al. 2019; Hight pers.comm.).

To date there has been no instance of parasitism by Apanteles opuntiarum on either lepidopteran non-target species or non-cactus-feeding insects in the Florida quarantine or in field collections in Argentina (Srivastava et al. 2019; Varone et al. 2015; Hight pers.comm.).

The scientists expected to complete host-specificity testing in the coming months, then submit a petition to APHIS requesting the release of the wasp as a biocontrol agent. Unfortunately, the project’s request for about $250,000 in the current year was not funded. This money would have funded completion of the host specificity testing, preparation of a petition to APHIS in support of release of the biocontrol agent into the environment, and preparation of the release plan.

Meanwhile, what can we expect regarding the probable efficacy of the anticipated biocontrol program?

Some of the wasp’s behavioral traits are encouraging. The wasp is widely present in the range of the cactus moth, and persisted in these areas over the years of the study. The wasp can deposit multiple eggs with each “sting”. Multiple wasps can oviposit into each cactus moth without detriment to the wasp offspring. Unmated wasp females produce male offspring only, whereas mated females produce mixed offspring genders. In the field, female wasps attack cactus moth larvae in a variety of scenarios: they wait at plant access holes to sting larvae when they come outside to defecate; they attack larvae when they are moving on the surface of the pads; they can sting the youngest cactus moth larvae through the thin plant wall of mined the pads; and they enter large access holes created by older larvae and attack larger larvae. The wasps are attracted by the frass (excrement) left on the outside of the cactus pads by cactus moth larvae (Varone et al. 2020).

However, I wonder about the extent to which the cactus moth is controlled by parasitoids in Argentina. Cactoblastis eggs are killed primarily by being dislodged during weather events (rain and wind) and by predation by ants. First instar larvae are killed primarily by the native Argentine cactus plants’ own defenses – thick cuticles and release of sticky mucilage when the young larvae chew holes into the pads where they enter and feed internally. As larvae feed and develop inside the pads, the primary cause of mortality is natural enemies.

Of all the parasitoid species that attack C. cactorum, A. opuntiarum is the most abundant and important. When the larvae reach their final state (6th instars), they leave the pads and find pupation sites in plant litter near the base of the plants. It is at this stage that the parasitism from A. opuntiarum is detected in the younger larvae that were attacked while feeding inside pads. As the moth larva begins to spin silk into which to pupate, larvae of the wasp erupt through the skin of the caterpillar and pupate within the silk spun by the moth. Predation by generalists (ants, spiders, predatory beetles) accounted for high mortality of the unprotected last instar and pupae (Varone et al. 2019).

Finally, the cactus moth has three generations per year when feeding on O. stricta in the subtropical and tropical coastal areas of the Americas and the Caribbean. In Argentina, on its native host, the moth completes only two generations per year (Varone et al. 2019).

 How to Get the Program Support Needed

Opuntia in Big Bend National Park
Photo by Cookie Ballou,
National Park Service

To date, no organized constituency has advocated for protection of our cacti from non-native insect pests. Perhaps now that the Cactoblastis moth is in Texas, the threat it represents to our desert ecosystems will become real to conservationists and they will join the struggle. The first step is to resolve the funding crisis so that the agencies can complete testing of the biocontrol agent and gain approval for its release. So now there is “something people can do” – and I hope they will step forward.

I hope Americans are not actually indifferent to the threat that many cacti in our deserts will be killed by non-native insects. Many are key components of the ecosystems within premier National Parks, and other protected areas. Cacti also are beautiful treasures in botanical gardens. I hope conservationists will agree that these threats must be countered, and will help to ensure funding of the final stages of the biocontrol tests.

Sources

Mengoni Goñalons, C., L. Varone, G. Logarzo, M. Guala, M. Rodriguero, S.D. Hight, and J.E. Carpenter. 2014. Geographical range & lab studies on Apanteles opuntiarum (hymenoptera: braconiDae) in AR, a candidate for BC of Cactoblastis cactorum (Lepidoptera: Pyralidae) in North America. Florida Entomologist 97(4) December 2014

Simonson, S.E., T. J. Stohlgren, L. Tyler, W. Gregg, R. Muir, and L. Garrett. 2005. Preliminary assessment of the potential impacts and risks of the invasive cactus moth, Cactoblastis cactorum Berg, in the U.S. and Mexico. Final Report to the International Atomic Energy Agency, April 25, 2005 © IAEA 2005

Srivastava, M., P. Srivastava,  R. Karan, A. Jeyaprakash, L. Whilby, E. Rohrig, A.C. Howe,  S.D. Hight, and L. Varone. 2019. Molecular detection method developed to track the koinobiont larval parasitoid Apanteles opuntiarum (Hymenoptera: Braconidae) imported from Argentina to control Cactoblastis cactorum (Lepidoptera: Pyralidae). Florida Entomologist 102(2): 329-335.

Varone, L., C.M. Goñalons, A.C. Faltlhauser, M.E. Guala, D. Wolaver, M. Srivastava, and S.D. Hight. 2020. Effect of rearing Cactoblastis cactorum on an artificial diet on the behavior of Apanteles opuntiarum. Applied Entomology DOI: 10.1111/jen.12731.

Varone, L., G. Logarzo, J.J. Martínez, F. Navarro, J.E. Carpenter, and S.D. Hight. 2015. Field host range of Apanteles opuntiarum (Hymenoptera: Braconidae) in Argentina, a potential biocontrol agent of Cactoblastis cactorum (Lepidoptera: Pyralidae) in North America. Florida Entomologist — Volume 98, No. 2 803

Varone, L., M.B. Aguirre, E. Lobos, D. Ruiz Pérez, S.D. Hight, F. Palottini, M. Guala, G.A. Logarzo. 2019. Causes of mortality at different stages of Cactoblastis cactorum in the native range. BioControl (2019) 64:249–261

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.

Growing Pathogen Threat from Southeast Asia – US Unprotected

APHIS can protect our native & agricultural plants – but will it?

Imports of large numbers of plants for planting from Southeast Asia represents a significant biosecurity risk for forestry, horticulture, and natural ecosystems in North America and Europe.  This threat is likely to grow unless APHIS takes action under its emergency authorities.

Recent pest introductions and related studies indicate that Southeast Asia is a newly-discovered center of origin for plant pathogens. Places of particular concern are Vietnam, southern Yunnan Province and Hainan Island of China, northern Laos, the eastern Himalayas, and Taiwan. Significant pathogens and associated insects apparently centered in these areas include the sudden oak death pathogen (Phytophthora ramorum) and other Phytophthora species; and several ambrosia beetles and associated fungi, including the laurel wilt fungus (Raffaelea lauricola) and its primary vector (Xyleborus glabratus), and the polyphagous (Euwallacea whitfordiaodendrus) and Kuroshio shot hole borers (Euwallacea kuroshio).

Southeast Asia is attractive to the plant trade because of the region’s high floral diversity, including such sought-after families as Ericaceae (rhododendrons). Indochina has more than 10,350 vascular plant species in 2,256 genera – equaling more than 20% of the world’s plant species (Jung et al. 2019).

Pathogens are notoriously difficult to detect during inspections at the time of shipment. One-time inspections of high volume imports are especially weak and prone to failure.

How do we protect America’s flora?

APHIS could — but has not yet — developed requirements that these countries institute integrated pest management procedures for their exporting nurseries – as provided under amendments to APHIS’ Q-37 regulation and ISPM#36. In any case, it is unlikely that such procedures would minimize the risk because many of the plants that would be imported would probably be wild-collected.

APHIS has – and should use – far more effective means to minimize risk. These are the Federal orders and listing process known as “not authorized for importation pending pest risk assessment” or NAPPRA. If – despite the scientific evidence – APHIS continues to allow high volumes of dangerous imports, the agency should immediately institute new phytosanitary controls to its inspection process. These include relying on risk-based inspection regimes and molecular high-through-put detection tools.

Supporting Material

SOD-killed tanoaks in Big Sur; photo provided by Matteo Garbelotto, UC Berkeley

Phytophthora species

A team of European pathologists, led by Thomas Jung and including Clive Brasier and Joan Webber (see full citation at the end of this blog) surveyed Phytophthora species by sampling rhizosphere soils in 25 natural and semi-natural forest stands, isolations from naturally fallen leaves, and waters in 16 rivers in temperate and subtropical montane and lowland regions of Vietnam during 2016 and 2017.

These studies detected 13 described Phytophthora species, five informally designated taxa, and 21 previously unknown taxa. Detections were made from soil samples taken from 84% of the forest stands and from all rivers.

As I reported in am earlier blog, P. ramorum and P. cinnamomi were among those species detected. Both the A1 and A2 mating types of both P. ramorum and P. cinnamomi co-occurred.

The survey also detected at least 15 species in other genera of oomycetes.

The scientists conclude that most of the 35 forest Phytophthora species detected are native to Vietnam or nearby surrounding areas, attributing species in Phytophthora clades (taxonomically related groups) 2, 5, 6, 7, 8, 9, and 10 as native to Indochina. Different clades were detected in high-elevation vs. lowland rivers, cooler (subtropical) vs. tropical streams, and in soils vs. streams. Given the relatively limited number and diversity of the sampled sites and ecosystem types, it is likely that the true Phytophthora diversity of Vietnam is markedly higher (Jung et al. 2019)  

Worrying diversity of Phytophthora has been detected in other areas of Southeast Asia. A 2013 survey in natural forests and streams of Taiwan detected 10 described species and 17 previously unknown taxa of which 9 were of hybrid origin. In three areas in northern Yunnan, a Chinese province adjacent to northern Vietnam, eight Phytophthora species were isolated from streams running through sclerophyllous oak forests; two were recovered from forest soil samples. In montane forests of the tropical island Hainan, located in the South China Sea close to Vietnam, six Phytophthora species were found (Jung et al. 2019).

