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.
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
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/
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.
Steelhead trout
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
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/
tanoak killed by P. ramorum; photo by F.T. Campbell
I am belatedly catching up on the situation with regard to Phytophthora ramorum – sudden oak death – in the US and other countries.
For a general factsheet on this plant disease, see profile here. Here, I’m summarizing more detailed information contained in the February, May, and August 2024 newsletters of the California Oak Mortality Task Force (COMTF) (Newsletters for earlier months are posted here.)
To obtain the most recent information, you can attend the Fall 2024 virtual meeting of the Task Force on Tuesday, October 29, 2024, from 1 pm to 3 pm PDT. Speakers will focus on the status of P. ramorum in California and Oregon wildlands.
On the next day, Wednesday, October 30, the Phytophthoras in Native Habitats Work Group will discuss “Threats to California Native Plants” including from viruses and excessive heat, along with other concerns.
Participation is free, but registration is required. Complete agendas and more information will be available soon here. Sessions will be recorded and posted to the same site. Questions? Contact Janice Alexander.
More in-depth information à Matteo Garbelotto’s UC Berkeley class, “Ecology and Impacts of Emergent Forest Diseases in California,” is now available free and online. Recommended reading, lecture recordings, slides, even essay topic suggestions are posted. Subjects covered include several high impact forest diseases, molecular diagnostics, disease control, and prevention.
I note that the recent detections of new outbreaks in forests and nurseries support the importance of weather in promoting or hindering establishment and spread of Phytophthora ramorum.
Phytophthora ramorum in North American Forests
In Oregon, P. ramorum continued to spread in 2023 and the first half of 2024.
These outbreaks were detected through extensive surveillance. Aerial monitoring (in cooperation with the USDA Forest Service) and high-resolution imagery covered more than half a million acres in Curry County — the region between the California border and the Coos County line. Ground surveys covered 860 acres. Sampling included 518 trees; 117 tested were positive for the fungus. Stream baits were deployed to 63 sites; 26 tested positive at least once (COMTF newsletter, February 2024; includes maps).
By summer 2024, 23 new P. ramorum infestations had been detected at or beyond the Generally Infested Area (GIA; the area where the pathogen is most commonly found) since 2021. Some of these involve one of the newly detected clonal lineages. Oregon officials are expecting to expand the state’s quarantine area to 901 square miles – 45% of Curry County. The designated GIA would also be enlarged to 178 square miles(COMTF newsletter, August 2024; contains maps).
Oregon continues trying to treat high-priority infestations. In 2023, the state treated 165 acres infested by one of the newly detected clonal lineages, NA2, in the Humbug Mountain area and 347 acres in the Port Orford infestation. Since 2001, Oregon’s Department of Forestry has completed eradication treatments on more than 9,000 acres at an estimated cost of over $37 million. Federal lands comprised 28% of treated acres; the remainder were private and state lands. Still, more than 1,000 high-priority acres have not been treated because neither state nor federal agencies could provide sufficient funds (COMTF newsletter, February 2024).
The stream baiting program in 64 stream drainages has – so far – detected six positive streams. Ground surveys are planned for the new positive drainages along the north bank of the Rogue River and a stream that drains into the Elk River east of Port Orford (COMTF newsletter, August 2024).
In California, recent wet winters have prompted several new detections. The first was in Del Norte County near previously detected sites. The UC Berkeley-coordinated “SOD Blitz” plans intensive surveys in this region in coming months (COMTF newsletter May 2024; contains map).
Somewhat later, new infestations were detected farther south, in Humboldt Redwoods State Park. The new sites were outside the formerly detected sites, on the north side of the creek and up to the top of the ridge (COMTF newsletter, August 2024).
Scientists have realized another concern: several other pathogens cause symptoms on bay laurel, tanoak, and madrone that are almost indistinguishable from SOD. This development will complicate monitoring (COMTF newsletter for August 2024; see below for more details).
Meanwhile, scientists determined that sites where the P. ramorum epidemic is driven by higher bay laurel (Umbellularia californica) densities sustained a higher genotypic diversity of P. ramorum. While tanoak (Notholithocarpus densiflorus) doesn’t contribute much to infection of true oaks (Quercus spp.) it can infect bay laurel, thus perpetuating the infection. Infected oaks and tanoaks maintain host-specific pathogen genotypes (Kozanitas et al. 2024)
The USDA Forest Service program that monitors streams in the East to detect P. ramorum placed baits in 63 streams in 10 eastern states: Alabama, Florida, Georgia, Illinois, Maryland, Mississippi, North Carolina, Pennsylvania, South Carolina, and Texas. In 2023, positive findings for P. ramorum were detected from two streams in Alabama, and one each in Mississippi and North Carolina. All sites are associated with nurseries that had previously tested positive for P. ramorum. Over the last five years – since 2019 – eight streams in four states have tested positive at least once: five in Alabama, and one each in Mississippi, North Carolina, and South Carolina. The detection in South Carolina is new. Vegetation in the watershed has been sampled multiple times; all samples collected so far — plant, soil, and run-off water – have been negative. The pathogen belongs to the NA1 lineage – the one established in forests in West Coast states. [COMTF newsletter February 2024]
from D.J. Haller & M.C. Wimberly. 2020
Situation in Europe
The February 2024, the COMTF newsletter summarized the situation in Great Britain. In England, aerial surveillance covered more than 31,000 ha of larch (Larix kaempferi)plantations. Follow-up investigations detected considerably fewer infested sites than the approximately 200 detected in 2022. Most remain in the southwest and northwest of the country. Weather conditions in 2023 were less conducive for sporulation in 2021 and 2022, which seemed to lead to a reduced level of disease in 2022 and 2023.
In Scotland, widespread aerial and ground surveillance detected a number of sites similar to those found since 2018. Scottish authorities note that where positive findings are not quickly followed by tree removal, localized spread occurred.
In Wales, four helicopter surveillance flights identified around 150 sites deserving further investigation. About 60 of these sites held infected trees, mainly larch, but some noble fir (Abies procera). The COMTF newsletter contains a map showing infested locations. This year’s infection level might be less than in previous years, but this might reflect the fact that the infections are in smaller forest blocks. However, the wet and mild weather in autumn/winter 2023 provided optimal conditions for sporulation, so the scientist expected higher infection rates in 2024. The Welsh Government is working on a new strategy for managing P. ramorum.
In Northern Ireland, P. ramorum was described as still active and spreading. Only two surveys were flown. They identified 49 locations for follow-up, many in forests where the pathogen had been found previously. At two locations, follow-up inspections and sampling of larch confirmed infection by a different pathogen, Phytophthora pseudosyringae. So in 2024, larch samples will be tested for both P. ramorum and P. pseudosyringae.
Other Phytopthoras in Europe
English scientists are trying to determine how damaging P. pseudosyringae is on larch. Infections have been observed at several locations in the north of England, as well as in Northern Ireland (COMTF newsletter February 2024).
Mullet et al. (2024) report that P. pseudosyringae is a self-fertile pathogen of woody plants, especially tree species in the genera Fagus, Notholithocarpus, Nothofagus and Quercus. It is found across Europe and in parts of North America and Chile. Genetic studies show that the North American population originated from Europe. P. pseudosyringae can infect roots; the stem collar region; bark; twigs and stems; as well as leaves. They report it is causing particular damage in Great Britain and western North America. Mullet et al. call for investigation of differences in life history traits between the two main population clusters, including their virulence and host ranges.
Nothofagus obliqua; photo by Line1 via Wikimedia
Chile (COMTF newsletter May 2024)
Concerned about decades of mortality of Nothofagus trees in native forests in Chile, González et al. 2024 sought to understand which other native plants might be reservoirs of inoculum of the pathogen Phytophthora pseudosyringae — which is a documented causal agent of partial defoliation and bleeding cankers on two native tree species,Nothofagus obliqua and N. alpina. P. pseudosyringae can sporulate on lesions on Cryptocarya alba, Nothofagus dombeyi and N. obliqua leaves. On Sophora macrocarpa, sporulation occurs on both asymptomatic tissues and on lesions. S. macrocarpa is a common understory species in Nothofagus forests, so it might be an inoculum reservoir for epidemic events in them.
Look-alikes on California Bay Laurel (COMTF newsletter May 2024)
Similar symptoms from a wide variety of pathogenic organisms were detected on bay laurels after last year’s wet winter. Among the pathogens — the list is not exhaustive — includes P. cinnamomi, Neofusicoccum nonquaesitum, Ganoderma brownie, P. pseudosyringae, P. nemorosa, Botryosphaeria dothidea, Armillaria gallica, Diplodia corticola, and others.
Foliar symptoms tend to look identical on bay laurel leaves. Two foliar pathogens cause particular concern. The first is an “anthracnose” disease of bay laurel caused by a species of Kabatiella. Although known to be present for ~80 years, this organism did not seem to cause problems until 2023. In multiple locations around the San Francisco Bay area, it has caused extensive browning defoliation of bay laurel crowns. Whether the trees will die is uncertain.
The second focus is on a recently named species, Calonectria californiensis. This organism produces P. ramorum-like similar symptoms on a wide variety of native plants, including bay laurel, tanoak, salal, mock-orange, Oregon-grape, and rhododendron. On most of these plants this fungus causes black spots that can grow to kill entire leaves, but apparently C. californiensis is not a pathogen of woody plant parts. Initial symptoms of infection on bay laurel appear identical to those caused by the SOD pathogen (Phytopthora ramorum). C. californiensis does not appear (yet) to lead to lasting debilitating disease or tree mortality.
