EAB-killed ash in Ontario; photo by Michael Hunger
Lately I have become aware of articles discussing how silviculturists and timber managers in the East are responding to the threat from introduced pests.
As Holt et al. (2022; full citation at end of blog) point out, private landowners control 56% of U.S. forestland – and 80% in the East. Their collective decisions about managing those forests are one of two factors that largely determine the composition and structure of the forested landscape and the ecosystem services those woodlands provide. The second determining factor is invasive pests. If an invasive pest prompts many landowners across the East to harvest their timber, the collective impact will be enormous. In this way, invasive species carry a double threat: direct mortality of one or more tree species or genera; and stimulation of removal of the host species from the forest by land managers trying to maximize or protect their current and future monetary investment.
Projections suggest that the number of non-native woodborers established in North America will increase three- or four-fold by 2050. If these prove true (see Leung et al. 2016), the impact on eastern North America forests and associated ecosystem services would be profound.
Holt et al. explore how private landowners have responded to an actual invasive species, the emerald ash borer (EAB). They analyze the influence of EAB’s presence on:
(1) annual probability that a landowner would decide to harvest timber on his/her own lands;
(2) intensity of any such harvest (percentage of trees cut); and
(3) diameter of harvested trees.
They examined harvesting of both the host (ash) and non-host species that co-occur.
Using data from U.S. Forest Service permanent inventory plots, they compared harvest levels in counties in which EAB was detected before 2007 to harvest levels in counties that were infected after 2012. To simplify, they omitted counties in which EAB was detected during the period 2007–2012. They excluded plots that did not contain any ash trees; and plots owned by federal or state agencies. They also excluded trees with diameters less than 12.7 cm (5 inches) dbh.
Ash harvests were apparently less widespread than non-ash harvests. Ash trees were harvested on 6% of the USFS Forest Inventory and Analysis (FIA) plots compared to 9% of plots for harvests of non-ash trees. However, a higher proportion of ash basal area was removed in these harvests — 63% of ash basal area versus 32% of non-ash basal area (remember, ash trees were present in all plots).
The presence of EAB resulted in
an increased amount of biomass harvested – by approximately 25% of basal area;
harvests contained greater quantities of ash, relative to non-ash species.
harvested trees in EAB-infested areas had smaller diameters, on average; this was true of both ash and non-ash species.
Two demographic variables were analyzed. Higher median household income resulted in a lower probability of non-ash harvest. Human population density had no significant effect.
Holt et al. say their findings indicate that a wave of ash removals will follow EAB spread with a potential to alter forest development trajectories and change structural legacies, with consequences for ecosystem services and biodiversity. They consider tree species that co-occur with ash, and that are preferred timber species, are the most likely to be removed in excessive numbers as a result of EAB-induced harvest.
Holt et al. note that ash removals were perhaps underestimated by the study because landowners might have cut their ash before EAB actually was detected in their county.
Managing the Northern Forest – Emphasis on reducing the beech component
Meantime, two other groups are suggesting how forest managers should respond to current challenges, including invasive pests. Both suggest steps to reverse – or at least slow – trends under which American beech (Fagus grandifolia) is becoming more dominant. (Given beech’s ecological importance, this stance bothers me! I don’t quarrel that many timber-oriented people don’t want more beech.) Neither of these studies considers the possible impact of beech leaf disease and beech leaf miner. I recently posted a blog link reporting Reed et al.’s (2022) analysis of interactions between BBD and BLD.
Rogers et al. (2022), the first group, note that successful silviculture is the art and science of managing forests intended to achieve human defined goals. Usually this means assuring the “desired” species composition and structure. However, to succeed, silviculture must also consider site conditions, including competing vegetation and changing climates.
They focus on the northern hardwood forest – also called the beech-birch-maple forest. It is broadly defined by the dominance of sugar maple (Acer saccharum), yellow birch (Betula alleghaniensis), and American beech. The northern hardwood forest occupies about 20 M ha across northern United States and southern Canada. From a traditional management perspective, maple and birch are the desired species; American beech is widely considered undesirable.
Unfortunately, from the timber point of view, Rogers et al. expect the abundance of sugar maple and yellow birch to decrease and American beech to increase. Important factors in this trend are soil types; deer numbers and preference for tree species other than beech; and high number of root sprouts stimulated by beech bark disease (BBD). Rogers et al. call for modification of traditional silvicultural approaches in the region. They call specifically for “adaptation planting” (also called “assisted migration”). They note that increased canopy openings – e.g., “irregular shelterwood system” — are important for establishing shade intolerant and mid-tolerant species, among them white ash (Fraxinus americana). They do mention the threat from emerald ash borer.