These studies are being conducted in the context of scientists discovering numerous new species of Phytophthora in recent decades. Since 1999, the number of described species and informally designated taxa of Phytophthora has tripled. World-renowned experts Clive Brasier anticipates that between 200 and 600 species of Phytophthora are extant in natural ecosystems around the world (Jung et al. 2019).

In the Vietnam survey, P. ramorum was the most widespread species. While genetic studies indicate ancestral connections to the four P. ramorum lineages (genetic strains) introduced to North America or Europe, further studies are under way to clarify these relationships (Jung et al. 2019).

Jung and colleagues found P. cinnamomi to be the most common soilborne Phytophthora species at elevations above 700 m. Two genotypes of the P. cinnamomi A2 mating type are causing epidemics in numerous natural and managed ecosystems worldwide. There was some evidence that the more frost sensitive A2 mating type might be spreading into higher altitudes in Vietnam (Jung et al. 2019).

Most of the Phytophthora species detected in the rhizosphere were not associated with obvious disease symptoms. (The principal exception was the A2 mating type of P. cinnamomi in montane forests in northern Vietnam.) (Jung et al. 2019) This lack of disease greatly reduces the chances of detecting the oomycetes associated with any plants exported from the region – there are no symptoms.

Since southern Yunnan, northern Laos, and the eastern Himalayas belong to the same biogeographic area those areas might also harbor endemic P. ramorum populations. Further surveys are needed to confirm this hypothesis (Jung et al. 2019).

Phytophthora lateralis – causal agent of Port-Orford cedar root rot – also probably originated in the area, specifically Taiwan (Vettraino et al. 2017).

Implications for phytosanitary measures

Many of the native Asian forest Phytophthora species have co-evolved with a variety of tree genera also present in Europe and North America, including Fagaceae, Lauraceae, Aceraceae, Oleaceae, and Pinaceae. Numerous examples demonstrate a strong potential that trees in these families that have not previously been exposed to these Phytophthora species might be highly susceptible. Scientists have begun an extensive host range study of Phytophthora species from Asia and South and Central America. One part of this study found that five Asian Phytophthora species caused significant rot and loss of fine roots and lateral roots in three European species of chestnut and oak (Jung et al. 2019).

Other pathogens

Studies by separate groups of scientists have concluded that several beetle-fungus disease complexes are native to this same region.

Sassafras – photo by David Moynihan

Both the laurel wilt fungus Raffaelea lauricola and its primary vector Xyleborus glabratus probably originated in Southeast Asia; there are probably different strains or genetic makeups across their wide ranges. For example, Dreaden et al. 2019 found that the fungus population from Myanmar differed genetically from those found in Japan, Taiwan, and the United States. Others had already expressed concern about the possibility that new strains of R. lauricola might be introduced (Wuest et al. 2017, cited in Cognato et al. 2019).

Cognato et al. 2019 found that the beetle occurs in deciduous forests from southern Japan to Northeast India, so genetic variation across this range is likely. In fact, they have separated the species X. glabratus into three species. They found that some of the beetles might thrive at 40o North – the latitude of central Illinois, Indiana, and Ohio and southern Pennsylvania. The ability of the vector of laurel wilt disease to spread so far north poses an alarming threat to sassafras (Sassafras albidum) – which is a major understory tree in forests of these regions.

It is unknown whether these new species and X. glabratus lineages are associated with different fungal strains. In company with the pathologists cited above, Cognato et al. 2019 warn that preventing introduction of the three beetle species to other regions is prudent. Cognato et al. 2019 point out that if other beetle lineages from the southern extent of their range can tolerate hotter and drier conditions, they might pose a greater risk to host species in the more arid areas of California and Mexico. In addition, Central America is at great risk because of the numerous plant species in the vulnerable Lauraceae found there.

Also from the region are two beetle-fungus combinations killing trees in at least seven botanical families, including maples, oaks, and willows, in southern California. The polyphagous shot hole borer (Euwallacea whitfordiaodendrus) apparently is native to Vietnam (Eskalen et al. 2013) and the closely related Kuroshio shot hole borer (Euwallacea kuroshio) to Japan, Indonesia, and Taiwan (Gomez et al. 2018).  

What you can do

Getting APHIS to act

1) communicate concern about the risk to APHIS leadership and ask that the agency take action under its NAPPRA authority

2) communicate the same to intermediaries who can influence APHIS:

  • State phytosanitary agency – especially through regional plant boards and National Plant Board
  • Your Congressional representative and senators (especially if one or more serves on Agriculture or Appropriations committee)
  • Professional societies – American Phytophathological Society, Mycological Society, American Society of Entomologists, Society of American Foresters …

3) communicate the same to university leadership and ask that their lobbyists advocate to USDA

4) communicate the same to the media

2) Research on extent of North American tree species’ vulnerability to the Oomycetes and other associated microorganisms

Jung et al. 2019 say that studies are under way to identify potential pest-host relationships with important tree species. However, all the authors are Europeans. Is anyone carrying out tests on North American trees in the apparently most vulnerable families — Fagaceae, Lauraceae, Aceraceae, Oleaceae, and Pinaceae?

1) Communicate with colleagues, scientific societies, APHIS, Agriculture Research Service, National Institute of Food and Agriculture, and USFS to determine whether such tests are under way or planned.

2) In those cases where no studies are planned, work with above to initiate them.

Sources

Cognato, A.I., SM. Smith, Y. Li, T.H. Pham, and J. Hulcr. 2019. Genetic Variability Among Xyleborus glabratus Populations Native to Southeast Asia (Coleoptera: Curculionidae: Scolytinae: Xyleborini) and the Description of Two Related Species. Journal of Economic Entomology XX(XX), 2091, 1 – 11.

Dreaden, T.J., M.A. Hughes, R.C. Ploetz, A. Black and J.A. Smith. 2019. Genetic Analyses of the Laurel wilt Pathogen, Raffaelea lauricola, in Asia Provide Clues on the Source of the Clone that is Responsible for the Current USA Epidemic. Forests 2019, 10, 37

Eskalen, A., Stouthamer, R. Lynch, S.C., Twizeyimana, M., Gonzalez, A., and Thibault, T. 2013. Host range of Fusarium dieback and its ambrosia beetle (Coleoptera Scolytinae) vector in southern California. Plant Disease 97938-951.

Gomez, D.F., J. Skelton, M.S. Steininger, R. Stouthamer, P. Rugman-Jones, W. Sittichaya, R.J. Rabaglia, and J. Hulcr1/ 2018. Species Delineation Within the Euwallacea fornicatus (Coleoptera: Curculionidae) Complex Revealed by Morphometric and Phylogenetic Analyses. Insect Systematics and Diversity, (2018) 2(6): 2; 1–11

Jung, T., B. Scanu, C.M. Brasier, J. Webber, I. Milenkovic, T. Corcobado, M. Tomšovský, M. Pánek, J. Bakonyi, C. Maia, A. Baccová, M. Raco, H. Rees, A. Pérez-Sierra & M. Horta Jung. 2020. A Survey in Natural Forest Ecosystems of Vietnam Reveals High Diversity of both New and Described Phytophthora Taxa including P. ramorum. Forests, 2020, 11, 93   https://gcc02.safelinks.protection.outlook.com/?url=https%3A%2F%2Fwww.mdpi.com%2F1999-4907%2F11%2F1%2F93%2Fpdf&data=02%7C01%7C%7Cfcd843919a3348a4a56108d7974039ab%7Ced5b36e701ee4ebc867ee03cfa0d4697%7C0%7C1%7C637144174418121741&sdata=WayrZsxp3P9Kj0h1aDPZnzu4yjDGA2ZEuH9NZITFQF4%3D&reserved=

Vettraino,  A.M., C.M. Brasier, J.F. Webber, E.M. Hansen, S. Green, C.Robin, A. Tomassini, N. Bruni, A. Vannini. 2017. Contrasting microsatellite diversity in the evolutionary lineages of Phytophthora lateralis. Fungal Biology Vol. 121, Issue 2, February 2017, pp. 112-126

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.

Solutions Suggested by 30 Years’ Work

Faith Campbell receives award for activism from National Association of State Foresters; 2016

For nearly 30 years I have documented bioinvasion threats and gaps, first in three Fading Forests reports (available here), then in five years of blogging. Here I pull together that information and suggest — in most cases reiterate — steps to address these threats and gaps. I list sources of discussion of the underlying issues – other than my reports and blogs – in references at the end of this blog.

My first premise is: robust federal leadership is crucial:

  1. The Constitution gives primacy to federal agencies in managing imports and interstate trade.
  2. Only a consistent approach can protect trees (and other plants) from non-native pests.
  3. Federal agencies have more resources than state agencies individually or in any likely collective effort — despite decades of budget and staffing cuts.

My second premise is: 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 not been adequate or stable.

My third premise is that programs’ effectiveness needs to be measured, not just effort (see the NECIS document referenced at the end of the blog).

SPECIFICS 

Preventing new introductions continues to be the most effective action. Mitigating options decrease and damages increase once a non-native pest has entered the country – much less become established (see Lovett et al. 2016 and Roy et al. 2014). I recognize that preventing new introductions poses an extremely difficult challenge given the volume and speed of international trade and the strong economic forces supporting free trade. These challenges have been exacerbated over several decades by the political zeitgeist – the anti-regulatory ideology, the emphasis on “collaborating” with “clients” rather than imposing requirements through regulations. Although the current “America First” policy might reduce import volumes and therefore reduce the invasive species threat to some extent, the anti-regulatory stance has only strengthened.

containers at the Port of Long Beach, California

Decades of cutting key agencies’ budgets and personnel are another factor. However, the damage to America’s natural systems is so great that we must try harder to find more effective strategies (See the Fading Forest reports; my previous blogs; Lovett et al. 2016; and APHIS annual reports – e.g., the 2019 report here)

Prevention

  1. Despite adoption and implementation of new international and national regulations to stem pest introductions, introductions continue – although probably at a lower level than would otherwise be the case. Delays in adoption of regulations (documented in Fading Forests II and III and my two recent 30-years-in-review blogs have facilitated damaging introductions and spread.

