March 2022 – Center for Invasive Species Prevention

What Do Invasive Species Cost?

brown tree snake *Boiga irregularis*; via Wikimedia; one of the species on which the most money is spent on preventive efforts

In recent years a group of scientists have attempted to determine how much invasive species are costing worldwide. See Daigne et al. 2020 here.

Some of these scientists have now gone further in evaluating these data. Cuthbert et al. (2022) [full citation at end of blog] see management of steadily increasing numbers of invasive, alien species as a major societal challenge for the 21st Century. They undertook their study of invasive species-related costs and expenditures because rising numbers and impacts of bioinvasions are placing growing pressure on the management of ecological and economic systems and they expect this burden to continue to rise (citing Seebens et al., 2021; full citation at end of blog).

They relied on a database of economic costs (InvaCost; see “methods” section of Cuthbert et al.) It is the best there is but Cuthbert et al. note several gaps:

Only 83 countries reported management costs; of those, only 24 reported costs specifically associated with pre-invasion (prevention) efforts.
Data comparing regional costs do not incorporate consideration of varying purchasing power of the reporting countries’ currencies.
Data available are patchy so global management costs are probably substantially underestimated. For example, forest insects and pathogens account for less than 1% of the records in the InvaCost database, but constitute 25% of total annual costs ($43.4 billion) (Williams et al., in prep.) .

Still, their findings fit widespread expectations.

These data point to a total cost associated with invasive species – including both realized damage and management costs – of about $1.5 trillion since 1960. North America and Oceania spent by far the greatest amount of all global money countering bioinvasions. North America spent 54% of the total expenditure of $95.3 billion; Oceania spent 30%. The remaining regions each spent less than $5 billion.

Cuthbert et al. set out to compare management expenditures to losses/damage; to compare management expenditures pre-invasion (prevention) to post-invasion (control); and to determine potential savings if management had been more timely.

Economic Data Show Global Efforts Could Be – But Aren’t — Cost-Effective

The authors conclude that countries are making insufficient investments in invasive species management — particularly preventive management. This failure is demonstrated by the fact thatreported management expenditures ($95.3 billion) are only 8% of total damage costs from invasions ($1.13 trillion). While both cost or losses and management expenditures have risen over time, even in recent decades, losses were more than ten times larger than reported management expenditures. This discrepancy was true across all regions except the Antarctic-Subantarctic. The discrepancy was especially noteworthy in Asia, where damages were 77-times higher than management expenditures.

Furthermore, only a tiny fraction of overall management spending goes to prevention. Of the $95.3 billion in total spending on management, only $2.8 billion – less than 3% – has been spent on pre-invasion management. Again, this pattern is true for all geographic regions except the Antarctic-Subantarctic. The divergence is greatest in Africa, where post-introduction control is funded at more than 1400 times preventive efforts. It is also significant for Asia and South America.

Even in North America – where preventative actions were most generously funded – post-introduction management is funded at 16 times that of prevention.

Cuthbert et al. worry particularly about the low level of funding for prevention in the Global South. They note that these conservation managers operate under severe budgetary constraints. At least some of the bioinvasion-caused losses suffered by resources under their stewardship could have been avoided if the invaders’ introduction and establishment had been successfully prevented.

While in the body of the article Cuthbert et al. seem uncertain about why funding for preventive actions is so low, in their conclusions they offer a convincing (to me) explanation. They note that people are intrinsically inclined to react when impact becomes apparent. It is therefore difficult to motivate proactive investment when impacts are seemingly absent in the short-term, incurred by other sectors, or in different regions, and when other demands on limited funds may seem more pressing. Plus efficient proactive management will prevent any impact, paradoxically undermining evidence of the value of this action!

*Aedes aegypti* mosquito; one of the species on which the most money is spent for post-introduction control; photo by James Gathany; via Flickr

Delay Costs Money

The reports contained in the InvaCost database indicate that management is delayed an average of 11 years after damage was first been reported. Cuthbert et al. estimate that these delays have caused an additional cost of about $1.2 trillion worldwide. Each $1 of management was estimated to reduce damage by $53.5 in this study. This finding, they argue, supports the value of timely invasive species management.

They point out that the Supplementary Materials contain many examples of bioinvasions that entail large and sustained late-stage expenditures that would have been avoided had management interventions begun earlier.

Although Cuthbert et al. are not as clear as I would wish, they seem to recognize also that stakeholders’ varying perceptions of whether an introduced species is causing a detrimental “impact” might also complicate reporting – not just whether any management action is taken

Cuthbert et al. are encouraged by two recent trends: growing investments in preventative actions and research, and shrinking delays in initiating management. However, these hopeful trends are unequal among the geographic regions.

Which Taxonomic Groups Get the Most Money?

About 42% of management costs ($39.9 billion) were spent on diverse or unspecified taxonomic groups. Of the costs that were taxonomically defined, 58% ($32.1 billion) was spent on invertebrates [see above re: forest pests]; 27% ($14.8 billion) on plants; 12% ($6.7 billion) on vertebrates; and 3% ($1.8 billion) on “other” taxa, i.e. fungi, chromists, and pathogens. For all of these defined taxonomic groups, post-invasion management dominated over pre-invasion management.

When considering the invaded habitats, 69% of overall management spending was on terrestrial species ($66.1 billion); 7% on semi-aquatic species ($6.7 billion); 2% on aquatic species ($2.0 billion); the remainder was “diverse/unspecified”. For pre-invasion management (prevention programs), terrestrial species were still highest ($840.4 million). However, a relatively large share of investments was allocated to aquatic invaders ($624.2 million).

Considering costs attributed to individual species, the top 10 targetted for preventive efforts were four insects, three mammals, two reptiles, and one alga. Top expenditures for post-invasion investments went to eight insects [including Asian longhorned beetle], one mammal, and one bird.

Asian longhorned beetle

Just two of the costliest species were in both categories: insects red imported fire ant(Solenopsis invicta) and Mediterranean fruitfly (Ceratitis capitate). None of the species with the highest pre-invasion investment was among the top 10 costliest invaders in terms of damages. However, note the lack of data on fungi, chromists, and pathogens. (I wrote about this problem in an earlier blog.)

Discussion and Recommendations

Cuthbert et al. conclude that damage costs and post-invasion spending are probably growing substantially faster than pre-invasion investment. Therefore, they call for a stronger commitment to enhancing biosecurity and for more reliance on regional efforts rather than ones by individual countries. Their examples of opportunities come from Europe.

Drawing parallels to climate action, the authors also call for greater emphasis on during decision-making to act collectively and proactively to solve a growing global and inter-generational problem.