Nurseries and Managed Landscapes
In administering APHIS’ cooperative program aimed at minimizing spread of P. ramorum via interstate trade in plants, California’s Department of Agriculture (CDFA) relies – at least in part – on funds from USDA. CDFA received $1,308,771 from APHIS in 2023. More than 300 establishments in California are regulated under the program. They submitted ~ 7,400 P. ramorum regulatory samples to the CDFA in 2023. Seventy-eight of the samples were positive (COMTF newsletter February 2024).
At the end of 2023, seven California nurseries that had tested positive for the presence of P. ramorum were operating under the APHIS regulation governing positive nurseries. This was an increase over previous years; zero in 2022, three in 2021 (COMTF newsletter February 2024 Table 4). During 2024 five nurseries were confirmed as positive. Three of these had tested positive in previous years. Two retail nurseries were newly positive; one of these was apparently infected when it brought in plants from another nursery (COMTF newsletter August 2024). I wonder whether the very wet winters California has experienced lately have enhanced the pathogen’s ability to grow – and be detected.
In Oregon, in 2023 the Department of Agriculture regulated five interstate shippers under federal compliance agreements and a sixth intrastate shipper regulated under state requirements (COMTF newsletter February 2024). Spring compliance surveys tested 1,228 foliar samples; ten were positive. After this nursery incinerated all nearby plants, none of the 1,664 foliar samples tested in the fall was positive.
In 2023, the Washington State Department of Agriculture processed more than 300 plant, soil, and water samples; all were negative. Washington also inspected five of the nine nurseries that had ‘opted-out’ of the Federal program so they can no longer ship interstate. Host material appeared free of symptoms so no samples were collected (COMTF newsletter February 2024).
Washington nurseries and regulators frequently encounter the problem of infected plants being shipped into the state from outside. (P. ramorum has been found in 33 Washington nurseries since 2003.) During 2023, the Washington State Department of Agriculture conducted three trace-forward investigations. Fortunately no infestations were detected (COMTF newsletter February 2024). In March 2024, Washington faced another trace-forward involving plants sold to homeowners (COMTF newsletter May 2024). Thirteen tissue samples and two soil samples all tested negative (COMTF newsletter August 2024)
Finally, Washington conducted stream baiting. In 2023, none of the 66 samples was positive (COMTF newsletter February 2024)
Infested Plants
Most of the plant species on which P. ramorum was detected during these years are the usual ones: Rhododendron, Viburnum, Pieris, Arbutus, Prunus, Camellia, Loropetalum. I think the several Cornus species might be somewhat unusual. Disease was confirmed on a new Cornus species, C. capitata (evergreen dogwood). One taxon — Arbutus x ‘Marina’ — is not yet listed by APHIS as a host because Koch’s postulates have not been completed (COMTF newsletters for February 2024 and August 2024).
Research (summarized in the February 2024 newsletter)
Two studies found evidence of seasonal and weather factors influenced establishment of P. ramorum. One study found a clear seasonal pattern of pathogen incidence in the western US, plus a link to the El Niño-Southern Oscillation (ENSO) (Xuechung et al. 2024. The second study looked at a Japanese larch plantation in Scotland (Dun et al. 2024).
In both Scotland (above) and France (Beltran et al. 2024 2024), scientists demonstrated that prompt action helps to suppress P. ramorum establishment.
APHIS Updates its Regulations
In March 2024, APHIS revised the P. ramorum “Domestic Regulatory Program Manual.” The agency said it updated figures and definitions, clarified operational steps, and revised the Retail Nursery Dealer Protocol (COMTF newsletter for May 2024).
Funding
In Fiscal Year 2024, under the Plant Protection Act Section 7721 program, APHIS funded $1 million worth of projects focused on P. ramorum and related species. This was out of a total $62 million in funds dispersed for pest survey, research, mitigation, and outreach programs. This money funded nursery surveys in 11 states. Also, it paid for a project to evaluate the threat of the NA2 & EU2 lineages to nurseries and forests (COMTF newsletter May 2024).
SOURCES
Beltran, A.; Laubray, S.; Ioos, R.; Husson, C.; Marçais, B. 2024. Low persistence of Phytophthora ramorum in western France after implementation of eradication measures. Annals of Forest Science. 81: 7. https://doi.org/10.1186/s13595-024-01222-1
Dun, H.F.; MacKay, J.J. & Green, S. 2024. Expansion of natural infection of Japanese larch by Phytophthora ramorum shows trends associated with seasonality & climate. Plant Pathology. 73(2): 419-430).
González, M.P.; Mizubuti, E.S.G.; Gonzalez, G.; Sanfuentes, E. 2024. Uncovering the hidden hosts: Identifying inoculum reservoirs for Phytophthora pseudosyringae in Nothofagus forests in Chile. Plant Pathology. 73(4): 937-947. https://doi.org/10.1111/ppa.13855. (Summarized in COMTF newsletter February 2024.)
Kozanitas, M.; Knaus, B.J.; Tabima, J.F.; Grünwald, N.J.; Garbelotto, M. 2024. Climatic variability, spatial heterogeneity & the presence of multiple hosts drive the population structure of the pathogen P ram & the epidemiology of Sudden Oak Death. Ecogeography. https://doi.org/10.1111/ecog.07012. (Summarized in COMTF newsletter May 2024.)
Mullet, M.S.; Harris, A.R.; Scanu, B. [and others]. 2024. Phylogeography, origin & population structure of the self-fertile emerging plant pathogen Phytophthora pseudosyringae. Molecular Plant Pathology. https://doi.org/10.1111/mpp.13450. (Summarized in COMTF newsletter for May 2024.)
Xuechung, K.; Wei, C.; Siliang, L.; Tiejun, W.; Le, Y. & Singh, R. 2024. Spatiotemporal distribution of sudden oak death in the US & Europe. Agricultural & Forest Meteorology. 346: 109891)
beech leaf disease symptoms; photo by Matthew Borden via Flickr
Beech leaf disease (BLD) came to attention in 2012 near Cleveland. It has since spread to the Atlantic – Maine to New Jersey and northern Delaware; south into Virginia; north in Ontario; and west to eastern Michigan.
Scientists have scrambled to understand the disease – how it hijacks the tree’s metabolism; & here its impacts on seedlings, saplings, and mature trees; how it spreads, locations at greatest risk.
Now Bartlett Tree Research Laboratories – the research arm of Bartlett Tree Experts – has announced development of Integrated Pest Management (IPM) strategies to treat individual trees – sadly not yet beech in the forest. The project is led by Dr. Andrew Loyd and Dr. Matthew Borden.
Seeing the disease’s impacts on a tree species with aesthetic and ecological values not easily replaced, and its rapid spread, scientists at Bartlett Tree Research Laboratories began testing fungicides and nematicides registered under the Federal Insecticide, Fungicide and Rodenticide Act (FIFRA) by the U.S. Environmental Protection Agency (EPA) to see whether they might be effective against the causal nematode Litylenchus crenatae ssp mccannii.
As Drs. Loyd and Borden note, managing BLD presents numerous challenges:
1. The disease was discovered recently, so there were many unknowns, including how it spreads and the causal organism’s novel life cycle.
2. The damage occurs in leaf buds during winter dormancy. There has been little previous research on such a system. It is difficult for chemicals to reach the tissues.
3. Mature trees are large, so reaching the vulnerable leaves in the canopy is difficult.
4. Treatment efficacy cannot be evaluated until nearly a year after application.
5. Few chemicals are registered for use against foliar nematodes or for trees in forest, nursery, or landscape settings.
6. Obtaining funding is difficult because protecting beech is a low priority among many of the usual sources.
Fortunately, the leadership at Bartlett – the company’s research department, the New England Division leadership, and especially Robert A. Bartlett, Jr. (head of the family-owned company) – saw the importance of protecting beech and have supported this research. The USDA Forest Service has also funded some of studies exploring soil drenches. Cameron McIntire reports that these studies do not yet have results.
Furthermore, Bartlett has chosen to make the science easily available to all interested parties. Three posters explaining experiments to date are available at ResearchGate. They have also published a study on the early tests of Fuopyram as a foliar spray. It is open-access. Additional publications presenting data on experiments with both spray (Fluopyram) and injection (Thiabendazole/Arbotect) are in preparation.
I summarize briefly here their findings as of August 2024.
In all the trials, the scientists judged efficacy of treatments by counting the number of viable nematodes in leaves, viable nematodes in overwintering buds, and BLD symptom severity at appropriate times before and after treatment (spray or injection).
Tests of foliar sprays on small to medium sized trees
The first tests of foliar applications that resulted in BLD suppression were carried out in Ohio starting in 2021, then expanded to other field sites in Ohio and several states in New England in 2022 and 2023 seasons. In early trials, trees were sprayed four times starting in mid to late July at 21-day intervals. The scientists say that recent trials focus on application timing and rate. They hope that optimizing these factors will help generate new recommendations that are more sustainable while maintaining efficacy.