Solutions 

  1. Stakeholders press USDA leadership to initiate rules intended to strengthen phytosanitary protection and expedite their completion
  2. APHIS promote and facilitate analysis of current programs and policies by non-agency experts to ensure the agency is applying most effective strategies (Lovett et al. 2016).
  • Adoption of insufficiently protective regulations (documented in FFII, FFIII, two 30-years-in-review blogs) – adopted in part because APHIS is trying to “balance” trade facilitation and phytosanitary protection – has further contributed to damaging pests’ introduction and spread.

Solutions:

  1. Boost priority of preventing pest introductions by amending the Congressional finding in the Plant Protection Act [7 USC 7701(3)] as follows

Existing language: “[I]t is the responsibility of the Secretary [of Agriculture] to facilitate exports, imports and interstate commerce in . . . commodities that pose a risk of harboring plant pests or noxious weeds in ways that will reduce, to the extent practicable, as determined by the Secretary, the risk of dissemination of plant pests and noxious weeds .… “

Amend to read as follows: “…. in ways that will reduce prevent, to the greatest extent practicable feasible, as determined by the Secretary, …” [emphasis added]

  • Adopt several actions to strengthen phytosanitary protections at the point of origin (Lovett et al. 2016)
  • Expand pre-clearance partnerships — as authorized for plants under Q-37 regulations and ISPM-36
  • Expand sentinel tree programs
  • Promote voluntary substitution of packaging made from materials other than solid wood.
  • APHIS doesn’t use the enforcement powers that it has under Plant Protection Act (see several of my past blogs)

Solutions:

CBP inspectors search for pests in a pallet; CBP photo
  1. APHIS follow the lead of Customs and Border Protection and begin penalizing importers on the first instance of their wood packaging not being in compliance with ISPM#15 (see blog here).
  2. APHIS prohibit use of wood packaging by countries and importers of categories of imports that – over the 13 years since implementation – have developed a record of frequent violations of ISPM#15.
  3. APHIS use its authority per revised Q-37 regulations to negotiate with countries that export plants to the U.S. to establish “integrated measures” programs aimed at minimizing the risk of associated pests being transported to the U.S.
  4. APHIS use its authority per revised Q-37 to place in the “Not Authorized for Import Pending Pest Risk Assessment (NAPPRA) “limbo” category genera containing North American “woody” plants  (see Roy et al. 2014; Lovett et al. 2016).

Spread within the U.S.

  1. The United States lacks a coordinated system to prevent pest spread within the country (see Fading Forests III Chapter 5). Even our strictest methods, like APHIS’s quarantines regulating interstate movement of goods, have failed to curtail spread of significant pests. The most obvious example is the emerald ash borer.

The regulations governing movement of the sudden oak death pathogen in the nursery trade have also failed: there have been periodic outbreaks in which the pathogen has been spread to nurseries across the country. Between 2003 and 2011, a total of 464 nurseries located in 27 states tested positive for the pathogen, the majority as a result of shipments traced from infested wholesalers. In 2019, plants exposed to the pathogen were again shipped to 18 states; eight of those states have confirmed that their plant retailers received infected plants (see my blog from summer here).

Another serious gap is the frequent failure of APHIS and states to adopt official programs targetting bioinvaders that will be difficult to control because of biological characteristics or cryptic natures – even when severe impacts are demonstrated. Recent examples include the laurel wilt disease complex, goldspotted oak borer, polyphagous and Kuroshio shot hole borers and associated pathogens, and even the spotted lanternfly (although the last has received significant funds from APHIS.)

redbay killed by laurel wilt disease, Georgia; photo by Scott Cameron

Solutions:

  1. APHIS apply much more stringent regulations to interstate movement, based on a heightened priority for prevention in contrast to facilitating interstate trade. E.g., prohibit nurseries on the West Coast from shipping P. ramorum hosts to states where the pathogen is not established.
  2. APHIS encourage states to adopt quarantines and regulations aimed at preventing spread of invasive pests to regions of the state that are not yet infested. For example, the sudden oak death pathogen in California and Oregon; the borers in southern California.
  3. APHIS abandon plans to deregulate emerald ash borer and step up its support for state  regulations on firewood.
  4. APHIS stop dumping pests it no longer wants to regulate onto the states through the “Federally Recognized State Manage Phytosanitary (FRSMP) program”.
  5. APHIS revise its policies so that the “special needs exemption” [7 U.S.C. 7756] actually allows states to adopt more stringent regulations to prevent introduction of APHIS-designated quarantine pests (see Fading Forests III Chapter 3).

To help fill the gaps, the states are trying to coordinate their regulations in some important areas. The most advanced example is the voluntary Systems Approach to Nursery Certification, or SANC program. APHIS has supported this initiative, including by funding from the Plant Pest and Disease Management and Disaster Program (see below). However, it is a slow process; USDA funds first became available in 2010. The states are trying to coordinate on firewood, but we don’t yet know what the process will be.

  • Funding shortfalls (See the three Fading Forests reports, my blogs about appropriations)
  • Increase APHIS’ access to emergency funds from the Commodity Credit Corporation by  amending the Plant Protection Act [7 U.S.C. 7772 (a)] to include this new definition of “emergency”:

the term “emergency” means any outbreak of a plant pest or noxious weed which directly or indirectly threatens any segment of the agricultural production of the United States and for which the then available appropriated funds are determined by the Secretary to be insufficient to timely achieve the arrest, control, eradication, or prevention of the spread of such plant pest or noxious weed.

  • Although APHIS has the most robust prevention program of any federal agency, its funding is still inadequate. Stakeholders should lobby the Congress in support of higher annual appropriations.

The Plant Pest and Disease Management and Disaster Program (now under Section 7721 of the Plant Protection Act) has provided at least $77 million for tree-pest programs (excluding NORS-DUC & sentinel plant programs and other programs) since FY 2008. Much useful work has been carried out with these funds. However, these short-term grants cannot substitute for stable, long-term funding. I reiterate my call for stakeholders to lobby the Congress to provide larger appropriations to the APHIS Plant Protection program and Forest Service Forest Health Protection and Research programs.

Long-term Responses to Bioinvasive Challenge

More stakeholders are advocating raising the priority of – and providing adequate resources to – such long-term solutions as biocontrol and breeding trees resistant to pests and restoring them to our forests. Advocates include the state forestry agencies of the Northeast and Midwest, some non-governmental organizations, some academics, and individual USFS scientists. One effort resulted in inclusion of language in the 2018 Farm Bill (see blog here) – although this approach has apparently run into a dead end. The new emphasis on breeding has so far not been supported by agency or Congressional leaderships.

test planting of an American chestnut bred to be resistant to chestnut blight

Solutions:

  1. USFS convene workshop of the federal, state, National Academy, academic, and NGO groups promoting resistance breeding in order to develop consensus on priorities  and general structure of program.

Explicitly include evaluation of the CAPTURE Project’s (see blog here) efforts to set priorities to guide funding allocations and policies; and proposals for providing needed supportive infrastructure – facilities, trained staff in various disciplines. (See my blogs here.)

Report results of meeting to USDA leadership, Congress, and stakeholders

Then ensure implementation of the accepted approach by both Research and Development and Forest Health Protection programs. Include provisions to provide sustainable funding.

These proposed actions still do not address ways to correct the provisions of the international phytosanitary agreements (World Trade Organization and International Plant Protection Convention) that complicate – or preclude – efforts to prevent introduction of pests currently unknown to science. This  issue is discussed in Fading Forests II. A current example is beech leaf disease (described here).

Continuing inadequate engagement by stakeholders

Most constituencies that Americans expect to protect our forests don’t press decision-makers to fix the problems I have identified above: inadequate resources, weak and tardy phytosanitary measures. Some of these stakeholders are other federal agencies, or state agencies – or their staffs. They face restrictions on how “political” they can be. But where are the professional and scientific associations, representatives of the wood products industry, forest landowners, environmental NGOs and their funders, urban tree advocates Efforts by me, USDA, and others to better engage these groups have had disappointing results.

As I have documented, groups of USFS scientists have made several attempts to document the extent of invasive species threats and impacts and to set priorities. So far, they have not gained much traction. Another USFS attempt, Poland et al. in press, will appear at the end of the year. Will this be more successful?

I detect growing attention to educating citizen scientists for early detection; but if there is an inadequate – or no – official response to their efforts won’t people become discouraged?

SOURCES

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

National Environmental Coalition on Invasive Species “Tackling the Challenge.”

Poland, T.M., Patel-Weynand, T., Finch, D., Miniat, C. F., and Lopez, V. (Eds) (2019), Invasive Species in Forests and Grasslands of the United States: A Comprehensive Science Synthesis for the United States 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 P&Ps requires new trade regulations. Front Ecol Environ 2014; 12(8): 457–465

Thirty Years of Tree Pest Analysis, Continued – Risks of New Introductions

It is widely recognized that invasions of non-native species occur as a consequence of international trade (see Seebens et. al. 2017 – full citations at the end of this blog). This is as true for non-native forest pests as for any other bioinvader – see Aukema et al. 2010; Liebhold et al. 2012, Lovett et al. 2016. In fact, gross domestic product – as an indicator of levels of trade — is a better predictor of the number of forest pest invasions in a given country than the country’s amount of forested land (Roy et al. 2014).

shipping containers at port of Long Beach, California

As I noted in my previous blog, I began studying and writing about the threat to North America’s forests from non-native insects and pathogens in the early 1990s. I reported my analyses of the evolving threat in the three “Fading Forests” reports – coauthored by Scott Schlarbaum – in 1994, 2003, and 2014. These reports are available here.