Cuthbert et al. focus many of their recommendations on improving reporting. One point I found particularly interesting: given the uneven and rapidly changing nature of invasive species data, they think it likely that future invasions could involve a new suite of geographic origins, pathways or vectors, taxonomic groups, and habitats. These could require different management approaches than those in use today.

As regards data and reporting, Cuthbert et al. recommend:

1) reducing bias in cost data by increasing funding for reporting of underreported taxa and regions;

2) addressing ambiguities in data by adopting a harmonized framework for reporting expenditures. For example, agriculture and public health officials refer to “pest species” without differentiating introduced from native species. (An earlier blog also discussed the challenge arising from these fields’ different purposes and cultures.)

3) urging colleagues to try harder to collect and integrate cost information, especially across sectors;

4) urging countries to report separately costs and expenditures associated with different categories, i.e., prevention separately from post-invasion management; damage separately from management efforts; and.

5) creating a formal repository for information about the efficacy of management expenditures.

While the InvaCost database is incomplete (a result of poor accounting by the countries, not lack of effort by the compilers!), analysis of these data points to some obvious ways to improve global efforts to contain bioinvasion. I hope countries will adjust their efforts based on these findings.

SOURCE

Cuthbert, R.N., C. Diagne, E.J. Hudgins, A. Turbelin, D.A. Ahmed, C. Albert, T.W. Bodey, E. Briski, F. Essl, P. J. Haubrock, R.E. Gozlan, N. Kirichenko, M. Kourantidou, A.M. Kramer, F. Courchamp. 2022. Bioinvasion costs reveal insufficient proactive management worldwide. Science of The Total Environment Volume 819, 1 May 2022, 153404

Seebens, H. S. Bacher, T.M. Blackburn, C. Capinha, W. Dawson, S. Dullinger, P. Genovesi, P.E. Hulme, M.van Kleunen, I. Kühn, J.M. Jeschke, B. Lenzner, A.M. Liebhold, Z. Pattison, J. Perg, P. Pyšek, M. Winter, F. Essl. 2021. Projecting the continental accumulation of alien species through to 2050. Glob Change Biol. 2021;27:970-982.

Williams, G.M., M.D. Ginzel, Z. Ma, D.C. Adams, F.T. Campbell, G.M. Lovett, M. Belén Pildain, K.F. Raffa, K.J.K. Gandhi, A. Santini, R.A. Sniezko, M.J. Wingfield, and P. Bonello 2022. The Global Forest Health Crisis: A Public Good Social Dilemma in Need of International Collective Action. submitted

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

Global Loss of Floristic Uniqueness

Hakalau Forest, Hawai“i; nearly 90% of Hawaiian flora is unique to the Islands

A recent article by Yang et al. 2021 (full citation at the end of this blog) seeks to determine the extent to which introduced plants reduce the uniqueness of regional floras. They analyzed data from 658 regions covering about 65.7% of the Earth’s ice-free land surface and about 62.3% of the planet’s known plant species.

They found strong homogenization of plant species’ taxonomic and phylogenetic diversity results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.

There are two aspects of a region’s floral uniqueness. One is the number of species that it shares with other regions. This is taxonomic uniqueness. The other is the distinctiveness of the evolutionary history of the region. When several species are endemic to a region’s flora, and lack close relatives in other regions, that equals phylogenetic uniqueness.

The effect of a species introduction differs depending on which of these aspects one focuses on. Thus, naturalization of a species closely related to native species (e.g., a congeneric species) will have less impact on the phylogenetic floristic uniqueness of the region than naturalization by a distantly related species. Taxonomic uniqueness, however, will be affected to the same degree, irrespective of the phylogenetic distance between the naturalized and native species.

Yang et al. found strong homogenization of plant diversity. They found that species introductions increased the taxonomic similarity in 90.7% of all regional pairs and phylogenetic similarity in 77.2% of all region pairs. Most homogenization results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.

This loss of regional biotic uniqueness or distinctiveness changes biotic interactions and species assemblages. These, in turn, have ecological and evolutionary consequences at larger scales and higher levels.

The degree of homogenization between regions’ floras depends on three factors:

1) The distance between the donor and recipient regions. Since nearby regions share more species, an introduction from a more distant origin is more likely to be a novel species and so contribute to homogenization of “donor” and “receiving” floras.

2) Climatic similarity, especially temperature. A plant species introduced from a climatically similar but geographically distant place is more likely to establish than a species from a different climatic zone. As a result, the recipient area’s flora is changed to more closely resemble the flora of the donor region with which it shares climatic conditions – regardless of the distance between them.

3) The level of exchange of goods and people between two regions. The higher the rate of exchange between two regions, the greater the chance that a species will be introduced and become established. Yang et al. used the existence of current or past administrative relationships (e.g., colonial relationship) between two regions as a proxy for intensity of trade and transport between donor and recipient regions. They found that floras of regions with current or past administrative links have taxonomically become more similar to each other than the floras of regions with no such links.

flora of the Cape Floral Kingdom – South Africa; photo from Michael Wingfield

Establishment of introduced species can increase floristic similarity of the donor and recipient regions (= floristic homogenization) when the species is native to one of the two regions and naturalizes in the other, or when it is not native to both regions and naturalizes in both. On the other hand, a species introduction can decrease the floristic similarity of the two regions (i.e., enhance floristic differentiation) when the species is not native to both regions but naturalized in only one.

Homogenization hotspots differed slightly depending on whether one focused on taxonomic or phylogenetic aspects.

The regions with the greatest average increase in taxonomic similarity with other regions due to naturalized alien species were New Zealand, portions of Australia, and many oceanic islands. The Australasian situation probably reflects its long biogeographic isolation from other parts of the globe and its highly unique native flora. As a result, nearly all non-native plants introduced to Australasia strongly increase levels of its floristic similarity to the rest of the world. Oceanic islands have species-poor floras with large proportions of unique endemics. They have also received high numbers of naturalized alien plants.

Hotspots of phylogenetic homogenization on continents are the same as those for taxonomic homogenization, but this is not true for islands. Yang et al. think this is because islands’ native floras were established by natural colonization from nearby continental floras so – despite subsequent speciation – they retain their phylogenetic relationship to the donor areas’ floras.

Yang et al. concede that they lacked high-quality data on native and naturalized alien species lists for a third of Earth’s ice-free terrestrial surface, especially Africa, Eastern Europe, and tropical Asia. They believe, however, that data from these regions are unlikely to change the overall finding. (Scientists are beginning to compile lists of forest pests in Africa). link to blog

Yang et al. note that introduction and naturalization of alien species are likely to increase in the future, thusaccelerating floristic homogenization. The ecological, evolutionary and socioeconomic consequences are largely unknown.They call for stronger biosecurity regulations of trade and transport and other measures to protect native vegetation.

