forest pest insects – Center for Invasive Species Prevention

Scientists: Introduced forest pest reshaping forests, with many bad consequences … will regulators step up?

The number of introduced forest pathogens are increasing – creating a crisis that is recognized by more scientists. These experts say tree diseases are reshaping both native and planted forests around the globe. The diseases are threatening biodiversity, ecosystem services, provision of products, and related human wellbeing. Some suggest that bioinvasions might threaten forests as much as climate change, while also undermining forests’ role in carbon sequestration.

Unfortunately, I see little willingness within the plant health regulatory community to tackle improving programs to slow introductions. Even when the scientists documenting the damage work for the U.S. Department of Agriculture – usually the U.S. Forest Service — USDA policy-makers don’t act on their findings. [I tried to spur a conversation with USDA 2 years ago. So far, no response.]

counties where beech leaf disease has been detected

What the scientists say about these pests’ impacts

Andrew Gougherty (2023) – one of the researchers employed by the USDA Forest Service – says that emerging infectious tree diseases are reshaping forests around the globe. Furthermore, new diseases are likely to continue appearing in the future and threaten native and planted forests worldwide. [Full references are provided at the end of the blog.] Haoran Wu (2023/24) – a Master’s Degree student at Oxford University – agrees that arrival of previously unknown pathogens are likely to alter the structure and composition of forests worldwide. Weed, Ayers, and Hicke (2013) [academics] note that forest pests — native and introduced — are the dominant sources of disturbance to North American forests. They suggest that, globally, bioinvasions might be at least as important as climate change as threats to the sustainability of forest ecosystems. They are concerned that recurrent forest disturbances caused by pests might counteract carbon mitigation strategies.

Scientists have proclaimed these warnings for years. Five years ago, Fei et al. (2019) reported that the 15 most damaging pests introduced to the United States — cumulatively — had already caused tree mortality to exceed background levels by 5.53 teragrams of carbon per year. As these 15 pests spread and invasions intensify, they threaten 41.1% of the total live forest biomass in the 48 coterminous states. Poland et al. (2019) (again – written by USFS employees) document the damage to America’s forest ecosystems caused by the full range of invasive species, terrestrial and aquatic.

Fei et al. and Weed, Ayers, and Hicke (2013) also support the finding that old, large trees are the most important trees with regard to carbon storage. This understanding leads them to conclude that the most damaging non-native pests are the emerald ash borer, Dutch elm disease fungi, beech bark disease, and hemlock woolly adelgid. As I pointed out in earlier blogs, other large trees, e.g., American chestnut and several of the white pines, were virtually eliminated from much of their historical ranges by non-native pathogens decades ago. These same large, old, trees also maintain important aspects of biological diversity.

It is true that not all tree species are killed by any particular pest. Some tree genera or species decrease while others thrive, thus altering the species composition of the affected stands (Weed, Ayers, and Hicke). This mode of protection is being undermined by the proliferation of insects and pathogens that cumulatively attack ever more tree taxa. And while it is true that some of the carbon storage capacity lost to pest attack will be restored by compensatory growth in unaffected trees, this faster growth is delayed by as much as two or more decades after pest invasions begin (Fei et al.).

ash forest after EAB infestation; Photo by Nate Siegert, USFS

Still, despite the rapid rise of destructive tree pests and disease outbreaks, scientists cannot yet resolve critical aspects of pathogens’ ecological impacts or relationship to climate change. Gougherty notes that numerous tree diseases have been linked to climate change or are predicted to be impacted by future changes in the climate. However, various studies’ findings on the effects of changes in moisture and precipitation are contradictory. Wu reports that his study of ash decline in a forest in Oxfordshire found that climate change will have a very small positive impact on disease severity through increased pathogen virulence. Weed, Ayers, and Hicke go farther, making the general statement that despite scientists’ broad knowledge of climate effects on insect and pathogen demography, they still lack the capacity to predict pest outbreaks under climate change. As a result, responses intended to maintain ecosystem productivity under changing climates are plagued by uncertainty.

Clarifying how disease systems are likely to interact with predicted changes in specific characteristics of climate is important — because maintaining carbon storage levels is important. Quirion et al. (2021) estimate that, nation-wide, native and non-native pests have decreased carbon sequestration by live forest trees by at least 12.83 teragrams carbon per year. This equals approximately 9% of the contiguous states’ total annual forest carbon sequestration and is equivalent to the CO₂ emissions from more than 10 million passenger vehicles driven for one year. Continuing introductions of new pests, along with worsening effects of native pests associated with climate change, could cause about 30% less carbon sequestration in living trees. These impacts — combined with more frequent and severe fires and other forest disturbances — are likely to negate any efforts to improve forests’ capacity for storing carbon.

Understanding pathogens’ interaction with their hosts is intrinsically complicated. There are multiple biological and environmental factors. What’s more, each taxon adapts individually to the several environmental factors. Wu says there is no general agreement on the relative importance of the various environmental factors. The fact that most forest diseases are not detected until years after their introduction also complicates efforts to understand factors affecting infection and colonization.

The fungal-caused ash decline in Europe is a particularly alarming example of the possible extent of such delays. According to Wu, when the disease was first detected – in Poland in 1992 – it had already been present perhaps 30 years, since the 1960s. Even then, the causal agent was not isolated until 2006 – or about 40 years after introduction. The disease had already spread through about half the European continent before plant health officials could even name the organism. The pathogen’s arrival in the United Kingdom was not detected until perhaps five years after its introduction – despite the country possessing some of the world’s premier forest pathologists who by then (2012) knew what they to look for.

Clearly, improving scientific understanding of forest pathogens will be difficult. In addition, effective policy depends on understanding the social and economic drivers of trade, development, and political decisions are primary drivers of the movement of pathogens. Wu calls for collaboration of ecologists, geneticists, earth scientists, and social scientists to understand the complexity of the host-pathogen-surrounding system. Bringing about this new way of working and obtaining needed resources will take time – time that forests cannot afford.

However, Earth’s forests are under severe threat now. Preventing their collapse depends on plant health officials integrating recognition of these difficulties into their policy formulation. It is time to be realistic: develop and implement policies that reflect the true level of threat and limits of current science.

Background: Rising Numbers of Introductions

Gougherty’s analysis of rising detections of emerging tree diseases found little evidence of saturation globally – in accord with the findings of Seebens et al. (2017) regarding all taxa. Relying on data for 24 tree genera, nearly all native to the Northern Hemisphere, Gougherty found that the number of new pests attacking these tree genera are doubling on average every 11.2 years. Disease accumulation is increasing rapidly in both regions where hosts are native and where they are introduced, but more rapidly in trees’ native ranges.This finding is consistent with most new diseases arise from introductions of pathogens to naïve hosts.

Gougherty says his estimates are almost certainly underestimates for a number of reasons. Countries differ in scientific resources and their scientists’ facility with English. Scientists are more likely to notice and report high-impact pathogens and those in high-visibility locations. Where national borders are closer, e.g., in Europe, a minor pest expansion can be reported as “new” in several countries. New pathogens in North America appear to occur more slowly, possibly because the United States and Canada are very large. He suggests that another possible factor is the U.S. (I would add Canada) have adopted pest-prevention regulations that might be more effective than those in place in other regions. (See my blogs and the Fading Forest reports linked to below for my view of these measures’ effectiveness.)

Wu notes that reports of tree pathogens in Europe began rising suddenly after the 1980s. He cites the findings by Santini et al. (2012) that not only were twice as many pathogens detected in the period after 1950 than in the previous 40 years, the region of origin also changed. During the earlier period, two-thirds of the introduced pathogens came from temperate North America. After 1950, about one-third of previously unknown disease agents were from temperate North America. Another one-third was from Asia. By 2012, more than half of plant infectious diseases were caused by introduction of previously unknown pathogens.

What is to be done?

Most emerging disease agents do not have the same dramatic effects as chestnut blight in North America, ash dieback in Europe, or Jarrah dieback in Australia. Nevertheless, as Gougherty notes, their continued emergence in naïve biomes increases the likelihood of especially damaging diseases emerging and changing forest community composition.

Gougherty calls for policies intended to address both the agents being introduced through trade, etc., and those that emerge from shifts in virulence or host range of native pathogens or changing environmental conditions. In his view, stronger phytosanitary programs are not sufficient.

Wu recommends enhanced monitoring of key patterns of biodiversity and ecosystem functioning, He says these studies should focus on the net outcome of complex interactions. Wu also calls for increasing understanding of key “spillover” effects – outcomes that cannot be currently assessed but might impact the predicted outcome. He lists several examples:

the effects of drought–disease interactions on tree health in southern Europe,
interaction between host density and pathogen virulence,
reproductive performance of trees experiencing disease,
effect of secondary infections,
potential for pathogens to gain increased virulence through hybridization.
potential for breeding resistant trees to create a population buffer for saving biological diversity. Wu says his study of ash decline in Oxfordshire demonstrates that maintaining a small proportion of resistant trees could help tree population recovery.

Quirion et al. provide separate recommendations with regard to native and introduced pests. To minimize damage from the former, they call for improved forest management – tailored to the target species and the environmental context. When confronting introduced pests, however, thinning is not effective. Instead, they recommend specific steps to minimize introductions via two principal pathways, wood packaging and imports of living plants. In addition, since even the most stringent prevention and enforcement will not eliminate all risk, Quirion et al. advocate increased funding for and research into improved strategies for inspection, early detection of new outbreaks, and strategic rapid response to newly detected incursions. Finally, to reduce impacts of established pests, they recommend providing increased and more stable funding for classical biocontrol, research into technologies such as sterile-insect release and gene drive, and host resistance breeding.

Remember: reducing forest pest impacts can simultaneously serve several goals—carbon sequestration, biodiversity conservation, and perpetuating the myriad economic and societal benefits of forests. See Poland et al. and the recent IUCN report on threatened tree species.

SOURCES

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

Clark, P.W. and A.W. D’Amato. 2021. Long-term development of transition hardwood and Pinus strobus – Quercus mixedwood forests with implications for future adaptation and mitigation potential. Forest Ecology and Management 501 (2021) 119654

Fei, S., R.S. Morin, C.M. Oswalt, and A.M. 2019. Biomass losses resulting from insect and disease invasions in United States forests. Proceedings of the National Academy of Sciences. www.pnas.org/cgi/doi/10.1073/pnas.1820601116

Gougherty AV (2023) Emerging tree diseases are accumulating rapidly in the native and non-native ranges of Holarctic trees. NeoBiota 87: 143–160. https://doi.org/10.3897/neobiota.87.103525

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

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, S.L. Ladeau, and T. Weldy. 2016. Nonnative forest insects and pathogens in the United States: Impacts and policy options. Ecological Applications, 26(5), 2016, pp. 1437-1455

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.

Quirion, B.R., G.M. Domke, B.F. Walters, G.M. Lovett, J.E. Fargione, L. Greenwood, K. Serbesoff-King, J.M. Randall, and S. Fei. 2021 Insect and Disease Disturbance Correlate With Reduced Carbon Sequestration in Forests of the Contiguous US. Front. For. Glob. Change 4:716582. [Volume 4 | Article 716582] doi: 10.3389/ffgc.2021.716582

Weed, A.S., M.P. Ayers, and J.A. Hicke. 2013. Consequences of climate change for biotic disturbances in North American forests. Ecological Monographs, 83(4), 2013, pp. 441–470

Wu, H. 2023/24. Modelling Tree Mortality Caused by Ash Dieback in a Changing World: A Complexity-based Approach MSc/MPhil Dissertation Submitted August 12, 2024. School of Geography and the Environment, Oxford University.