I document here that both introduction and spread of pests within the country have continued apace. While significant efforts have been made to prevent introductions (described briefly under the “Invasives 101” tab of the CISP website), they have fallen short. As I noted in Fading Forests III, programs aimed at preventing spread of pests within the country remain fragmented and often are unsuccessful.

The Challenge: Huge Volumes of goods are moving, providing opportunities for pests

Since 1990, volumes of imported goods more than quintupled. Within the U.S., a total of 17,978 million tons of goods were transported in 2015; 10,776 million tons of this total by truck. About one-third of this total – 5,800 million tons – was moved farther than 250 miles. These vehicles moved on a public roads network of 4,154,727 miles (US DOT FFA). Consequently, once a pest enters the U.S., it can be moved quickly into every corner of the country.

Introductions

By and large, establishment of tree-killing pests has occurred at a fairly steady rate of about 2.5 per year, with “high-impact” insects and pathogens accumulating at 0.43 per year (Aukema et al. 2010). Since introductions did not rise commensurately with rising import volumes, Lovett et al. (2016) concluded that the recently adopted policies for preventing introductions referenced above are having positive effects but are insufficient to reduce the influx of pests in the face of ever-growing global trade volumes. The study’s authors went on to say that absent more effective policies, they expect the continued increase in trade will bring many new establishments of non-native forest pests.

One group of forest pests did not enter at a steady rate, but rather entered at a higher rate since 1985 – wood-boring insects. Experts concluded that the increase probably reflected increases in containerized shipping (Lovett et al. 2016). At the global level, the rate of fungal invasions has also recently been reported to be increasing rapidly (Roy et al. 2014).

Asian longhorned beetle

Geography of trade patterns also matters. Opening of trade with China (in 1979) offered opportunities for pests from a new source country which has a similar climate and biology. Roy et al. describe the importance of phylogenetic relatedness of pests and of tree hosts in explaining tree species’ vulnerability to introduced pests. The most vulnerable forests are those made up of species similar to those growing in the source of the traded goods – i.e., the temperate forests of the northeastern U.S. – when goods are imported from similar forested areas of Europe and Asia. Chinese-origin wood-boring pests began to be detected around 1990. This already short interval probably underestimates how quickly pests began arriving; detection methods were poor in those years, so a pest was often present for close to a decade before detection.

Between 1980 and 2016, at least 30 non-native species of wood- or bark-boring insects in the Scolytinae / Scolytidae were newly detected in the United States (Haack and Rabaglia 2013; Rabaglia et al. 2019).  Over the same period, approximately 20 additional tree pests were introduced to the continental states (Wu et al. 2017; Digirolomo et al. 2019; R. Haack, pers. comm.) plus about seven to America’s Pacific islands. Not all of the new species are highly damaging, but enough are. See my previous blog here.

Many of the tree-killing pests were probably associated with pathways other than wood packaging. These include 6 of the 7 Agrilus species, sudden oak death pathogen, three pests of palm trees, the spotted lanternfly, beech leaf disease; and the pests introduced to America’s Pacific Islands.

HIGH-RISK PATHWAYS OF INTRODUCTION

Already in the 1990’s it was evident that better preventing pest introductions would depend on shutting down the variety of pathways by which they move around the world.   At that time, attention focused on imports of logs and nursery stock (nursery stock makes up one component of a broader category called by phytosanitary agencies “plants for planting”). Both logs and “plants for planting” had well-established histories of transporting pests and import volumes were expected to grow. We have since learned that there are many more pathways!

Plants for Planting

Imports of “plants for planting” (phytosanitary agencies’ term, which encompasses nursery stock, roots, bulbs, seeds, and other plant parts that can be planted) have long been recognized as a dangerous pathway for introduction of forest pests. For example, this risk was the rationale for adopting the 1912 Plant Quarantine Act. Charles Marlatt, Chairman of USDA’s Federal Horticultural Board (see “Then and Now” in Fading Forests III here), wrote about the risk in National Geographic in April 1911 (urging adoption of the 1912 law) and again in August 1921. See also Brasier (2008), Roy et al. (2014), Liebhold et al. (2012), Jung et al. (2016).

Japanese cherry trees being burned because of scale infestation
January 28, 1910; Agriculture Research Service

Of the 91 most damaging non-native forest pest species in the U.S. (Guo et al. 2019), about 62% are thought to have entered North America with imports of live plants. These include nearly all the sap-feeding insects, almost 90% of the foliage-feeding insects, and approximately half of the pathogens introduced during the period 1860-2006 (Liebhold et al. 2012). Specific examples include chestnut blight, white pine blister rust, Port-Orford-cedar root disease, balsam woolly adelgid, hemlock woolly adelgid, beech scale, butternut canker, dogwood anthracnose, and sudden oak death. In more recent years, introductions via this pathway possibly include ‘ōhi‘a rust, rapid ‘ōhi‘a death pathogens, and beech leaf disease. The gypsy moth, while a foliage feeder, was not introduced via imports of live plants.

The APHIS annual report for 2018 reported that in that year we imported 18,502 shipments containing  more than 1.7 billion plant units (plants, bulbs, in vitro materials, etc.).

Liebhold et al. 2012, relying on 2009 data, found that about 12 percent of incoming plant shipments had symptoms of pests – a rate more than 100 times greater than that for wood packaging. Worse, a high percentage of the pests associated with a shipment of plants is not detected by the federal inspectors. The meaning of this finding is unclear because the study did not include any plant genera native to temperate North America and APHIS points out that infestation rates varied considerably among genera in the study. However, APHIS has not conducted its own analysis to document the “slippage rate” on imports of greatest concern to forest conservationists, i.e., imports of woody plants. I provide details on pests detected on imports of woody plants in recent in my blog here.

Clearly the risk of pest introductions continued at least until recently. I reviewed an APHIS database listing pests newly detected in the country during the period 2009-2013. I concluded that approximately 37 of the 90 “new” pests listed in the database (viruses, fungi, aphids and scales, whiteflies, mites) were probably introduced via imports of plants, cuttings, or cut foliage or flowers. I discussed these matters in greater detail here.

Adoption of a new regulatory regime governing imported plants for planting  (Q-37 regulation) in 2018 is too recent to for us to see its impact. But the new regulation sets up a process under which APHIS can impose more protective regulations on specific types of plants or plants from certain countries of origin to counter a perceived concerning level of risk. Until APHIS begins activating its new powers by negotiating more protective regulations governing plant imports from high-risk sources, it seems unlikely there will be any meaningful change in the introduction rates.

Crates, Pallets, and Other Forms of wood packaging (solid wood packaging, or SWPM)

Recognition of the risk associated with wood packaging is much more recent. In 1982, a USDA risk assessment concluded that the wood boring insects found in crates and pallets were not of great concern (USDA APHIS and Forest Service, 2000). However, contradictory indications were quickly documented – including from APHIS’ own port interception data – which the agency began collecting in 1985. Over the 16-year period 1985-2000, 72% of the 6,825 bark beetles (Scolytidae) intercepted by APHIS were found on SWPM (Haack 2002). Cerambycids (longhorned beetles) and buprestids (jewel beetles) make up nearly 30% of insects detected in wood packaging over the last 30 years (Haack et al. 2014).

Detection of outbreaks of the Asian longhorned beetle and other woodborers in the mid-1990s made it clear that wood packaging was, indeed, a high-risk pathway.

Of the 91 most damaging non-native pest species in the US, 30% probably arrived with wood packaging material or other wood products (Liebhold et al. 2012). This group includes many of the most damaging pests, the deadly woodborers – Asian longhorned beetle, emerald ash borer, redbay ambrosia beetle,  possibly the polyphagous and Kuroshio shot hole borers.

CBP agents inspecting a pallet

As noted above, introductions of wood borers have risen in recent decades, widely accepted as associated with the rapid increase in containerized shipping after 1980. In 2009 it was estimated that 75% of maritime shipments were packaged in crates or pallets made of wood (Meissner et al. 2009). A good history of the global adoption of containerized shipping is Levinson, M. The Box: How the Shipping Container Made the World Smaller and the World Economy Bigger (Princeton University Press 2008)

The simultaneous opening of trade with China (in 1979) offered opportunities for pests from a new source country which has a similar climate and biology. Chinese-origin wood-boring pests began to be detected around 1990. This already short interval probably underestimates how quickly pests began arriving; detection methods were poor in those years, so a pest was often present for close to a decade before detection.

I have already documented numerous times that, despite the U.S.’ implementation of the International Standard of Phytosanitary Measures (ISPM) #15 in 2006, live quarantine pest woodborers continue to enter the U.S. in wood packaging. The best estimate is that 0.1% of wood packaging entering the United States is infested with wood-borers considered to be quarantine pests (Haack et al. 2014). More than 22 million shipping containers entered the U.S. via maritime trade in 2017 (US DoT). As noted, an estimated 75% of sea-borne containers include wood packaging. Applying the 0.1% estimate to these figures results in an estimate that as many as 17,650 containers per year (or 48 per day) transporting tree-killing insects enter the U.S.

Over a period of nine years – Fiscal Years 2010 through 2018 – U.S. Customs and Border Protection (CBP) detected more than 28,600 shipments with wood packaging that did not comply with ISPM#15 (Harriger presentations to the annual meetings of the Continental Dialogue on Non-Native Forest Insects and Diseases). While most of the non-compliant shipments were wood packaging that lacked the required mark showing treatment per ISPM#15, in 9,500 cases the wood packaging actually harbored a pest in a regulated taxonomic group.