At the annual meeting of the American Phytopathological Society in July 2023, Bartlett announced that Fluopyram is an effective management tool to combat BLD – on smaller trees that can be treated using foliar application. There are several EPA-registered products, though only one, Broadform, has been so far been granted a section 2(EE) recommendation “For Control of Beech Leaf Disease on Beech Trees.”
Treatments are less effective in situations where the inoculum load is very high (for example, a very dense stand of infected trees); or where mature, untreated canopies hang over treated understory beech.
They suggest that managers focus treatments on high-value specimen beech, collection preservation, and potentially uncrowded mixed natural stands.
Treatments should be made by certified pesticide applicators who are familiar with the disease and treatment specifications. For the injection treatment, technical training and specialized equipment is needed. Bartlett arborists and plant health care specialists in locations affected by BLD have all been trained to perform the treatments, and some other arborists are doing BLD treatments as well using the same products.
Soil drench
Matt Borden said that they tested drenches with three different chemicals. The approach did not reduce nemtatode numbers sufficiently. However, as noted above, the Forest Service is funding additional tests exploring possible combinations of drenches with other actions, such as thinning. Discovering management options across a range of application methods (e.g., foliar, injection, drench) and modes of action is vital for a disease that covers such a broad range of locations and tree sizes and forms.
a macroinjection demonstration; photo by Matthew Borden via Flickr
Injections
Scientists injected Thiabendazole (TBZ) into beech on private land in three locations in Ohio and New Jersey. They tested two application rates and three application timings. They have two years of follow-up data for one site, one year for the others.
Key findings:
nematode numbers in buds in late winter consistently reflected foliar symptoms when the leaves opened.
Injections made before mid-July provided the greatest reduction in nemtatode numbers and best canopy improvement. Trees injected late in the season (30 August), after the nematode has begun dispersing from leaves to buds, exhibited some BLD symptoms the next year, but suffered less canopy dieback than controls.
Margery Daughtrey of Cornell said during a discussion of these finding that the trees’ persistence suggests that trees can tolerate some level of symptoms. Among other things, it might be possible to treat the trees less frequently than annually.
TBZ appears to provide at least two seasons of nematode suppression
Bartlett continues to monitor these trees to see how long the injected chemical suppresses nematode numbers and how long the tree remains healthy. They are also establishing new field sites to further optimize rate and timing.
TBZ – in a product called Arbotect 20-S – has been used to manage Dutch elm disease and sycamore anthracnose since the 1970s. However, it is also a well-known nematicide, previously used as an anti-parasitic drug in human and veterinary medicine. Once injected, TBZ protects the tree for more than one season. The injection technology (MACRO-Injection) has also been used for decades. It infuses the chemical directly into the tree’s vascular system; it does not rely on root uptake. Matt says injection does require take technical skill and the right equipment. To minimize the risk of the wound cracking and weeping, the injection should be done low on the side of the root flare, not on top.
While Arbotect 20-S has been registered for use in 48 states for many years, new labeling is required for its use in beech trees and against BLD. Special Local Needs labels, 24(C)s, have been granted by eight states – Connecticut, Massachusetts, Maine, New Jersey, New York, Pennsylvania, and Virginia. Registration in a ninth – Maryland – is in progress and Bartlett scientists are prepared to apply for several more. The problem is that only a limited number of these “special needs” labels may be issued, and BLD has expanded so far, and so rapidly, that it is already infesting beech in more states than may be covered by 24(C)s. Furthermore, 24(C) labels expire if not renewed. Most current 24(C)s will be active through 2028 – not ideal for a disease that will likely be with us long into the future. The product manufacturer (Syngenta) and distributor (Rainbow Ecoscience) are drafting a change to the main Arbotect 20-S label to add beech and the new nematode pest, but warn that EPA review and approval of amendments can take a very long time. Until then, we must resort to limited special local needs labels, and some states will miss out.
contrasting canopy transparency in beech treated with TBZ v. untreated controls; photo by Matthew Borden
One of the key scientists who developed these treatments for Dutch elm disease, R. Jay Stipes, professor emeritus at Virginia Tech, is quoted by Bartlett rejoicing that his work might help protect another tree species.
Matt believes the treatments will be effective if applied every 2-3 years. This approach would also spread out the cost – which will depend on the arborist but Dave Anderson of Rainbow Ecoscience estimated to be about $25 / inch of dbh.
It is always best to obtain an accurate diagnosis before treatment. The next step is talking through your options with a certified arborist or tree disease specialist. The “good” thing about BLD is that it is a progressive disease and will not kill a tree in a single year. Therefore, waiting until you know the disease is present or active locally is generally recommended.
Tree injection is better than foliar application where the latter is impractical (e.g., the tree is tall) or to reduce runoff, particularly near streams. Bartlett recommends treating any beech larger than 10 cm dbh by injection; smaller trees by foliar spray.
Treated trees should be sound, without serious decay, girdling roots, or other conditions that curtail uptake. Based on research results to date, they recommend treating the tree before mid-July. Bartlett is testing the results of injecting the shortly after full leaf expansion – early to mid-June. Bartlett scientists are testing several application rates to determine how long a single injection will suppress BLD. So far they have had good results from both low and moderate label rates (0.4-1.6 fl oz/inch DBH).
All the technical information re: research into treatments and recommendations for applying either the foliar or injection treatments has been provided by Dr. Matthew Borden of Bartlett Tree Research Laboratories. He can be reached at
Dr. Borden says he is immensely grateful for the support that allows him and Dr. Loyd to travel widely to establish the BLD research sites and spend weeks collecting data each year with their team. Company founder Francis A. Bartlett established the Bartlett Tree Research Laboratories as a separate entity within the company, where capital is reinvested directly into stable, long-term support of scientific tree research and preservation. The model is well-suited to provide the flexibility and freedom needed to rapidly respond to emerging invasive species issues.
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/
Flighted spongy moths infesting a ship’s superstructure
USDA’s Animal and Plant Health Inspection Service (APHIS) has issued its annual report for Fiscal Year 2023. The report is part of an enhanced outreach effort that I believe is an effort to persuade the Congress to provide additional funds. However, as I describe below, at this summer’s annual meeting of the National Plant Board, link APHIS’ leadership stated that funding shortfalls are forcing them to curtail many programs. These include ones important to those of us concerned about threats to North American trees. I applaud this action and hope it succeeds!
The report contains some good news but I consider the overall approach depressing. Tree-killing pests continue to receive little attention. The report also emphasizes APHIS’ efforts to facilitate export of agricultural products – an understandable stance given American politics.
The opening summarizes the agency’s activities includes:
Examples of programs targetting pests abroad, before they can reach the U.S. All are fresh fruits and vegetables;
APHIS or staff at U.S. borders:
Approved (cleared) 27,235 shipmentscontaining over 1.87 billion plant units (e.g., a single plant or cutting, or vial of tissue culture plantlets) and 670,811 kilograms of seeds. They intercepted 2,176 quarantine pests. (APHIS carry out these inspections at Plant Inspection Stations – separate from the port environment where DHS Customs and Border Protection (CBP) staff inspects other cargo.)
Identified approximately 92,000 pestsfound during CBP inspections of cargo, mail, and express carrier shipments and took quick action to prevent those of concern from entering the U.S.
Facilitated entry of regulated agricultural cargo by monitoring more than 62,000 treatments of various kinds, that is, fumigations, cold or heat treatments, and irradiation.
Examples of APHIS’ efforts to slow pests’ spread within the country cited plant pest surveys — with coordinated responses — for approximately 45 pests. Also APHIS described funding to help citrus growers combat citrus greening.
The report has separate subreports on the following programs: risk analysis, pest detection, “specialty crop” pests, and tree and wood pests. The last two contain information specific to our interests.
Tree and Wood Pests
This program protects forests, private working lands, and natural resources. It targets specific pests: the Asian longhorned beetle, emerald ash borer, spongy moth, and most recently the invasive shot hole borers. The report notes that numerous native, widespread hardwood tree species are vulnerable to these pests. APHIS asserts an economic justification for the program: conserving forests enhances rural communities’ economic vitality, supports forest-related industries, and maintains the ecosystem services provided by urban trees.
Unfortunately, at this summer’s annual meeting of the National Plant Board APHIS leadership said funding shortfalls forced them to pull back on all these programs.
ALB eradication aims to protect the 30% of U.S. trees that are ALB hosts. These trees support multi-billion-dollar maple syrup, timber, tree nursery, trade, and tourism industries. After reviewing the history of ALB detections, starting in Brooklyn in August 1996, the report describes APHIS’ eradication strategy as comprising surveys, regulatory inspections and quarantine restrictions, removal of infested and high-risk trees, and chemical treatment applications. In FY 2023, the program surveyed more than 763,000 trees across the four regulated areas: New York, Massachusetts, Ohio, and South Carolina. Each program is summarized.
Good news at two locations. On Long Island: only 11 new infested trees were found after a survey of 43,480 trees. In Worcester County, Massachusetts, no new infested trees were found after surveying nearly 360,000 trees. However, in Tate Township, Ohio, surveys detected 163 new infested trees. And in
South Carolina, the program is at an earlier stage — surveying a portion of the quarantine area. The program surveyed nearly 140,000 trees and removed 1,700 in FY 2023.
At the National Plant Board Meeting, Deputy Administrator Mark Davidson explained that the FY2024 appropriation cut $3.6 million from the “tree and wood pests” account. This required the agency to reduce funding for the ALB eradication program.