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 https://treeimprovement.tennessee.edu/

www.fadingforests.org

New Attention to Threats to Trees — While They Worsen

ohia (*Metrosideros polymorpha*) — one subspecies designated as Vulnerable due to restricted range
The species is under attack by rapid ohia death [https://www.dontmovefirewood.org/pest_pathogen/ceratocystis-wilt-ohi-html/]

I welcome new attention to the threats posed to tree species around world.

Last week, at the conclusion of Conference of the Parties (COP) to the Convention on Biodiversity (CBD), the International Union for the Conservation of nature (IUCN) released its most recent iteration of the Red List of Threatened Species. The headline was that 38% of the world’s trees are at risk of extinction.

This is the finding of a decade-long Global Tree Assessment. The assessment was led by Botanic Gardens Conservation International and IUCN’s Species Survival Commission Global Tree Specialist Group. Partners in the effort included Conservation International, NatureServe, Missouri Botanical Garden and Royal Botanic Gardens, Kew. The project was funded primarily by Fondation Franklinia. The foundation was formed in 2005 expressly to conserve threatened tree species! I regret that I had not heard about it before.

At least 16,425 of the 47,282 tree species assessed are at risk of extinction. Trees now account for over one quarter of species on the IUCN Red List, and the number of threatened trees is more than double the number of all threatened birds, mammals, reptiles and amphibians combined. Tree species are at risk of extinction in 192 countries around the world.

No surprise: the highest proportion of threatened trees is found on islands. Island trees are at particularly high risk due to deforestation for urban development, conversion to agriculture, invasive species, pests and diseases. Climate change is increasingly threatening trees, especially in the tropics, through sea-level rise and stronger, more frequent storms.

The COP was held in Cali, Columbia. This is fitting because South America is home to the greatest diversity of trees in the world. Twenty-five percent – 3,356 out of 13,668 assessed species are at risk of extinction. Forest clearance for crop farming and livestock ranching are the largest threats on the continent. Dr Eimear Nic Lughadha, Senior Research Leader in Conservation Assessment and Analysis at the Royal Botanic Gardens, Kew, said this percentage is sure to increase as many additional tree species are described for science.

IUCN spokespeople emphasized that the loss of trees is a major threat to thousands of other plants, fungi and animals. Cleo Cunningham, Head of Climate and Forests at Birdlife International pointed out that over two-thirds of globally threatened bird species are dependent on forests. Speakers also noted that people depend on trees; over 5,000 of the tree species on the Red List are used in construction, and over 2,000 species provide medicines, food and fuels.

Sam Ross, Sustainable Business Project Analyst at ZSL, noted that “Despite growing pressure to halt worldwide deforestation by 2030, … most of the world’s 100 most significant tropical timber and pulp companies have made limited progress in disclosing their zero deforestation and traceability commitments. We must all do more to safeguard these vital forest ecosystems, especially consumer goods manufacturers, financial institutions funding forestry, and agriculture companies.”

IUCN and the Red List Partners are launching a global social media campaign to raise awareness and funds to accelerate species assessments and reassessments. The campaign will culminate at the IUCN World Conservation Congress in Abu Dhabi, in October 2025.

Impacts from Pathogens Continue to Increase

Meanwhile, in North America and elsewhere, infections by tree-killing pathogens are spreading and intensifying.

tanoak at Big Sur killed by *P. ramorum*

Phytophthora ramorum (sudden oak death)

In California, P. ramorum the statewide rate of tree infection in 2024 doubled from 2023. Expansions were most obvious in Mendocino and Del Norte counties. Worse, California has now detected a third strain of P. ramorum in its forests. The NA2 strain was first detected in Del Norte County in 2020. Now it has been found in five sites closer to the “core” of the infestation closer to San Francisco Bay. Dr. Matteo Garbelotto believes the strain – formerly known only in nurseries – had been present for some years. It appears to be more aggressive than the strain long present in forests – NA1 – and might be favored by warmer temperatures. [The EU1 strain was detected in Del Norte County in 2021.]

Oregon has been wrestling with the EU1 strain since 2015; the NA2 strain since 2021. Beginning in late 2022, authorities have discovered multiple disease outbreaks between the Rogue River and Port Orford (farther north than the area previously known to be infected). Many of these new outbreaks are the EU1 lineage. The state is struggling to carry out eradication treatments using funds from state legislative appropriations, support from USDA Forest Service and USDI Bureau of Land Management, USDA Agriculture Research Service, and direct Congressional appropriations. The last resulted from assertive lobbying!

The Government Accountability Office is studying interactions between climate change and agricultural pests; sudden oak death is one of four focal pests. The report is expected to be released in 2025.

[Most of this information is from the California Oak Mortality Task Force (COMTF) webinar on 29 October, 2024. Recording available here.]

limber pine in Rocky Mountain National Park; photo by F.T. Campbell

Cronartium ribicola White Pine Blister Rust

Limber pine (Pinus flexilis) is heavily infected by blister rust in Alberta; in its U.S. range

the disease is increasing. Scientists had been cheered by the presence of major gene resistance (MGR) in limber pine to the rust. However, a strain of blister rust in Alberta has been determined to be virulent despite this gene (Liu et al. 2024). Scientists might have to launch a breeding program to try to enhance quantitative disease resistance (QDR) in the species. Unfortunately, the frequency and level of partial resistance in limber pine has been very low in trees tested so far. Scientists now must test more limber pines to see whether some have higher levels of QDR.

Southwestern white pine (Pinus strobiformis) presents the same problem; the MGR gene might even be the same gene. Some some populations of SWWP have higher partial or quantitative disease resistance.

Beech leaf disease

BLD continues to be detected in new sites. According to Matthew Borden of Bartlett Tree Research Laboratories, since 2021, BLD has been detected in five counties in Virginia:

Prince William County — Prince William Forest Park;
Fairfax County: southern Fairfax County on the border with Prince William County (Fountainhead Park, Hemlock Overlook Park, and Meadowood Special Recreation Area), somewhat farther north (Burke Lake Park), and northern edge (Great Falls);
Loudoun County;
Stafford County – just outside the city of Fredricksburg and along the Spotsylvania river
New Kent County in Wahrani Natural Preserve

Several of these outbreaks – e.g., southern Fairfax County, Stafford County, and Loudoun County – are 20 miles or more away from other known outbreaks. Virginia Department of Agriculture staff are monitoring the disease. All these sites are near water – although the Potomac River in Loudoun County is above the fall line so narrower than at the other sites.

SOURCE

Liu, J-J., R.A. Sniezko, S. Houston, G. Alger, J. Krakowski, A.W. Schoettle, R. Sissons, A. Zamany, H. Williams, B. Rancourt, A. Kegley. 2024. A New Threat to Limber Pine (Pinus flexilis) Restoration in Alberta and Beyond: First Documentation of a Cronartium ribicola race (vcr4) Virulent to Cr4-Controlled Major Gene Resistance. Phytopathology. Published Online:25 Sep 2024 https://doi.org/10.1094/PHYTO-04-24-0129-R

Posted by Faith Campbell

www.fadingforests.org

Non-Native Moths in England: Ever Upward

*Platyperigea kadenii* — one of the moth species that feeds on native plant species introduced recently to Great Britain. Photo by Tony Morris via Flickr

Will phytosanitary agencies and the international system respond to continuing introductions of non-native species?

A new study confirms that introductions of insects continue apace, links this pattern to the horticultural trade, and examines the role of climate change in facilitating introductions. This study focuses on moths introduced to the United Kingdom (Hordley et al.; full citation at the end of the blog). The study sought to detect any trends in numbers of species establishing and the relative importance of natural dispersal vs. those assisted – intentionally or inadvertently – by human activities.

The authors determined that moths continue to be introduced by both processes; there is no sign of “saturation”. This finding agrees with that of Seebens and 44 others (2017; citation below), which analyzed establishments of all types of non-native species globally. The British scientists found that rapidly increasing global trade is the probable driver of the recent acceleration of human-assisted introductions. They emphasize the horticultural trade’s role specifically. Climate change might play a role in facilitating establishment of species entering the UK via human activities.

Hordley et al. found that long-term changes in climate, not recent rapid anthropogenic warming, was important in facilitating introductions of even those moth species that arrived without human assistance. As they note, temperatures in Great Britain have been rising since the 17^th Century. These changes in temperature have probably made the British climate more suitable for a large number of Lepidoptera. The data show that the rate of natural establishments began rising in the 1930s, 60 years before anthropogenic changes in temperatures became evident. Hordley et al. point out that an earlier study that posited a more significant role for climate change did not distinguish between insect species which have colonized naturally and those benefitting from human assistance.

The authors expect introductions to continue, spurred by ongoing environmental and economic changes. Fortunately, very few of the introduced moths had any direct or indirect negative impacts. (The box-tree moth (Cydalima perspectalis) is the exception. [Box-tree moth is also killing plants in North America.]

boxtree moth; photo by Tony Morris via Flickr

Still, they consider that introductions pose an ongoing potential risk to native biodiversity and related human interests. Therefore, they advocate enhanced biosecurity. Specifically, they urge improved monitoring of natural colonizations and regulation of the horticultural trade.

Hordley et al. estimated the rate of establishment during the period 1900 – 2019 for (i) all moth species; (ii) immigrants (i.e., those introduced without any human assistance); (iii) immigrants which feed on native hosts; (iv) immigrants which feed on non-native hosts; (v) adventives (i.e., species introduced with human assistance); (vi) adventives which feed on native hosts; and (vii) adventives which feed on NIS hosts.

Their analysis used data on 116 moth species that have become established in Great Britain since 1900. Nearly two-thirds of these species – 63% – feed on plant species native to Great Britain; 34% on plant species that have been imported – intentionally or not. Data were lacking on the hosts of 3 species.

Considering the mode of introduction, the authors found that 67% arrived through natural colonization; 33% via human assistance. Sixty-nine percent of the 78 species that were introduced through natural processes (54 species) feed on plant species native to Great Britain; 31% (24 species) feed on non-native plants. Among the 38 species whose introduction was assisted by human activities, one-half (19 species) feed on native plant species; 42% (16 species) feed on introduced hosts.

Regarding trends, they found that when considering all moth species over the full period, 21.5% more species established in each decade than in the previous decade. This average somewhat obscured the startling acceleration of introductions over time: one species was reported as established in the first decade (1900–1909) compared to 18 species in the final decade (2010–2019).

The rate of introduction for all immigrant (naturally introduced) species was 22% increase per decade. Considering immigrant species that feed on native plants, the rate of establishment was nearly the same – 23% increase per decade – when averaged over the 120-year period. However, a more detailed analysis demonstrated that these introductions proceeded at a steady rate until 1935, then accelerated by 11% per decade thereafter. In contrast, immigrants that feed on non-native plants have maintained a steady rate of increasing establishments – 13% per decade since 1900.

Adventive species (those introduced via human assistance) increased by 26% per decade. The data showed no signs of saturation. The rates of introduction were similar for adventives that feed on both native plants (22%) and non-native hosts (26%). Again, additional analysis demonstrated a break in rates for adventives that feed on native hosts. The rate was steady until the 1970s, then significantly increased during the years up to 2010. (The scientists dropped data from the final decade since lags in detection might artificially suppress that number.)

