Center for Invasive Species Prevention

Hawaiian Efforts to Restore Threatened Trees

ʻŌhiʻa trees killed by ROD; photo by Richard Sniezko, USFS

Several Hawaiian tree species are at risk due to introduced forest pests. Two of the Islands’ most widespread species are among the at-risk taxa. Their continuing loss would expose watersheds on which human life and agriculture depend. Habitats for hundreds of other species – many endemic and already endangered – would lose their foundations. These trees also are of the greatest cultural importance to Native Hawaiians.

I am pleased to report that Hawaiian scientists and conservationists are trying to protect and restore them.

Other tree species enjoy less recognition … and efforts to protect them have struggled to obtain support.

1) koa (Acacia koa)

Koa is both a dominant canopy tree and the second-most abundant native tree species in Hawai`i in terms of areas covered. The species is endemic to the Hawaiian archipelago. Koa forests provide habitat for 30 of the islands’ remaining 35 native bird species, many of which are listed under the U.S. Endangered Species Act. Also dependent on koa forests are native plant and invertebrate species and the Islands’ only native terrestrial mammal, the Hawaiian hoary bat. Finally, koa forests protect watersheds, add nitrogen to degraded soils, and store carbon [Inman-Narahari et al.]

Koa forests once ranged from near sea level to above 7000 ft (2100 m) on both the wet and dry sides of all the large Hawaiian Islands. Conversion of forests to livestock grazing and row-crop agriculture has reduced koa’s range. Significant koa forests are now found on four islands – Hawai’i, Maui, O‘ahu, and Kauaʻi. More than 90% of the remaining koa forests occur on Hawai`i Island (the “Big Island) [Inman-Narahari et al.]

In addition to its fundamental environmental role, koa has immense cultural importance. Koa represents strength and the warrior spirit. The wood was used traditionally to make sea-going canoes. Now Koa is widely used for making musical instruments, especially guitars and ukuleles; furniture, surfboards, ornaments, and art [Inman-Narahari et al.]

Koa timber has the highest monetary value of any wood harvested on the Islands. However, supplies of commercial-quality trees are very limited (Dudley et al. 2020). Harvesting is entirely from old-growth forests on private land. [Inman-Narahari et al.]

Koa forests are under threat by a vascular wilt disease caused by Fusarium oxysporum f. sp. koae (FOXY). This disease can kill up to 90% of young trees and – sometimes — mature trees in native forests. The fungus is a soil-dwelling organism that spreads in soil and infects susceptible plants through the root system (Dudley et al. 2020).

Conservation and commercial considerations have converged to prompt efforts to breed koa resistant to FOXY. Conservationists hope to restore native forests on large areas where agriculture has declined. The forestry industry seeks to enhance supplies of the Islands’ most valuable wood. Finally, science indicated that a breeding program would probably be successful. Field trials in the 1990s demonstrated great differences in wilt-disease mortality among seed sources (the proportion of seedlings surviving inoculation ranged from 4% to 91.6%) [Sniezko 2003; Dudley et al. 2009].

In 2003, Dudley and Sniezko outlined a long-term strategy for exploring and utilizing genetic resistance in koa. Since then, a team of scientists and foresters has implemented different phases of the strategy and refined it further (Dudley et al. 2012, 2015, 2017; Sniezko et al. 2016]

First, scientists determined that the wilt disease is established on the four main islands. Having obtained more than 500 isolates of the pathogen from 386 trees sampled at 46 sites, scientists tested more than 700 koa families from 11 ecoregions for resistance against ten of the most highly virulent isolates (Dudley et al. 2020).

The Hawaiian Agricultural Research Center (HARC), supported by public and private partners, has converted the field-testing facilities on Hawai`i, Maui, and Oahu into seed orchards. The best-performing tree families are being grown to maturity to produce seeds for planting. It is essential that the seedlings be not just resistant to FOXY but also adapted to the ecological conditions of the specific site where they are to be planted [Dudley et al. 2020; Inman-Narahari et al. ] Locally adapted, wilt-resistant seed has been planted on Kauaʻi and Hawai`i. Preparations are being made to plant seed on Maui and O‘ahu also. Scientists are also exploring methods to scale up planting in both restoration and commercial forests [R. Hauff pers. comm.].

Restoration of koa on the approximately half of lands in the species’ former range that are privately owned will require that the trees provide superior timber. Private landowners might also need financial incentives since the rotation time for a koa plantation is thought to be 30-80 years. [Inman-Narahari et al.]

Plantings on both private and public lands will need to be protected from grazing by feral ungulates and encroachment by competing plants. These management actions are intensive, expensive, and must be maintained for years.

Some additional challenges are scientific: uncertainties about appropriate seed zones, efficacy of silvicultural approaches to managing the disease, and whether koa can be managed for sustainable harvests. Human considerations are also important: Hawai`i lacks sufficient professional tree improvement or silvicultural personnel, a functioning seed distribution and banking network — and supporting resources. Finally, some segments of the public oppose ungulate control programs. Inman-Narahari et al.

Finally, scientists must monitor seed orchards and field plantings for any signs of maladaptation to climate change. (Dudley et al. 2020).

2) ʻŌhiʻa Metrosideros polymorpha)

ʻŌhiʻa lehua is the most widespread tree on the Islands. It dominates approximately 80% the biomass of Hawaii’s remaining native forest, in both wet and dry habitats. ʻŌhiʻa illustrates adaptive radiation and appears to be undergoing incipient speciation. The multitude of ecological niches and their isolation on the separate islands has resulted in five recognized species in the genus Metrosideros. Even the species found throughout the state, Metrosideros polymorpha, has eight recognized varieties (Luiz et al. (2023) (some authorities say there are more).