The report summarizes the spread of EAB since its first detection in 2002 in Michigan to 37 states and the District of Columbia (APHIS does not mention EAB’s presence in five Canadian provinces.)
Saying that EAB has spread beyond what a regulatory program can control, the report notes that APHIS ended the regulatory program in FY 2021. In FY 2023 the agency continued the transition to a program relying primarily on biocontrol. In FY2023, APHIS provided parasitoids to 155 release sites – three in Canada, the rest in 122 counties in 25 states. APHIS and cooperators continue to assess their impacts on EAB populations and tree health at release sites and nearby areas. Field evaluations indicate the EAB parasitoid wasps and other EAB natural enemies (woodpeckers) are protecting regenerating sapling ash from EAB.
At the National Plant Board Meeting, Deputy Administrator Mark Davidson explained that the FY2024 appropriation cut $3.6 million from the “tree and wood pests” account. This required the agency to reduce funding for the EAB containment program – probably the biocontrol component.
Spongy moths (the species formerly called European gypsy moths) are established in all or parts of 20 eastern and midwestern states, plus the District of Columbia. APHIS and state cooperators regulate activities in the quarantine area to prevent the moths’ human-assisted spread to non-quarantine (non-infested) areas – primarily West coast states. To address the moths’ natural spread, APHIS PPQ monitors the 1,200-mile-long border of the quarantine area and adds newly infested areas to the regulated area. The USDA Forest Service – APHIS – Slow-the-Spread Foundation program has greatly reduced the moth’s rate of spread and has eradicated isolated populations.
Another component of the program aims to prevent introduction of members of the flighted spongy moth complex link from Asia. The Asian species have broader host ranges and the females can fly, so they could spread faster. A multi-nation cooperative program is designed to prevent the moths’ hitchhike on vessels coming from Asia. link APHIS supports this program through negotiations and support of CBP’s offshore vessel inspection, certification, and cleaning requirements. Canada participates in the same program.
In FY 2023, APHIS and state cooperators continued efforts to delimit possibly introductions in Washington State (no additional moths detected); and California and Oregon (initial detections in FY 2020).
At the National Plant Board Meeting, Deputy Administrator Mark Davidson explained that the FY2024 appropriation cut $3.6 million from the “tree and wood pests” account. This required the agency to reduce funding for the flighted spongy moth program.
California sycamore infested by polyphagous shot hole borer; photo by Beatriz Nobua-Behrmann UC Cooperative Extension
The report notes that various non-native shot hole borers have been detected in several states. Their hosts include trees in forests and urban landscapes, tea plantations, and avocado orchards. The program’s focus was apparently on the polyphagous and Kuroshio shot hole borers devastating riparian habitats in southern California and urban areas in other parts of California. At California’s request, APHIS and the USDA Forest Service helped establish a working group, led by USFS, with the goal of strategically addressing both shot hole borers in California. In FY 2023, APHIS’ helped with foreign explorations for possible biocontrol agents, as well as host specificity testing.
APHIS leadership told the National Plant Board in July 2024 that it had dropped this entire program due to funding shortfalls.
Specialty Crop Pests
While much of this report concerns pests of agricultural crops (e.g., grapes, citrus, potatoes), it also summarized efforts re: Phytophthora ramorum (sudden oak death) and spotted lanternfly. APHIS says its efforts protected nursery stock production worth approximately $1.3 billion as of 2019, and tree fruit production worth approximately $1.7 billion in 2021.
map showing areas of the Eastern United States at risk to P. ramorum – developed by Gilligan of Cambridge University
The report states that APHIS seeks to limit P. ramorum’s spread from affected nurseries. The agency does this via regulatory strategies. During FY 2023, 16 nurseries were governed by more stringent rules under the federal program which are imposed on nurseries that have been determined in past years to harbor P. ramorum-infected plants.
In addition, Oregon officials continued surveys of an area outside its quarantine zone because of a detection the previous year. APHIS will adjust the federal quarantine depending on the state’s findings.
The APHIS report does not discuss several pertinent events that occurred in FY2023. [For more details, read the California Oak Mortality Task Force newsletters for 2023 – posted here.
First, APHIS does not mention or discuss the implications of detection of two new strains of P. ramorum — EU1 & NA2 — in west coast forests. The presence of EU1 in a new California county (Del Norte) was confirmed in Feb 2023.
Second, the report said that Oregon is trying to determine the extent of the P. ramorum infection detected outside the state’s quarantine zone. However, it does not mention that this outbreak involves the new NA2 lineage – and that NA2 was known to be present in nurseries in the region since 2005.
The report also does not clarify that three nurseries to added to the more stringent program were so treated because SOD-infected plants were found on their premises.
Nor does the report note that at least two new naturally-infected hosts of P. ramorum were identified: Western sword fern (Polystichum munitum) and Arbutus x ‘Marina’.Koch’s postulates need to be completed on the latter so it has not yet been added to APHIS’ official host list.
Spotted Lanternfly (SLF) was found in 16 states in FY 2023. APHIS’ program enjoyed funding provided through Specialty Crop Pests and from the Plant Protection Act’s Section 7721 link ($6 million from the latter).
The report notes that APHIS still does not have enough data to determine SLF’s impacts on agriculture. Thus far, vineyards have been the most adversely affected agricultural segment, mostly due to SLF acting as a stressor to vines. Also, the sticky, sugary “honeydew” produced by SLF attracts other insects and promotes sooty mold growth. These can ruin the fruit and further damage the plant.
SLF populations are strongly linked to major transportation pathways, such as railroads and interstate highways. APHIS targets treatments and, in some areas, removes SLF’s preferred host plant (tree of heaven [Ailanthus]), from transportation hubs. The aim is to reduce the risk of SLF’ spread to new areas and to eradicate isolated infestations. In FY 2023, APHIS and cooperators treated 4,637 properties covering 6,455 acres in affected areas. However, during the National Plant Board meeting both state and APHIS officials complained to me that managers of these transportation hubs raise many barriers to their access, sharply limiting the program’s chance of success.
Ailanthus altissima – drive of spotted lanternfly invasion
The program was expanded after National Environmental Policy Act-mandated environmental review. This allowed APHIS to conduct treatments in four additional states—Indiana, Massachusetts, Michigan, and Rhode Island. In addition, program cooperators identified three potential biological control organisms, one that targets the tree of heaven and two that target SLF. APHIS will continue to evaluate them and develop methods to rear them in the laboratory.
Finally, in fiscal year 2023, APHIS joined the National Association of State Departments of Agriculture and the National Plant Board to develop a national strategic plan outlining the future direction of the SLF program. With the strategic plan, PPQ aims to harmonize the approach across states to slow SLF’s spread, develop consistent outreach messaging for a nationwide audience, and more effectively use existing state and Federal resources. Continued spread of SLF despite the huge effort, rising costs of the program, and new scientific findings spurred reconsideration of the strategy.
To summarize, I hope that APHIS’ annual report will – in the future – help members of Congress and their staff understand the agency’s programs’ purpose and past successes. This increased understanding might make it easier to advocate for more funding. I am troubled, however, by the agency’s glossing over significant problems.
Posted by Faith Campbell
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/
Breeding Port-Orford cedar for resistance to Phytophthora lateralis; photo by Richard Sniezko, USDA Forest Service
At the annual meeting of the National Plant Board in July, I learned that two new Phytophthora species have been detected in the United States. Questions remain about how each arrived.
Phytophthora austrocedrii
This species was detected in a nursery in Oregon, then traced back to a supplier in Ohio. Officials are trying to determine how it entered the country – and then spread.
junipers in Great Britain killed by P. austrocedri; Forestry Research
In the United Kingdom, P. austrocedri has killed trees in the Juniperus and Cupressus genera. Damage is particularly significant at two sites in northern Scotland and in England’s Lake District. The principal host, Juniperus communis, is an important native species. It is already considered vulnerable. P. austrocedri has also been detected in Argentina, where it is killing the native Chilean or Patagonian cedar (Austrocedrus chilendris). The cedar species is the only one in the genus. Evidence indicates the pathogen was introduced to both Britain and Argentina; but its origin is unknown. Indeed, the species was first isolated by scientists as an unknown Phytopthora taxon on a juniper in an import/export nursery in Germany. All reported hosts are members of the Cupressaceae family (UK forest research website).
Of greater concern to Americans, P. austrocedri has also infected individual trees of Port-Orford cedar (Chamaecyparis lawsoniana). (UK forest research website).
Port-Orford cedar is a species endemic to a small range in southwestern Oregon and Northwestern California.
POC populations have been severely reduced over the past century by a different non-native Phytophthora, P. lawsonii. US Forest Service scientists recently announced that they have bred trees resistant to this pathogen – and offered seedlings for widespread planting.
Possible hosts in the Pacific Northwest – other than Port Orford cedar – include Juniperus californica, Juniperus grandis, Juniperus occidentalis, and Juniperus maritima – although the junipers might be limited to arid environments, where they would presumably be less vulnerable. https://plants.usda.gov/home/classification/15147
Research in Great Britain shows that P. austrocedri spreads in water and by movement of infected plants and contaminated soil. Footwear, camping equipment, and vehicle tires can all carry the pathogen. This makes the pathogen particularly difficult to control (this is another similarity with P. lawsonii).