SOURCE

Yang, Q., P. Weigelt, T.S. Fristoe, Z. Zhang, H. Kreft, A. Stein, H. Seebens, W. Dawson, F. Essl, C. König, B. Lenzner, J. Pergl, R. Pouteau, P. Pyšek, M. Winter, A.L. Ebel, N. Fuentes, E.L.H. Giehl, J. Kartesz, P. Krestov, T. Kukk, M. Nishino, A. Kupriyanov, J.L. Villaseñor, J.J. Wieringa, A. Zeddam, E. Zykova and M. van Kleunen. 2021. The global loss of floristic uniqueness. NATURE COMMUNICATIONS (2021) 12:7290. https://doi.org/10.1038/s41467-021-27603-y

Posted by Faith Campbell

Urban Forests at Risk: Thousands of Communities, Millions of Trees, & Tens of Millions of Dollars

EAB-killed ash tree lying on highway in Fairfax County, Virginia; photo by F.T. Campbell

A recent study (Hudgins, Koch, Ambrose & Leung 2022; full citation at end of blog) projects that, by 2050, 1.4 million street trees in urban areas and communities will be killed by introduced insect pests. This represents 2.1- 2.5% of all urban street trees. Nearly all of this mortality will occur in a quarter of the 30,000 communities evaluated. Additional urban trees – in parks, other plantings, on homeowners’ properties, and in urban woodlands – are also expected to die.

Loss of these trees will undercut all the ecosystem services provided by urban trees.

The principal cause of mortality will be the emerald ash borer (EAB). Already, an estimated 230,000 ash trees have been killed by EAB. The authors predict that 6,747 communities not yet affected by the EAB will suffer the highest losses between now and 2060. Most of these communities are in a 350,000 square mile area of the northeast and central states. However, the risk is far wider, reaching as far as Seattle.

This ash tree has been standing – dead – since 2016. When will it fall?

In the top ‘hotspot cities’ projected mortality is in the range of 5,000–25,000 street trees. These include Milwaukee; the Chicago area (Chicago / Aurora / Naperville / Arlington Heights); Cleveland; and Indianapolis. As in previous studies, the highest insect impacts are in the Northeast. Pests impacting this region – in addition to the emerald ash borer – include the spongy moth (formerly called gypsy moth) and hemlock woolly adelgid.

Because insect-killed trees must be treated or removed to minimize the risk to human life and property, the pest risk represents an economic as well as ecological threat. Removing and replacing just the street trees is projected to cost cities $30 million per year. Considering the cities I mentioned above, Milwaukee faces costs estimated at $13 million; Warwick, RI $2.5 million; Baltimore $1.7; Richmond and Virginia Beach $7.3 million and $700,000 respectively; and three New Jersey cities (Jersey City, Elizabeth City, and Patterson) $1.6 million combined.

USDA APHIS ended the federal quarantine for EAB in 2021. Therefore these cities and states are on their own to protect themselves from not only this and other damaging insects but also their extraordinarily high economic costs.

The study evaluated the risk to 48 genera of trees in about 30,000 communities. The most widely planted genera are maples (Acer spp.) and oaks (Quercus spp.). Consequently, they will die in largest numbers. An estimated 26.5 million maples and 5.9 million oaks are at risk, primarily in the East. As noted above, EAB is expected to kill 99% of ash trees in 6,747 communities across the country. In the Southwest, there are 3.4 million pines (Pinus spp.); the threat to them is not woodborers, but scale insects (San Jose scale [Quadraspidiotus perniciosus] and calico scale [Eulecanium cerasorum]).

As we know, urban forests are easily invaded because they are close to ports of entry and are often composed primarily of highly susceptible species. Hudgins, Koch, Ambrose and Leung analyzed the potential risk associated with introduction of a new woodboring insect from Asia – which they point out is the source of most imported goods. They determined that if such an introduced pest were to attack maples or oaks, it could kill 6.1 million trees and cost American cities $4.9 billion over 30 years. The risk would be highest if this pest were introduced via a port in the South.

In an earlier blog I reported that the U.S. is currently importing about 20 million shipping containers filled with goods from Asia per year. I have often blogged about the pest risk associated with wood packaging accompanying these imports. The number of containers from Asia entering Southeastern ports rose by more than 10% from December to January.

Hudgins, Koch, Ambrose & Leung combined four sources of information to produce these estimates:

a model of spread for 57 species of introduced insect pests already determined to cause significant damage to trees;
the distribution of genera of urban street trees across 30,000 US communities;
a model of host mortality in response to each insect-host combination; and
the cost of removing and replacing dead trees, linked to tree size (dbh).

They excluded several categories of pests. One of the most damaging, Asian longhorned beetle, was excluded because scientists have already developed control methods to limit its spread. Also excluded were species present in the U.S. for less than five years; species with no known economic impacts; and species for which no hosts in natural North American forests have been identified. Also excluded – although the authors do not mention this – are species that did not qualify for inclusion in the Aukema et al. study (see reference at end of this blog) because they have been introduced from nearby portions of North America, e.g., goldspotted oak borer. Finally, the study does not include pathogens. Some pathogens have caused huge losses of urban trees in the past, e.g., Dutch elm disease; some are causing losses now, e.g., sudden oak death. The authors do mention the Fusarium disease vectored by polyphagous (and Kuroshio) shot hole borers in southern California.

Consequently, the study’s estimate of 1.4 million street trees dead and costs of $30 million per year are underestimates.

The study has generated considerable media interest, including in the Washington Post.

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

Hudgins, E.J., F.H. Koch, M.J. Ambrose, B. Leung. 2022. Hotspots of pest-induced US urban tree death, 2020–2050. Journal of Applied Ecology 2022;00:1-11 DOI: 10.1111/1365-2664.14141

Posted by Faith Campbell

Restoring Port-Orford cedar – a role for you!!!!

Port-Orford cedar; photo by Julie Kierstead

I report here on recent developments on breeding resistant trees. These include both promising results from decades-long efforts and also a promising start to addressing a new challenge.

These programs have benefited from major commitments by the USDA Forest Service. I hope they encourage similar commitments for other priority species – such as those named by the CAPTURE program.

Port-Orford cedar – ready to be planted in the forest!

Scientists who have been working for decades to breed seedlings of Port-Orford cedar (POC) trees resistant to the root rot caused by Phytophthora lateralis https://www.dontmovefirewood.org/pest_pathogen/port-orford-cedar-root-disease-html/now say that they have seedlings ready for planting in the forest. They made this case in a webinar in late February. It can be viewed here. The full webinar runs somewhat over two hours.