In an earlier blog I noted that the second group, Clark and D’Amato(2021), called for silvicultural management of New England forests (part of the same northern hardwood forest). Their goal was to maximize carbon sequestration. They advised management to promote retention of eastern white pine (Pinus strobus) and slow takeover by American beech and eastern hemlock (Tsuga canadensis). They say these species will fare poorly in warmer climates. Of course, all these species face non-native pests. See above for beech; hemlock is being decimated by hemlock woolly adelgid. Eastern white pine has apparently survived its own non-native pest, white pine blister rust.
I hope these pest-related hindrances to traditional timber-focused forestry will help convince the U.S. Department of Agriculture and Congressional agriculture and natural resource committees that non-native pests are a significant threat. Clearly past documentation of impacts to biological diversity and native ecosystems have not prompted them to adopt adequate protective measures or to respond effectively to established invaders. See earlier blogs, my recent article, and the Fading Forests reports (link at end of blog) for suggestions on what actions should be taken.
SOURCES
Clark, P.W. and A.W. D’Amato. 2021. Long-term development of transition hardwood and Pinus strobus – Quercus mixedwood forests with implications for future adaptation and mitigation potential. Forest Ecology and Management 501 (2021) 119654
Leung, B., M.R. Springborn, J.A. Turner, E.G. Brockerhoff. 2014. Pathway-level risk analysis: the net present value of an invasive species policy in the US. The Ecological Society of America. Frontiers of Ecology.org
Rogers, N.S., AW. D’Amato, C.C. Kern, S. B`edardd. 2022. Northern hardwood silviculture at a crossroads: Sustaining a valuable resource under future change
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
black berry eating hawthorn berries; photo by Paul D. Vitucci
Articles by Evan Fricke and colleagues remind us to look more broadly at bioinvasion to consider the impact on ecosystem function and evolution. They focus on animal interactions with plants in the shared environment, especially animals’ role as seed dispersers.
The authors also remind us that natural barriers explain why there are different species in different areas and thus how evolution and speciation follow different paths in different places. Think of Galapagos finches evolving in isolation from a few ancestors that somehow made it over the ocean from mainland South America.
These points are made in two recent articles.
In the first, Fricke and Svenning 2020 (full citation at end of this blog) note that about half of all plant species depend on animals to disperse their seeds. Animal seed dispersal is influenced by several drivers of global change, including local or generalized extinction (= defaunation); bioinvasion; and habitat fragmentation. The decline of large vertebrates has a particularly important role in these interactions.
Their study focused on fleshy-fruited plants that are dispersed by animals. (The study does not include nuts, e.g., acorns, which are presumably subject to some of the same pressures.) They expect evolution of the affected plants and animals to proceed differently as a result of the new partnerships, but they did not study any such interactions.
Their study covered animal seed-dispersal interactions with plants at 410 locations. The data encompassed 24,455 unique animal-plant pairs involving 1,631 animal and 3,208 plant species. Three quarters of the animals were birds; most of the rest were mammals, primarily bats and primates. Only 1% were in other animal groups – lizards, tortoises, or fish.
fruit bats on Luzon, Philippines; photo by Francesco Vernonesi; Flickr.com
They found that introduced plants and animals are twice as likely as native species to interact with introduced partners. The resulting interactions are likely to amplify biotic homogenization in future ecosystems. Already, introduced species have largely replaced missing native frugivore species in some places. In fact, mutualisms in which either or both the plant and animal is an introduced species are now about seven times higher than decades ago.
These mutual-benefit interactions of introduced species are even more prevalent in areas where human modification of the environment is greater. The proportion of introduced species and of novel interactions caused by introduced plant or animal species was higher for oceanic island systems than for continental bioregions. This finding adds a new dimension to the already recognized heightened susceptibility of remote islands to invasion and their loss of native species. Continental bioregions’ networks typically had few introduced animals and a greater prevalence of intro plants than animals.
Fricke and colleagues think plant-frugivore networks are likely to increasingly favor a relatively few introduced generalists over many native species, reducing the uniqueness of future biotas. The result might be to reduce resilience of terrestrial ecosystems by, first, allowing perturbations to propagate more quickly; and, second, by exposing disparate ecosystems to similar drivers. They called for giving higher priority to managing increasing ecological homogenization.
In the second article, Fricke, Ordonez, Rogers, and Svenning (2022) note that climate change requires many plant species to shift their populations hundreds of meters to tens of kilometers per year to track their climatic niche. Earth is also experiencing the formation of novel communities as species introductions and shifting ranges result in co-occurrence of species that do not share co-evolutionary history. They conclude that the novel mutualistic interaction networks will influence whether certain plant species persist and spread.
These authors examined four scenarios to assess how current long-distance dispersal has been affected by past defaunation and invasion and how it is threatened by species endangerment. These scenarios are as follows:
1st scenario (current scenario) = natural and introduced ranges of extant species today.