In summary, Hordley et al. found no significant differences in trends between

the number of species that established naturally (20%) vs. adventives (26%).
immigrant or adventive species that feed on native vs. non-native hosts.

The authors discuss the role of climate change facilitating bioinvasion by spurring natural dispersal, changing propagule pressure in source habitats, changing the suitability of receiving habitat, and changing in pathways for natural spread, e.g., altered wind and ocean currents. They recognize that the two modes of colonization – adventives and immigrants – can interact. They stress, however, that the two colonization modes require different interventions.

Although their findings don’t support the premise that a surge of natural colonizers has been prompted by anthropogenic warming, Hordley et al. assert that climate clearly links to increased moth immigration to Britain and increased probability of establishment. They note that even so assisted, colonists still must overcome both the natural barrier of the English Channel and find habitats that are so configured as to facilitate breeding success. They report that source pools do not appear to be depleted — moth species richness of neighboring European countries greatly exceeds that in Great Britain.

I would have liked to learn what factors they think might explain the acceleration in both natural and human-assisted introductions of species that feed on plant species native to Great Britain. In 2023 I noted that scientists have found that numbers of established non-native insect species are driven primarily by diversity of plants – both native and non-indigenous.

Hordley et al. assert that Great Britain has advantages as a study location because as a large island separated from continental Europe by the sea – a natural barrier – colonization events are relatively easy to detect. However the English Channel is only 32 km across at its narrowest point. I wonder, whether this relatively narrow natural barrier might lead to a misleadingly large proportion of introduced species being natural immigrants. I do agree with the authors that moths are an appropriate focal taxon because they are sensitive to climate and can be introduced by international trade. Furthermore, Britain has a long tradition of citizen scientists recording moth sightings, so trends can be assessed over a long period.

Hordley et al. stress that they measured only the temporal rate of new species’ establishments, not colonization pressure or establishment success rate. They had no access to systematic data regarding species that arrived but failed to establish. Therefore, they could not deduce whether the observed increase in establishment rates are due to:

(1) more species arriving—due either to climate-driven changes in dispersal or to accessibility of source pools; or

(2) higher establishment success due to improved habitat and resource availability; or

(3) both.

Hordley et al. noted two limitations to their study. First, they concede that there is unavoidably some subjectivity in classifying each species as colonizing naturally or with human assistance. They tried to minimize this factor by consulting two experts independently and including in the analysis only those species on which there was consensus.

Second, increases in detection effort and effectiveness might explain the recent increases in establishment rates. They agree that more people have become “citizen scientists” since 1970. Also, sampling techniques and resources for species identification have improved considerably. They note, however, that Seebens et al. (2018) tested these factors in their global assessment and found little effect on trends.

Hordley et al. believe that they have addressed a third possible limitation – the lag between introduction and detection – by running their analyses both with and without data from final decade (2010-2019). The results were very similar qualitatively.

SOURCE

Hordley, L.A., E.B. Dennis, R. Fox, M.S. Parsons, T.M. Davis, N.A.D. Bourn. 2024. Increasing rate of moth species establishment over 120 years shows no deceleration. Insect Conserv. Divers. 2024;1–10. DOI: 10.1111/icad.12783

Seebens, H. et al. 2017. No saturation in the accumulation of alien species worldwide. Nature Communications. January 2017. DOI: 10.1038/ncomms14435

Seebens, H. et al. 2018. Global rise in emerging IAS results from increased accessibility of new source pools. Proceedings of the National Academy of Sciences. www.pnas.org/cgi/doi/10.1073/pnas.1719429115

Posted by Faith Campbell

www.fadingforests.org

Forest Regeneration — Need to See Holistic Picture

Research scientists in the USFS Northern Region (Region 9) – Maine to Minnesota, south to West Virginia and Missouri – continue to be concerned about regeneration patterns of the forest and the future productivity of northern hardwood forests.

The most recent study of which I am aware is that by Stern et al. (2023) [full citation at the end of this blog]. They sought to determine how four species often dominant in the Northeast (or at least in New England) might cope with climate change. Those four species are red maple (Acer rubrum), sugar maple (Acer saccharum), American beech (Fagus grandifolia), and yellow birch (Betula alleghaniensis). The study involved considerable effort: they examined tree ring data from 690 dominant and co-dominant trees on 45 plots at varying elevations across Vermont. The tree ring data allowed them to analyze each species’ response to several stressors over the 70-year period of 1945 to 2014.

In large part their findings agreed with those of studies carried out earlier, or at other locations. As expected, all four species grew robustly during the early decades, then plateaued – indicative of a maturing forest. All species responded positively to summer and winter moisture and negatively to higher summer temperatures. Stern et al. described the importance of moisture availability in non-growing seasons – i.e., winter – as more notable.

The American Northeast and adjacent areas in Canada have already experienced an unprecedented increase of precipitation over the last several decades. This pattern is expected to continue or even increase under climate change projections. However, Stern et al. say this development is not as promising for tree growth as it first appears. The first caveat is that winter snow will increasingly be replaced by rain. The authors discuss the importance of the insulation of trees’ roots provided by snow cover. They suggest that this insulation might be particularly necessary for sugar maple.

The second caveat is that precipitation is not expected to increase in the summer; it might even decrease. Their data indicate that summer rainfall – during both the current and preceding years – has a significant impact on tree growth rates.

Stern et al. also found that the rapid rise in winter minimum temperatures was associated with slower growth by sugar maple, beech, and yellow birch, as well as red maple at lower elevations. Still, temperature had less influence than moisture metrics.

Stern et al. discuss specific responses of each species to changes in temperatures, moisture availability, and pollutant deposition. Of course, pollutant levels are decreasing in New England due to implementation of provisions of the Clean Air Act of 1990.

They conclude that red maple will probably continue to outcompete the other species.

In their paper, Stern et al. fill in some missing pieces about forests’ adaptation to the changing climate. I am disappointed, however, that these authors did not discuss the role of biotic stressors, i.e., “pests”.

They do report that growth rates of American beech increased in recent years despite the prevalence of beech bark disease. They note that others’ studies have also found an increase in radial growth for mature beech trees in neighboring New Hampshire, where beech bark disease is also rampant.

For more specific information on pests, we can turn to Ducey at al. – also published in 2023. These authors expected American beech to dominate the Bartlett Experimental Forest (in New Hampshire) despite two considerations that we might expect to suppress this growth. First, 70-90% of beech trees were diseased by 1950. Second, managers have made considerable efforts to suppress beech.

Stern et al. say specifically that their study design did not allow analysis of the impact of beech bark disease. I wonder at that decision since American beech is one of four species studied. More understandable, perhaps, is the absence of any mention of beech leaf disease. In 2014, the cutoff date for their growth analysis, beech leaf disease was known only in northeastern Ohio and perhaps a few counties in far western New York and Pennsylvania. Still, by the date of publication of their study, beech leaf disease was recognized as a serious disease established in southern New England.

counties where beech leaf disease has been confirmed

Eastern hemlock (Tsuga canadensis) and northern red oak (Quercus rubra) are described as common co-occurring dominant species in the plots analyzed by Stern et al. Although hemlock woolly adelgid has been killing trees in southern Vermont for years, Stern et al. did not discuss the possible impact of that pest on the forest’s regeneration trajectory. Nor did they assess the possible effects of oak wilt, which admittedly is farther away (in New York (& here) and in Ontario, Canada, west of Lake Erie).

In contrast, Ducey at al. (2023) did discuss link to blog 344 the probable impact of several non-native insects and diseases. In addition to beech bark disease, they addressed hemlock woolly adelgid, emerald ash borer, and beech leaf disease.

Non-native insects and pathogens are of increasing importance in our forests. To them, we can add overbrowsing by deer, proliferation of non-native plants, and spread of non-native earthworms. There is every reason to think the situation will only become more complex. I hope forest researchers will make a creative leap – incorporate the full range of factors affecting the future of US forests.

I understand that such a more integrated, holistic analysis might be beyond any individual scientist’s expertise or time, funding, and constraints of data availability and analysis. I hope, though, that teams of collaborators will compile an overview based on combining their research approaches. Such an overview would include human management actions, climate variables, established and looming pest infestations, etc. I hope, too, that these experts will extrapolate from their individual, site-specific findings to project region-wide effects.

Some studies have taken a more integrative approach. Reed, Bronson, et al. (2022) studied interactions of earthworm biomass and density with deer. Spicer et al. (2023) examined interactions of deer browsing and various vegetation management actions. Hoven et al. (2022) considered interactions of non-native shrubs, tree basal area, and forest moisture regimes.

See also my previous blogs on studies of regeneration in New Hampshire, North Carolina, National parks in the East, Allegheny Plateau and Ohio, and the impact of deer.

SOURCE

Stern, R.L., P.G. Schaberg, S.A. Rayback, C.F. Hansen, P.F. Murakami, G.J. Hawley. 2023. Growth trends and environmental drivers of major tree species of the northern hardwood forest of eastern North America. J. For. Res. (2023) 34:37–50 https://doi.org/10.1007/s11676-022-01553-7

Posted by Faith Campbell

www.fadingforests.org

Two Non-Native Insects Threaten Forest, Salmonid, and Waterway Conservation in Pacific Northwest

Oregon ash dominate wetlands of Ankeny NWR; photo by Wyatt Williams, Oregon Department of Forestry

One of these insects is the emerald ash borer (EAB). We easterners have “been there & done that”. However, programs aimed at conserving wetlands and riparian areas of the Western states – and the associated species — are at least as vulnerable to loss of ash. Worse, other tree taxa, specifically oaks, and the open woodlands they inhabit — are also under threat. The ecological tragedies continue to affect ever more forests.

|Emerald Ash Borer in Oregon and British Columbia

The emerald ash borer (EAB; Agrilus planipennis) was detected in Oregon in June 2022. Officials had been expecting an introduction and had begun preparations. Unsurprisingly, the infestation is more widespread than known at first: detections in two new locations, fairly close to the original in Forest Grove, mean the infested area now occupies three neighboring counties — Washington, Yamhill, and Marion counties.

Oregon officials are trying to slow spread of EAB by removing infested trees. Surveys in Washington County had identified 190 infested ash trees; 80 were removed in April 2024. They treated healthy ash trees in Washington County with injections of the systemic insecticide emamectin benzoate. The effort was already a daunting task: the survey had disclosed 6,500 ash trees in the vicinity. The city of Portland – only 25 miles away – has 94,000 ash trees (Profita 2024).

In May, 2024 EAB was detected in the city of Vancouver in British Columbia. This detection in the sixth Canadian province adds to the threat to the ecosystems of the region. The Canadian Food Inspection Agency (CFIA) now regulates the movement of all ash material such as logs, branches, and woodchips, and all species of firewood, from the affected sites.

The CFIA is also conducting surveillance activities to determine where EAB might be present, and is collaborating with the City of Vancouver, the Vancouver Board of Parks and Recreation, the Province of British Columbia, and other stakeholders to respond to the detections and slow the spread of this pest.