Loss of this iconic species could result in significant changes to the structure, composition, and potentially, the function, of forests on a landscape level. High elevation ‘ohi‘a forests protect watersheds across the state. ʻŌhiʻa forests shelter the Islands’ one native terrestrial mammal (Hawaiian hoary bat), 30 species of forest birds, and more than 500 endemic arthropod species. Many species in all these taxa are endangered or threatened (Luiz et al. 2023). The increased light penetrating interior forests following canopy dieback facilitates invasion by light-loving non-native plant species, of which Hawai`i has dozens. There is perhaps no other species in the United States that supports more endangered taxa or that plays such a geographical dominant ecological keystone role [Luiz et al. 2023]

For many Native Hawaiians, ‘ōhi‘a is a physical manifestation of multiple Hawaiian deities and the subject of many Hawaiian proverbs, chants, and stories; and foundational to the scared practice of many hula. The wood has numerous uses. Flowers, shoots, and aerial roots are used medicinally and for making lei. The importance of the biocultural link between ‘ōhi‘a and the people of Hawai`i is described by Loope and LaRosa (2008) and Luiz et al. (2023).

In 2010 scientists detected rapid mortality affecting ‘ōhi‘a on Hawai‘i Island. Scientists determined that the disease is caused by two recently-described pathogenic fungi, Ceratocystis lukuohia and Ceratocystis huliohia. The two diseases, Ceratocystis wilt and Ceratocystis canker of ʻōhiʻa, are jointly called “rapid ‘ōhi‘a death”, or ROD. The more virulent species, C. lukuohia, has since spread across Hawai`i Island and been detected on Kaua‘i. The less virulent C. huliohia is established on Hawai`i and Kaua‘i and in about a dozen trees on O‘ahu. One tree on Maui was infected; it was destroyed, and no new infection has been detected [M. Hughes pers. comm.] As of 2023, significant mortality has occurred on more than one third of the vulnerable forest on Hawai`i Island, although mortality is patchy.

[ʻŌhiʻa is also facing a separate disease called myrtle rust caused by the fungus Austropuccinia psidii; to date this rust has caused less virulent infections on ‘ōhi‘a.]

rust-killed ‘ōhi‘a in 2016; photo by J.B. Friday

Because of the ecological importance of ‘ōhi‘a and the rapid spread of these lethal diseases, research into possible resistance to the more virulent pathogen, C. lukiohia began fairly quickly, in 2016. Some ‘ōhi‘a survive in forests on the Big Island in the presence of ROD, raising hopes that some trees might possess natural resistance. Scientists are collecting germplasm from these lightly impacted stands near high-mortality stands (Luiz et al. 2023). Five seedlings representing four varieties of M. polymorpha that survived several years’ exposure to the disease are being used to produce rooted cuttings and seeds for further evaluation of these genotypes.

Encouraged by these developments, and recognizing the scope of additional work needed, in 2018 stakeholders created a collaborative partnership that includes state, federal, and non-profit agencies and entities, ʻŌhiʻa Disease Resistance Program (‘ODRP) (Luiz et al. 2023). The partnership seeks to provide baseline information on genetic resistance present in all Hawaiian taxa in the genus Metrosideros. It aims further to develop sources of ROD-resistant germplasm for restoration intended to serve several purposes: cultural plantings, landscaping, and ecological restoration. ‘ODRP is pursuing screenings of seedlings and rooted cuttings sampled from native Metrosideros throughout Hawai`i while trying to improve screening and growing methods. Progress will depend on expanding these efforts to include field trials; research into environmental and genetic drivers of susceptibility and resistance; developing remote sensing and molecular methods to rapidly detect ROD-resistant individuals; and support already ongoing Metrosideros conservation. If levels of resistance in wild populations prove to be insufficient, the program will also undertake breeding (Luiz et al. 2023).

To be successful, ‘ODRP must surmount several challenges (Luiz et al. 2022):

increase capacity to screen seedlings from several hundred plants per year to several thousand;
optimize artificial inoculation methodologies;
determine the effects of temperature and season on infection rates and disease progression;
find ways to speed up seedlings’ attaining sufficient size for testing;
develop improved ways to propagate ʻōhiʻa from seed and rooted cuttings;
establish sites for field testing of putatively resistant trees across a wide range of climatic and edaphic conditions;
establish seed orchard, preferably on several islands;
establish systems for seed collection from the wide variety of subspecies/varieties;
if breeding to enhance resistance is appropriate, it will be useful to develop high-throughput phenotyping of the seed orchard plantings.

[See DMF profile for more details.]

Developing ROD-resistant ‘ōhi‘a is only one part of a holistic conservation program. Luiz et al. (2023) reiterate the importance of quarantines and education to curtail movement of infected material and countering activities that injure the trees. Fencing to protect these forests from grazing by feral animals can drastically reduce the amount of disease. Finally, scientists must overcome the factors there caused the almost complete lack of natural regeneration of ‘ōhi‘a in lower elevation forests. Most important are competition by invasive plants, predation by feral ungulates, and the presence of other diseases, e.g., Austropuccinia psidii.

Hawaii’s dryland forests are highly endangered: more than 90% of dry forests are already lost due to habitat destruction and the spread of invasive plant and animal species. Two tree species are the focus of species-specific programs aimed at restoring them to remaining dryland forests. However, support for both programs seems precarious and requires stable long-term funding; disease resistance programs often necessitate decades-long endeavors.

naio in bloom; photo by Forrest & Kim Starr via Creative Commons

1) naio (Myoporum sandwicense)

Naio grows on all of the main Hawaiian Islands at elevations ranging from sea level to 3000 m. While it occurs in the full range of forest types from dry to wet, naio is one of two tree species that dominate upland dry forests. The other species is mamane, Sophora chrysophylla. Naio is a key forage tree for two endangered honeycreepers, palila (Loxioides bailleui) and `akiapola`au (Hemignathus munroi). The tree is also an important host of many species of native yellow-face bees (Hylaeus spp). Finally, loss of a native tree species in priority watersheds might lead to invasions by non-native plants that consume more water or increase runoff.

The invasive non-native Myoporum thrips, Klambothrips myopori, was detected on Hawai‘i Island in December 2008 (L. Kaufman website). In 2018 the thrips was found also on Oahu (work plan). The Myoporum thrips feeds on and causes galls on plants’ terminal growth. This can eventually lead to death of the plant.