Disturbingly, 97% of the shipments that U.S. CBP found with infested wood packaging bear the ISPM#15 mark certifying that wood had been fumigated or heat-treated (Harriger 2017). CBP inspectors tend to blame this on widespread fraud in use of the mark. On the other hand, one study found that larvae can survive both treatments – although the frequency of survival was not determined. It was documented that twice as many larvae reared from wood treated by methyl bromide fumigation survived to adulthood than larvae reared from heat-treated wood; the reason is unclear (Nadel et al. 2016).

The APHIS’ record of interceptions for the period FYs 2011 – 2016 contained 2,547 records for insect detections on wood packaging. The insects belonged to more than 20 families. Families with the highest numbers of detections were Cerambycids – 25% of total; Curculionidae – 23% (includes Dendroctonus, Ips, Orthotomicus, Scolytinae, Xyleborus, Euwallacea); Scolytidae – 17% (includes true weevils such as elm bark beetles); Buprestids – 11%; and Bostrichidae – 3%. Not all of the insects in these groups pose a threat to North American plant species.

One encouraging data point is that since 2010, there have been no detections of species of bark and ambrosia beetles new to North America in the traps deployed by the USDA Forest Service Early Detection and Rapid Response program (Rabaglia 2019). The 2014 recognition of the Kuroshio shothole borer apparently did not result from this trapping program.

There have been several changes in the wood packaging standard and its implementation by CBP since 2009, the year Haack et al. 2014 analyzed the “pest approach rate”. APHIS has not carried out a study to determine whether these recent changes have reduced the approach rate below Haack’s estimate of 0.01%.  Consequently, we do not know whether these changes have reduced the risk of pest introductions.

Other Pathways That Transport  Fewer Pests –  Some of Which Have High Impacts

Insects that attach egg masses to hard surfaces can be transported by ship superstructures, containers, and hardsided cargoes such as cars, steel beams, and stone. While relatively few species have been moved in this way, some have serious impacts. The principal examples are the gypsy moths from Asia, which feed on 500 species of plants (Gibbon 1992). 

The United States and Canada have a joint program – under the auspices of the North American Plant Protection Organization (see RSPM #33) aimed at preventing introduction of species of Asian gypsy moths. The NAPPO standard originally went into force in March 2012. Under its terms, ships leaving ports in those countries during gypsy moth flight season must be inspected and cleaned before starting their voyage.

Gypsy moth populations rise and fall periodically; it is much more likely that egg masses will be attached to ships during years of high moth population densities. These variations are seen in U.S. and Canadian detection reports – as reported here.

While most AGM detections are at West Coast ports, [here;  and here] the risk is not limited to that region. AGM have been detected at Wilmington, NC; Baltimore, MD; Charleston, SC; Savanna and Brunswick, GA; Jacksonville, FL; New Orleans, LA; Houston and Corpus Christi, TX; and even McAlester, OK.  

Nor is the risk limited to the ships themselves. In 2014, more than 500 Asian gypsy moth egg masses were found on four shipments of imported steel slabs arriving at ports on the Columbia River in Washington.

Between 1991 and 2014, AGM was detected and eradicated on at least 20 occasions in locations across the United States (USDA AGM pest alert). Additional outbreaks have been discovered and eradication efforts undertaken in more recent years.

A second example is the spotted lanternfly (SLF) (Lycorma delicatula), which was first detected in southeast Pennsylvania in autumn 2014. It is native to Asia; it is believed to have entered the country as egg masses on imported stone.

While SLF is clearly a pest of agriculture – especially grapes and tree fruits – its importance as a forest pest is still unclear. Many native forest trees appear to be hosts during the insect’s early stages, including maples, birches, hickories, dogwoods, beech, ash, walnuts, tulip tree, tupelo, sycamore, poplar, oaks, willows, sassafras, basswood, and elms. Adult lanternflies strongly prefer the widespread invasive species tree of heaven (Ailanthus altissima).

As of August 2019, SLF was established in parts of five states: Delaware, Maryland, New Jersey, Pennsylvania, and Virginia. It was detected as having spread to a 14th county in Pennsylvania; five new counties in New Jersey. APHIS is working with state departments of Agriculture in these states, as well as supporting surveys in New York, North Carolina, and West Virginia (USDA APHIS DA-2019-20, August 7, 2019). Apparently the detections of a few adults – alive or dead – in Connecticut and New York had not evolved into an outbreak. See description and map here.

Imports of logs – roundwood – seem inherently risky. Certainly Dutch elm disease was introduced via this pathway. However, there have been few pest introductions linked to this pathway in recent years, probably because we import most of our unprocessed lumber from Canada. (I provide considerable data on U.S. roundwood imports in Fading Forests III here.)

Decorative items and furniture made of unprocessed wood certainly have the potential to transport significant pests (USDA APHIS 2007). Examples include boxes and baskets; wood carvings; birdhouses; artificial Christmas trees or other plants; trellises; lawn furniture. To date, apparently, no high-impact pest has been introduced via this pathway, although pests intercepted on shipments have included Cerambycids from Asia, e.g., velvet longhorned beetle and here.

Alarmed by high numbers of infested shipments from China, APHIS first suspended imports of such items temporarily; then adopted a regulation (finalized in March 2012 – USDA APHIS 2012).

APHIS has not taken action to prevent introductions on such items imported from other countries – although the North American Plant Protection Action adopted a regional standard making the case for such action and outlining a risk-based approach (NAPPO RSPM#38).

Snails on Shipping Containers

Snails have been detected on shipping containers and wood packaging for decades. In 2015, APHIS stepped up its efforts to address this risk through bilateral negotiations with Italy and launching regional and international efforts to develop guidance for ensuring pest-free status of shipping containers (Wendy Beltz, APHIS, presentation to National Plant Board, 2018 annual meeting).

SPREAD WITHIN THE UNITED STATES

Major pathways for human-assisted spread of pests within the country are sales of plants for planting, movement of unprocessed wood – especially firewood, and hitchhiking on transport vehicles. Since most forest pests are not subject to federal quarantine, any regulatory programs aimed at preventing spread depend on cooperation among the 50 states. None of these pathways is regulated adequately to prevent pests’ spread. See Chapter 5 of Fading Forests III here.

And since neither federal nor state agencies do significant enforcement of existing regulations, preventing spread often depends upon pest awareness of, and voluntary compliance by, individuals and companies.

Even pests subject to a federal quarantine are not prevented from spreading. Plants exposed to the sudden oak death pathogen were shipped to 18 states in spring 2019.

SOD-infected rhododendron plant; Indiana Department of Natural Resources

A collaborative effort by the nursery industry, APHIS, and states (Systems Approach to Nursery Certification, or SANC) is striving to close gaps linked to the standard practice of inspecting plants at the time of shipping, but full implementation of this voluntary program is still years away.

Transport of firewood has been responsible for movement of pests both short distances, e.g., goldspotted oak borer in southern California; and long distances – e.g., emerald ash borer to Colorado. APHIS attempted to develop a certification program but the industry was unable to put one together (see Chapter 5 of Fading Forests III). Current federal and state regulations of firewood are tied to the emerald ash borer quarantine, which APHIS has proposed to terminate. Wood for turning and woodworking has also been linked to movement of pests, e.g., walnut twig beetle/thousand cankers disease from the west to Pennsylvania.

emerald ash borer

Truck transport of a variety of goods has transported European gypsy moths from the infested areas in the east to the west coast. Transport of stone probably moved spotted lanternfly from southeastern Pennsylvania to Winchester, Virginia.

SOURCES

Aukema, J.E., D.G. McCullough, B. Von Holle, A.M. Liebhold, K. Britton, & S.J. Frankel. 2010. Historical Accumulation of Nonindigenous Forest Pests in the Continental United States. Bioscience. December 2010 / Vol. 60 No. 11

Brasier, C.M. 2008. The biosecurity threat to the UK and global environment from international trade in plants.  Plant Pathology (2008) 57, 792-808

Bray, A.M., L.S. Bauer, T.M. Poland, R.A. Haack, A.I. Cognato, J.J. Smith. 2011. Genetic analysis of emerald ash borer (Agrilus planipennis Fairmaire) populations in Asia and North America. Biol. Invasions (2011) 13:2869-2887

Gibbon, A. 1992. “Asian Gypsy Moth Jumps Ship to United States.” Science. Vol. 235. January 31, 1992.

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

Haack R. A. and J.F. Cavey. 1997. Insects Intercepted on Wood Articles at United States Ports-of-Entry and Two Recent Introductions: Anoplophora glabripennis and Tomicus piniperda. In press in International forest insect workshop proceedings, 18 – 21 August 1997, Pucon, Chile. Corporacion National Forestal, Santiago, Chile.

Haack, R.A., F. Herard, J. Sun, J.J. Turgeon. 2010. Managing Invasive Populations of Asian Longhorned Beetle and Citrus Longhorned Beetle:A Worldwide Perspective. Annu. Rev. Entomol. 2010. 55:521-46.

Haack, R.A. and R.J. Rabaglia. 2013. Exotic Bark and Ambrosia Beetles in the USA: Potential and Current Invaders. CAB International 2013. Potential Invasive Pests of Agricultural Crops (ed. J. Peña) 

Haack R.A., Britton K.O., Brockerhoff, E.G., Cavey, J.F., Garrett., L.J., 2014. Effectiveness of the International Phytosanitary Standard ISPM No. 15 on Reducing Wood Borer Infestation Rates in Wood Packaging Material Entering the United States. PLoS ONE 9(5): e96611. doi:10.1371/journal.pone.0096611

Haack, R.A., F. H´erard, J. Sun, and J.J. Turgeon. 2010. Managing Invasive Populations of Asian Longhorned Beetle and Citrus Longhorned Beetle: A Worldwide Perspective. Annu. Rev. Entomol. 2010. 55:521–46

Harriger, K. Department of Homeland Security Bureau of Customs and Border Protection, presentation to the Continental Dialogue on Non-Native Forest Insects and Diseases, November 2017.