Phytophthora abietivora
P. abietivora was originally found on a diseased Christmas tree (Fraser fir, Abies fraseri) in Connecticut in 2019. It has since been reported in Pennsylvania and Virginia; and in forest nurseries and Christmas tree plantations in Quebec and Ontario. The Canadians report that it has not caused disease (Canadian website). However, the Canadian representative at the National Plant Board meeting expressed concern and asked USDA APHIS to clarify what actions it is taking regarding this species.
(Natural populations of Fraser fir have been severely reduced over the past century by the balsam woolly adelgid.)
Fraser fir killed by balsam woolly adelgid; Clingman’s Dome, Great Smoky Mountains National Park
Several additional hosts have been identified, including balsam fir (Abies balsamea) and eastern hemlock (Tsuga canadensis); and deciduous or hardwood species: hickory (Carya sp.), flowering dogwood (Cornus florida), American witch hazel (Hamamelis virginiana), mountain holly (Ilex montana), red maple (Acer rubrum), silver birch (Betula lenta), American beech (Fagus grandifolia); and several oaks: white (Quercus alba), chestnut (Q. montana) and northern red oak (Q. rubra) (Canadian fact sheet).
According to the Canadian website, P. abietivora causes root rot and subsequent foliar chlorosis, discoloration, stem cankers, and sometimes tree decline and death. Determining which Phytophthora species is the causal agent of a tree’s symptoms requires laboratory testing. The Canadian fact sheet reports that wet, cool conditions provide ideal environments for P. abietivora. Like other Phytophthora species, P. abietivora can be spread through soil and water, as well as via infected plant material or pots or trays (particularly if soil remains on the equipment). The Canadian fact sheet has several photographs illustrating symptoms and additional sources.
Liriodendron tulipifera; photo by Evelyn Simak via Geograph
Phytophthora kernoviae
P. kernoviae was first detected in southwestern England in 2003. link In England, this pathogen has caused significant diseases in native Fagus sylvatica (European beech) and lesions on trunks of a European oak, Quercus robur. More worrying are the trunk lesions on the North American native yellow or tulip poplar (Liriodendron tulipifera) and lesions on foliage of Monterey pine (Pinus radiate), giant sequoia(Sequoiadendron giganteum), and several North American native shrubs, Rhododendron macrophyllum (Pacific rhododendron), R. occidentale (western rhododendron), R. catawbiense(Catawba rosebay) and Umbellularia californica (California bay laurel).
The infestation in Cornwall is sustained by heavy sporulation on the non-native shrub Rhododendron ponticum, which is invasive in woodlands. Worrying for Americans is the fact that P. kernoviae sporulates on three plant species native to West coast forests — Rhododendron macrophyllum, R. occidentale, and Umbellularia californica – as well as on R. catawbiense, which is native to the southern Appalachians.
USDA APHIS requested adoption of a “response plan” targetting P. kernoviae under the National Plant Disease Recovery System (NPDRS). This plan was adopted in 2008 and updated in 2015.
The recovery plans found the areas at highest risk are eastern slopes of the Appalachian Mountains because this area combines a native sporulating host and residential landscaping choices that are likely to include hosts that could transport the pathogen. A lower risk was identified for West Coast forests.
Because of this status, P. kernoviae is also a “priority” pest for surveys under the Cooperative Agricultural Pest Survey (CAPS) program. According to Purdue University’s “pest tracker” website four states have reported carrying out surveys for P. kernoviae in one or more years since 2016: Oregon, Tennessee, Pennsylvania, and Virginia. Surveys in Oregon were carried out in 2018 – 2020. In 2020 the counties surveyed included Curry County, where three strains of P. ramorum link have become established. The Purdue list is not certified as accurate or complete. To date, no surveys have detected P. kernoviae in the United States or – I believe – in Canada.
For details on existence of two clonal lineages of Phytophthora austrocedrii, see Henricot, B. A. Perez-Sierra, A.C. Armstrong, P.M. Sharp, and S. Green. Phytopathology 2017. 107:12, 1532-1540.
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/
The House and Senate Agriculture committees are edging toward adopting the next Farm Bill, which is a year past due. Farm bills set policy, funding levels, and more, for 5 years. Each covers a wide range of subjects, including crop subsidies and insurance; food stamps; rural development (including wifi access); forestry policy; and research.
As you might remember, CISP aims to improve USDA’s programs — not only to prevent introductions of non-native tree killing pests and pathogens but also to better respond to those that enter the US and become established. I summarize here what the Senate and House bills have in common and how they differ on these issues.
I understand that the minorities, that is, House Democrats and Senate Republicans, have not accepted all aspects of the majorities’ drafts. So let’s take the opportunity to ask for better bills.
Both the House and Senate bills would “simplify” the USDA Forest Service’s obligations to prepare environmental assessments under the National Environmental Policy Act (NEPA). I have not analyzed which bill weakens NEPA more.
The Senate Bill: The Rural Prosperity and Food Security Act of 2024
The Senate bill addresses forest pest species in several places: Title II — Conservation, Title VII — Research, and Title VIII — Forestry. Here, I describe relevant sections, beginning with the section that partially addresses CISP’s proposal.
Title VIII — Forestry. Section 8214 requires the USDA Secretary to establish a national policy to counter threats posed by invasive species to tree species and forest ecosystems and identify areas for interagency cooperation.
This mandate falls far short of what we sought in a previous bill (S. 1238). However, depending on the exact wording of the bill and accompanying report, perhaps we can succeed in building a stronger program.
It is most important to obtain funding for applied, directed research into resistance breeding strategies, “bulking up,” and planting seedlings that show promise. Please contact your senators and ask them to work with the sponsors – Peter Welch [D-VT], Maggie Hassan [D-NH], and Mike Braun [R-IN] – to try to incorporate more of S. 1238 in the final bill.
The Senate bill contains other provisions that might be helpful for invasive species management – although not part of what CISP and our partners asked for.
‘ōhi‘a trees killed by rapid ‘ōhi‘a death; photo by Richard sniezko, USFS
Title VIII — Forestry. In Section 8506, the Senate bill would require that the US Departments of Agriculture and Interior continue working with Hawai`i to address the pathogen that causes rapid ‘ōhi‘a death. The section authorizes $5 million for each of the coming five fiscal years to do this work. Unfortunately, authorization does not equal funding. Only the Senate and House Appropriations Committees can make this funding available. Hawai`i’s endemic ‘ōhi‘a trees certainly face a dire threat. CISP is already advocating for funding to support resistance breeding and other necessary work.
Title VIII — Forestry. Sections 8247 and 8248 support USDA Forest Service’s nursery and tree establishment programs. My hesitation in fully supporting these provisions is that I fear the urge to plant lots of trees in a hurry will divert attention for the need to learn how to propagate many of the hardwood tree species that have been decimated by non-native pests. However, I agree that the U.S. lacks sufficient nursery capacity to provide anything close to the number of seedlings sought. Perhaps this program can be adjusted to assist the “planting out” component of our request.
Title VII — Research. Section 7208 designates several high-priority research initiatives. On this list are spotted lanternfly, and “invasive species”. A number of forest corporations have been urging Members of Congress to upgrade research on this broad category, which I believe might focus more on invasive plants than the insects and pathogens on which CISP focuses. How the two ideas are integrated will be very important.
Another high-priority initiative concerns the perceived crisis in failed white oak regeneration.
Title VII — Research.Section 7213 mandates creation of four new Centers of Excellence at 1890 Institutions. These are historically Black universities that are also land-grant institutions]. These centers will focus on: 1) climate change, 2) forestry resilience and conservation; 3) food safety, bioprocessing, and value-added agriculture; and, 3) food and agricultural sciences and the social sciences.
Title II — Conservation. Section 2407 provides mandatory funding (which is not subject to annual appropriations) of $75 million per year to the national feral swine eradication/control program (run by USDA APHIS’ Wildlife Service Division). I discuss this program in a separate blog.
The Senate bill also mandates use of several conservation and other programs to address the causes and impacts of climate change. This requirement is directly countered by the House Agriculture Committee’s bill (see below).
Title VIII — Forestry. This section contains none of the provisions CISP’ sought to USDA’s management of tree-killing non-native insects and diseases.
Instead, the House bill calls on the USFS to establish a comprehensive approach to addressing the demise of the giant sequoia trees.
Title VII — Research The House bill, like the Senate’s, lists the invasive species and white oak research initiatives as high priority. The House, unlike the Senate, does not include spotted lanternfly.
Title II — Conservation. As I noted above, the House bill explicitly rescinds all unobligated conservation funding from the Inflation Reduction Act. It reallocates these funds to the traditional conservation programs, e.g., the Environmental Quality Incentive Program and Watershed Protection and Flood Prevention. The bill would use these funds to support “orphan” programs – naming specifically the national feral swine eradication/control program. The House bill provides $150 million – apparently across the five years covered by the Farm Bill, so $30 million per year. Finally, the House allocates 60% of the hog management funds to APHIS, 40% to the Natural Resources Conservation Service.
spotted lanternfly – target of at least 11 projects funded through APHIS’ the Plant Pest and Disease Management and Disaster Prevention Program in FY24. Photo by Holly Raguza, Pennsylvania Department of Agriuculture
Title X —Horticulture, Marketing, and Regulatory Reform. The House’s summary says it is taking steps to protect plant health. It does this by increasing funding for the grant program under the Plant Pest and Disease Management and Disaster Prevention Program – §7721 of the last (2018) Farm Bill. The increase would raise the amount of money available each year from the current level of $70 million to $90 million. These funds are mandatory; they are not subject to annual appropriations. Research, development, and outreach projects funded by this program have certainly added to our understanding of plant pests, hence to their effective management. However, they are usually short-term projects. Therefore they are not suitable for the long-term commitment required for resistance breeding programs. See here and here.