2nd scenario (natural scenario) = mammal and bird ranges as they would be if unaffected by extinctions, range contractions, or introductions.
3rd scenario (extinction scenario) = those bird and mammal species listed as vulnerable or endangered by the IUCN go extinct.
4th scenario (extirpation of introduced species scenario) = introduced species are extirpated.
Fricke and colleagues estimate that extinction of at least local populations of seed-dispersing mammals and birds has already reduced the capacity of plants to track climate change by 60% globally. The effect is strongest in temperate regions and regions with little topographic complexity. Two examples are eastern North America and Europe. These regions face a double threat: rapid climate change and loss of large mammals that provided long-distance dispersal.
The extinction scenario is most evident in Southeast Asia and Madagascar. The remaining animal seed dispersers are already threatened or endangered. Fricke and colleagues project that future loss of vulnerable and endangered species from their current ranges would result in a further reduction of 15% in the capacity of plants to track climate change.
The contrary situation is found on islands which have few native mammals. Introduced species are now important long-distance seed dispersers. In some cases, the introduced animals are dispersing invasive plant seeds, e.g., on Hawai`i feral hogs are spreading the invasive plant strawberry guava (Psidium cattleianum).
strawberry guava on Maui; photo by Forest and Kim Starr
People’s actions have resulted in ecoregions disproportionately losing the species that provide long-distance seed dispersal function, i.e., large mammals. In other words, human activities have caused not only rapid climate change—requiring broad-scale range shifts by plants—but also defaunation of the birds and mammals needed by plants to do so. Habitat fragmentation and other land-use changes will likely amplify existing constraints on plant range shifts.
Fricke and colleagues say their findings emphasize the importance of not only promoting habitat connectivity to maximize the functional potential of current seed dispersers but also restoring biotic connectivity through the recovery of large-bodied animals to increase the resilience of vegetation communities under climate change.
SOURCES
Fricke, E. C., & Svenning, J. C. (2020). Accelerating homogenization of the global plant–frugivore meta-network. Nature, 585(7823), 74-78. https://www.nature.com/articles/s41586-020-2640-y
Fricke, E. C., Ordonez, A., Rogers, H. S., & Svenning, J. C. (2022). The effects of defaunation on plants’ capacity to track climate change. Science, 375(6577), 210-214. https://www.science.org/doi/full/10.1126/science.abk3510
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
nearly pure stand of Oregon ash in Ankeny National Wildlife Refuge, Oregon; photo by Wyatt Williams, Oregon Department of Forestry
While Michiganders document the impacts of the emerald ash borer (EAB) there, conservationists on the West Coast are jump-starting efforts to save their regional species, Oregon ash (Fraxinus latifolia). Earlier field tests in the Midwest showed that EAB will attack Oregon ash (press release) – something West Coast state would like to counter as early and effectively as possible.
Oregon ash is a wide-ranging species, occurring from California to Washington and possibly into British Columbia. The species has not been studied extensively (it is not a timber species!), but it is clearly an imponearlrtant component of riparian forests. In wetter parts of the Willamette Valley, ash is the predominant tree species. See the photo of the riparian forest in the Ankeny National Wildlife Refuge; this forest is nearly 100% Oregon ash (ODA/ODF EAB Response Plan).
As is true in the Midwest, ash provides important food and habitat resources along creeks and rivers where seasonally high water-tables can exclude nearly all other tree species. Standing and fallen dead ash biomass can alter soil chemistry and affect rates of decomposition, nutrient, and water cycling, i.e., nutrient resource availability for the remaining trees. Gaps in tree canopy can increase soil erosion, stormwater runoff and elevated stream temperatures. In dense stands of Oregon ash, understory vegetation is often sparse, consisting primarily of sedges. The authors of the Response Plan anticipate invasion by non-native plants into canopy gaps caused by the loss of ash trees as a result of an EAB invasion. In Michigan, though, it is the sedges that dominate these gaps.
The Oregon Department of Forestry, the state Department of Agriculture, and other entities have actively participated in “don’t move firewood” campaigns for at least a decade. The Departments of Forestry and Agriculture also led a team that prepared the EAB Response Plan in 2018 (full citation at the end of this blog). It lays out in considerable detail the roles of both government agencies and non-governmental stakeholders. Oregon’s quarantine is broad, covering all insects not on an approved list (Williams, pers. comm.)
California has inspected incoming firewood for years. In April 2021 – after APHIS terminated the federal quarantine on EAB — California Department of Food and Agriculture established a state quarantine on the beetle and articles that could transport it into the state. In doing so, CDFA noted that commercially grown olive trees might also be at risk to EAB.