Jung T, Orlikowski  L, Henricot B, et al. 2016. Widespread Phytophthora infestations in European nurseries put forest, semi-natural and horticultural ecosystems at high risk of Phytophthora diseases. Forest Pathology 46: 134–163.

Liebhold, A.M., E.G. Brockerhoff,  L.J. Garrett, J.L.Parke, and K.O Britton. 2012. Live plant inports: the major pathway for forest insect and  pathogen invasions of the US. Frontiers in Ecology.

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. Ayers, C.D. Canham, D.R. Foster, S.L. LaDeau, and T. Weldy. 2016.  Non-native forest insects and pathogens in the United States: Impacts and  policy options. Eological Applications, 26(5) pp. 1437-1455.

Meissner, H., A. Lemay, C. Bertone, K. Schwartzburg, L. Ferguson, and 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 Plant Epidemiology and Risk Analysis Laboratory (PERAL) / CPHST. June 4, 2009

Morin, R. presentation at Northeastern Forest Pest Council 81st Annual Meeting, March 12 – 14, 2019,  West Chester, Pennsylvania

Nadel, N., S. Myers, J. Molongoski, Y. Wu, S. Linafelter, A. Ray S. Krishnankutty, and A. Taylor. 2016. Identificantion of Port Interceptions in Wood Packaging Material: Cumulative Progress Report, April 2012 – August 2016

North American Plant Protection Organization Regional Standard for Phytosanitary Measures#24 https://www.nappo.org/english/products/regional-standards/regional-phytosanitary-standards-rspms/rspm-24/

North American Plant Protection Organization Regional Standard for Phytosanitary Measures#38 https://www.nappo.org/english/products/regional-standards/regional-phytosanitary-standards-rspms/rspm-38/

Rabaglia, R.J., A.I. Cognato, E. R. Hoebeke, C.W. Johnson, J.R. LaBonte, M.E. Carter, and J.J. Vlach. 2019. Early Detection and Rapid Response. A Ten-Year Summary of the USDA Forest Service Program of Surveillance for Non-Native Bark and Ambrosia Beetles. American Entomologist Volume 65, Number 1 

Roy, B.A., H.M. Alexander, J. Davidson, F.T. Campbell, J.J. Burdon, R. Sniezko, and C. Brasier.  2014. Frontiers in Ecology 12(8): 457-465

Seebens et. al. 2017. No saturation in the accumulation of alien species world wide. Nature Communications. January 2017 http://www.nature.com/articles/ncomms14435

U.N. Food and Agriculture Organization International Plant Protection Convention. 2012. International Standards for Phytosanitary Meaures No. 36 Integrated Measures for Plants for planting. Rome. Online at https://www.ippc.int/  Accessed December 7, 2012.

United States Department of Agriculture Animal and Plant Health Inspection Service and Forest Service, 2000. Pest Risk Assessment for Importation of Solid Wood Packing Materials into the United States.

United States Department of Agriculture Animal and Plant Health Inspection Service 2009. Proposed Rule Importation of plants for planting: establishing a category of plants for planting not authorized for importation pending pest risk assessment. Federal Register 74(140): 36403-36414 July 23, 2009.

United States Department of Agriculture Animal and Plant Health Inspection Service 7 CFR Parts 318, 319, 330, 340, 360, and 361. Federal Register Rules and Regulations Vol. 83, No. 53. Monday, March 19, 2018

United States Department of Agriculture, Animal and Plant Health Inspection Service. 2014. Asian gypsy moth pest alert https://www.aphis.usda.gov/publications/plant_health/content/printable_version/fs_phasiangm.pdf and pers. comm.

United States Department of Agriculture Animal and Plant Health Inspection Service 2007. Pests and mitigations for manufactured wood décor and craft products from China for importation into the United States. Revision 6. July.

United States Department of Agriculture Animal and Plant Health Inspection Service. 2012. Importation of wooden handicrafts from China. Final rule. Federal Register 77(41): 12437-12444. March 1. Online at http://www.gpo.gov/fdsys/pkg/FR-2012-03-01/pdf/2012-4962.pdf. Accessed August 2, 2013.

United States Department of Transportation Bureau of Transportation Statistics Freight Facts and Figures

https://www.bts.dot.gov/sites/bts.dot.gov/files/docs/FFF_2017.pdf  accessed 19/8/13

United States Department of Transportation, Maritime Administration, U.S. Waterborne Foreign Container Trade by U.S. Customs Ports (2000 – 2017) Imports in Twenty-Foot Equivalent Units (TEUs) – Loaded Containers Only

at https://ops.fhwa.dot.gov/freight/freight_analysis/nat_freight_stats/docs/06factsfigures/fig2_9.htm

Williams, L.H. and J.P. La Fage. 1979. Quarantine of Insects Infesting Wood in International Commerce. in J.A. Rudinksy, ed. Forest Insect Survey and Control Fourth Edition 1979

Wu,Y., N.F. Trepanowski, J.J. Molongoski, P.F. Reagel, S.W. Lingafelter, H. Nadel1, S.W. Myers & A.M. Ray. 2017. Identification of wood-boring beetles (Cerambycidae and Buprestidae) intercepted in trade-associated solid wood packaging material using DNA barcoding and morphology  Scientific Reports 7:40316

30 years of Analyzing Forest Pest Issues

dead whitebark pine in Crater Lake National Park
photo by F.T. Campbell

I began studying and writing about the threat to North America’s forests from non-native insects and pathogens in the early 1990s – nearly 30 years ago. I reported my analyses of the evolving threat in the three “Fading Forests” reports – coauthored by Scott Schlarbaum – in 1994, 2003, and 2014. These reports are available here.

So what has changed over those 30 years? What remains the same? Why have both the changes and the stasis occurred? What can we do to fix the gaps, close unaddressed pathways, strengthen flabby policies? I will address these issues in this and following blogs.

experimental American chestnut planted in Fairfax County, VA
photo by F.T. Campbell

What has changed since the early 1990s:

  • Adoption and implementation of significant new international and national regulations and programs aimed at preventing introductions of non-native invasive species.
  • Despite the welter of new regulations, an alarming increase in numbers of highly damaging forest pests established in the country.  (By my count, about 50 new species have established on the continent, six on Pacific islands; see details below.)
  • Alarming spread of established pests to new geographic regions and new hosts (e.g., emerald ash borer in 35 states and 5 provinces; laurel wilt disease across the range of redbay and swamp bay; rapid ‘ōhi‘a death on three of the main Hawaiian islands).
  • Introductions via unexpected pathways and vectors far removed from phytosanitary agencies’ usual targets, e.g., ship superstructures, imported steel and stone …

What has remained the same since the early 1990s:

  • Inadequate resources provided to response and recovery efforts.
  • Available funding focused on only a few of the more than 90 species causing damage.
  • Adoption of insufficiently protective regulations that have failed to prevent introduction and spread of tree-killing pests.
  • Lengthy delays in implementing programs that tighten controls – another factor in continuing introductions and spread.
  • Continued importance of expected pathways – nursery stock and raw wood, especially crates, pallets, and other forms of wood packaging.
  • Federal and state agencies still choose not to take action on pests e.g., goldspotted oak borer, polyphagous and Kuroshio shothole borers, beech leaf disease.
  • Inadequate coordination despite several efforts to set priorities.
  • Spurts of attention by media and political decision-makers, contrasted by lengthy periods of inattention.
  • Failure of most stakeholders to support efforts to prevent and respond to introductions of tree-killing pests. 

Details: The Situations Then and Now

(Many of the individual species mentioned here are described more fully here.  Full citations of sources are at the end of blog.)

American elm on the National Mall, Washington, D.C.

photo by USDA Agricultural Research Service

In 1993:

  • The number of non-native forest pest species established in the U.S. was estimated at between 300 (Millers et al. 1993) and 380 (Mattson et al., 1994; Liebhold et al., 1995) .
  • The area suffering the greatest numbers and impacts was the Northeast.
  • Several highly damaging pests that had been established for decades, including chestnut blight, white pine blister rust, Port-Orford-cedar root disease, Dutch elm disease, hemlock woolly adelgid, butternut canker, and dogwood anthracnose were receiving some attention but continued to spread.
  • USDA Forest Service funding for management of exotic pest infestations was crisis-oriented, with “… priorities … set under political pressures for immediate answers, with too much regard for short-term problems and too little consideration for broader management objectives.” (NAS 1975)
  • Since few high-profile pests had been introduced in recent years, APHIS was not actively engaged. In FY92, APHIS spent $20 million on efforts to eradicate the Asian gypsy moth. The narrow focus is illustrated by the fact that in FY93, more than two-thirds of all USDA tree pest control funds were devoted to efforts to suppress or eradicate the European gypsy moth (See FFI).
  • Concern about possible new introductions had grown; it focused on proposals to import unprocessed wood from Siberia, New Zealand, and Chile. The USDA Forest Service, academic scientists, and therefore APHIS emphasized the risks of known Asian pests, e.g., Asian gypsy moth, to western coniferous forests (See FFI). While individual scientists had expressed concern about wood packaging material, there was little public discussion of this threat.
  • We would learn later that several of the most damaging pests were already present in the country but not yet recognized – Asian longhorned beetle, sudden oak death pathogen, probably emerald ash borer.

beech leaf disease

photo by John Pogacnik

In 2019:

  • Numbers of non-native insects and pathogens attacking trees in North America approach 500 species.  (In Fading Forests III, I calculated that by the first decade of the 21st Century, the number had risen to at least 475. Several more have been detected since 2014. More than 181 exotic insects that feed on woody plants had established in Canada. (Source: USDA APHIS. 2000. Wood packaging risk assessment.)
  • Of these, 91 are considered “serious” threats (Guo et al. 2019). This estimate excludes pests native to portions of North America that are causing severe damage in naïve hosts – e.g., goldspotted oak borer; pests of palms; and pests attacking trees on U.S. Pacific and Caribbean islands.
  • Introductions had continued.
    • Between 1980 and 2016, at least 30 non-native species of wood- or bark-boring insects (Scolytinae / Scolytidae) were newly detected in the U.S. (Haack and Rabaglia 2013; Rabaglia et al.  2019). A few of these are highly damaging, e.g. redbay ambrosia beetle, polyphagous and Kuroshio shothole borers.
    • In addition to these 30 new pests, other highly damaging tree-killing pests probably introduced since the 1980s include (on the continent):
      • Eight Cerambycids such as Asian longhorned beetle (Wu et al. 2017)
      • 7 Agrilus, including emerald ash borer and soapberry borer; plus goldspotted oak borer transported from Arizona to California (Digirolomo et al. 2019; R. Haack, pers. comm.)
      • Sirex woodwasp
      • Pests of palm trees, e.g., red palm mite, red palm weevil, South American palm weevil
      • Spotted lanternfly
      • Beech leaf disease
    • Also not included in the above estimate and lists are tree-killing pests on America’s Pacific Islands :
      • ‘ōhi‘a rust
      • Cycad scale
      • Cycad blue betterfly
      • Erythrina gall wasp
      • two Ceratocystis pathogens that cause rapid ‘ōhi‘a death
      • Coconut rhinoceros beetle
    • Authorities also carried out approximately 25 eradication programs targetting introductions of the Asian gypsy moth (USDA Pest Alert Asian Gypsy Moth plus additional outbreaks since 2014).
  • Impacts of exacerbated tree mortality rates linked to these introduced pests are seen across wide swaths of the country, and affect widespread species, genera, and families. 
dead redbay in Claxton, Georgia
photo by Scott Cameron

I will discuss the risk of continuing new introductions in a separate blog.

Trying to Develop the Big Picture and Set Priorities

In recent years, USDA Forest Service scientists have made several attempts to provide nation-wide assessments of the impact of these pests and criteria for establishing priorities.

The National Insect and Disease Forest Risk Assessment predicted the loss of basal area to various pests over the 15-year time period 2012 – 2027. The assessment predicted the following losses for specific species: 90% for redbay; 60% for whitebark pine; more than 40% for limber pine; 24% for tanoak; 11% for coast live oak; 6% for eastern and Carolina hemlock; 27% for eight species of ash; 20% for American elm; 19% for red oak; 18% for American beech (Krist et al. 2014).

A separate group of scientists found that, nation-wide, non-native forest pests are causing an approximate 5% increase in total mortality by tree volume (Randy Morin at NEFPC). For details on Dr. Morin’s findings, see my blog here.

A third approach to developing a nation-wide picture, Project CAPTURE, (and my blog here) utilized FIA data to develop priorities for conservation action. Fifteen species were placed in the highest priority category, including Florida torreya (Torreya taxifolia), American chestnut and Allegheny and Ozark chinquapins, redbay, five species of ash, two species of hemlock, Port-Orford cedar, tanoak, and butternut (Potter et al. 2019(b).

According to Project CAPTURE, the non-native pests affecting the largest number of hosts are the European gypsy moth, which attacks 65 hosts; and oak wilt (Bretziella fagacearum), which infects 61 hosts. The Asian longhorned beetle attacks 43 hosts (Potter et al. 2019(b).

I note that several other non-native pests also have high numbers of host species. In the Project CAPTURE study, these pests are ranked lower because the project limited its evaluation to the five agents with the greatest effect on any particular host. Thus, of the 18 native tree species that host one or both of the invasive shothole borers and associated Fusarium disease complex (PSHB website), the project included only six. Of the 22 tree species listed by APHIS as hosts of Phytophtora ramorum, the project included 12 (K. Potter, pers. comm. April 17, 2019).

SOD-killed tanoak on the Big Sur peninsula, California
photo by Matteo Garbelotto, University of California Berkeley

More extensive discussions of non-native pests’ impacts are provided in Lovett et al. 2006, Lovett et al. 2016, and Potter et al. 2019. A book-length discussion of invasive species impacts – ranging from feral hogs to invasive plants, is expected in December; look for Poland et al. (in press).

SOURCES

Aukema, J.E., D.G. McCullough, B. Von Holle, A.M. Liebhold, K. Britton, & S.J. Frankel. 2010. Historical Accumulation of Nonindigenous Forest Pests in the Continental United States. Bioscience. December 2010 / Vol. 60 No. 11

Digirolomo, M.F., E. Jendek, V.V. Grebennikov, O. Nakladal. 2019. First North American record of an unnamed West Palaearctic Agrilus (Coleoptera: Buprestidae) infesting European beech (Fagus sylvatica) in New York City, USA. European Journal of Entomology. Eur. J. Entomol. 116: 244-252, 2019

Guo, Q., S. Fei, K.M. Potter, A.M. Liebhold, and J. Wenf. 2019. Tree diversity regulates forest pest invasion. Proceedings of the National Academy of Sciences of the United States of America. www.pnas.org/cgi/doi/10.1073/pnas.1821039116

Haack, R.A. and R.J. Rabaglia. 2013. Exotic Bark and Ambrosia Beetles in the USA: Potential and Current Invaders. CAB International 2013. Potential Invasive Pests of Agricultural Crops (ed. J. Peña) 

Krist, F.J. Jr., J.R. Ellenwood, M.E. Woods, A. J. McMahan, J.P. Cowardin, D.E. Ryerson, F.J. Sapio, M.O. Zweifler, S.A. Romero 2014. National Insect and Disease Forest Risk Assessment. United States Department of Agriculture Forest Service Forest Health Technology Enterprise Team FHTET-14-01

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

Liebhold, A. M., W. L. MacDonald, D. Bergdahl, and V. C. Mastro.  1995.  Invasion by exotic forest pests:  a threat to forest ecosystems.  Forest Sci., Monograph 30. 49 pp.

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

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

Mattson, W. J., P. Niemela, I. Millers, and Y. Ingauazo.  1994. Immigrant phytophagous insects on woody plants in the United States and Canada: an annotated list.  USDA For. Ser. Gen. Tech. Rep. NC-169, 27 pp.

Millers, I. United States Department of Agriculture, Forest Service Entomologist, Forest Health Protection Northeastern Area State and Private Forestry. Durham, NH. Personal communication to F.T. Campbell, 1993.

Morin, R. presentation at Northeastern Forest Pest Council 81st Annual Meeting, March 12 – 14, 2019,  West Chester, Pennsylvania

National Academy of Sciences. 1975. Forest Pest Control. Washington, D.C.

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

Polyphagous shothole borer website https://ucanr.edu/sites/pshb/overview/About_PSHB/

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

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 insect and disease threats, Global Ecology and Conservation (2019), doi: https://doi.org/10.1016/

Rabaglia, R.J., A.I. Cognato, E. R. Hoebeke, C.W. Johnson, J.R. LaBonte, M.E. Carter, and J.J. Vlach. 2019. Early Detection and Rapid Response. A Ten-Year Summary of the USDA Forest Service Program of Surveillance for Non-Native Bark and Ambrosia Beetles. American Entomologist Volume 65, Number 1 

USDA, Animal and Plant Health Inspection Service. 2014. Asian gypsy moth pest alert https://www.aphis.usda.gov/publications/plant_health/content/printable_version/fs_phasiangm.pdf and pers. comm.

U.S. Department of Agriculture, Animal and Plant Health Inspection Service.  2009.  Risk analysis for the movement of wood packaging material (WPM) from Canada into the US.

Wu,Y., N.F. Trepanowski, J.J. Molongoski, P.F. Reagel, S.W. Lingafelter, H. Nadel1, S.W. Myers & A.M. Ray. 2017. Identification of wood-boring beetles (Cerambycidae and Buprestidae) intercepted in trade-associated solid wood packaging material using DNA barcoding and morphology  Scientific Reports 7:40316

Collapse of Biodiversity – Causes and What We Can Do

frogs in California killed by chytrid fungus
photo by Rick Kyper, US Fish and Wildlife Service

I expect you have heard about the report issued on May 6 by the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services. The executive summary is available here

Based on thousands of scientific studies, the report concludes that the biosphere, upon which humanity as a whole depends, is being altered to an unparalleled degree across all spatial scales. The trends of decline are accelerating. As many as 1 million species (75% of which are insects) are threatened with extinction, many within decades.

Humans dominate Earth: natural ecosystems have declined by 47% on average. Especially hard-hit are inland waters and freshwater ecosystems: only 13% of the wetland present in 1700 remained by 2000. Losses have continued rapidly since then.

The report lists the most important direct drivers of biodiversity decline – in descending order – as habitat loss due to changes in land and sea use; direct exploitation of organisms; climate change; pollution; and invasive species. The relative importance of each driver varies across regions.

If you have been paying attention, these conclusions are not “news”.

However, the report serves two valuable purposes. First, it provides a global overview, a compilation of all the data and trends. Second, the report ties the direct drivers to underlying causes which are in turn underpinned by societal values and behaviors. Specifically mentioned are production and consumption patterns, human population dynamics and trends, trade, technological innovations, and governance (decision making at all levels, from local to global).

The report goes to great lengths to demonstrate that biological diversity and associated ecosystem services are vital for human existence and good quality of life – especially for supporting humanity’s ability to choose alternative approaches in the face of an uncertain future. The report concludes that while more food, energy and materials than ever before are now being supplied to people, future supplies are undermined by the impact of this production and consumption on Nature’s ability to provide.   

The report also emphasizes that both the benefits and burdens associated with the use of biodiversity and ecosystem services are distributed and experienced inequitably among social groups, countries and regions. Furthermore, benefits provided to some people often come at the expense of other people, particularly the most vulnerable.  However, there are also synergies – e.g., sustainable agricultural practices enhance soil quality, thereby improving productivity and other ecosystem functions and services such as carbon sequestration and water quality regulation.