Title III — Trade. Here, the House bill exacerbates the current imbalance between trade promotion and phytosanitary protection. The bill doubles the authorized funding for USDA’s Market Access and Foreign Market Development programs. I concede that this measure probably does reflect a bipartisan consensus in the Congress to support robust programs for promoting agricultural exports.
Also under this Title, the House bill requires the USDA Secretary to conduct regular assessments to identify risks to critical infrastructure that supports food and agriculture sector. This might be helpful – although it is not clear that this assessment would include to threats to forest or urban trees not used commercially (e.g., for timber).
At a recent forum on biological control sponsored by the National Association of State Foresters (NASF), it was reported that participants noted several problems: insufficient funding, significant delays in refilling positions, inadequate research capacity, lack of brick-and-mortar infrastructure, and declining college enrollments in biocontrol-related studies. The NASF Forest Science Health Committee is developing a “Statement of Needs” document that NASF and others can use to lobby for funding to fill these gaps. I hope you will join them in doing so!
salt cedar (Tamarix sp.) attacked by biocontrol agent; photo by J.N. Stuart via Flickr
However, as I note above, empowering resistance breeding programs requires a long-term commitment, that is, a comprehensive alteration of policies and infrastructure – beyond annual appropriations.
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
Sakhalin knotweed (Fallopia (Reynoutria) sachalinensis) – an invasive plant widespread in Europe; photo by Katrin Schneider [korina.info] via Wikimedia
There is growing evidence that invasive plants – as distinct from invasive species of animals, microbes, etc. – play a significant role in causing the loss of floristic uniqueness at the local or regional level. I provide full citations of all sources at the end of this blog.
Less Diversity. More Similarity
Several studies show that plant invasions have a bigger impact than extinction in the homogenization of Earth’s flora. A major driver is sheer numbers. Daruet al. point out that 10,138 plant species have become naturalized to a region outside their native ranges while only 1,065 species have gone extinct. Even under a scenario in which all species currently included in IUCN Red List as “threatened” become extinct, non-native plant species naturalizations are by far the stronger contributor to biotic reorganization.
Winter et al. report that in Europe since AD 1500, plant invasions have greatly exceeded extinctions, resulting in increased taxonomic diversity (i.e., species richness) on the Continent but increased taxonomic and phylogenetic similarity among European regions. In other words, floras of individual European countries became phylogenetically and taxonomically impoverished. This situation is likely to worsen in the future because introductions continue.
Winter et al. conclude, more broadly, that a focus on species richness can be misleading because it does not capture the important effects of taxonomic or phylogenetic distinctiveness.
Yanget al. (2021) considered the situation globally. They divided most of Earth’s ice-free land surface into 658 regions. They found that introduction of non-native plants has increased the taxonomic similarity between any two of these regions in 90.7% of the time. Introductions increased phylogenetic similarity in 77.2% of those pairs. Australasia illustrates the situation. The region has a large proportion of endemic species, reflecting its unique evolutionary history and exhibiting high floristic diversity. However, the region has also received large numbers of non-native plants from other regions. The result is that the Australasian flora has lost much of its original uniqueness.
rubbervine (Cryptostegia madagascariensis) – one of the worst invasive plants in Australia; photo by Tatters via Flickr
Introduced plant species rarely cause outright extinction of members of the native flora of the receiving ecosystem – at least at the scale of a continent. Winter et al. found that in Europe, extinction usually occurs to plant species with small numbers that are limited to localized habitats. Often, however, the same species continue to thrive elsewhere on the continent. The “losing” country finds its flora becoming more similar to that of other European countries. It loses some uniqueness because it lost one or more components of its flora. However, for Europe as a whole, there is no loss. The homogenization of the “losing” country’s flora is exacerbated by the fact that more than half of plant species listed as invading a particular European country are from other European regions. Winter et al. say a similar pattern has been found in North America.
The picture is more complex for small isolated ecosystems. Carvallo and Castro (2017), writing about isolated volcanic islands in the southeastern Pacific Ocean, introduction of large numbers of non-native plant species has not caused extinction of native plant species. It has, however, resulted in the homogenization of the islands’ floras.
These authors worry that this reduction in phylogenetic diversity could have detrimental impacts for ecosystem function and ecosystem services. (Interestingly, at the level of order or family rather than species or genus, the combined effects of species introductions and extinctions did not change the islands’ taxonomic diversity. They don’t explicitly say whether that fact might mitigate effects on ecosystem function.)
What is the situation in Hawai`i? The Islands are the “capital” of both extinction and invasion. I know the Hawaiian flora has very high levels of endemism and of endangerment. In addition, naturalized non-native plant taxa constitute up to 54% of the archipelago’s flora (Potter et al. 2023). However, it is probably extremely difficult to distinguish the impacts of introduced plants separate from the impacts of the many non-native animals, e.g., feral hogs.
Extinction by Introduction
It has been reported that invasive species have caused the extinction of at least seven species of plants on the Cape of Good Hope and endangered another 14% (Houreld 2024). Unfortunately, the report doesn’t specify whether the non-native species are plants or animals. Either way, this is a tragedy. I remind you that the Cape Floral Kingdom is Earth’s smallest Plant Kingdom in geographic size (78,555 km2), and extremely important in uniqueness. According to the article in The Washington Post, two-thirds of the 20,400 plant species growing in South Africa are endemic – found nowhere else on Earth.
Nearly a decade ago, Downey and Richardson objected to measuring the impact of introduced plant species by considering only total extinction of native plant species. They complain that this approach fails to recognize that plants experience a long decline before reaching extinction. They divide this decline into six “thresholds”. Downey and Richardson say there is abundant evidence of invasive plants driving native plants along this extinction trajectory. For example, increases in non-native plant cover or density that result in decreased native plant species diversity or richness equates, under their hierarchy, to the native species crossing from the first to second threshold. They note there are also examples of species causing “extinction debts”. That is, the extinction occurs long after the invader is introduced and initiates a native species’ decline. They call for conservationists to intervene earlier in that trajectory.
The Global Assessment on Biodiversity and Ecosystem Services was recently published by the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services. This report notes that there are at least 1,061 invasive plants on Earth. In terrestrial systems, invasive plants are the taxonomic group most frequently reported as having negative impacts, especially in cultivated areas, plus temperate and boreal forests. As I have noted above, non-native plant taxa constitute a particularly high proportion of the flora on islands. The assessment found that the number of non-native plants exceeds the total number of native plants on more than one quarter of the Earth’s islands. However, this report does not distinguish the number of species endangered by plant invasions from the number of species endangered by invasive species of all taxonomic groups.
Tiburon mariposa lily (Calochortus tiburnensis) – a federally Threatened species in California; photo by T.J Gehring via Flickr
None of the experts denies the impact of extinction on biodiversity. Extinction represents the loss of phylogenetically and taxonomically unique organisms. This loss is exacerbated if some taxonomic groups are at disproportionately higher risk of extinction. Introduced non-native species compensate for these losses only to a point (Daru et al.). In Europe, Winter et al. found that extinctions usually befall specialized endemic or rare species, often from species-poor families. On the other hand, successful invaders are often ecological generalists with large ranges, often belonging to species-rich families. This results in the pronounced decrease of phylogenetic and taxonomic ß-diversity within and between regions to which the rare species are unique.
All these experts agree that species declines — short of extinction — have severe impacts on ecosystem functions, and even evolution.
Yang et al. emphasize that the rapid and accelerating loss of regional biotic uniqueness changes biotic interactions and species assemblages, with probable impairment of key ecosystem functions. Daru et al. stress that biotic homogenization— declining ß-diversity—reduces trait and phylogenetic differences between regions. Conceding that the consequences of this global biotic reorganization on ecosystems are poorly understood, Daru et al. cite increasing evidence that biotic heterogeneity provides insurance for the maintenance of ecosystem functioning in a time of rapid global change. They assert that declining ß-diversity is a more characteristic feature of the Anthropocene than species loss.
Winter et al. also state that the phylogenetic structure of a species assemblage represents the evolutionary history of its members and reflects the diversity of genetic and thus morphologic, physiologic, and behavioral characteristics. High phylogenetic diversity might enable rapid adaptation to changing environmental conditions.
According to Daru et al., the loss of 14 billion years of evolutionary history has affected some regions particularly. The most disturbed biotas include those of California and Florida; Mesoamerica; the Amazon; the Himalaya-Hengduan region; Southeast Asia; and Southwest Australia. These are regions that experienced spectacular taxonomic radiation over time, and now have both high levels of threat and also species invasion.
Carvallo and Castro, focused on the Pacific islands, call for integrating the two parallel channels of conservation that currently operate separately: those focused on reversing plant extinctions and those focused on reducing invasions. They call for a biogeographical approach that addresses all causes of phylogenetic homogenization.