Importance of Oregon ash (Fraxinus latifolia)

The Oregon ash is the only ash species native to the Pacific Northwest. Its range stretches from southern British Columbia to so California, where it has hybridized with velvet ash (F. velutina). It is highly susceptible to EAB attack; there is a high probability that Oregon ash could be rendered functionally extinct (Maze, Bond and Mattsson 2024). This vulnerability prompted the International Union for Conservation of Nature (IUCN) to classify Oregon ash as “near threatened” as long ago as 2017 (Melton et al. 2024).

Oregon ash typically grows in moist, bottomland habitats. There it is a late-successional climax species. In Oregon’s Willamette Valley and Washington’s Puget Trough, the tree improves streams’ water quality by providing shade, bank stabilization, and filtration of pollutants and excess nutrients. Maintaining these ecological services is particularly important because these streams are crucial to salmonids (salmon and trout) and other native aquatic species (Maze, Bond and Mattsson 2024).

So it is not surprising that one component of Oregonians’ pre-detection preparations was an analysis of the likely impact of widespread ash mortality on populations of salmon, trout, and other aquatic species. I summarize the key findings of Maze, Bond and Mattsson here.

According to this study, salmonids and other cold-water aquatic species suffer population declines and health effects when stream water temperatures are too warm. A critical factor in maintaining stream temperatures is shade – usually created by trees. In the Pacific Northwest many streams’ temperatures already exceed levels needed to protect sensitive aquatic species. A key driver of increased stream temperatures – at least in the Willamette Basin – is clearing of forests to allow agriculture.

Decreasing streams’ temperatures is not only a good thing to do; it is legally required by the Endangered Species Act because several salmon and steelhead trout species are listed. In one response, the Oregon Department of Environmental Quality recommends restoration and protection of riparian vegetation as the primary methods for increasing stream shading and mitigating increased stream temperatures in the lower Willamette Basin.

The forests shading many low-elevation forested wetlands and tributaries of the Willamette and lower Columbia rivers are often composed exclusively of Oregon ash. Loss of these trees’ shade will affect not just the immediate streams but also increase the temperature of mainstem waterways downstream.

Oregon ash – EAB detection site; photo by Wyatt Williams, Oregon Department of Forestry

Replacements for Oregon Ash?

The magnitude of the ecological impacts of ash mortality in the many forested wetlands in the Willamette Valley will largely be determined by what plant associations establish after the ash die. Oregon ash is uniquely able to tolerate soils inundated for extended periods. No native tree species can fill the void when the ash die. Oregon white oak (Quercus garryana), black cottonwood (Populus trichocarpa), and the alders (Alnus rubra and A. rhombifolia), are shade intolerant and unlikely to persist in later seral stages in some settings.

If non-native species fill the gaps, they will provide inferior levels of ecosystem services – I would think particularly regarding wildlife habitat and invertebrate forage. Maze, Bond and Mattsson expect loss of ash to trigger significant physical and chemical changes. These will directly impact water quality and alter native plant and animal communities’ composition and successional trajectories.

The authors cite expectations of scientists studying loss of black ash (F. nigra) from upper Midwestern wetlands. There, research indicates loss of ash from these systems is likely to result in higher water tables and a conversion from forested to graminoid- or shrub-dominated systems. Significant changes follow: to food webs, to habitat structure, and, potentially, to nitrogen cycling.

Maze, Bond and Mattsson expect similar impacts in Willamette Valley wetlands and floodplains, especially those with the longest inundation periods and highest water tables. That is, there will probably be a broad disruption of successional dynamics and, at many sites, a conversion to open, shrub-dominated systems or to wetlands invaded by exotic reed canary grass (Phalaris arundinacea), with occasional sedge-dominated (Carex obnupta) wetlands. They think this change is especially likely under canopies composed of Oregon white oak (see below). The authors admit some uncertainty regarding the trajectories of succession because 90 years of water-control projects has almost eliminated the possibility of high-intensity floods.

Oregon Ash and Salmonids

Maze, Bond and Mattsson point out that all salmonids that spawn in the Willamette basin and the nearly 250,000 square mile extent of the Columbia basin upstream of Portland pass through the two wooded waterways in the Portland area that they studied. Applying a model to simulate disappearance of ash from these forests, the authors found that the reduced shade would raise the “solar load” on one waterway, which is wide and slow-moving, by 1.8%. On the second, much narrower, creek (mean channel width of 7 m), solar load was increased by of 23.7%.

Maze, Bond and Mattsson argue that even small changes can be important. Both waterbodies already regularly exceed Oregon’s target water temperature throughout the summer. Any increase in solar loading and water temperatures will have implications for the fish – and for entities seeking to comply with Endangered Species Act requirements. These include federal, state, and local governments, as well as private persons.

The Willamette and lower Columbia Rivers, and their tributaries, traverse a range of elevations. Ash trees comprise a larger proportion of the trees in the low elevation riparian and wetland forests. Consequently, Maze, Bond and Mattsson expect that EAB-induced loss of Oregon ash will have significant impacts on these rivers’ water quality and aquatic habitats. The higher water temperatures will affect aquatic organisms at multiple trophic levels.

They conclude that the EAB invasion West of the Cascade Mountain range constitutes an example of the worst-case forest pest scenario: the loss of a dominant and largely functionally irreplaceable tree species that provides critical habitat for both ESA-listed and other species, along with degradation of ecosystem services that protect water quality.

Breeding Oregon Ash … Challenges to be Overcome

According to Melton et al. (2024), Oregon ash does not begin to reproduce until it is 30 years old. Such an extended reproductive cycle could complicate breeding efforts unless scientists are able to accelerate flowering or use grafting techniques to speed up reproduction – as suggested by Richard Sniezko, USFS expert on tree breeding.

Melton et al. (2024) note that the IUCN has recently highlighted the importance of maintaining a species’ genetic variation in order to maintain its evolutionary potential. Consequently, they examined genetic variation in Oregon ash in order to identify the species’ ability to adjust to both the EAB threat and climate change. The authors sequenced the genomes of 1,083 individual ash trees from 61 populations. These spanned the species’ range from Vancouver Island to southern California. The genetic analysis detected four genetic clusters:

British Columbia;
Washington to central Oregon – including the Columbia River and its principal tributaries;
Southwest Oregon and Northwest California — the Klamath-Siskiyou ecoregion; and
all other California populations.

Connectivity between populations (that is, the potential corridors of movement for pollen and seeds and hence, genetic flow) was greatest in the riparian areas of the Columbia River and its tributaries in the center to the species’ range. Despite this evidence of connectivity, nucleotide diversity and effective population size were low across all populations. This suggests that the patchy distribution of Oregon ash populations might reduce its long-term evolutionary potential. As average temperatures rise, the regional populations will become more distinct genetically. The species’ ability to adjust to future climate projections is most constrained in populations on Vancouver Island and in smaller river valleys at the eastern and western edges of the range. Populations in southern California might be “pre-adapted” to warmer temperatures.

The resulting lower effective population size might exacerbate risks associated with EAB. The authors warned that although seeds from more than 350 maternal parent trees have been preserved since 2019, these collections do not cover the full genomic variation across Oregon ash’s range. Some genomic variation that represents adaptive variation critical to the species’ long-term evolution might be missing. They advocate using the genetic data from their study to identify regions where additional collections of germplasm are needed for both progeny trials and for long-term conservation.

Oregon white oak with symptoms of Mediterranean oak borer infestation; photo by Christine Buhl, Oregon Department of Forestry

Oregon White Oak (Quercus garryana) and the Mediterranean Oak Borer

The U.S. Department of Interior has been working with regional partners for 10 years to protect oak and prairie habitat for five ESA-listed species, two candidate species, and numerous other plant and animal species of concern. In August 2025 the Department announced creation of the Willamette Valley Conservation Area. It becomes part of the Willamette Valley National Wildlife Refuge Complex. These units are managed predominantly to maintain winter habitat for dusky geese (a separate population of Canada geese). Other units in the Complex are William L. Finley National Wildlife Refuge, Ankeny National Wildlife Refuge, and Baskett Slough National Wildlife Refuge.

These goals too face threats from non-native forest pests. First, the forested swamps of Ankeny NWR are composed nearly 100% of ash.

Second, Oregon white oak now confronts its own non-native pest – the Mediterranean oak borer (Xyleborus monographus). This Eurasian ambrosia beetle has been introduced to the northern end of the Willamette Valley (near Troutville, Oregon). It is likely that infestations are more widespread. Authorities are surveying areas near Salem. A separate introduction has become established in California, north of San Francisco Bay plus in Sacramento County in the Central Valley. Oregon white oak is vulnerable to at least one of the fungi vectored by this borer – Raffaelea montety. https://www.dontmovefirewood.org/pest_pathogen/mediterranean-oak-borer/

SOURCES

Maze, D., J. Bond and M. Mattsson. 2024. Modelling impacts to water quality in salmonid-bearing waterways following the introduction of emerald ash borer in the Pacific Northwest, USA. Biol Invasions (2024) 26:2691–2705 https://doi.org/10.1007/s10530-024-03340-3

Melton, A.E., T.M. Faske, R.A. Sniezko, T. Thibault, W. Williams, T. Parchman, and J.A. Hamilton. 2024. Genomics-driven monitoring of Fraxinus latifolia (Oregon Ash) for conservation and emerald ash borer resistance breeding. https://link.springer.com/article/10.1007/s10530-024-03340-3

Profita, C. April 26, 2024. State crews remove trees in Washington County to slow spread of emerald ash borer. Oregon Public Broadcasting. https://www.opb.org/article/2024/04/26/oregon-invasive-beetle-emerald-ash-borer-infestation-tree-removal/#:~:text=It%20was%20first%20detected%20in%20Oregon%20in%20Forest%20Grove%20in%20June%202022.&text=This%20week%2C%20crews%20removed%20dozens,ash%20trees%20from%20the%20area.

Posted by Faith Campbell

www.fadingforests.org

What I learned at the NPB meeting

The National Plant Board’s members are the lead plant health officials of the states and territories. Many federal officials also attend – from APHIS and DHS Bureau of Customs and Border Protection. Representatives of other North American phytosanitary entities participate – i.e., Canada, Mexico, and the North American Plant Protection Organization (NAPPO). Some stakeholder groups participate, especially the nursery industry. I have attended these meetings for over a decade because they provide an overview of pest issues and programs plus an unparalleled opportunity to network. The Nature Conservancy’s Leigh Greenwood also attends. We are the only representatives of the species conservation community to attend – others are missing great opportunities.

Here, I’ve listed 10 items that are among the most important the group discussed.

1) The funding situation for APHIS is worse than I realized

APHIS Administrator Mike Watson and Deputy Administrator (for plants) Mark Davidson both spoke about the need to cut programs to stay within the limits set by congressional appropriations. Funding for APHIS, as a whole, was cut only 1% for the current year (Fiscal Year 2024), cost-of-living salary increases mean less money for programs. (I believe Dr. Watson said $41 million less for FY24). If FY25 funding is the same, Congressionally mandated additional payraises will mean an another $20 million decrease in program funding.

Dr. Davidson said that the plant programs (Plant Protection and Quarantine) had been cut by 5% in FY24. However, Congress did not finalize the funding levels until about half-way through the fiscal year – so staying within the limits required even more severe cuts to programs in the remainder of FY24. To stay within these limits, APHIS cut several programs, among them a $3.6 million cut from the “tree and wood pest” program. This meant loss of funds to manage the polyphagous and Kuroshio shot hole borers, smaller cuts for programs managing Asian longhorned beetle and emerald ash borer, and perhaps the Asian flighted spongy moth. They anticipate additional cuts in these programs in FY25. The one bright light is the Section 7721 Plant Pest & Disease Management & Disaster Prevention Program. It provides steady funding for a range of plant health programs. The House version of the still-pending Farm Bill calls for increasing funding for this program by $15 million each year.