The scientist who led early studies of POC and the root disease, Don Zobel, Professor Emeritus, Oregon State University, described the ecological requirements that should guide planting programs. POC produces high-calcium litter. It grows from the sea coast to 1950 meters elevation, on sand dunes, fens, soils with hardpans; mafic & ultramafic rocks (serptentines) and fertile soils on some sedimentary rocks. POC is less shade tolerant than western hemlock but more fire tolerant. It can form a secondary canopy under Douglas-fir and supercede other conifers when fire occurs repeatedly. The tree needs surface water, e.g., seepages and stream sides; but the water must be flowing, not stagnant. Seedlings are especially vulnerable to drying during winter.

[I posted a separate blog about other trees native to this region, including serpentine soils, here.]

One purpose of the webinar was to encourage owners and managers of lands within POC’s historic range (see the map under Dr. Zobel’s presentation) to begin planting the species in appropriate sites. With this in mind, Dr. Zobel emphasized criteria for selecting sites:

Climates in coastal areas of the range are less likely to change under climate change
Quartenary marine terraces are the best geologic type; Lookingglass and Roseburg geologic types are also acceptable
Availability of water during summer, e.g., streamside and seepage areas. Try planting beneath alder. However, avoid interior valley stream corridors if the soils are not ultramafic. And avoid stagnant water.

a POC tree in a bog next to the endemic pitcher plant of southern Oregon, *Darlingtonia* *californica*; photo by Richard Sniezko

Dr. Zobel also says one should plant pathogen-resistant genotypes and pay attention to local genetic varieties (which have largely been determined).

Dr. Richard Sniezko of the USFS Dorena Genetic Resource Center described the Center’s 30-year effort to find and exploit resistance to the pathogen. Funding has come from the USFS Forest Health Protection program, other parts of the USFS, and the Bureau of Land Management (BLM). The goal all along has been to produce seedlings for restoration to the forest – meaning not just resistant to the pathogen but also adapted to various local conditions. The program can now provide resistant seedlings in large quantities for planting by landowners and public land managers.

Dr. Sniezko emphasizes that success depends on engagement of four sets of people: research by university scientists; application of that research and development of propagule growing methods by the Dorena Center; support from USFS leaders to continue the program; involvement of land managers who choose to plant the resistant seedlings.

USFS and BLM staff described efforts to determine where POC grows on land under their management, the status of disease in those areas, and efforts to slow the spread of the disease, especially along roadsides and as result of timber or engineering projects. Some of this sanitation work has been funded by USFS Forest Health Protection program — not the National Forest System.

Richard Sniezko stated that the seedlings’ quantitative disease resistance means that some seedlings will die. He expects 40-50% survival of seedlings from many of the breeding zones. This is well above the level of resistance in un-improved populations.

Both BLM and the Rogue-River-Siskiyou National Forest have planted tens of thousands of resistant seedlings in recent years and plan to continue. Funding provided by COVID-19 legislation might allow increased effort. [See Dr. Sniezko’s presentation on the webinar for photos from some plantings.]

POC seedlings at Dorena; photo by Richard Sniezko

Norma Kline of the Oregon State University extension program has distributed more than 10,000 seedlings to small/non-industrial landowners. Many of the recipients shared seedlings with neighbors or are coordinating their planting over a large area. They were motivated primarily by conservation concerns. Her monitoring showed that the POC seedlings survived but did not thrive under dense tanoak canopy. They did well in competition with grass in areas near the coast where there was more moisture. They also did well under Douglasfir as long as there was dappled sunlight.

The non-governmental organization American Forests is likely to participate actively in the planting effort.

In an email to me, Dr. Sniezko asks that people who have planted POC outside its native range inform him where the tree(s) is/are thriving. This information would enhance scientists’ understanding of the species’ environmental tolerances.

Posted by Faith Campbell

Rapid ‘Ohi‘a Death – Comprehensive New Program Begins – Challenges to Achieving Success

‘Ohi‘a (Metrosideros polymorpha) is the most abundant native forest tree in Hawai`i and of enormous ecological, cultural, and economic importance. Five species of endemic Metrosideros are recognized on the Hawaiian islands. Only one — M. polymorpha — is found throughout the state. Eight varieties are recognized. These varieties inhabit different environments and have adapted to selective pressures characteristic of these locations. There are at least five other species in the Metrosideros genus, each endemic to one or a few nearby islands. Blaine et al. (2022) [full citation at end of this blog] provide a helpful summary of the tree’s ecological importance and its apparently on-going speciation.

‘Ohi‘a provides habitat for endemic birds, insects, and plants, many of which are endangered. Thus, conservation of this species — and all Hawaiian Metrosideros – is vital for the conservation of countless other taxa. In addition, high elevation ʻōhiʻa forests protect vitally important watersheds across the state. For Native Hawaiians, ʻōhiʻa is a physical manifestation of multiple Hawaiian deities so is the subject of many proverbs, chants, stories, and a foundation of scared hula. Finally, the tree is beautiful!

Native Hawaiian forests face multiple threats — invasive animals and plants, wildfire, and land-use changes. Due to such threats, natural ʻōhiʻa regeneration is largely absent in most lower-elevation forests. In this case, competition with invasive species and the presence of diseases such as ʻōhiʻa rust (Austropuccinia psidii) are probably the specific causes. Multiple government and non-governmental entities have made substantial effort to mitigate these threats.

ROD-infected ʻōhiʻa; photo by J.B. Friday

The disease Rapid ‘Ohi‘a Death (ROD) is an unprecedented threat to this species and the forests it constitutes. The disease is caused by two newly described fungal pathogens: Ceratocystis lukuohia and C. huliohia. The disease caused by C. lukuohia is more severe. To date it has been detected on the two islands farthest apart in the chain — Hawai`i (the Big Island) and Kaua‘i. C. huliohia causes a canker disease that kills trees more slowly. It is more widespread, found on Maui and O‘ahu in addition to Hawai`i and Kaua‘i. Blaine et al. (2022) and the profile here describe the two diseases’ epidemiologies, progression, impacts, and challenges.

Because of the clear threat to Hawaiian ecosystems, ecosystem services, and cultural assets, considerable effort has put into delimitation and research on possible mitigation actions since ROD was discovered in 2010. The first strategic plan covered the period 2017–2019. It focused on expanded efforts to map outbreaks, research on the epidemiology of the pathogens, and most-promising management practices. The second strategic plan covers 2020–2024. It provides for continued surveillance and improvement of these technologies; expanding outreach and public engagement; research on possible vectors of the pathogens; collection and preservation of seeds for research and future restoration; and comprehensive evaluation and development of disease resistance in ʻōhiʻa.