Aware of thrips-caused death of plants in the Myoporum genus in California, the Hawaii Department of Lands and Natural Resources Division of Forestry and Wildlife and the University of Hawai‘i began efforts to determine the insect’s distribution and infestation rates, as well as the overall health of naio populations on the Big Island. This initiative began in September 2010, nearly two years after the thrips’ detection. Scientists monitored nine protected natural habitats for four years. This monitoring program was supported by the USFS Forest Health Protection program. This program is described by Kaufman.

naio monitoring sites from L. Kaufman article

The monitoring program determined that by 2013, the thrips has spread across most of Hawi`i Island, on its own and aided by human movement of landscaping plants. More than 60% of trees being monitored had died. Infestation and dieback levels had both increased, especially at medium elevation sites. The authors feared that mortality at high elevations would increase in the future. They found no evidence that natural enemies are effective controlling naio thrips populations on Hawai`i Island.

Kaufman was skeptical that biological control would be effective. She suggested, instead, a breeding program, including hybridizing M. sandwicensis with non-Hawaiian Myoporum species that appear to be resistant to thrips. Kaufman also called for additional programs: active monitoring to prevent thrips from establishing on neighboring islands; and collection and storage of naio seeds.

Ten years later, in February 2024, DLNR Division of Forestry and Wildlife adopted a draft work plan for exploring possible resistance to the Myoporum thrips. Early steps include establishing a database to record data needed to track parent trees, associated propagules, and the results of tests. These data are crucial to keeping track of which trees show the most promise. Other actions will aim to hone methods and processes. Among practical questions to be answered are a) whether scientists can grow even-aged stands of naio seedlings; b) identifying the most efficient resistance screening techniques; and c) whether K. myopori thrips are naturally present in sufficient numbers to be used in tests, or – alternatively – whether they must be augmented. [Plan]

Meanwhile, scientists have begun collecting seed from unaffected or lightly affected naio in hotspots where mortality is high. They have focused on the dry and mesic forests of the western side of Hawai`i (“Big”) Island, where the largest number of naio populations still occur and are at high risk. Unfortunately, these “lingering” trees remain vulnerable to other threats, such as browsing by feral ungulates, competition with invasive plants, drought, and reduced fecundity & regeneration.

Hawai`i DLNR has secured initial funding from the Department of Defense’s REPI program to begin a pest resistance project and is seeking a partnership with University of Hawai`i to carry out tests “challenging” different naio families’ resistance to the thrips [R. Hauff pers. comm.]

2) wiliwili (Erythrina sandwicensis)

Efforts to protect the wiliwili have focused on biological control. The introduced Erythrina gall wasp, Quadrastichus erythrinae (EGW) was detected on the islands in 2005. It immediately caused considerable damage to the native tree and cultivated nonnative coral trees.

A parasitic wasp, Eurytoma erythrinae, was approved for release in November 2008 – only 3 ½ years after EGW was detected on O‘ahu. The parasitic wasp quickly suppressed the gall wasp’s impacts to both wiliwili trees and non-native Erythrina. By 2024, managers are once again planting the tree in restoration projects.

However, both the gall wasp and a second insect pest – a bruchid, Specularius impressithorax – can cause loss of more than 75% of the seed crop. This damage means that the tree cannot regenerate. By 2019, Hawaiian authorities began seeking permission to release a second biocontrol gent, Aprostocitus nites.Unfortunately, the Hawai’i Department of Agriculture still has not approved the release permit despite five years having passed. Once they have this approval, the scientists will then need to ask USDA Animal and Plant Health Inspection Service (APHIS) for its approval [R. Hauff, pers. comm.]

SOURCES

www.RapidOhiaDeath.org

Dudley, N., R. James, R. Sniezko, P. Cannon, A. Yeh, T. Jones, & Michael Kaufmann. 2009? Operational Disease Screening Program for Resistance to Wilt in Acacia koa in Hawai`i. Hawai`i Forestry Association Newsletter August 29 2009

Dudley, N., T. Jones, K. Gerber, A.L. Ross-Davis, R.A. Sniezko, P. Cannon & J. Dobbs. 2020. Establishment of a Genetically Diverse, Disease-Resistant Acacia koa Seed Orchard in Kokee, Kauai: Early Growth, Form, & Survival. Forests 2020, 11, 1276; doi:10.3390/f11121276 www.mdpi.com/journal/forests

Friday, J. B., L. Keith, and F. Hughes. 2015. Rapid ʻŌhiʻa Death (Ceratocystis Wilt of ʻŌhiʻa). PD-107, College of Tropical Agriculture and Human Resources, University of Hawai‘i, Honolulu, HI. URL: https://www.ctahr.HI.edu/oc/freepubs/pdf/PD-107.pdf Accessed April 3, 2018.

Friday, J.B. 2018. Rapid ??hi?a Death Symposium -West Hawai`i (“West Side Symposium”) March 3rd 2018, https://vimeo.com/258704469 Accessed April 4, 2018 (see also full video archive at https://vimeo.com/user10051674)

Inman-Narahari, F., R. Hauff, S.S. Mann, I. Sprecher, & L. Hadway. Koa Action Plan: Management & research priorities for Acacia koa forestry in Hawai`i. State of Hawai`i Department of Land & Natural Resources Division of Forestry & Wildlife no date

Kaufman, L.V, J. Yalemar, M.G. Wright. In press. Classical biological control of the erythrina gall wasp, Quadrastichus erythrinae, in Hawaii: Conserving an endangered habitat. Biological Control. Vol. 142, March 2020

Loope, L. and A.M. LaRosa. 2008. ‘Ohi’a Rust (Eucalyptus Rust) (Puccinia psidii Winter) Risk Assessment for Hawai‘i.

Luiz, B.C. 2017. Understanding Ceratocystis. sp A: Growth, morphology, and host resistance. MS thesis, University of Hawai‘i at Hilo.