Tetragonia tetragonoides – the most widespread invasive plant on these Pacific islands; photo by Jake Osborn via Flickr
I believe all these experts, in all their papers, have made the case for such integration world-wide.
Invasive plants’ impact on non-plant species
While I have focused here – and in most of my blogs more broadly — on impacts on wild, native plant communities, it is clear that alterations to floristic communities influence other taxonomic groups. A couple of years ago I summarized findings by Douglas Tallamy and colleagues on what happens to insects – and their predators – when a landscape is dominated by introduced plant species.
In short, domination by non-native plants – whether invasive or just widely planted – suppresses the numbers and diversity of native lepidopteran caterpillars. One study cited in the blog found that 75% of all lepidopteran species were found exclusively on native plant species. Non-native plants in the same genus as native plants often support a similar but depauperate subset of the native lepidopteran community. Tallamy and colleagues conclude that a reduction in the abundance and diversity of insect herbivores will probably cause a concomitant reduction in the insect predators and parasitoids of those herbivores – although few studies have measured this impact beyond spiders, which are generalists. Tallamy focuses on birds.
In the same blog I reviewed publications by Lalk and colleagues, which examined interactions between invasive woody plants and arthropod communities more broadly. They decried the insufficient data about most of these interactions.
A few weeks ago I saw a report of an unexpected impact of invasive plants: roots of beach naupaka [beach cabbage or sea lettuce] (Scaevola sericea) are penetrating sea turtle nests so aggressively that they kill the unhatched turtles. Apparently this is happening at several sites in the Caribbean, where the plant is not native (Houreld 2024). I could find no scientific reports of this phenomenon. One reference noted that a related species (S. taccada) can form large, dense stands that might prevent adult sea turtles’ access to nesting areas (Swensen et al. 2024).
Sources:
Daru, B.H., T.J. Davies, C.G. Willis, E.K. Meineke, A. Ronk, M. Zobel, M. Pärtel, A. Antonelli, and C.C. Davis. 2021. Widespread homogenization of plant communities in the Anthropocene. NATURE COMMUNICATIONS (2021) 12:6983. https://doi.org/10.1038/s41467-021-27186-8
www.nature.com/naturecommunications
Downey, P.O. and D.M. Richardson. 2016. Alien plant invasions and native plant extinctions: a six-threshold framework. AoB Plants, 2016; 8: plw047 DOI: 10.1093/aobpla/plw047; open access, available at http://aobpla.oxfordjournals.org/
Houreld, K. 2024. “Parched Cape Town copes with climate change by cutting down trade.”. The Washington Post. February 29, 2024.
Potter, K.M., C.Giardina, R.F. Hughes, S. Cordell, O. Kuegler, A. Koch, and E. Yuen. 2023. How invaded are Hawaiian forests? Non-native understory tree dominance signals potential canopy replacement. Landsc Ecol https://doi.org/10.1007/s10980-023-01662-6
Yang, Q., P. Weigelt, T.S. Fristoe, Z. Zhang, H. Kreft, A. Stein, H. Seebens, W. Dawson, F. Essl, C. König, B. Lenzner, J. Pergl, R. Pouteau, P. Pyšek, M. Winter, A.L. Ebel, N. Fuentes, E.L.H. Giehl, J. Kartesz, P. Krestov, T. Kukk, M. Nishino, A. Kupriyanov, J.L. Villaseñor, J.J. Wieringa, A. Zeddam, E. Zykova. and M. van Kleunen. 2021. The global loss of floristic uniqueness. NATURE COMMUNICATIONS (2021) 12:7290.
Winter, M., O. Schweiger, S. Klotz, W. Nentwig, P. Andriopoulos, M. Arianoutsou, C. Basnou, P. Delipetrou, V. Didz.iulis, M. Hejdah, P.E. Hulme, P.W. Lambdon, J. Pergl, P. Pys.ek, D.B. Roy, and I. Kuhn. 2009. Plant extinctions and intros lead to phylogenetic and taxonomic homogenization of the European flora PNAS Vol 106 # 51 December 2009
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
white flowered trillium (Trillium grandiflorum); via PICRYL One of the “charismatic wildflowers” mentioned by Blossey and colleagues
Bernd Blossey, Darragh Hare, and Don Waller have published a plea that America’s federal government take the lead in formulating a new national program on managing deer. Otherwise, they fear that deer populations will not be reduced to ecologically sustainable levels. I find their argument convincing and well-sourced. I agree that Americans need to figure out how to address this threat. (The full citation is at the end of this blog).
First, Blossey and colleagues describe the damage caused by overabundant deer:
severe declines in populations of many native forest herbs and shrubs, probably including disappearing wildflowers;
their replacement by non-native species that are less palatable;
poor regeneration of many canopy hardwood species;
decreased forest resilience, lowering forests’ ability to adapt to stressors, especially climate change;
decreased ability of forests to deliver benefits that are of increasing value to many people;
increased prevalence of wildlife and human diseases associated with the spread and size of growing tick populations; and
people – and deer — killed vehicle accidents on roads.
The widespread impacts of white-tailed deer (Odocoileus virginianus) in forests of the East are well-documented (see my previous blogs for a few examples; scroll below the “Archives” to find “Categories”). Blossey and colleagues note examples of similar impacts in the West, attributed to elk (Cervus elaphus) and black-tailed and mule deer (Odocoileus hemionus).
The authors review the decimation of deer populations in earlier centuries and the efforts of state wildlife agencies to rebuild their populations during the 20th Century. The problem, in their view, is that federal and — especially — state wildlife agencies have retained their traditional focus on managing wildlife for recreational hunters. However, recreational hunters make up a small and shrinking proportion of all Americans. Many more people now engage in “non-consumptive” enjoyment of wildlife.
lack of regeneration in Rock Creek Park, Washington D.C.; photo by Sam Sheline, NatureServe, via Flickr
State agencies’ narrow focus might partly arise from fragmented authorities. Agencies other than wildlife departments are responsible for addressing some repercussions of overabundant deer. These include threats to human health, loss of agricultural crops.
For several reasons, Blossey and colleagues call for federal leadership. They think that only a national strategy can address, in a holistic way, the interrelated deer, human health, forest, and biodiversity crises. The strategy’s goal should be to protect species that are in decline because of over-browsing by deer and to avoid further declines in environmental and human health.
The authors reason that states are tied to traditional constituencies. Also, they have difficulty acting across jurisdictional boundaries. Second, few state wildlife agencies have authority to protect plant and invertebrate species. Yet these are the taxa most directly affected by overabundant deer. Blossey and colleagues point out that, of the ~1,300 species listed under the federal Endangered Species Act, 942 are plants and 287 are invertebrates.
They point out that deer also suffer the effects of overpopulation. Chronic wasting disease is spreading. It causes a slow, agonizing death of affected animals. Another 2.1 million deer are killed each year in vehicle crashes. [According to the World Animal Foundation, the current number of deer killed in traffic crashes is 1.8 million — 300,000 fewer.) Again, these deaths are often gruesome. Finally, the principal population “control” now is death by starvation in winter. This, too, is cruel.
Blossey and colleagues say that return of large predators, even where feasible, will not result in sufficient reduction in deer populations. Nor will encouragement of greater hunting pressure on does.
They note that the federal government owns nearly 30% of the United States’ terrestrial surface area. Management is divided among many agencies – National Park Service, Fish and Wildlife Service, Bureau of Land Management, USDA Forest Service, Department of Defense, and many smaller agencies. Management approaches vary. However, it would be possible to bring them into agreement – although, in some cases, this would require new legislation.
Another issue requires resolution: federal agencies’ authority to manage wildlife on federal land.. The states have repeatedly claimed constitutional and legal authority to manage (vertebrate) wildlife on the federal lands within their borders. This assertion was countered years ago by Nie et al. (2017):
‘Federal land management agencies have an obligation, not just the discretion, to manage and conserve fish and wildlife on federal lands. … [M]ost states have not addressed the conservation obligations inherent in trust management; rather, states wish to use the notion of sovereign ownership as … a source of unilateral power but not of public responsibility. Furthermore, the states’ trust responsibilities for wildlife are subordinate to the federal government’s statutory and trust obligations over federal lands and their integral resources.’
Blossey and colleagues assert that managing wildlife (typically defined as mammals, birds, and fish) is much broader than establishing hunting seasons or methods. Furthermore, the concept of “public trust resources” means resources should be managed for all citizens, not just the fewer than 10% of US residents who hunt. A growing proportion of society expects this management to support healthy and diverse environments.
The authors stress that reducing deer overpopulations is necessary to meet numerous policy goals. These include fulfilling obligations under international treaties related to climate change, invasive species, and threatened species; restoring and conserving the nation’s forests to provide habitat; and adopting “nature-based” climate adaptations, such as carbon sequestration. They express the hope that recent presidential mandates to better quantify and value natural assets will increase awareness of the harm caused by deer overpopulation. Their proposed national strategy would develop goals and metrics to match specific environmental and human health outcomes.
Of course, management of deer must extend beyond federal property lines. This will require cooperation among states, Tribes, and private landowners.
The paper proposes the North American Waterfowl Management Plan as a model. Under this scheme the US Fish and Wildlife Service works with states, tribal governments, Mexico, and Canada to ensure accurate information on waterfowl populations a to calculate harvest levels. States implement their assigned quotas through their own regulations. Waterfowl hunters purchase Duck Stamps to fund the monitoring efforts. This program has worked well for most species covered by the program. Waterfowl are one of the few bird groups that have not declined dramatically.