Soon after the causal agents were clarified, the USDA Agriculture Research Service (ARS) began screening for disease resistance. By 2016, ARS had demonstrated that five individuals from two varieties of M. polymorpha had survived inoculation by the more virulent pathogen, C. lukuohia. Their survival raised hopes that natural resistance might be present in wild populations of at least some varieties. However, more comprehensive screening of trees from throughout the species’ range is needed to provide an accurate baseline on the frequency, level, and distribution of genetic resistance to both pathogens. The goal is to produce material resistant to both pathogens that can be used to preserve the ecology, culture, and biotic communities that are dependent on this tree species.

To carry the expanded effort forward, in 2018 a collaborative partnership of state, federal, and non-profit groups was formed. Participants in the ‘Ohi‘a Disease Resistance Program (‘ODRP) include: the Akaka Foundation for Tropical Forests; USDA’s Forest Service and Agriculture Research Service; the state’s Division of Forestry and Wildlife and Agriculture Research Center; programs of the University of Hawai‘i at Manoa and at Hilo; Purdue and Arizona State universities; the Tropical Hardwood Tree Improvement and Regeneration Center; and Kalehua Seed Conservation Consulting.

Blaine et al. (2022) have now outlined a framework to guide the overall effort to identify and develop ROD resistance in M. polymorpha and, possibly, all Hawaiian Metrosideros species. The framework calls for the following activities:

(1) evaluating and operationalizing methods for inoculation-based screening and greenhouse-based production of test plants; and

(2) short-term greenhouse screenings of seedlings and rooted cuttings sampled from native Metrosideros throughout Hawai’i.

Once these tasks have been achieved, the effort is expected to expand to address:

(3) establishing field trials to validate the short-term greenhouse assays and monitor durability and stability of resistance;

(4) understanding environmental (climate, soils) and genetic (vascular architecture, wound response)

drivers of susceptibility and resistance to characterize the durability and stability of genetic resistance to ROD;

(5) developing remote sensing and molecular methods to rapidly detect ROD-resistant individuals;

(6) if necessary, conducting breeding to increase the efficacy of resistance and improve durability of ROD resistance; and

(7) supporting already established and ongoing Metrosideros conservation, including state-wide seed collection and banking, with information on not only genotypes resistant to ROD but also production of ROD-resistant seed.

Blaine et al. (2022) outline how to proceed on each step, and describe the challenges that must be overcome. Challenges range from building growing and screening capacity to handle the thousands of plants required, to developing the remote sensing tools to identify diseased trees in the forest, to identifying sites for seed orchards. Actions by ‘ODRP will focus on Stage II screening in the field to examine the durability of resistance under the wide variety of ecological conditions in which ʻōhiʻa grows and in the presence of a potentially evolving pathogen. Resistance studies must expand beyond M. polymorpha varieties from only one island (the Big Island) to include the other Hawaiian Metrosideros taxa.

Once ROD-resistant M. polymorpha trees are discovered and groundwork has been laid to satisfy initial needs for resistant tree seedlings for forest restoration, scientists can begin research into the genetic basis of ROD resistance. This knowledge will assist breeding efforts which might be necessary if resistance to one of the pathogens does not confer resistance to the other, since the goal is to provide seedlings that are resistant to both.

Blaine et al. (2022) note that the state and others continue efforts to address other aspects of ROD management. These include

1) controlling the spread of the pathogen through local quarantines on movement of infected material and increased public education on bio-sanitation for forest users;

2) testing repellants to reduce beetle attack on infected trees and subsequent frass production.

3) reducing wounding of trees by fencing more pristine forests and removing feral ungulates

SOURCE

Blaine C. Luiz, Christian P. Giardina, Lisa M. Keith, Douglass F. Jacobs, Richard A. Sniezko, Marc A. Hughes, James B. Friday, Philip Cannon, Robert Hauff, Kainana Francisco, Marian M. Chau, Nicklos Dudley, Aileen Yeh, Gregory Asner, Roberta E. Martin, Ryan Perroy, Brian J. Tucker, Ale.alani Evangelista, Veronica Fernandez, Chloe Martins-Keli.iho.omalu, Kirie Santos, Rebekah Ohara. 2022. A framework for establishing a rapid ‘Ohia death resistance program. New Forests. https://doi.org/10.1007/s11056-021-09896-5

See also the video at https://www.bigislandvideonews.com/2019/06/16/video-to-save-ohia-a-genetic-resistance-program-will-be-built/

Posted by Faith Campbell

Integrating Invasion & Phytosanitary Sciences & Practice

vegetation killed by *Phytophthora cinnamomi* in West Australia

Some invasive species practitioners have been trying to develop a standardized framework for describing bioinvasions. Their goal is to overcome disparities in approaches developed by scientists working with various taxonomic groups in hopes of improving understanding of, and communication about, bioinvasions. Prominent among these efforts is the “Unified Framework for Bioinvasion” published by Blackburn et al. in 2011 (full citation at end of blog).

Now several forest pathologists (Paap et al; full citation at end of blog) say that this framework does not adequately integrate forest pathogens. This omission is particularly unfortunate given the prominence of forest pathogens as damaging invaders – e.g., chestnut blight in Europe and North America; white pine blister rust in North America; sudden oak death in North America and Great Britain; myrtle rust and Phytophthora cinnamomi in Australia. (See profiles of all these pathogens here; I note additional examples in North America, such as laurel wilt disease.)

Paap et al think that this omission impedes understanding of both forest pests and invasive species in general. Also, they say that integrating microorganisms into the broader Blackburn framework would help forest pathologists better understand how and why invasions occur, where they occur, and how they can be stopped or mitigated.

Furthermore, they note the importance of integrating the diverging terminologies used by invasive species practitioners and plant pathologists and their separate regulatory bodies – the Convention on the Conservation of Biological Diversity (CBD) and the International Plant Protection Convention (IPPC). I concur, since nations’ programs regulating plant diseases and their vectors operate under the IPPC rubric.

Figure 2 and Table 1 lay out Paap et al.’s proposed modification of Blackburn’s framework, and detail strategies linked to management goals appropriate for the stages of plant disease development.

Tanoak mortality in southern Oregon caused by *P. ramorum* – a pathogen completely unknown until it was introduced to North America and Europe; photo by Oregon Department of Forestry

However, such integration will be impeded by many difficulties (I have re-ordered these points):

1) The first – which underlies all others — is the paucity of data on microbial taxa, which undermines the pest risk analyses and other systems developed for assessing and managing other types of invasive species. That is,

Many of the vast number of microbial taxa have not yet been described.
Even species that have been describe often cannot be ascribed to a specific geographic origin. This information gap undercuts efforts to determine whether a disease outbreak is caused by an “introduced” organism.

2) Microbial species are usually detected only when disease impacts become obvious. However, an outbreak might not signal a new or spreading “introduction”. While invasive species must—by definition—cross a geographic boundary (through the assistance of human actions), pathogens can cause disease outbreaks through breaching a wider range of boundaries, including ecological and evolutionary ones. Thus, the disease outbreak doesn’t always fit the definition of “invasive species”.