Luiz, B.C., C.P. Giardina, L.M. Keith, D.F. Jacobs, R.A. Sniezko, M.A. Hughes, J.B. Friday, P. Cannon, R. Hauff, K. Francisco, M.M. Chau, N. Dudley, A. Yeh, G. Asner, R.E. Martin, R. Perroy, B.J. Tucker, A. Evangelista, V. Fernandez, C. Martins-Keli’iho.omalu, K. Santos, R. Ohara. 2023. A framework for establishlishing a rapid ‘Ohi‘a death resistance program New Forests 54, 637–660. https://doi.org/10.1007/s11056-021-09896-5

Additional information on the koa resistance program is posted at http://www.harc-hspa.com/forestry.html

Sniezko, R.A., N. Dudley, T. Jones, & P. Cannon. 2016. Koa wilt resistance & koa genetics – key to successful restoration & reforestation of koa (Acacia koa). Acacia koa in Hawai‘i: Facing the Future. Proceedings of the 2016 Symposium, Hilo, HI: www.TropHTIRC.org , www.ctahr.HI.edu/forestry

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

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

www.fadingforests.org

Impacts of introduced rust on unique flora — New Zealand’s expectations

predicted community vulnerability from A. psidii mediated mortality of Kunzea ericoides & Leptospermum scoparium; from McCarthy et al.

Scientists in New Zealand have recently completed a study of the probable impact of myrtle rust – caused by Austropuccinia psidii – on plants in the plant family Myrtaceae. McCarthy et al. say their results should guide management actions to protect not only the unique flora of those islands but also on Australia and Hawai`i – other places where key dominant tree species are susceptible to myrtle rust. The disease attacks young tissue; susceptible Myrtaceae become unable to recruit new individuals or to recover from disturbance. Severe cases can result in tree death & localized extinctions

[I note that myrtle rust is not the only threat to the native trees of these biologically unique island systems. New Zealand’s largest tree, kauri (Agathis australis), is threatened by kauri dieback (caused by Phytophthora agathidicida). On Hawai`i, while the most widespread tree, ‘ōhi‘a (Metrosideros polymorpha) is somewhat vulnerable to the strain of rust introduced to the Islands, the greater threat is from a different group of fungi, Ceratocystis lukuohia and C. huliohia, collectively known as rapid ‘ōhi‘a death. On Australia, hundreds of endemic species on the western side of the continent are being killed by Phytophthora dieback, caused by Phytophthora cinnamomi. [I note the proliferation of tree-kiling pathogens; I will blog more about this in the near future.]

Myrtle rust arrived in New Zealand in 2017, probably blown on the wind from Australia (where it was detected in 2010). In New Zealand, myrtle rust infects at least 12 of 18 native tree, shrub, and vine species in the Myrtaceae plant family. Several of these species are important in the structure and succession of native ecosystems. They also have enormous cultural significance.

McCarthy et al. note that species differ in their contribution to forest structure and function. They sought to determine where loss of vulnerable species might have the greatest impact on community functionality. They also explored whether compensatory infilling by co-occurring, non-vulnerable species in the Myrtaceae would reduce the community’s vulnerability. Even when co-occurring Myrtaceae are relatively immune to the pathogen, only some of them – the fast-growing species – are likely to fill the gaps. They might lack the functional attributes of the decimated species.

To identify areas at greatest risk, McCarthy et al. took advantage of a nationwide vegetation plot dataset that covers all the country’s native forests and shrublands. The plot data enabled McCarthy et al. to determine which plant species not vulnerable to the rust are present and so are likely to replace the rust host species as they are killed.

*Leptospermum scoparium*; photo by Alyenaa Buckles via Flickr

McCarthy et al. concluded that forests and shrublands containing Kunzea ericoides and Leptospermum scoparium are highly vulnerable to their loss. Ecosystems with these species are found predominantly in central and southeastern North Island, northeastern South Island, and Stewart Island. While compensatory infilling by other species in the Myrtaceae would moderate the impact of the loss of vulnerable species, if these co-occurring species were unable to respond for various reasons, such as also being infected by the rust pathogen, community vulnerability almost always increased. In these cases the infilling species would probably have different functional attributes. In many areas the species most likely to replace the rust-killed native species would be non-native shrubs. Consequently, early successional woody plant communities, where K. ericoides and L. scoparium dominate, are at most risk.

Because the risk of A. psidii infection is lower in cooler montane and southern coastal areas, parts of inland Fiordland, the northwestern South Island and the west coast of the North Island might be less vulnerable.

Austropuccinia psidii has been spreading in Myrtaceae-dominated forests of the Southern Hemisphere since the beginning of the 21^st Century. It was detected in Hawai`i in 2005; in Australia in 2010; in New Caledonia in 2013, and finally in New Zealand in 2017. Within 12 months of its first detection in the northern part of the North Island it had spread to the northern regions of the South Island.

Specific types of Threat

Succession

The ecosystem process most at risk to loss of Myrtaceae species to A. psidii is succession. About 10% of once-forested areas of New Zealand are in successional shrublands, mostly dominated by Kunzea ericoides and Leptospermum scoparium. Both species are wind dispersed, grow quickly, are resistant to browsing by introduced deer, and are favored by disturbance, especially fire. Both are tolerant of exposure and have a wide edaphic range (including geothermal soils). Still, K. ericoides prefers drier, warmer sites while L. scoparium tolerates saturated soils, frost hollows and subalpine settings.

*Kunzea ericoides*; photo by Tony Foster via Flickr

Loss of these two species would result in a considerable change in stand-level functional composition across a wide variety of locations. Their extensive ranges mean that it would be difficult for other species – even if functionally equivalent – to expand sufficiently quickly. Second, non-native species are common in these communities. All of these invaders – Ulex europaeus, Cytisus scoparius and species of Acacia, Hakea and Erica – promote fire. Some are nitrogen fixers. While they can facilitate succession, the resulting native forest will differ from that formed via Leptospermeae succession. Furthermore, compensatory infilling by the invasive species might also reduce carbon sequestration. Successional forests dominated by K. ericoides are significant carbon sinks owing to the tree’s size (up to 25 m under favorable conditions), high wood density, and long lifespan (up to ~150 years). In contrast, shrublands dominated by at least one of the non-native species, U. europaeus, are significant carbon sources.