Nearly 100% ash trees in Oregon wetland — exposed to spreading EAB. Photo by Wyatt Williams, Oregon Department of Forestry

Remember this when I ask you to lobby for appropriations! If we don’t advocate for funding the programs dealing with “our” pests, they will shrink.

Watkins and Davidson also worry that whoever is the next secretary of USDA might not support the agency when it seeks to withdraw funds to cover emergencies from the Commodity Credit Corporation – as Secretary Vilsack has.

APHIS and the DHS Customs and Border Protection (CBP) both praised a recent regulatory action that increases user fees for importers having goods cleared at ports. Kevin Harriger, CPB official in charge of agriculture programs, said the new funds would allow CBP to hire 700 new agricultural inspectors (currently there are 2,800 agricultural officials). That sounds great, but … when trade and passenger volumes crashed early in the COVID pandemic, things looked dicey for a while. Plus – as I have argued repeatedly – real protection against pest introductions will come from stronger policies, not ramped-up inspections.

Pathologist Bruce Moltzan reported on the USFS Forest Health Protection program. He pointed out that the USFS has a very limited toolbox. In this fiscal year, the program has about $48 million, after salaries, to support its activities. Projects targetting insects receive 70% of the funding; those targetting pathogens 15%.

2) Invasive hornets

Washington State has not found any new nests of the Northern (formerly Asian) giant hornet (Vespa mandarinia). Miraculous!

However, Georgia detected another species, the yellow-legged hornet (Vespa velutina), near Savannah in August 2023. The Georgia Department of Agriculture, APHIS, and the University of Georgia are working to find nests – which are located at the top of tall pine trees in residential areas. Five nests were found in 2023; another four so far in 2024. Georgia hopes to place traps 100 miles out from each detection site. Like the northern hornet, V. velutina preys on honey bees. It was probably transported by ship or with its cargo.

A third species, V. tropica, has been introduced on Guam.

3) Better Federal-State Cooperation — Sometimes

APHIS and the state phytosanitary officials have set up structures – e.g., Strategic Alliance/Strategic Initiative, or SASI – to work together more closely. CBP joins the coordinating meetings. One program described at the meeting is the effort to contain spread of the box tree moth (Cydalima perspectalis). This effort came out of discussions at last year’s Plant Board meeting, with follow-up gatherings of APHIS, the states, and the nursery industry. The moth is known to be present in New York, Massachusetts, Michigan, Ohio, and now Delaware – plus several Canadian provinces.

A second project concerns how much data to share about state detections of pests – which are recorded in the National Plant diagnostic Network database. These data have accrued over 20 years … and are sought by both other states and academic researchers. States are often reluctant to allow public review of detection data because they fear it will cause other states or private parties to avoid buying plants or other goods from the infested area. While the project team has not yet decided how to deal with these conflicts, they said they were more inclined to share “nonconsequential data” – meaning data on a pest everyone already knows is present, not a pest under regulation or a new detection. In other words, “consequential” seems to pertain to industry profits, not damage to agricultural or natural resources.

EAB-killed ash along Mattawoman Creek, Maryland. Photo by Leslie A. Brice

4) Update: 20 years of tackling the emerald ash borer

Craig Kellogg, APHIS’ chief plant health representative in Michigan, reviewed 20 years of dealing with the emerald ash borer (EAB). He is optimistic about the impact of the biocontrol agents that have now been released in 32 states and four provinces. The larval parasitoids are dispersing and EAB densities are coming down. He conceded that over-story and mature ash are still dying, but says ash in long-infested areas are regenerating well. Scientists agree (see Wilson et al. 2024; full citation at end of the blog). Woodpeckers are still the most effective biocontrol agent of EAB for over-story ash, especially in locations where introduced parasitoids are not established. So far, the growing numbers of biocontrol agents are still parasitizing too few EAB larvae to prevent decline of over-story ash trees.

5) Flighted Spongy Moths

APHIS reported on recent detections of flighted spongy moth from Asia on ships coming to U.S. ports. The program covers four closely related species of Lymantria. All have much broader host ranges than Lymantria dispar, plus the females are capable of sustained flight, so they spread more rapidly.

The principal strategy to prevent their introduction is to require ships that call at ports along the Pacific coast in Russia, China, Japan, and North and South Korea to ensure that the ships’ superstructures and cargo are clean. Before arriving at U.S. ports, the ship’s captain must inform CBP where it has called over the last 24 months. Then, CBP conducts an inspection. If CBP inspectors find a small number of egg masses, they remove the eggs and spray pesticide. If the inspectors detect a large number of egg masses, the ship is ordered to leave port, clean itself, and undergo re-inspection before it can return.

Four countries in the Americas – the U.S., Canada, Chile, and Argentina – and also New Zealand have very similar programs.

Detections follow natural changes in population levels in the exporting regions. APHIS’ program leader, Ingrid Asmundsson, reported on an unfortunate coincidence in 2014. A huge moth population outbreak occurred simultaneously with very low fuel prices in Russia. The latter attracted many ships to call there. An even bigger population surge occurred in 2019. Asmundsson expects another high-moth period this year.

flighted spongy moths infesting a ship superstructure

APHIS is working on putting this program on a regulatory foundation; this would allow the agency to be more specific in its requirements and to impose penalties (other than expulsions from ports). I’ll let you know when the proposed rule is published for comment.

6) Regional Reports: Old Pests, New Pests

Representatives of the regional plant boards informed us of their “new pest” or other concerns.

Gary Fish, president of the Eastern Plant Board, mentioned

the need for additional research on management of beech leaf disease
concern about impact of box tree moth and vascular streak dieback on the nursery industry (the latter is a threat to dogwood and redbud)
spread of elm zig-zag sawfly in Vermont and Connecticut
awareness that laurel wilt is moving into Virginia and maybe farther north.

elm zig-zag sawfly; photo by Gyorgy Csoka *via* Bugwood

There was a more general discussion of beech leaf disease. What can be done, given that the disease is so widespread that no one is regulating movement of beech. Gary Fish advised outreach and efforts to reach agreement on management approaches. Chris Benemann, of Oregon, suggested informing other states so that they can decide whether to take regulatory action. A representative of CBP urged engaging stakeholders by asking for their help.

Chris Benemann, President of the Western Plant Board, expressed concern about APHIS’ reduced funding for spongy moth detection and control efforts. She also worries about the recently detected population of Phytophthora austrocedrii in an Oregon nursery. The western states are also focused on several longstanding pest problems – grasshoppers, Japanese beetle; and a new beetle from Australia that is attacking almonds, pistachios, and walnuts.

tree infested by hemlock woolly adelgid; photo by F.T. Campbell

Megan Abraham of Indiana reported that members of the Central Plant Board are concerned about

funding decrease for APHIS and USFS efforts to slow the spread of spongy moth and manage Asian longhorned beetle and emerald ash borer;
Invaders associated with Christmas decorations, e.g., hemlock woolly adelgid, elongate hemlock scale, and balsam woolly adelgid
spotted lanternfly finding new ways to travel;
invasion by beech leaf disease and possibly by sudden oak death;
elm zig-zag sawfly entering the region;
boxwood blight, potato pests, honeybee pests, and invasive plants – e.g., Hydrilla.

She noted that nursery stock is increasingly coming from more distant – and cheaper – producers, raising the risk of new pests being introduced.

Finally, Abraham expressed concern about decreased funding at the same time as the need is growing – and asked with whom states should collaborate in order to reverse this trend.

Kenny Naylor of Oklahoma, Vice President of the Southern Plant Board, concurred that funding levels are a major concern. He mentioned specifically the spongy moth Slow the Spread program and eradication of the Asian longhorned beetle outbreak in South Carolina. Another concern is the Georgia hornet outbreak.

7) Phasing Out Post-Entry Quarantine

APHIS and the NPB have agreed to phase out the post-entry quarantine (PEQ) program. A program review revealed several problems, some of which astound me. When examining plants in quarantine the scientists still relying on visual inspection! And they are looking for pests identified 45 years ago (1980)! While I think PEQ programs can be valuable in preventing introduction of disease agents, as implemented in recent decades it does not. Twenty years ago, citrus longhorned beetles escaped from a “quarantine” area in a commercial nursery in Washington state. These Cerambycids are more than an inch long!

citrus longhorned beetle; photo by Art Wagner, USDA *via* Bugwood

Part of this phase-out is to transfer plant species harboring pests of concern to the Not Authorized for Importation Pending Pest Risk Assessment (NAPPRA). While the APHIS speaker said that NAPPRA allows the agency to act quickly when it detects evidence of pest risk, I have found lengthy delays. The third round of proposals was published in December 2019! The fourth round of species proposed for NAPPRA listing should be published soon; a fifth round is now in draft inside the agency.

8) Christmas Greens – Spreading Pests

Officials from Oregon, Maine, and Illinois described their concerns about pests being spread by shipments of various forest or plant products, especially Christmas greens. Mentioned were spongy moths, link hemlock woolly adelgid, link elongate hemlock scale, balsam woolly adelgid, link and box wood moth. Part of the challenge is that the vectoring items are often sold by unregulated outlets – multiple stores, Christmas tree lots – and through on-line or catalog outlets. There are also extreme demands on the regulatory enforcement staff during the brief holiday sales season. Several states are unsure whether they have authority over decorative products – although others pointed out that they are regulating the pest, regardless of the object for sale or type of store.

9) Pests in Firewood

Leigh Greenwood of The Nature Conservancy noted that the state agencies that issue firewood regulations – often the plant protection organization (state department of agriculture) — do a good job alerting the public about the risks and rules. However, the public looks first to state parks agencies for information about camping – and those agencies have less robust educational efforts. It is important to put the message where the public can find it when they don’t know it exists – before they include firewood from home in their camping gear.

10) Projects of the North American Plant Protection Organization

The North American Plant Protection Organization (NAPPO) is working on several projects of interest to those of us concerned about tree-killing pests. One project is evaluating risks associated with wood products, especially how well one international regulation, ISPM#15 is working for dunnage. Another projects is testing the efficacy of the heat treatment specified by ISPM#15 (50^o C for 30 minutes). A third project — almost completed – is evaluating fumigants that can be alternatives to methyl bromide.

In conclusion, each annual meeting of the National Plant Board is packed with new information, updates on current pests, and comments on by the state agencies who suggest new approached to APHIS and hold the agency to account. It is well worth attending. Information about upcoming meetings of both the national and four regional plant boards is posted on the NPB website, https://www.nationalplantboard.org/

Signatories to the APHIS-NPB strategic alliance

SOURCE

Wilson, C.J., T.R. Petrice, T.M. Poland, and D.G. McCullough. 2024. Tree species richness and ash density have variable effects on emerald ash borer biological control by woodpeckers & parasitoid wasps in post-invasion white ash stands. Environmental Entomology.