The report contains vast amounts of data on the recent explosion of human numbers and – especially – consumption – of agricultural production, fish harvests, forest products, bioenergy production … and on the associated declines in “regulating” and “non-material contributions” ecosystem services. In consequence, the report concludes, these recent gains in material contributions are often not sustainable.

While invasive species rank fifth as a causal agent of biodiversity decline globally, alien species have increased by 40% since 1980, associated with increased trade and human population dynamics and trends. The authors report that nearly 20% of Earth’s surface is at risk of bioinvasion. The rate of invasive species introduction seems higher than ever and shows no signs of slowing.

The report notes that the extinction threat is especially severe in areas of high endemism. Invasive species play a more important role as an extinction agent in many such areas, especially islands. However, some bioinvaders also have devastating effects on mainlands; the report cites the threat of the pathogen Batrachochytrium dendrobatidis to nearly 400 amphibian species worldwide.

The report also mentions that the combination of species extinctions and transport of species to new ecosystems is resulting in biological communities – both managed and unmanaged — becoming more similar to each other — biotic homogenization.

The report notes that human-induced changes are creating conditions for fast biological evolution of species in all taxonomic groups. The authors recommend adopting conservation strategies designed to influence evolutionary trajectories so as to protect vulnerable species and reduce the impact of unwanted species (e.g., weeds, pests or pathogens).

The report says conservation efforts have yielded positive outcomes – but they have not been sufficient to stem the direct and indirect drivers of environmental deterioration. Since 1970, nations have adopted six treaties aimed at protection of nature and the environmental, but few of the strategic objectives and goals adopted by the treaties’ parties are being realized. One objective that is on track to partial achievement is the Aichi Biological Diversity Target that calls for identification and prioritization of invasive species. 

That might well be true – but I would not consider global efforts to manage invasive species to be a success story in any way. I have blogged often about studies showing that introductions continue unabated … and management of established bioinvaders only rarely results in measurable improvements.   [For example, see here and here.]

The report gives considerable attention to problems caused by some people’s simultaneous lack of access to material goods and bearing heavier burden from pollution and other negative results of biodiversity collapse. Extraction of living biomass (e.g. crops, fisheries) to meet the global demand is highest in developing countries whereas material consumption per capita is highest in developed countries. The report says that conservation of biodiversity must be closely linked to sustainable approaches to more equal economic development. The authors say both conservation and economic goals can be achieved – but this will require transformative changes across economic, social, political and technological factors.

One key transformation is changing people’s conception of a good life to downplay consumption and waste. Other attitudinal changes include emphasizing social norms promoting sustainability and personal responsibility for the environmental impacts of one’s consumption. Economic measures and goals need to address inequalities and integrate impacts currently considered to be “economic externalities”. The report also calls for inclusive forms of decision-making and promoting education about the importance of biodiversity and ecosystem services.

Economic instruments that promote damaging, unsustainable exploitation of biological resources (or their damage by pollution) include subsidies, financial transfers, subsidized credit, tax abatements, and commodity and industrial goods prices that hide environmental and social costs. These need to be changed.

Finally, limiting global warming to well below 2oC would have multiple co-benefits for protecting biodiversity and ecosystem services. Care must be exercised to ensure that large-scale land-based climate mitigation measures, e.g., allocating conservation lands to bioenergy crops, planting of monocultures, hydroelectric dams) do not themselves cause serious damage to biodiversity or other ecosystem services.

The threats to biodiversity and ecosystem services are most urgent in South America, Africa and parts of Asia. North America and Europe are expected to have low conversion to crops and continued reforestation.

Table SPM.1 lays out a long set of approaches to achieve sustainability and possible actions and pathways for achieving them. The list is not exhaustive, but rather illustrative, using examples from the report.

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.

Promising Biocontrol to Protect Some Cacti

Photo of infested cactus at Cabo Rojo National Wildlife Refuge, Puerto Rico. Taken August 20, 2018 by Yorelyz Rodríguez-Reyes

Three and a half years ago, I blogged about the threat to columnar cacti in Puerto Rico from the Harrisia cactus mealybug. The mealybug clearly threatens the endemic cacti of the Caribbean islands, and possibly some of the hundreds of other columnar cacti growing across two million square miles of desert ecosystems that straddle the U.S.-Mexico border region.

I am pleased to report that scientists continue efforts to find biocontrol agents to reduce this insect’s damage on Caribbean islands. Much of this work is being done by the Center for Excellence in Quarantine and Invasive Species at University of Puerto Rico. The team consists of Michael West Ortiz, Yorelys Rodrígues Reyes, Ferdinand Correa and Jose Carlos Verle Rodrigues.

As of February 2019, the Center is conducting host specificity tests on a primary parasitoid of the Harrisia Cactus mealybug — Anagyrus cachamai. This wasp was found as a result of almost a decade of searching in South America and other locations. It is native to Argentina and Paraguay (Triapitsyn et al. 2018; sources listed at the end of the blog).The Center also continues surveys and studies of other primary and secondary parasitoids of the mealybug.

The work to develop a biocontrol agent for the mealybug continues despite continuing uncertainty about the true species of the mealybug. At the time of its discovery on Puerto Rico, the mealybug was believed to belong to a species used as a biocontrol agent for invasive cacti in Australia and South Africa, designated as Hypogeococcus pungens. However, H. pungens is now thought to be a species complex, and the species in Puerto Rico differs from the earlier designation (Triapitsyn et al. 2018). 

Apparently the mealybug was introduced in Puerto Rico around 2000   — probably on the ornamental common purslane (Portulaca olerácea), an annual succulent. (Note: the introduction was on a host different from the vulnerable cacti.) Within five years of the first detection in San Juan, the mealybug was sighted on cacti on the other side of the island in the Guánica State Forest and Biosphere Reserve. By 2010, the mealybug was widely distributed in most dry districts. Surveys found it in all 11 municipalities surveyed in southern Puerto Rico. At some locations, infestation levels were extremely high – e.g., 86% of stems surveyed were infested at Guánica. Infestation rates were lower in other municipalities. As of 2010, infestations were estimated to be present on about 1,400 km2 on the southern coast; the rate of new infestations suggests that the mealybug was spreading rapidly (Segarra-Carmona et al. 2010).  I have been unable to obtain more recent estimates.

The mealybug impacts seven of 14 native cactus species occurring in dry forests of the island, including three endemic and two endangered species in the subfamily Cactoideae. The two endangered species are Harrisia portoricensis and Leptocereus grantianus (USDA ARS). The tissue damage caused by the mealybug interferes with sexual reproduction and can cause direct mortality of the plant (Triapitsyn et al. 2018).  These cacti provide food or shelter for endemic bats, birds, moths and other pollinators (Segarra & Ramirez; USDA ARS). The mealybug is also now killing native cacti on the U.S. Virgin Islands (H. Diaz-Soltero pers. comm. August 2015).

 USDA Funds Conservation Efforts Despite Apparent Absence of a Constituency Calling for Such Action

Efforts to identify and test possible biocontrol agents targetting the Harrisia cactus mealybug received significant funds from the Plant Pest and Disease Management and Disaster Prevention Program. This is a competitive grant program managed by APHIS. It is permanently funded and thus not subject to the vagaries of annual appropriations. Until last year, this program operated under Section 10007 of the 2014 Farm Bill. With passage of a new Farm Bill, it is now designated as Section 7721 of the Plant Protection Act.

Since Fiscal Year 2018, APHIS has had authority to spend more than $60 million per year on this program.  In Fiscal Year 2017, , the program provided $120,000 to an unspecified federal agency, $70,000 to an academic institution in Puerto Rico (presumably the Center), $15,000 to another academic institution in California, and $3,000 divided among two APHIS facilities – for a total of $208,000. The next round of funds came in FY19, when the program provided $277,267 to an unspecified federal agency to continue work on biocontrol. In addition, the program provided $78,507 to an unspecified federal agency to “safeguard[e] genetic diversity of native and listed cacti threatened by Harrisia cactus mealybug in Puerto Rico”.

No Apparent Action on Threats to Opuntia Cacti

In my earlier blog, I also described the threat to flat-padded Opuntia (prickly pear) cacti from the cactus moth Cactoblastis cactorum. Various federal, state, and academic entities received $463,000 from the permanent fund in Fiscal Year 2016 and another $100,000 in FY2017. No cactus moth programs have received funds in more recent years.

SOURCES

Segarra-Carmona, A.E., A. Ramirez-Lluch. No date. Hypogeococcus pungens (Hemiptera: Pseudococcidae): A new threat to biodiversity in fragile dry tropical forests.

Segarra-Carmona, A.E., A. Ramírez-Lluch, I. Cabrera-Asencio and A.N. Jiménez-López. 2010.  FIRST REPORT OF A NEW INVASIVE MEALYBUG, THE HARRISIA CACTUS MEALYBUG HYPOGEOCOCCUS PUNGENS (HEMIPTERA: PSEUDOCOCCIDAE). J. Agrie. Univ. RR. 94(1-2):183-187 (2010)

Triapitsyn, Aguirre, Logarzo, Hight, Ciomperlik, Rugman-Jones, Rodriguez. 2018. Complex of primary and secondary parasitoids (Hymenoptera: Encyrtidae and Signiphoridae) of Hypogeococcus spp. mealybugs (Hemiptera: Pseudococcidae) in the New World. Florida Entomologist Volume 101, No. 3 411

USDA Agriculture Research Service, Research Project: Biological Control of the Harrisia Cactus Mealybug, Hypogeococcus pungens (Hemiptera:pseudococcidae) in Puerto Rico Project Number: 0211-22000-006-10 Project Type: Reimbursable

West Ortiz, M. pers. comm. February 2019

Posted by Faith Campbell

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