Northern and central regions of the North Island and the northeastern and interior parts of the South Island are most vulnerable to the loss of these species since these successional shrub communities are widespread and the area’s climate is highly suitable for A. psidii infection. The southern regions of the South Island, including Stewart Island, are somewhat protected by the cooler climate.

Fortunately, neither Kunzea ericoides nor Leptospermum scoparium has yet been infected in nature. Laboratory trials indicate that some families of K. ericoides are resistant. Vulnerability also varies among types of tissue – i.e., leaf, stem, seed capsule.

*Metrosideros umbellata*; photo by Stan Shebs via Wikimedia

Forest biomass

Although from the overall community perspective loss of species in the Metrosidereae would have a lower impact than loss of those in the Leptospermeae, there would be significant changes associated with loss of Metrosideros umbellata. This species can grow quite large (dbh often > 2 m; heights up to 20 m). That size and its exceptionally dense wood means that M. umbellata stores high amounts of carbon. Also, its slow decomposition provides habitat for decomposers. Lessening the potential impact of loss of this species are two facts: its litter nutrient concentrations and decomposition rates do not differ from dominant co-occurring trees; and, most important, it grows primarily in the south, where weather conditions are less suitable for A. psidii infection. One note of caution: if A. psidii proves able to spread into these regions, not only M. umbellata but also susceptible co-occurring Myrtaceae species are likely to be damaged by the pathogen.

Highly specific habitats

McCarthy et al. note that their study might underestimate the impact of loss of species with unique traits that occupy specialized habitats. They focus on the climber Metrosideros excelsa. This is an important successional species that helps restore ecosystems following fire, landslides, or volcanic eruptions. The species’ tough and nutrient poor leaves promote later successional species by forming a humus layer and altering the microenvironment beneath the plant. Its litter has high concentrations of phenolics and decomposes more slowly than any co-occurring tree species. [They say its role is analogous to that of M. polymorpha in primary successions on lava flows in Hawai`i.] M. excelsa dominates succession on many small offshore volcanic islands, rocky coastal headlands and cliffs.

Another example is Lophomyrtus bullata, a small tree that is patchily distributed primarily in forest margins and streamside vegetation. This is the native species most affected by A. psidii; the pathogen is likely to cause its localized extinction. McCarthy et al. call for assessment of ex situ conservation strategies for this species.

Each of these species is represented in only seven of the plots used in the analysis, so community vulnerability to their loss might be underestimated.

Another habitat specialist, Syzygium maire, is found mostly in lowland forests, usually on saturated soils. It currently occupies only a fraction of its natural range due to deforestation and land drainage. Evaluating the impact of loss of S. maire is complicated by its poor representation in the database (only six plots), and the fact that many of the co-occurring species are also Myrtaceae.

Lack of data similarly prevents detailed assessment of the impacts from possible loss of other species, including M. parkinsonii, M. perforata and L. obcordata. McCarthy et al. say only that their disappearance will “take the community even further from its original state”.

McCarthy et al. warn that the risk could increase if more virulent strains of A. psidii were introduced or evolved through sexual recombination of the current pandemic strain. Other scientists have discovered strong evidence that the many strains of A. psidii attack different host species (see Costa da Silva et al. 2014).

New Zealand bell bird (*Anthornis melanura*); photo from https://animalia.bio/new-zealand-bellbird

McCarthy et al. note that other factors are also important in determining the impact of loss of a plant species. Especially significant is the host plant species’ association with other species. They say these relationships are poorly understood. One example is that only four Myrtaceae species produce fleshy fruits. Loss or decline of these four species might severely affect populations of native birds, many of which are endemic. Many invertebrates – also highly endemic — are dependent on nectar from other plants in the family.

In their conclusion, McCarthy et al. note that A. psidii has been introduced relatively recently so there is still time to reduce the disease’s potential consequences. They suggest such management interventions as identifying and planting resistant genotypes and applying chemical controls to protect important individual specimens. They hope their work will guide prioritization of both species and spatial locations. They believe such efforts have substantial potential to reduce myrtle rust’s overall functional impact to New Zealand’s unique ecosystems.

SOURCES

Costa da Silva, A; P.M. Teixeira de Andrade, A. Couto Alfenas, R. Neves Graca, P. Cannon, R. Hauff, D. Cristiano Ferreira, and S. Mori. 2014. Virulence and Impact of Brazilian Strains of Puccinia psidii on Hawaiian Ohia (Metrosideros polymorpha). Pacific Science 68(1):47-56. doi: https://dx.doi.org/10.2984/68.1.4

McCarthy, J.K., S.J. Richardson, I. Jo, S.K. Wiser, T.A. Easdale, J.D. Shepherd, P.J. Bellingham. 2024. A Functional Assessment of Community Vulnerability to the Loss of Myrtaceae From Myrtle Rust. Diversity & Distributions, 2024; https://doi.org/10.1111/ddi.13928

Posted by Faith Campbell

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

Phytophthora here, Phytopthora there … level of threat is unclear

Mt. Triglav – highest peak in the Slovenian (Julian) Alps; photo by Gunter Nuyts via Pexel

Scientists have discovered sizable diversity of pathogenic Phytophthora species in Europe, specifically in the Alps of northeastern Italy and western Slovenija. They have also named a new species, and noted the need to change the definition of species previously named. See Bregant et al. – full citation at the end of this blog – open access!

Two of its findings are especially important for the US

First, the authors document the vulnerability of alpine areas to 18 Phythophthora species. Most of the plant hosts they studied have congenerics in mountainous areas of North America: Acer, Alnus, Betula, Fagus, Fragaria, Fraxinus, Ilex, Juniperus, Larix, Lonicera, Lycopodium, Pinus, Populus, Quercus, Rhododendron, Rubus, Salix, Sorbus, Taxus, and Vaccinium.