Reducing deer populations will probably require lethal control. Studies indicate that at least 60% of does must be removed from a population to reduce herd sizes over time. Other means have been attempted at regional or larger landscape levels, such as sterilization, fertility control. These methods have failed even when paired with recreational hunting. Lethal approaches will probably distress many people. However, Blossey, Hare, and Waller believe the program would be supported if it is understood to be undertaken with the goal of improving the health of both humans and also the environment.
In the end, Blossey, Hare, and Waller say they are not willing to leave the killing to cars, disease, and starvation. They emphasize our moral responsibility to protect humans and the many other species that rely on diverse ecosystems. Our policies and choices created the problem, so we must try to correct it.
SOURCES
Blossey. B., D. Hare, and D.M. Waller, 2024. Where have all the flowers gone? A call for federal leadership in deer management in the US. Front. Conserv. Sci. 5:1382132. doi: 10.3389/fcosc.2024.1382132
Nie, M., C. Barns, J. Haber, J. Joly, K. Pitt and S. Zellmer. 2017. Fish and Wildlife Management on Federal Lands: Debunking State Supremacy. Environmental Law, Vol. 47, no. 4 (2017).
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
I have recently recent blogged several times about threats to regeneration of eastern forests. Most of the underlying studies stress the role of deer browsing as a major driver of suppression of native plant species and proliferation of non-native ones. Most studies discussed at a recent Northern Hardwood research forum (USDA, FS 2023b Proceedings) found that deer browsing overwhelms other disturbances, such as fire and canopy gaps that typically promote seedling diversity. Miller et al. also stressed the importance of the deer-invasive plant complex in interrupting regeneration in National parks. Reed et al. found that, on the Allegheny Plateau of western Pennsylvania, high deer densities at the time stands formed reduced tree species diversity, density, and basal area – changes that were still detectable decades later.
On the other hand, Hovena et al. found that at their study sites in Ohio, interaction between non-native shrubs and soil wetness overshadowed even the impact of deer herbivory on the species richness and abundance of seedlings.
Unlike the others, Ducey et al. don’t mention deer as a factor in their analysis of regeneration in a forest in the northern half of New Hampshire. They focus on the minimal impact of silvicultural management. Its effect is secondary to overall forest development as the forest ages. Is it possible that overabundant deer are not a factor in the Bartlett Experimental forest.
American elm (Ulmus americana); photo by F.T. Campbell
Some of the studies acknowledge the impacts of non-native insects and pathogens. The most thorough discussion is in Payne and Peet. They note that specialist pathogens have caused the loss of important tree species, specifically elms and dogwoods plus the impending widespread mortality of ash. Such mortality is resulting in drastic and long-lasting shifts in community dynamics.
Ducey et al. anticipate pest-driven reversals of increases over the decades of eastern hemlock (Tsuga canadensis) and American beech (Fagus grandifolia). Also, they expect that white ash (Fraxinus americana), which has a minor presence, will disappear.
Miller et al. also stressed the importance of emerald ash borer-induced suppression of ash regeneration in some National parks . The authors also noted the threat to beech trees from beech leaf disease in other parks. Hovena et al. state that the interaction between non-native shrubs and soil wetness was more influential than ash mortality in shaping woody seedling communities.
Reed et al. considered the role of non-native earthworm biomass on plant species’ growth.
But too many of the studies, in my view, make no mention of the probably significant role of non-native insects and pathogens.
It is perhaps understandable that they don’t address emerging pests that either have not yet or have barely reached their study sites. For example, Hovena et al. and Yacucci et al. [see below] noted growth of one native shrub, Lindera benzoin, in the face of the challenges presented by deer and invading plants. Neither acknowledges the approach of laurel wilt disease, which has not yet become established in Ohio (it has been detected on the Kentucky-Indiana border). Similarly, neither mentions beech leaf disease, although some of the plots studied by Hovena et al. are just east of Cleveland – where BLD was first detected. The location of the Yacucci et al. study is less than 50 miles away. The North Carolina forests studied by Payne and Peet are apparently too far east to be affected by beech bark disease and beech leaf disease is not yet established nearby.
Less understandable is the failure to mention loss of elms – which were abundant in riparian areas until killed off by Dutch elm disease – which was first detected in Cleveland!); or to discuss the impact of dogwood anthracnose. Their focus on the deciduous forest might explain why they don’t mention hemlock woolly adelgid – which is just now invading the area discussed by Reed et al. I suppose the demise of American chestnut was so many decades ago that it is truly irrelevant to current forest dynamics.
A new study raises anew these questions about whether inattention to the role of non-native pests has skewed past studies’ results. Yacucci et al. compared regeneration in a military installation (Camp Garfield), to the results in the surrounding second-growth forest. This choice allowed them to overcome one drawback of other studies: using deer exclosures that are small and of short durations. The military facility covers 88 km2. Inside it, deer populations have been controlled for 67 years at a density of 6.6 – 7.5 deer/km2. Outside, deer have been overabundant for decades. Populations have grown to densities estimated (but not measured) to be at least 30 deer/km2 – more than four times as high.
These authors established 21 experimental gaps in the low-deer-density area and 20 gaps outside the installation where deer densities are high. Some of the gaps in both low- and high-deer-density environs were located on wetter, seasonally flooded soils, some on drier sites. None of the forest sites had experience fire in recent decades.
Their findings support the importance of deer browsing as driver of changes to forest regeneration.
northern spicebush (Lindera benzoin); photo by R.A. Nonemacher via Wikimedia
They found that at low deer densities, gaps develop a vigorous and diverse native sapling layer, including oaks. Total stem density of red and pin oaks was 13 times higher in these gaps than in gaps in high-deer-density locations. Oak saplings were growing into the subcanopy – that is, above deer browse heights. Saplings of other species – i.e., tuliptree (Liriodendron tulipifera), red maple, and ash (Fraxinus spp.) were also flourishing. Also present were dogwood (Cornus florida) and two native shrubs — Lindera benzoin and Rubus allegheniensis. One non-native shrub, buckthorn (Rhamnus frangula), also thrived at low deer densities. Other non-native plant species were far fewer; their cover was 80% lower. Overall, abundance, richness, and diversity of native herbaceous and woody species were 37–65% higher at the low-deer-density study sites. On average tree species were more than twice as tall as in high-deer-density plots.
In high-deer-density plots, non-native species were six times more abundant while native species richness was 39% lower. Diversity was 27% lower. Most native tree species were short in stature and in low abundance. The one exception was black cherry (Prunus serotina), which deer avoid feeding on. The cherry was 95% more abundant in these high-deer-density plots.
There were several surprising results. In most cases, neither years since gap formation nor habitat type (wet vs. dry) had a significant impact on plant diversity, richness, or abundance. The exception was that non-native plant species were more abundant in older gaps where deer densities were high. Yacucci et al. warn that this phenomenon is a potential threat to biodiversity since high deer density is now the norm across eastern forests.
The authors also note that fire has probably never been a factor in these forests, which are primarily beech-maple forests. Certainly there have been no fires over the past 70 years, either inside or outside the military installation.
Yacucci et al. did not discuss past or possible future impacts of non-native insects or pathogens. They did not mention emerald ash borer or dogwood anthracnose – both of which had been present in Ohio for at least two decades when they completed their study. Although they said their study forest was a beech-maple forest, they did not discuss whether beech are present and – if so – the impact of beech bark disease or beech leaf disease. Both of these are spreading in Ohio. The latter was originally detected in 2012 near Cleveland, just 50 miles from the location of Camp Garfield (between Youngstown and Cleveland, Ohio). As noted above, they also did they mention that Lindera benzoin is susceptible to laurel wilt disease.
beech seedlings in Virginia; photo by F.T. Campbell
Proposed solutions to deer over-browsing
Given the combined threat from widespread deer overpopulation and invasions by non-native plants, Yacucci et al. propose enlisting those military posts that regularly cull deer into efforts to conserve and regenerate native plants. Otherwise, they say, the prognosis for regeneration is poor.
Bernd Blossey and colleagues propose a more sweeping solution: implementation of a national policy to reduce deer populations on all land ownerships. They point out that overabundant deer:
disrupt the plant communities of affected forests – from spring ephemerals to tree regeneration;
promote disease in wildlife and people; and
lead to miserable deaths of deer on our highways, through winter starvation, and disease.
They call for federal leadership of coordinated deer-reduction programs. I discuss their proposal in detail in a separate blog.
SOURCES
Ducey, M.J, O.L. Fraser, M. Yamasaki, E.P. Belair, W.B. Leak. 2023. Eight decades of compositional change in a managed northern hardwood landscape. Forest Ecosystems 10 (2023) 100121
Hovena, B.M., K.S. Knight, V.E. Peters, and D.L Gorchov. 2022. Woody seedling community responses to deer herbivory, intro shrubs, and ash mortality depend on canopy competition and site wetness. Forest Ecology and Management. 523 (2022) 120488
Payne, C.J. and R.K. Peet. 2023. Revisiting the model system for forest succession: Eighty years of resampling Piedmont forests reveals need for an improved suite of indicators of successional change. Ecological Indicators 154 (2023) 110679
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Posted by Faith Campbell
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For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