Posted by Faith Campbell

www.fadingforests.org

APHIS Annual Report Describes Helpful Programs … Since Cut Back Because of Funding Shortfalls

Flighted spongy moths infesting a ship’s superstructure

USDA’s Animal and Plant Health Inspection Service (APHIS) has issued its annual report for Fiscal Year 2023. The report is part of an enhanced outreach effort that I believe is an effort to persuade the Congress to provide additional funds. However, as I describe below, at this summer’s annual meeting of the National Plant Board, link APHIS’ leadership stated that funding shortfalls are forcing them to curtail many programs. These include ones important to those of us concerned about threats to North American trees. I applaud this action and hope it succeeds!

The report contains some good news but I consider the overall approach depressing. Tree-killing pests continue to receive little attention. The report also emphasizes APHIS’ efforts to facilitate export of agricultural products – an understandable stance given American politics.

The opening summarizes the agency’s activities includes:

Examples of programs targetting pests abroad, before they can reach the U.S. All are fresh fruits and vegetables;
APHIS or staff at U.S. borders:
- Approved (cleared) 27,235 shipmentscontaining over 1.87 billion plant units (e.g., a single plant or cutting, or vial of tissue culture plantlets) and 670,811 kilograms of seeds. They intercepted 2,176 quarantine pests. (APHIS carry out these inspections at Plant Inspection Stations – separate from the port environment where DHS Customs and Border Protection (CBP) staff inspects other cargo.)

Identified approximately 92,000 pestsfound during CBP inspections of cargo, mail, and express carrier shipments and took quick action to prevent those of concern from entering the U.S.
Facilitated entry of regulated agricultural cargo by monitoring more than 62,000 treatments of various kinds, that is, fumigations, cold or heat treatments, and irradiation.
Examples of APHIS’ efforts to slow pests’ spread within the country cited plant pest surveys — with coordinated responses — for approximately 45 pests. Also APHIS described funding to help citrus growers combat citrus greening.
The report has separate subreports on the following programs: risk analysis, pest detection, “specialty crop” pests, and tree and wood pests. The last two contain information specific to our interests.

Tree and Wood Pests

This program protects forests, private working lands, and natural resources. It targets specific pests: the Asian longhorned beetle, emerald ash borer, spongy moth, and most recently the invasive shot hole borers. The report notes that numerous native, widespread hardwood tree species are vulnerable to these pests. APHIS asserts an economic justification for the program: conserving forests enhances rural communities’ economic vitality, supports forest-related industries, and maintains the ecosystem services provided by urban trees.

Unfortunately, at this summer’s annual meeting of the National Plant Board APHIS leadership said funding shortfalls forced them to pull back on all these programs.

Programs as Described in the Report

Asian Longhorned Beetle

ALB eradication aims to protect the 30% of U.S. trees that are ALB hosts. These trees support multi-billion-dollar maple syrup, timber, tree nursery, trade, and tourism industries. After reviewing the history of ALB detections, starting in Brooklyn in August 1996, the report describes APHIS’ eradication strategy as comprising surveys, regulatory inspections and quarantine restrictions, removal of infested and high-risk trees, and chemical treatment applications. In FY 2023, the program surveyed more than 763,000 trees across the four regulated areas: New York, Massachusetts, Ohio, and South Carolina. Each program is summarized.

Good news at two locations. On Long Island: only 11 new infested trees were found after a survey of 43,480 trees. In Worcester County, Massachusetts, no new infested trees were found after surveying nearly 360,000 trees. However, in Tate Township, Ohio, surveys detected 163 new infested trees. And in

South Carolina, the program is at an earlier stage — surveying a portion of the quarantine area. The program surveyed nearly 140,000 trees and removed 1,700 in FY 2023.

At the National Plant Board Meeting, Deputy Administrator Mark Davidson explained that the FY2024 appropriation cut $3.6 million from the “tree and wood pests” account. This required the agency to reduce funding for the ALB eradication program.

Emerald Ash Borer

The report summarizes the spread of EAB since its first detection in 2002 in Michigan to 37 states and the District of Columbia (APHIS does not mention EAB’s presence in five Canadian provinces.)

Saying that EAB has spread beyond what a regulatory program can control, the report notes that APHIS ended the regulatory program in FY 2021. In FY 2023 the agency continued the transition to a program relying primarily on biocontrol. In FY2023, APHIS provided parasitoids to 155 release sites – three in Canada, the rest in 122 counties in 25 states. APHIS and cooperators continue to assess their impacts on EAB populations and tree health at release sites and nearby areas. Field evaluations indicate the EAB parasitoid wasps and other EAB natural enemies (woodpeckers) are protecting regenerating sapling ash from EAB.

Spongy Moths

Spongy moths (the species formerly called European gypsy moths) are established in all or parts of 20 eastern and midwestern states, plus the District of Columbia. APHIS and state cooperators regulate activities in the quarantine area to prevent the moths’ human-assisted spread to non-quarantine (non-infested) areas – primarily West coast states. To address the moths’ natural spread, APHIS PPQ monitors the 1,200-mile-long border of the quarantine area and adds newly infested areas to the regulated area. The USDA Forest Service – APHIS – Slow-the-Spread Foundation program has greatly reduced the moth’s rate of spread and has eradicated isolated populations.

Another component of the program aims to prevent introduction of members of the flighted spongy moth complex link from Asia. The Asian species have broader host ranges and the females can fly, so they could spread faster. A multi-nation cooperative program is designed to prevent the moths’ hitchhike on vessels coming from Asia. link APHIS supports this program through negotiations and support of CBP’s offshore vessel inspection, certification, and cleaning requirements. Canada participates in the same program.

In FY 2023, APHIS and state cooperators continued efforts to delimit possibly introductions in Washington State (no additional moths detected); and California and Oregon (initial detections in FY 2020).

California sycamore infested by polyphagous shot hole borer; photo by Beatriz Nobua-Behrmann UC Cooperative Extension

Shot Hole Borers

The report notes that various non-native shot hole borers have been detected in several states. Their hosts include trees in forests and urban landscapes, tea plantations, and avocado orchards. The program’s focus was apparently on the polyphagous and Kuroshio shot hole borers devastating riparian habitats in southern California and urban areas in other parts of California. At California’s request, APHIS and the USDA Forest Service helped establish a working group, led by USFS, with the goal of strategically addressing both shot hole borers in California. In FY 2023, APHIS’ helped with foreign explorations for possible biocontrol agents, as well as host specificity testing.

APHIS leadership told the National Plant Board in July 2024 that it had dropped this entire program due to funding shortfalls.

Specialty Crop Pests

While much of this report concerns pests of agricultural crops (e.g., grapes, citrus, potatoes), it also summarized efforts re: Phytophthora ramorum (sudden oak death) and spotted lanternfly. APHIS says its efforts protected nursery stock production worth approximately $1.3 billion as of 2019, and tree fruit production worth approximately $1.7 billion in 2021.

map showing areas of the Eastern United States at risk to *P. ramorum* – developed by Gilligan of Cambridge University

Phytophthora ramorum

The report states that APHIS seeks to limit P. ramorum’s spread from affected nurseries. The agency does this via regulatory strategies. During FY 2023, 16 nurseries were governed by more stringent rules under the federal program which are imposed on nurseries that have been determined in past years to harbor P. ramorum-infected plants.

In addition, Oregon officials continued surveys of an area outside its quarantine zone because of a detection the previous year. APHIS will adjust the federal quarantine depending on the state’s findings.

The APHIS report does not discuss several pertinent events that occurred in FY2023. [For more details, read the California Oak Mortality Task Force newsletters for 2023 – posted here.

First, APHIS does not mention or discuss the implications of detection of two new strains of P. ramorum — EU1 & NA2 — in west coast forests. The presence of EU1 in a new California county (Del Norte) was confirmed in Feb 2023.

Second, the report said that Oregon is trying to determine the extent of the P. ramorum infection detected outside the state’s quarantine zone. However, it does not mention that this outbreak involves the new NA2 lineage – and that NA2 was known to be present in nurseries in the region since 2005.

The report also does not clarify that three nurseries to added to the more stringent program were so treated because SOD-infected plants were found on their premises.

Nor does the report note that at least two new naturally-infected hosts of P. ramorum were identified: Western sword fern (Polystichum munitum) and Arbutus x ‘Marina’.Koch’s postulates need to be completed on the latter so it has not yet been added to APHIS’ official host list.

Spotted Lanternfly

Spotted Lanternfly (SLF) was found in 16 states in FY 2023. APHIS’ program enjoyed funding provided through Specialty Crop Pests and from the Plant Protection Act’s Section 7721 link ($6 million from the latter).

The report notes that APHIS still does not have enough data to determine SLF’s impacts on agriculture. Thus far, vineyards have been the most adversely affected agricultural segment, mostly due to SLF acting as a stressor to vines. Also, the sticky, sugary “honeydew” produced by SLF attracts other insects and promotes sooty mold growth. These can ruin the fruit and further damage the plant.

SLF populations are strongly linked to major transportation pathways, such as railroads and interstate highways. APHIS targets treatments and, in some areas, removes SLF’s preferred host plant (tree of heaven [Ailanthus]), from transportation hubs. The aim is to reduce the risk of SLF’ spread to new areas and to eradicate isolated infestations. In FY 2023, APHIS and cooperators treated 4,637 properties covering 6,455 acres in affected areas. However, during the National Plant Board meeting both state and APHIS officials complained to me that managers of these transportation hubs raise many barriers to their access, sharply limiting the program’s chance of success.

*Ailanthus altissima* – drive of spotted lanternfly invasion

The program was expanded after National Environmental Policy Act-mandated environmental review. This allowed APHIS to conduct treatments in four additional states—Indiana, Massachusetts, Michigan, and Rhode Island. In addition, program cooperators identified three potential biological control organisms, one that targets the tree of heaven and two that target SLF. APHIS will continue to evaluate them and develop methods to rear them in the laboratory.

Finally, in fiscal year 2023, APHIS joined the National Association of State Departments of Agriculture and the National Plant Board to develop a national strategic plan outlining the future direction of the SLF program. With the strategic plan, PPQ aims to harmonize the approach across states to slow SLF’s spread, develop consistent outreach messaging for a nationwide audience, and more effectively use existing state and Federal resources. Continued spread of SLF despite the huge effort, rising costs of the program, and new scientific findings spurred reconsideration of the strategy.

To summarize, I hope that APHIS’ annual report will – in the future – help members of Congress and their staff understand the agency’s programs’ purpose and past successes. This increased understanding might make it easier to advocate for more funding. I am troubled, however, by the agency’s glossing over significant problems.

Posted by Faith Campbell

www.fadingforests.org

Congress is considering the Farm Bill – help improve it!!!

The House and Senate Agriculture committees are edging toward adopting the next Farm Bill, which is a year past due. Farm bills set policy, funding levels, and more, for 5 years. Each covers a wide range of subjects, including crop subsidies and insurance; food stamps; rural development (including wifi access); forestry policy; and research.

As you might remember, CISP aims to improve USDA’s programs — not only to prevent introductions of non-native tree killing pests and pathogens but also to better respond to those that enter the US and become established. I summarize here what the Senate and House bills have in common and how they differ on these issues.

I understand that the minorities, that is, House Democrats and Senate Republicans, have not accepted all aspects of the majorities’ drafts. So let’s take the opportunity to ask for better bills.

Both the House and Senate bills would “simplify” the USDA Forest Service’s obligations to prepare environmental assessments under the National Environmental Policy Act (NEPA). I have not analyzed which bill weakens NEPA more.