3) Substantial differences exist in training and goals between fields. Forest pathologists are usually trained in plant pathology (often focused on crops) rather than in forestry or ecology. Their goal is to manage the pathogen. Invasion scientists tend to focus on natural ecosystems, study animal and plant invasions, and seek understanding of the invasion process.

4) A related issue is that the two fields operate under separate regulatory bodies that have different emphases and aims. Paap et al. note that while the IPPC ostensibly includes impacts on natural environments, its members’ priority is plants of economic importance. The World Trade Organization’s Agreement on the Application of Sanitary and Phytosanitary Measures (WTO SPS) seeks primarily to minimize disruption of trade resulting from plant health regulation. On the other hand, the CBD explicitly considers invasive species’ impact to the natural environment (Aichi Biodiversity Target 9). [To read my critique of the WTO SPS and IPPC, read the Fading Forests reports (link at end of this blog), especially FF II.]

They note that in 2004, the IPPC and CBD secretariats established a Memorandum of Cooperation to promote synergy and to avoid duplication. Paap et al. appear disappointed that despite development of joint work plans, phytosanitary programs are still focused largely on crop pathogens.

Disease development – a complex set of circumstances that makes risk assessment less reliable

Since I am not a pathologist (or even a biologist), I learned a lot about the complexities of plant pathology from Paap et al.

While I am certainly familiar with the “disease triangle” concept, I had not thought about certain implications. For example, pathogens can cause severe disease outbreaks by evading any one of three types of barriers: geographic, environmental, or evolutionary. Transport of the micro-organism to a new ecosystem (leaping the geographic barrier and meeting the definition of an “introduction” in invasive species terminology) certainly can facilitate disease outbreaks. However, evolutionary and environmental barriers might also be overcome in other ways.

The result is that a plant disease can develop under multiple scenarios following the introduction of an alien pathogen. These scenarios are:

disease on a coevolved host growing as an alien species in the new environment, for example plantations of trees grown for timber (pathogen reunion);
disease on a naïve host that is itself alien to the geographic region in question (host jump);
disease on an alien host (naïve or coevolved) which supports disease on a host native to the new geographic area that could not be sustained in the absence of the alien host;
disease on alien and native hosts; and
disease on a host native to the new geographic area but not on an alien host.

Countries’ efforts to conduct pest risk analyses are unlikely to be straightforward – or even possible – with so many disease scenarios

Paap et al. proceed to compare introductory pathways under the CBD categorization and plant pathology. In doing so they point out several aspects of introduction, establishment, and spread that are specific to pathogens. For example, trees’ long life spans and inability to adapt as rapidly as the micro-organism increase their vulnerability to devastating disease outbreaks following the arrival of a novel pathogen.

Participants in the Montesclaros meeting that drafted an early critique of international phytosanitary procedures

Paap et al. reinforce points made by other critics of current phytosanitary programs. (See my earlier blogs under the category “plants as pest vectors”.) In particular, they point out the weakness of visual inspection and note that new molecular assays can detect only known microorganisms. An additional complication is that DNA can persist in soil and plant tissue after death of the organism, leading to false positives. RNA is cannot yet be used as a viability marker.

Paap et al. provide three case studies to illustrate in greater depth several major challenges encountered when managing invasive forest pathogens. Most of these weaknesses are well known to forest pathologists.

1. The inconspicuous nature of microorganisms

As noted by Paap et al. and other authors, the difficulty detecting microbes is exacerbated by the huge volumes of goods, especially live plants, in international trade; the small proportion of those plants that can be inspected; the weakness of visual examination; application of fungicides and fertilizers before export that suppress symptoms. The chosen example is the oomycete genus Phytophthora, specifically P. ramorum.

2. Cryptic status of many species

Current biosecurity programs rely on naming the organism and its place of origin. This is actually impossible for many microorganisms. The tardy response to ash dieback (Hymenoscyphus fraxineus) in Europe illustrates the delay in determining the causal agent and its geographic origin. During this nearly two-decade period the possibility of preventing spread was lost.

3. Rapid evolution

Rapid evolution of the introduced pathogen can overcome resistance in a host. The example described is Cronartium ribicola (causal agent of white pine blister rust) on Western white pine (Pinus monticola) and sugar pine (P. lambertiana). They also mention the threat from hybridization between previously isolated populations, specifically Phytophthora x alni causing a devastating decline of black alder in Europe.

Sugar pine in Sequoia National Park; photo by S. Rae via Flickr

Paap et al. call for increased research to increase our knowledge of microbial diversity, especially in taxonomically rich and poorly studied ecosystems. They praise sentinel plantings as a powerful tool for early warning of pathogen threats.

SOURCES

Blackburn, T.M., P. Pysek, S. Bacher, J.T. Carlton, R.P. Duncan, V. Jarosik, et al. A proposed unified framework for biological invasions. Trends Ecol Evol. 2011; 26(7):333-9.

Paap, T., M.J. Wingfield, T.I. Burgess, J.R.U. Wilson, D.M. Richardson, A. Santini. 2022. Invasion Frameworks: a Forest Pathogen Perspective. FOREST PATHOLOGY https://doi.org/10.1007/s40725-021-00157-4

Posted by Faith Campbell

Forest Pest Threat to Africa

Eucalyptus plantation in Kwa-Zulu-Natal, South Africa; Kwa-Zulu-Natal Dept. of Transportation

Graziosi et al. (full citation at the end of the blog) point out that trees are crucial for Africa’s future. Eight hundred of the 4,500–6,000 indigenous tree species provide significant food. As elsewhere, trees provide wood and other extractive resources essential for economic growth. They also support biodiversity and mitigate current and impending climatic variations. Africa– especially the Sub-Saharan countries – is already considered highly vulnerable to climate change.

According to Graziosi et al., the cumulative economic impact of all invasive species in Africa is expected to exceed $1.2 billion per year. The total invasion cost as a proportion of GDP for many African countries is among the highest in the world. This raises the stakes for developing locally appropriate management strategies across the continent.

Responding effectively to this threat is hampered by gaps in data as well as some countries’ limited capacity for biosecurity. Graziosi et al. say that improved knowledge of taxonomy, distribution, and damage caused by these organisms is essential. Such knoledge will be crucial to develop continent-wide strategies to manage this emergency and to enhance capacity for country-level interventions.