Second, the paper discusses how junipers are at particular risk. I remind you that P. austrocedrii has recently been detected in nurseries in Ohio and Oregon. This is another non-native Phythophthora that attacks junipers. I hope authorities are actively seeking to determine whether P. austrocedrii is present in nurseries or natural systems in other parts of the country.

The genus Phytophthora includes many serious plant pathogens, from the one that caused the disastrous potato blight of Ireland (Phytophthora infestans) to globally important forest-destroying invasive species, e.g., P. cinnamomi and “sudden oak death” P. ramorum.

Bregant et al. surveyed 33 small tree, shrub, and herbaceous plant species in 54 sites on the Italian island of Sardinia and the Alps of both northeastern Italy and western Slovenija. Altitudes varied from the valley bottom (700 m) to above tree line (2100 m). Sites included typical forests, riparian ecosystems, and heathlands.

The 360 isolates taken from 397 samples belonged to 17 known Phytophthora species. Some species are widespread and well-known, e.g., P. pseudosyringae. Three isolates belonged to a putative new species described by Bregant et al. – Phytophthora pseudogregata sp. nov. This total of 18 taxa was unexpectedly high. Many of the species are able to cause aerial infections via production of caducous sporangia. These can infect various organs of the plant host: fruits, leaves, shoots, twigs and branches; and cause necrosis and rots. They detected 56 new host–pathogen associations. All are listed, by type of host, in Tables 4 – 6 of the paper.

The surprising diversity and detection of taxa previously described in Australia (see below) illustrate scientists’ still poor understanding of this genus. They also confirm fears that the global phytosanitary system is unable control intercontinental movement of Phytophthora.

The authors express concern because Alpine and subalpine regions are important hotspots for floral biodiversity. The great variation in altitude, aspect, moisture regimes, etc. – including extreme conditions – results in many different habitats on small spatial scales, with large numbers of both plant species and endemics in very confined spaces. The pathogens they discovered are spreading and compromising the biodiversity of these ecologically fragile habitats.

The authors say their study emphasizes the need to assess the full diversity of Phytophthora species and the factors driving the emergence and local spread of these invasive pathogens. They specify studying the Phytophthora communities on fallen leaves to evaluate host specificity, geographic distribution and survival strategies of the main Phytophthora species detected in this study. They report that scientists are currently mapping the distribution of the new species, P. pseudogregata, in the Alpine habitats and trying to establish its natural host range.

another view of the Julian Alps; photo via Rawpixl

Bregant et al. point out that increased scientific interest over the last 30 years has led to discovery of several previously unknown Phytophthora species and pathogen-host associations. They note that all but two of the taxa in one taxonomic grouping, Sub-clade 6b, have been described in the last 12 years. The majority of taxa have been described from forest ecosystems. This trend is depicted in Figure 8 of the article. This figure also displays which species were isolated from nurseries, agricultural systems, and forest ecosystems.

Results by Plant Type – Disease incidence was highest in shrub vegetation, alpine heathlands and along the mountain riparian systems. The most impacted ecosystems were heathlands dominated by common juniper & blueberry, and riparian systems dominated by alders. In these ecosystems, the Phytophthora-caused outbreaks had reached epidemic levels trend with a high mortality rate. On shrubs and heath formations, disease was initially observed in small areas and progressively spread in a concentric manner affecting more plant species.

Hosts and Diseases – Table 3 in the article lists the 33 host plant species, briefly describes the symptoms, and in some cases provides incidence and mortality rates. Those hosts described as suffering “sudden death” included Alnus viridis, Calluna vulgaris, Genista corsica, Juniperus communis, Lycopodium clavatum, Pinus mugo,Rhododendron ferrugineum, Salix alpine, Vaccinium myrtillus and Vaccinium vitis-idaea

Role of P. pseudosyringae – The most common and widespread species detected was P. pseudosyringae. It constituted more than half of the isolates (201 of the 360). Also, it infected the highest number of hosts (25 out of 33, including all three plant types). It was isolated at 36 of the 54 sites distributed throughout all geographic regions. Seventeen of the host–pathogen associations were new to science. (See Tables 4-6, in the paper.)

*Vaccinium myrtillis* – a vulnerable host; photo by Tatyana Prozovora via Wikimedia

P. pseudosyringae dominated disease agents in the shrub community, especially among high-altitude shrubs and heaths, e.g., blueberry, dwarf pine, juniper, rhododendron, and alpine willows. Bregant et al. note that these shrubs are extremely low-growing (an adaptation to high elevation conditions). This form might favor attack by Phytophthora sporangia and zoospores present in fallen leaves. Vaccinium myrtillus suffers particularly severe disease – as previously reported in Ireland. In their laboratory studies, Bregant et al. found P. pseudosyringae to be highly aggresse on common juniper (Juniperus communis), producing wood necrosis and shoot blight only four weeks after inoculation.

The importance of P. pseudosyringae in mountainous regions has been found in previous studies in Asia, Europe, and North and South America. However, the authors call for further study of certain aspects of the species. These regard infectivity and survival of the species’ sporangia in infected tissues fallen to the ground; and the ability of oospores to persist for years in environments subject to extreme low temperatures. The former could increase the risk of outbreaks and promote faster disease progression.

The authors suggest P. pseudosyringae’s survival stems from its production of very large and thick-walled chlamydospores. This reported feature is in contradiction with the original species description, which prompts Bregant et al. to call for a correction.

Other Species, Old and New – P. cactorum was the only Phytophthora species other than P. pseudosyringae detected on all three types of hosts (small trees, shrubs, and herbaceous plants). Phytophthora plurivora was the second-most isolated species. It was detected on 12 hosts in 24 sites.

The new putative species — Phytophthora pseudogregata sp. nov. – was detected on Alnus viridis, Juniperus communis, and Rhododendron ferrugineum. As noted above, scientists are now testing whether other plant species are also hosts. It was detected at two sites in Italy — Borso del Grappa and San Nicolò di Comelico; and one site in Slovenija.