The Senate Bill: The Rural Prosperity and Food Security Act of 2024

The Senate bill addresses forest pest species in several places: Title II — Conservation, Title VII — Research, and Title VIII — Forestry. Here, I describe relevant sections, beginning with the section that partially addresses CISP’s proposal.

Title VIII — Forestry. Section 8214 requires the USDA Secretary to establish a national policy to counter threats posed by invasive species to tree species and forest ecosystems and identify areas for interagency cooperation.

This mandate falls far short of what we sought in a previous bill (S. 1238). However, depending on the exact wording of the bill and accompanying report, perhaps we can succeed in building a stronger program.

It is most important to obtain funding for applied, directed research into resistance breeding strategies, “bulking up,” and planting seedlings that show promise. Please contact your senators and ask them to work with the sponsors – Peter Welch [D-VT], Maggie Hassan [D-NH], and Mike Braun [R-IN] – to try to incorporate more of S. 1238 in the final bill.

The Senate bill contains other provisions that might be helpful for invasive species management – although not part of what CISP and our partners asked for.

‘ōhi‘a trees killed by rapid ‘ōhi‘a death; photo by Richard sniezko, USFS

Title VIII — Forestry. In Section 8506, the Senate bill would require that the US Departments of Agriculture and Interior continue working with Hawai`i to address the pathogen that causes rapid ‘ōhi‘a death. The section authorizes $5 million for each of the coming five fiscal years to do this work. Unfortunately, authorization does not equal funding. Only the Senate and House Appropriations Committees can make this funding available. Hawai`i’s endemic ‘ōhi‘a trees certainly face a dire threat. CISP is already advocating for funding to support resistance breeding and other necessary work.

Title VIII — Forestry. Sections 8247 and 8248 support USDA Forest Service’s nursery and tree establishment programs. My hesitation in fully supporting these provisions is that I fear the urge to plant lots of trees in a hurry will divert attention for the need to learn how to propagate many of the hardwood tree species that have been decimated by non-native pests. However, I agree that the U.S. lacks sufficient nursery capacity to provide anything close to the number of seedlings sought. Perhaps this program can be adjusted to assist the “planting out” component of our request.

Title VII — Research. Section 7208 designates several high-priority research initiatives. On this list are spotted lanternfly, and “invasive species”. A number of forest corporations have been urging Members of Congress to upgrade research on this broad category, which I believe might focus more on invasive plants than the insects and pathogens on which CISP focuses. How the two ideas are integrated will be very important.

Another high-priority initiative concerns the perceived crisis in failed white oak regeneration.

Title VII — Research. Section 7213 mandates creation of four new Centers of Excellence at 1890 Institutions. These are historically Black universities that are also land-grant institutions]. These centers will focus on: 1) climate change, 2) forestry resilience and conservation; 3) food safety, bioprocessing, and value-added agriculture; and, 3) food and agricultural sciences and the social sciences.

Title II — Conservation. Section 2407 provides mandatory funding (which is not subject to annual appropriations) of $75 million per year to the national feral swine eradication/control program (run by USDA APHIS’ Wildlife Service Division). I discuss this program in a separate blog.

The Senate bill also mandates use of several conservation and other programs to address the causes and impacts of climate change. This requirement is directly countered by the House Agriculture Committee’s bill (see below).

The House Bill

Title VIII — Forestry. This section contains none of the provisions CISP’ sought to USDA’s management of tree-killing non-native insects and diseases.

Instead, the House bill calls on the USFS to establish a comprehensive approach to addressing the demise of the giant sequoia trees.

Title VII — Research The House bill, like the Senate’s, lists the invasive species and white oak research initiatives as high priority. The House, unlike the Senate, does not include spotted lanternfly.

Title II — Conservation. As I noted above, the House bill explicitly rescinds all unobligated conservation funding from the Inflation Reduction Act. It reallocates these funds to the traditional conservation programs, e.g., the Environmental Quality Incentive Program and Watershed Protection and Flood Prevention. The bill would use these funds to support “orphan” programs – naming specifically the national feral swine eradication/control program. The House bill provides $150 million – apparently across the five years covered by the Farm Bill, so $30 million per year. Finally, the House allocates 60% of the hog management funds to APHIS, 40% to the Natural Resources Conservation Service.

spotted lanternfly – target of at least 11 projects funded through APHIS’ the Plant Pest and Disease Management and Disaster Prevention Program in FY24. Photo by Holly Raguza, Pennsylvania Department of Agriuculture

Title X —Horticulture, Marketing, and Regulatory Reform. The House’s summary says it is taking steps to protect plant health. It does this by increasing funding for the grant program under the Plant Pest and Disease Management and Disaster Prevention Program – §7721 of the last (2018) Farm Bill. The increase would raise the amount of money available each year from the current level of $70 million to $90 million. These funds are mandatory; they are not subject to annual appropriations. Research, development, and outreach projects funded by this program have certainly added to our understanding of plant pests, hence to their effective management. However, they are usually short-term projects. Therefore they are not suitable for the long-term commitment required for resistance breeding programs. See here and here.

Title III — Trade. Here, the House bill exacerbates the current imbalance between trade promotion and phytosanitary protection. The bill doubles the authorized funding for USDA’s Market Access and Foreign Market Development programs. I concede that this measure probably does reflect a bipartisan consensus in the Congress to support robust programs for promoting agricultural exports.

Also under this Title, the House bill requires the USDA Secretary to conduct regular assessments to identify risks to critical infrastructure that supports food and agriculture sector. This might be helpful – although it is not clear that this assessment would include to threats to forest or urban trees not used commercially (e.g., for timber).

At a recent forum on biological control sponsored by the National Association of State Foresters (NASF), it was reported that participants noted several problems: insufficient funding, significant delays in refilling positions, inadequate research capacity, lack of brick-and-mortar infrastructure, and declining college enrollments in biocontrol-related studies. The NASF Forest Science Health Committee is developing a “Statement of Needs” document that NASF and others can use to lobby for funding to fill these gaps. I hope you will join them in doing so!

salt cedar (*Tamarix* sp.) attacked by biocontrol agent; photo by J.N. Stuart via Flickr

However, as I note above, empowering resistance breeding programs requires a long-term commitment, that is, a comprehensive alteration of policies and infrastructure – beyond annual appropriations.

Posted by Faith Campbell

www.fadingforests.org

Forest Regeneration Again … Deer!

I have recently recent blogged several times about threats to regeneration of eastern forests. Most of the underlying studies stress the role of deer browsing as a major driver of suppression of native plant species and proliferation of non-native ones. Most studies discussed at a recent Northern Hardwood research forum (USDA, FS 2023b Proceedings) found that deer browsing overwhelms other disturbances, such as fire and canopy gaps that typically promote seedling diversity. Miller et al. also stressed the importance of the deer-invasive plant complex in interrupting regeneration in National parks. Reed et al. found that, on the Allegheny Plateau of western Pennsylvania, high deer densities at the time stands formed reduced tree species diversity, density, and basal area – changes that were still detectable decades later.

On the other hand, Hovena et al. found that at their study sites in Ohio, interaction between non-native shrubs and soil wetness overshadowed even the impact of deer herbivory on the species richness and abundance of seedlings.

Unlike the others, Ducey et al. don’t mention deer as a factor in their analysis of regeneration in a forest in the northern half of New Hampshire. They focus on the minimal impact of silvicultural management. Its effect is secondary to overall forest development as the forest ages. Is it possible that overabundant deer are not a factor in the Bartlett Experimental forest.

American elm (*Ulmus americana*); photo by F.T. Campbell

Some of the studies acknowledge the impacts of non-native insects and pathogens. The most thorough discussion is in Payne and Peet. They note that specialist pathogens have caused the loss of important tree species, specifically elms and dogwoods plus the impending widespread mortality of ash. Such mortality is resulting in drastic and long-lasting shifts in community dynamics.

Ducey et al. anticipate pest-driven reversals of increases over the decades of eastern hemlock (Tsuga canadensis) and American beech (Fagus grandifolia). Also, they expect that white ash (Fraxinus americana), which has a minor presence, will disappear.

Miller et al. also stressed the importance of emerald ash borer-induced suppression of ash regeneration in some National parks . The authors also noted the threat to beech trees from beech leaf disease in other parks. Hovena et al. state that the interaction between non-native shrubs and soil wetness was more influential than ash mortality in shaping woody seedling communities.

Reed et al. considered the role of non-native earthworm biomass on plant species’ growth.

But too many of the studies, in my view, make no mention of the probably significant role of non-native insects and pathogens.

It is perhaps understandable that they don’t address emerging pests that either have not yet or have barely reached their study sites. For example, Hovena et al. and Yacucci et al. [see below] noted growth of one native shrub, Lindera benzoin, in the face of the challenges presented by deer and invading plants. Neither acknowledges the approach of laurel wilt disease, which has not yet become established in Ohio (it has been detected on the Kentucky-Indiana border). Similarly, neither mentions beech leaf disease, although some of the plots studied by Hovena et al. are just east of Cleveland – where BLD was first detected. The location of the Yacucci et al. study is less than 50 miles away. The North Carolina forests studied by Payne and Peet are apparently too far east to be affected by beech bark disease and beech leaf disease is not yet established nearby.

Less understandable is the failure to mention loss of elms – which were abundant in riparian areas until killed off by Dutch elm disease – which was first detected in Cleveland!); or to discuss the impact of dogwood anthracnose. Their focus on the deciduous forest might explain why they don’t mention hemlock woolly adelgid – which is just now invading the area discussed by Reed et al. I suppose the demise of American chestnut was so many decades ago that it is truly irrelevant to current forest dynamics.

A new study raises anew these questions about whether inattention to the role of non-native pests has skewed past studies’ results. Yacucci et al. compared regeneration in a military installation (Camp Garfield), to the results in the surrounding second-growth forest. This choice allowed them to overcome one drawback of other studies: using deer exclosures that are small and of short durations. The military facility covers 88 km². Inside it, deer populations have been controlled for 67 years at a density of 6.6 – 7.5 deer/km². Outside, deer have been overabundant for decades. Populations have grown to densities estimated (but not measured) to be at least 30 deer/km²– more than four times as high.

These authors established 21 experimental gaps in the low-deer-density area and 20 gaps outside the installation where deer densities are high. Some of the gaps in both low- and high-deer-density environs were located on wetter, seasonally flooded soils, some on drier sites. None of the forest sites had experience fire in recent decades.

Their findings support the importance of deer browsing as driver of changes to forest regeneration.

northern spicebush (*Lindera benzoin*); photo by R.A. Nonemacher via Wikimedia

They found that at low deer densities, gaps develop a vigorous and diverse native sapling layer, including oaks. Total stem density of red and pin oaks was 13 times higher in these gaps than in gaps in high-deer-density locations. Oak saplings were growing into the subcanopy – that is, above deer browse heights. Saplings of other species – i.e., tuliptree (Liriodendron tulipifera), red maple, and ash (Fraxinus spp.) were also flourishing. Also present were dogwood (Cornus florida) and two native shrubs — Lindera benzoin and Rubus allegheniensis. One non-native shrub, buckthorn (Rhamnus frangula), also thrived at low deer densities. Other non-native plant species were far fewer; their cover was 80% lower. Overall, abundance, richness, and diversity of native herbaceous and woody species were 37–65% higher at the low-deer-density study sites. On average tree species were more than twice as tall as in high-deer-density plots.