Native and alien pests. Indigenous and plantation trees

Africa’s trees and their services are threatened by both native pests and accelerating introductions of pests and diseases from elsewhere. Long-established and new invaders increasingly affect planted forests of exotic eucalypts, pines, and Australian acacias, as well as important indigenous trees. Graziosi et al. note that the U.N. Food and Agriculture Organization (FAO) in an annex to a report issued in 2009 recorded about100 species of forest pests affecting trees in planted and natural forests across Africa. Half are native insects and pathogens, a third are alien; about 15% are of unknown origin. Considering all pests, broadleaf trees (predominantly native) are most affected.

The result is damage from the local – e.g., to rural livelihoods – to the continental – e.g., to economic development and biological diversity across Africa. Moreover, pests exacerbate widespread loss of forest cover. Overall, African forests are shrinking at the rate of almost 0.5% annually. This deforestation is affecting particularly natural forests; planted forests are actually growing 1.3% annually.

Exotic plantation trees face severe threats. More than 47 native and 19 non-indigenous defoliators, sap-feeders, wood- and shoot-borers attack plantations of Acacia spp., Eucalyptus spp., Pinus spp., and teak (Tectona grandis). About 90% of pathogens of plantation forestry are either non-indigenous or of uncertain origin. Eucalyptus alone are severely damaged by 15 species of pathogens. These organisms are listed in Tables 1 and 2.

Numerous native insect species, known as pests of indigenous trees, have reportedly widened their host range and now damage exotic trees too. Some introduced insects appear to pose significant threats to native tree species. One example is the Cypress aphid Cinara cupressi, which is attacking both exotic cypress plantations and the native African cedar Juniperus procera. Some fungi in the family Botryosphaeriaceae are latent pathogens infecting a wide range of hosts including indigenous Acacia. Dieback of large baobab trees was recently reported from southern Africa. While various microorganisms are associated with these symptoms, the specific cause is still uncertain.

A baobab tree in Limpopo region of South Africa; Wikimedia

The risk currently appears to be particularly high in South Africa. The country’s flora is highly diverse and has a high level of endemism. In fact, South Africa is home to the Earth’s smallest floral kingdom, the Cape Floral Kingdom. It is also the apparent hot spot for pest introductions from overseas (see below). Phytophthora cinnamomi is attacking native Proteaceae in South Africa. According to Graziosi et al., an “incredible diversity” of Phytophthora taxa is present, portending threats to additional plant species. Other pathogens are attacking native conifers in the Podocarpus genus, Ekebergia capensis (Meliaceae), and Syzygium trees.

*Protea repens* and fynbos vegetation near Table Mountain; photo by Mike Wingfield

There is a clear pattern to further spread: pests first introduced to South Africa often spread. Examples include several insects and pathogens on Eucalyptus and the wood-boring pest of pine Sirex noctilio. This pattern is explained by two main factors. South Africa has a high capacity to detect introduced species. Also, there is an active plantation forestry sector that imports propagules. This offers opportunities for contaminating organisms to be introduced simultaneously.

Furthermore, as Graziosi et al. note, determining the geographic origin of significant proportion of pathogens is extremely difficult – an issue I will discuss in a separate blog based on a publication by primarily South African scientists. Some non-indigenous pathogens have been on the African continent for a long time. The Armillaria root rot pathogen apparently was introduced to South Africa with potted plants from Europe in the 1600s! They note also that many non-indigenous pathogens are probably already established on the continent but not yet detected due to the organisms’ cryptic nature and lagging detection abilities.

The future of African forests

African countries expect economic growth with associated increased trade with countries off-continent. The probable result will be to accelerate the rate of species introductions and spread. However, as climate change worsens, managers will find it increasingly difficult both to predict introduced species’ impact and to implement management programs.

This led Graziosi et al. to call for urgent improvements in plant biosecurity across the continent. They advocate improved coordination at regional and international levels. The list of needed actions is a familiar one: development and application of improved diagnostic tools, updated plant exchange regulations, and revised trade policies.

Graziosi et al. also call for development of effective control and management options. They suggest biocontrol, innovative silviculture practices, and selection of resistant trees. The good news is that African countries have already initiated programs to conserve tree germplasm and domesticate indigenous species, including establishment of field gene banks of high-priority indigenous trees. I have previously praised South African efforts, specifically reports here and here.

Mudada, Mapope, and Ngezimana (2022) describe the risk from introduced species to agriculture and human well-being in southern Africa beyond forestry. The region is already ravaged by food insecurities and hidden hunger. It would be devastated if the global average of crop loss due to plant diseases (10-16%) occurs there. They say these losses can be avoided with improved biosecurity mechanisms focused primarily on pest exclusion and plant quarantine regulations.

SOURCES

Graziosi, I. M. Tembo, J. Kuate, A. Muchugi. 2020 Pests and diseases of trees in Africa: A growing continental emergency. Plants People Planet DOI: 10.1002/ppp3.31

Mudada, N. Mapope, N., and Ngezimana, W. 2022 – The threat of transboundary plant pathogens to agricultural trade in Southern Africa: a perspective on Zimbabwe’s plant biosecurity – A review. Plant Pathology & Quarantine 12(1), 1–33, Doi 10.5943/ppq/12/1/1

Posted by Faith Campbell

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

Imports from Asia Continue to Surge; Awaiting Better Analysis

The surge in US imports from Asia that began in the second half of 2020 continued through 2021 and into January 2022. As of September 2021, import volumes from Asia averaged almost 20% higher than the historical monthly average for every month of 2021 (Mongelluzzo, October 13, 2021). The surge continued into 2022. In January 2022, US containerized imports from Asia hit the highest monthly total ever recorded — 1.7 million TEU. This was a 14.6% increase over December 2021 – and a 4.5% increase from a year earlier (January 2021). [Mongelluzzo Feb 23]

The 2022 increase in import volumes was on top of the record-breaking levels seen in 2021. For example, average monthly import volumes during 2021 at the principal ports for receipt of goods from Asia — Los Angeles-Long Beach — were 23% over the 2019 average (Mongeluzzo April 2021).

Increases in volume from December 2021 occurred at ports across the country. Pacific coast ports saw increases – 25.8% at the LA/LB complex (which handles ~50% of US imports from Asia); 39.1% at Northwest Seaport Alliance (Seattle and Tacoma); 19.7% at Oakland. So did ports in the Southeast – 12.7% in Savannah and 14.1% in Charleston. However, New York/New Jersey saw a decrease of 2.2% and Norfolk saw a decrease of 10.6%. [Mongelluzzo Feb 23] New York had seen a steep increase in mid-2021 (Angell Dec. 22, 2021), but apparently this did not hold up through the year.

The southern California ports report that ships leaving China in early March will – as expected – increase import volumes before the end of the month. Long Beach projected that numbers of arriving shipping containers will rise 34% in the week beginning March 20, compared with the week of March 7; Los Angeles projected an increase of 63% [Mongelluzzo March 10].