*Juniperus communis*; photo by Joan Simon via Flickr

Diseases of Juniper – Koch’s postulates have been fulfilled, demonstrating that eight Phytophthora species – the new P. pseudogregata sp. nov. as well as P. acerina, P. bilorang, P. gonapodyides, P. plurivora, P. pseudocryptogea, P. pseudosyringae, P. rosacearum – are pathogenic on common juniper (Juniperus communis). The lesions caused by P. pseudosyringae were significantly larger than those caused by other species. Lesions caused by P. pseudosyringae, P. plurivora and acerina progressively girdled the twigs causing shoot blight, browned foliage & wilting symptoms.

Most Threatening Phytophthora clades – The most-frequently isolated Phytophthora species belong mainly to clades 1 and 3 – including P. pseudosyringae. Bregant et al. say these species have several advantages for surviving in mountainous ecosystems: they produce caducous sporangia useful for aerial infections and they tolerate relatively low temperatures. Twoother species in clade 3 were isolated only from the mountains of Sardinia. One, P. psychrophila, was isolated from bleeding cankers on an oak species, Quercus pubescens. Its geographic distribution and impact are still unknown. A second species, P. ilicis, is a well-known pathogen on various hollies in Europe and North America.

Four species belonging to subclade 1a were isolated in the Alps of northeastern Italy and Slovenija. P. cactorum is a widespread polyphagous pathogen found from tropical to temperate climates. It has been responsible for severe diseases on agricultural crops and forest trees. Its occurrence in cold areas has recently been reported in Europe and Australia. The recently described P. alpina has the highest ability to survive in extremely cold conditions. It was detected on four hosts – Alnus viridis, Lonicera alpigena, Vaccinium myrtillus, and V. vitis-idaea.

Some species, e.g., P. hedraiandra and P. idaei, were reported for the first time in natural ecosystems in Europe. They have previously been linked to root and foliar disease in agricultural and ornamental nurseries.

The second-most common species in the Bregant et al. study, P. plurivora, was isolated from 54 symptomatic samples from 12 plant species; eight of the hosts are new. It is common in forest ecosystems of Central Europe – which is now considered to be its region of origin. Little is known about the closely related P. acerina. To date, the latter has been detected widely in agricultural systems, nurseries, forests, and ornamental trees in northern Italy and Sardinia. It is much more rarely found elsewhere. Both P. acerina and P. plurivora are already known to be primary pathogens involved in decline of common and grey alder in Italy.

Five of the Phytophthora species in this study, including the new species P. pseudogregata, are in Clade 6. These include pathogens very common in European forests, e.g., P. bilorbang and P. gonapodyides. Others have more limited or still unknown distributions, e.g., P. amnicola and P. rosacearum. These five species’ ability to cause aerial infections on mountain vegetation might warrant re-evaluation of the reputation of species in this clade being saprophytes or only occasional weak opportunistic pathogens.

P. pseudogregata – in sub-clade 6a – was originally described in 2011 in wet native forests in Australia and on dying alpine heathland vegetation in Tasmania. It has recently been reported in the Czech Republic and Finland. The related P. gibbosa is known to occur only in Australia, where it is associated with dying native vegetation on seasonally wet sites.

Two species of clade 8 — P. kelmanii & P. syringae — have a very limited distribution. A third – P. pseudocryptogea — is widespread in Italian ecosystems from Mediterranean areas to the tree line in the Dolomites. One species from clade 7 (P. cambivora) isolated, mainly from stem bleeding cankers of small trees and shrubs. It has two mating types; bothoccurr in the Alps of northeastern Italy and neighboring Slovenija — on Alnus incana, Laburnum alpinum and Sorbus aucuparia.

SOURCE

Bregant, C., G. Rossetto, L. Meli, N. Sasso, L. Montecchio, A. Brglez, B. Piškur, N. Ogris, L. Maddau, B.T. Linaldeddu. 2024. Diversity of Phytophthora Species Involved in New Diseases of Mountain Vegetation in Europe with the Description of Phytophthora pseudogregata sp. nov. Forests 2023, 14, 1515. https://doi.org/10.3390/f14081515 https://www.mdpi.com/journal/forests

Posted by Faith Campbell

www.fadingforests.org

A newly detected pathogen on elms

I learned at the beginning of August that Canadian scientists have discovered a new pathogen causing wilt disease on American elms (Ulmus americana). The pathogen is Plenodomus tracheiphilus, which is known primarily for causing serious disease in citrus.

P. tracheiphilus is described as common on Alberta’s elm trees, especially in the Edmonton area. It was found on 116 of 200 trees which were sampled – see map. The wilting had previously been blamed on Dothiorella ulmi. I have been unable to find a source for the geographic origin of Dothiorella ulmi; perhaps it is native to North America. It is reported to be present at least from Alberta to Texas. (Presumably if Plenodomus tracheiphilus were in Texas it would have caused obvious symptoms on that state’s citrus crops.)

poster prepared by Alberta Plant Health Lab, Alberta Agriculture & Irrigation, and Society to Prevent Dutch Elm Disease

I am unaware of any North American forest pathologists studying whether this pathogen is also established in the United States, or its possible effects. The discovery in Alberta is the first time this organisms has been associated with disease on elms; I have asked European and North American forest pathologists whether they are looking into possible disease on any of the European or North American elm species. So far, no one reports that s/he has been.

In the meantime, the California Department of Food and Agriculture has begun the process of assigning Plenodomus tracheiphilus the highest pest risk designation for the state. CDFA is worried primarily about damage to the state’s $2.2 billion citrus industry. CDFA is seeking comments on its proposed action; go here .