In high-deer-density plots, non-native species were six times more abundant while native species richness was 39% lower. Diversity was 27% lower. Most native tree species were short in stature and in low abundance. The one exception was black cherry (Prunus serotina), which deer avoid feeding on. The cherry was 95% more abundant in these high-deer-density plots.

There were several surprising results. In most cases, neither years since gap formation nor habitat type (wet vs. dry) had a significant impact on plant diversity, richness, or abundance. The exception was that non-native plant species were more abundant in older gaps where deer densities were high. Yacucci et al. warn that this phenomenon is a potential threat to biodiversity since high deer density is now the norm across eastern forests.

The authors also note that fire has probably never been a factor in these forests, which are primarily beech-maple forests. Certainly there have been no fires over the past 70 years, either inside or outside the military installation.

Yacucci et al. did not discuss past or possible future impacts of non-native insects or pathogens. They did not mention emerald ash borer or dogwood anthracnose – both of which had been present in Ohio for at least two decades when they completed their study. Although they said their study forest was a beech-maple forest, they did not discuss whether beech are present and – if so – the impact of beech bark disease or beech leaf disease. Both of these are spreading in Ohio. The latter was originally detected in 2012 near Cleveland, just 50 miles from the location of Camp Garfield (between Youngstown and Cleveland, Ohio). As noted above, they also did they mention that Lindera benzoin is susceptible to laurel wilt disease.

beech seedlings in Virginia; photo by F.T. Campbell

Proposed solutions to deer over-browsing

Given the combined threat from widespread deer overpopulation and invasions by non-native plants, Yacucci et al. propose enlisting those military posts that regularly cull deer into efforts to conserve and regenerate native plants. Otherwise, they say, the prognosis for regeneration is poor.

Bernd Blossey and colleagues propose a more sweeping solution: implementation of a national policy to reduce deer populations on all land ownerships. They point out that overabundant deer:

disrupt the plant communities of affected forests – from spring ephemerals to tree regeneration;
promote disease in wildlife and people; and
lead to miserable deaths of deer on our highways, through winter starvation, and disease.

They call for federal leadership of coordinated deer-reduction programs. I discuss their proposal in detail in a separate blog.

SOURCES

Ducey, M.J, O.L. Fraser, M. Yamasaki, E.P. Belair, W.B. Leak. 2023. Eight decades of compositional change in a managed northern hardwood landscape. Forest Ecosystems 10 (2023) 100121

Hovena, B.M., K.S. Knight, V.E. Peters, and D.L Gorchov. 2022. Woody seedling community responses to deer herbivory, intro shrubs, and ash mortality depend on canopy competition and site wetness. Forest Ecology and Management. 523 (2022) 120488

Payne, C.J. and R.K. Peet. 2023. Revisiting the model system for forest succession: Eighty years of resampling Piedmont forests reveals need for an improved suite of indicators of successional change. Ecological Indicators 154 (2023) 110679

Miller, K.M., S.J. Perles, J.P. Schmit, E.R. Matthews, and M.R. Marshall. 2023. Overabundant deer and invasive plants drive widespread regeneration debt in eastern United States national parks. Ecological Applications. 2023;33:e2837. https://onlinelibrary.wiley.com/r/eap Open Access

Reed, S.P., D.R. Bronson, J.A. Forrester, L.M. Prudent, A.M. Yang, A.M. Yantes, P.B. Reich, and L.E. Frelich. 2023. Linked disturbance in the temperate forest: Earthworms, deer, and canopy gaps. Ecology. 2023;104:e4040. https://onlinelibrary.wiley.com/r/ecy

United States Department of Agriculture, Forest Service. 2023a. Proceedings of the First Biennial Northern Hardwood Conference 2021: Bridging Science and Management for the Future. Northern Research Station General Technical Report NRS-P-211 May 2023

Yacucci, A.C., W.P. Carson, J.C. Martineau, C.D. Burns, B.P. Riley, A.A. Royo, T.P. Diggins, I.J. Renne. 2023. Native tree species prosper while exotics falter during gap-phase regeneration, but only where deer densities are near historical levels New Forests https://doi.org/10.1007/s11056-023-10022-w

Posted by Faith Campbell

www.fadingforests.org

Funding of tree pest programs through §7721

spotted lanternfly – target of many projects funded by USDA’s Plant Pest & Disease Management & Disaster Prevention Program; photo by Holly Raguza of Pennsylvania Department of Agriculture

I am belatedly reporting on the forest-pest projects funded by annual grants under the Plant Pest & Disease Management & Disaster Prevention Program ( of the Plant Protection Act). As usual, APHIS funded projects totaling $62.975 million in FY24. In total, 353 projects were funded. These projects represented 70% of the 504 project proposals submitted (the total funding sought was $106 M).

APHIS reserved $11 million for responding to P&P emergencies. I applaud this choice since the agency’s annual appropriation provides only a completely inadequate $1 million (or less) to cover emergencies.

APHIS notes that since initiation of the Plant Pest & Disease Management & Disaster Prevention Program in 2009, it has funded more than 5,500 projects with a total of nearly $870 million.

In FY24 the program funded 30 more projects than the 322 projects funded in FY23. blog 320 The FY24 allocation provides more than $1 million more for goal area 1S — Enhance Plant Pest/Disease Survey (from $14.4 million to $15.7 million). This was balanced by small decreases for the other goal areas: enhancing mitigation capabilities received $13.6 million; inspections at domestic sites important in invasive species’ spread received $6.3 million; pest identification and detection received $5.3 million; and outreach and education received $4.1 million. Projects safeguarding nursery production and those improving pest and disease analysis each received about $2 million.

By my calculation – subject to error! – about $7.5 million went to projects clearly dealing with forest pests [12% of total funding]. This is a welcome increase from FY23 – when funding of such projects reached about $6.5 million (a little over 10%). blog 320 Funding for tree pest projects might be higher. Some $1.9 million is allocated to surveys of grapevines and orchards — hosts of the spotted lanternfly (SLF). However, it is not clear whether these projects are focused on detecting and managing SLF; they might have a much broader goal. If we do include these projects, the total for tree-killing pests rises to $9.4 million — nearly 15% of the total.

Over both FY23 and FY24, the majority of funds went to similar topics: survey and management of sudden oak death in nurseries; surveys for bark beetles, Asian defoliators, and forest pests generally; and outreach programs targetting the spotted lanternfly. In FY24, just under $100,000 paid for efforts to develop tools for rapid detection of laurel wilt link to DMF in avocados – that is, in a crop rather than the natural environment.

No projects addressing tree or forest pests were funded in seven states or territories: Guam, Idaho, Nebraska, New Mexico, Rhode Island, South Dakota, and Utah. This was three fewer states than in FY23. In neither year do I know whether these states submitted proposals in this category that ended up not being funded.

In FY24, spotted lanternfly is by far the pest addressed by the most projects. As noted above, I can’t be precise about the number because of the lack of information about the 23 projects that fund pest surveys of grapes and/or tree crops that are SLF hosts. Eleven projects named SLF specifically. A final project (not included in above) is one funding registration of Verticillium nonalfalfae as a biocontrol for Ailanthus altissima – an invasive tree that is the preferred host of SLF.

The District of Columbia, Kansas, Missouri, and Oklahoma each had one tree pest project funded. In the cases of Kansas and Missouri, the single project was surveys for thousand cankers disease of walnut. Three other states — Iowa, Maryland, and Pennsylvania — also obtained funding to survey for TCD.

The single Oklahoma project concerned efforts to ensure that the sudden oak death pathogen(Phytopthora ramorum) is not present in nurseries. (An Oklahoma wholesaler was one of the hubs of this pathogen’s spread to 18 states in 2019). Eleven other states were also funded to survey their nurseries for P. ramorum: Alabama, Kentucky, Louisiana, Nevada, North Carolina, Ohio, Pennsylvania, South Carolina, Virginia, and West Virginia. P. ramorum is a “program pest” in 2024. That is, APHIS had designated it as a regulated pest for which the agency wished to fill knowledge gaps about its distribution. I note that last year APHIS published a risk assessment that downplayed the likelihood that P. ramorum would establish in the eastern states. Is APHIS seeking more information to test this conclusion?

In a separate case, Oregon received $76,000 to evaluating the threat to nurseries and forests arising from the presence in the state’s forests of two strains or lineages of P. ramorum that previously had not been extant in the environment of North America.

Another approximately 53 projects fund surveys for tree pests other than spotted lanternfly; these are often fairly general surveys, such as for woodborers or “Asian defoliators”. About ten projects fund management efforts – including evaluation of the efficacy of emerald ash borer biocontrol programs.

Last year I noted that two states – Mississippi and Nevada — had projects to survey the “palm commodity”. Hawai`i joined this group in FY24. The project descriptions don’t specify which pests are the targets. The South American palm weevil (Rhynchophorus palmarum) seems most probable; it is established in far southern California and neighboring Mexico. APHIS prepared a risk assessment on the species in 2012. link? In Hawa`ii, concern probably focuses on the coconut rhinoceros beetle (Oryctes rhinoceros). link? There are other threats to palms, e.g., the red palm weevil (Rhynochophorus ferrugineus), link? and a deadly Fusarium wilt. link?

native palms in the desert at Anza-Borrego, California; photo by F.T. Campbell

California has native palms (Washingtonia filifera); southern states from Texas to at least South Carolina have native palmettos. Of course, many species of palms are important ornamental plants in these states, and dates are raised commercially.

Another “program pest” that I have blogged about in the past is box tree moth. link to blog 287 In FY24 five projects addressed this pest, including surveys and efforts to develop better control tools.

beavertail cactus (*Opuntia basilaris*) in Anza-Borrego, California; photo by F.T. Campbell

I am pleased by continued funding of projects trying to utilize biocontrol agents to protect two groups of cactus severely threatened by non-native insects: lepidoptera that attack flat-padded prickly pear cacti (Opuntia spp.) link to DMF and the mealybug that attacks columnar cacti of Puerto Rico and the Virgin Islands. link to DMF

vulnerable cactus on St. John, US Virgin Islands; photo by F.T. Campbell

I applaud the decision to fund projects focused on determining the efficacy of biocontrol projects. As noted above, three projects are asking these questions in the case of the emerald ash borer. link to DMF Another project funds production, release, and efficacy evaluation of biocontrol agents targetting Brazilian peppertree in Florida & Texas.

I am also pleased that three projects assist Washington State in its efforts to eradicate the invasion by giant hornets from Asia. link to blogs & Hornet Herald – no detections in 2023 … A company in California also received funding to developing hornet detection tools.

Nineteen projects funded outreach efforts, including continued funding for the “Don’t Move Firewood” program. In addition to those focused on spotted lanternfly, such projects also included other firewood programs, Asian longhorned beetle awareness, and the nursery industry.

I note that while California received funding for 27 projects, none dealt with any of several deadly tree pests extant in the state – goldspotted oak borer, polyphagous and Kuroshio shot hole borers, Mediterranean oak borer, and the palm weevils. Nor did Hawai`i obtain funding to address rapid ohia death. Did no one submit proposals to address any of the many issues impeding management of these killers?

South American palm weevil; photo by Allan Hopkins via Flickr

Posted by Faith Campbell