Volumes Will Probably Continue to Rise Along the Gulf

Containerized imports from Asia through US Gulf ports had risen 27.2% to 1.14 million TEU in 2021. At the port of Mobile, specifically, imports from Asia last year rose 25% from 2020 to 230,347 TEU in 2021. Imports from Asia through Houston jumped 34 % to 807,376 TEU in 2021 [Mongelluzzo Feb 2 2022]

Increasing manufacturing and distribution industries in the Gulf region are probably an important factor in rising import volumes there. Mongelluzzo Feb 2 2022 notes the presence of a Hyundai factory in Alabama, a Tesla factory and Amazon fulfillment center near Austin, as well as several retail chains’ distribution centers near Houston. Many of these facilities opened in 2021.

Import volumes entering via Gulf and Southeastern ports are expected to continue growing in coming months and years. Several carriers have announced new direct Asia-to-US-east coast transport services. These include South Korea’s HMM (to Houston); CMA CGM; and Maersk (Vietnam and China to Houston and Norfolk; China and Indonesia to Charleston and Newark)

Those who follow shipping expect import volumes to drop in February because many factories in Asia were closed for two weeks or more for the Lunar New Year holidays, which began on Febrary 1. Imports should surge again in March. [Mongelluzzo Feb 23]

The Risk

Remember, Asia is the origin of many of the most damaging forest pests. These include Asian longhorned beetle, emerald ash borer, redbay ambrosia beetle, phytophagous and Kuroshia shot hole borers (for profiles of each visit here). Indeed, 15 of 16 non-native bark beetles in the Xyleborini (a tribe of ambrosia beetles) detected in the United States since 2000 are from Asia (Bob Rabaglia, USFS Forest Health Protection, presentation at IUFRO meeting in Prague, September 2021).

It seems to me that the beetles native to southeast Asia, e.g., the phytophagous and Kuroshio shot hole borers, are likely to find the climate along the Gulf of Mexico to their liking. Indeed, the redbay ambrosia beetle profile already has!

dead redbay in Georgia killed by laurel wilt disease

Li et al. (2021) assessed fungi associated with Eurasian bark and ambrosia beetles and their potential to impact North American trees. They assessed 111 fungal associates of 55 beetle species. They found that none was “highly virulent” on four important pines or oaks of the Southeast. However, I note two caveats. First, they tested only four host species – two pines (Pinus taeda and P. elliottii var. elliottii) and two oaks (Quercus shumardii and Q. virginiana). They did not test against the many other tree species that comprise important components of forests of the region. Second, their bar for concern was extremely high: to qualify as “highly virulent,” the pathogens had to be as damaging as laurel wilt disease or Dutch elm disease! Both have had extremely damaging impacts on their hosts across North America.

Updated Haack Analysis

As has been documented repeatedly (e.g., my blogs), the current approach to curtailing pest introductions associated with wood packaging is not sufficiently effective. Customs officials continue to detect live quarantine pests in wood packaging as it enters the country. However, the exact level of this threat is unclear since the only assessment was based on data from 2009 (Haack et al., 2014). I eagerly await the results of Bob Haack’s updated analysis, which I hope will be published by mid-year.

SOURCES

Angell, M. NY-NJ vessel backlog creeps back up amid bigger ship calls. Journal of Commerce. Dec. 22, 2021 https://www.joc.com/maritime-news/ny-nj-vessel-backlog-creeps-back-amid-bigger-ship-calls_20211222.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%2012/23/21%20SUBSCRIBER%20%28Copy%29_PC00000_e-production_E-122936_KB_1223_0617

Angell, M. Maersk to debut new Houston, Norfolk trans-Pac service in March. Journal of Commerce. Feb. 10, 2022 https://www.joc.com/maritime-news/container-lines/maersk-line/maersk-debut-new-houston-norfolk-trans-pac-service-march_20220210.html?utm_campaign=CL_JOC%20Ports%202%2F16%2F22%20_PC00000_e-production_E-127385_TF_0216_0900&utm_medium=email&utm_source=Eloqua

Haack, R.A., K.O. Britton, E.G. Brockerhoff, J.F. Cavey, L.J. Garrett. 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

Li, Y., C. Bateman, J. Skelton, B. Wang, A. Black, Y-T. Huang, A. Gonzalez, M.A. Jusino, Z.J. Nolen, S. Freeman, Z. Mendel, C-Y. Chen, H-F. Li, M. Kolařík, M. Knížek, J-H. Park, W. Sittichaya, P. H. Thai, S. Ito, M. Torii, L. Gao, A.J. Johnson, M. Lu, J. Sun, Z. Zhang, D.C. Adams, J. Hulcr. 2021. Pre-invasion assessment of exotic bark beetle-vectored fungi to detect tree-killing pathogens. Phytopathology. https://doi.org/10.1094/PHYTO-01-21-0041-R

Mongeluzzo, B. Additional port capacity alone can’t solve congestion issues: LA-LB. Journal of Commerce. April 2021 https://www.joc.com/port-news/us-ports/additional-port-capacity-alone-can%E2%80%99t-solve-congestion-issues-la-lb_20210407.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%204%2F8%2F21_PC00000_e-production_E-95420_KB_0408_0837

Mongelluzzo, B. September imports show no relief for stressed US ports. Journal of Commerce. Oct. 12, 2021. https://www.joc.com/port-news/us-ports/september-imports-show-no-relief-stressed-us-ports_20211013.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%2010%2F14%2F21_PC00000_e-production_E-116084_KB_1014_0617

Mongelluzzo, B. Gulf Coast import growth propels regional warehousing boom. Journal of Commerce. Feb. 2, 2022. https://www.joc.com/port-news/us-ports/port-mobile/gulf-coast-import-growth-propels-regional-warehousing-boom_20220202.html?utm_campaign=CL_JOC%20Ports%202%2F9%2F22%20%20_PC00000_e-production_E-126647_TF_0209_0900&utm_medium=email&utm_source=Eloqua

Mongelluzzo, B. Asian imports to US surged to new record in January. Journal of Commerce. Feb 23, 2022

https://www.joc.com/maritime-news/container-lines/asian-imports-us-surged-new-record-january_20220223.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%202%2F24%2F22%20NONSUBSCRIBER_PC015255_e-production_E-128466_KB_0224_0617

Mongelluzzo, B. Coming LA-LB cargo surge to rebuild vessel backlog, say terminals. Journal of Commerce. March 10, 2022. https://www.joc.com/port-news/us-ports/coming-la-lb-cargo-surge-rebuild-vessel-backlog-say-terminals_20220310.html?utm_source=Eloqua&utm_medium=email&utm_campaign=CL_JOC%20Daily%203/11/22%20NONSUBSCRIBER_PC015255_e-production_E-130350_KB_0311_0617