CDFA points out that despite awareness of the disease on economically important citrus since at least 1900 and efforts by phytosanitary agencies, it has spread to most citrus-growing countries around the Mediterranean and Black seas and parts of the Middle East. The primary mode of spread is movement of infected plant material, e.g., rootstocks, grafted plants, scions, budwood, and even fruit peduncles and leaves. Transmission is possible from latently infected, asymptomatic material. Once established at a site, the conidia produced on diseased plant parts can be spread over relatively short distances by rain-splash, overhead irrigation, water surface flow, or wind-driven rain. Transport by birds and insects is also suspected. The pathogen can survive on pruned material or in soil containing infected plant debris for up to four month.

The report from Canada does not speculate on how a disease associated with plants in a Mediterranean climate was transported to Alberta, which has a cold continental climate. Nor is there any information on the possible presence of the disease on elms in warmer parts of Canada.

U.S. elms appear to be at high risk because phytosanitary restrictions leave dangerous gaps.

First, under the Not Authorized for Importation Pending Pest Risk assessment (NAPPRA) program, USDA APHIS has prohibited importation of plants in the Ulmus genus from all countries except Canada. Second, importation of cut greenery is allowed from all countries – and the CDFA analysis indicates that the pathogen can be transported on leaves. Third, it appears to me that it is probable that this pathogen survives on plants in additional taxa.

See this profile for a description of other threats to North American elms.

SOURCES

Poster prepared by Alberta Plant Health Lab, Alberta Agriculture & Irrigation, and Society to Prevent Dutch Elm Disease https://www.alberta.ca/system/files/agi-plenodomus-poster.pdf

Yang, Y., H. Fu, K. Zahr, S. Xue, J. Calpas, K. Demilliano, et al. 2024. Plenodomus tracheiphilus, but not Dothiorella ulmi, causes wilt disease on elm trees in Alberta, Canada. European Journal of Plant Pathology 169(2):409-420. Last accessed August 1, 2024, from https://link.springer.com/article/10.1007/s10658-024-02836-x

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

Invasive fungus ==> bat populations fall. Insecticides go up, as does infant mortality

little brown bat with WNS; photo by US FWS

Those of us striving to increase action! to curb bioinvasion have had a hard time demonstrating socio-economic costs sufficiently compelling to prompt adoption of more effective policies. However, see

Donovan, et al. 2013. The Relationship Between Trees and Human Health. American Journal of Preventive Medicine. Volume 44 Issue 2.
Fantle-Lepczyk, et al. 2022. Economic costs of biological invasions in the United States. Science of the Total Environment 806 (2022) 151318

Now, two new papers show how high the costs can be under some circumstances.

Eyal G. Frank compared the rate of infant mortality in counties across the U.S. where insectivorous bats had been severely reduced by whitenose syndrome to that in counties where bat populations were not affected. He found that “internal” infant mortality – defined as deaths not caused by accidents or homicides — rose, on average, 7.9%. An estimated additional 1,334 infants died. [A related finding not directly pertinent to this blog’s purpose: Frank notes that real-world use levels of insecticides have a detrimental impact on health, even when used within regulatory limits.]

Whitenose syndrome (WNS) is caused by Pseudogymnoascus destructans, a fungus native to Europe. It was introduced to North America around the beginning of the 21^st Century. It has spread quickly since its initial detection in 2006. By 2024, populations of 12 of ~ 50 insectivorous bat species in the US have been negatively affected. The fungus has caused an estimated decline of more than 90% in bat populations monitored in hibernating caves (Larson, Engst, and Noack 2024).

map showing distribution of WNS in 2013 (a decade ago); from https://databasin.org/datasets/0d755891b0714c9fbce21fbc279608f9/

This population crash is devastating, especially where pest control is supplied by bats. Individual bats are voracious predators. But, for pest control to succeed, the total population must be high. .

Frank compiled information at the county level on:

the spread of white nose syndrome to new counties;
rates of pesticide application, presuming that higher rates reflected increased insect presence on crops; and
infant mortality rates.

Larson, Engst, and Noack (2024) call Frank’s findings on infant mortality “shockingly large”. The increase in insecticide application rates was just 2.7 kg/km². These findings show that technological substitutes for suppressed biological services can markedly and adversely affect human well-being.

Both Frank and Larson, Engst, and Noack emphasize the value in demonstrating that declines in biological diversity do have repercussions for human well-being. They call for more expansive and intensive monitoring of biodiversity trends, especially among non-charismatic taxa such as insects. Furthermore, there should be more multidisciplinary studies that integrate social, natural, and health datasets and research methods to distill information of policy relevance. The Science authors’ expectation is that quantifying these relationships will guide better decisions about conservation policies.

All the authors devote considerable attention to the difficulty in establishing these links because scientists cannot manipulate large-scale ecosystems to conduct experiments. They recommend taking advantage of natural experiments – such as tracking the spread of a newly introduced disease that kills large proportions of a taxon that provides demonstrated ecosystem services. Frank studied the loss of insectivorous bats. Larson, Engst, and Noack (2024) mention an earlier study of the collapse of amphibian populations in Central America caused by the fungal pathogen Batrachochytrium dendrobatidis. One result in this case was an increase in the incidence of malaria.

Frank and Larson, Engst, and Noack clearly hope that this approach will promote stronger and more targetted biodiversity conservation policies and programs. I hope they are right! But did enough Americans hear about these results? I heard a report on “BBC America” radio. I searched and found a print report published by Vox – which connected me to Science. How do we expand media coverage of this type of information?

SOURCES

Frank, E.G. 2024. The economic impacts of ecosystem disruptions: Costs from substituting biological pest control. Science. 6 Sep 2024 Vol 385, Issue 6713 DOI: 10.1126/science.adg0344

The economic impacts of ecosystem disruptions: Costs from substituting biological pest control | Science The econ impacts of ecosystem disruptions: Costs from substituting biological pest control | Science

Larson, A.E., Engist, D., Noack, F. 2024. The long shadow of Biodiversity loss: Technological substitutes are poor proxies for functioning ecosystems. Science 5 Sep 2024 Vol 385, Issue 6713 pp. 1042-1044 DOI: 10.1126/science.adq2373 The long shadow of BD loss | Science

Posted by Faith Campbell