Washington State operates a statewide trapping program for invasive insects. There has also been significant attention to non-native insect threats to urban forests. These have included a study in 2016 led by the Washington Invasive Species Council (WISC). It involved a partnership of WISC with the Washington Department of Natural Resources Urban and Community Forestry Program as well as and statewide stakeholder meetings [Bush, pers. comm.].
Of these various state-wide initiatives, the institutions in Oregon appear to be most pro-active. The Tualatin Soil and Water Conservation District provided $10,000 to fund some of the genetics work and testing for EAB resistance. Other funding came from the USDA Forest Service Forest Health Protection unit of State and Private Forestry (not from USFS’ Research Program). As described by USFS geneticist Richard Sneizko in an article in the publication TreeLine (full citation at end of blog), participants hope to find at least some level of genetic resistance to EAB. Any such resistance might be deployed in several ways: 1) promoting reproduction by resistant trees to enhance their numbers before EAB gets to Oregon; 2) using seeds from resistant trees for restoration of natural areas; or 3) cross-breeding resistant trees to build genetically diverse stocks of resistant trees for future restoration.
Participants think it is vitally important to work from seeds collected over much of the range of Oregon ash – first, to search for probably very rare resistant trees; and second, to preserve the full diversity of the tree species’ genome so that restored ash will be adapted to the wide variety of conditions in which ash grow.
Participants in this effort include the forest genetics/tree improvement community – specifically, the USDA Forest Service Dorena Genetic Resource Center (located in Cottage Grove, Oregon) and Washington State University at Puyallup Research & Extension Center. Also engaged is the public gardens community, specifically the Huntington Botanical Gardens in San Marino, Los Angeles County. The garden is collecting seed of Oregon and other western ashes from California and Washington State.
The first step in assessing resistance is collecting seed from ash trees across the range of Oregon ash. This began in 2019. Carried out by, inter alia, some USFS and Interior’s Bureau of Land Management units, Oregon State University, citizen scientists [Sniezko] and the Oregon Department of Forestry [press release & Sniezko pers. comm.] Also, some seeds were collected in Washington State in 2020. Additional collections in Oregon are scheduled for 2022.
The collected seeds have been evaluated for vitality and stored by the USFS Dorena Center and at the USFS National Seed Lab (Macon, GA).
Oregon ash planting at Dorena; photo by Emily Boes
The USFS Dorena Center and Washington State University have begun germinating and growing some of the seedlings for various tests of possible resistance. There is concern that the 2021 drought might have killed some of the seedlings in Oregon; those in Washington are not affected. The initial seedlings are mostly from Oregon but there is space to add additional families from a wider geographical area. Experimenters plan to collect data annually on bud break, yearly growth, and any diseases or pests that develop on the trees. (Chastagner pers. comm.)
The next step is systematic testing whether some of the ash show genetic resistance to EAB. Richard Sneizko has sent seedlings of 17 ash families to USFS colleague Dr. Jennifer Koch. She operates a breeding facility in northern Ohio where they can be tested for resistance. Testing is expected to begin this year. [Tree Line]
The Dorena Center is also helping a researcher at Penn State University, Dr. Jill Hamilton, to set up a landscape genomics project. She will evaluate the genetic variability in the species by using leaf samples from about 20 trees from many populations across the Oregon ash’s range (California to British Columbia). This potentially includes a collection from the Dorena population of ash in late Spring 2022. [Sniezko]
These various ash plantings can also be “sentinel” plantings to assist in early detection of newly arriving EAB. [Tree Line]
SOURCES
Bush J. Executive Coordinator | Washington Invasive Species Council
ODF and ODA Emerald Ash Borer Readiness and Response Plan. 2018.
The newsletter is issued by Bonneville Environmental Foundation for a consortium of conservation agencies
Sniezko pers comm Feb 2022 22-2/24
A video explaining the campaign to save Oregon ash is at https://youtu.be/uZmfLrxEA7g or https://youtu.be/S8y-XK285S8
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
results of EAB infestation; photo by Nate Siegert, USFS
A series of studies by Patrick J. Engelken, M. Eric Benbow, Deborah G. McCullough, Nate Siegert, Randall Kolka, Melissa Youngquist and others examine the status of ash (Fraxinus spp.) in the aftermath of the emerald ash borer (EAB) invasion. Initial studies documented the crash of biomass supporting EAB numbers when the large ash trees died (Siegert, Engelken, McCullough. 2021; full citation at end of blog.) More recent studies have focused on bogs and forests in the riparian areas where ash were especially numerous and arguably ecologically most important. I posted a blog about black ash bogs earlier.
I will focus here on the studies in riparian areas of southern and northwest Michigan. Information about impacts in forests of southern Michigan are from Engelken, Benbow and McCullough (2020); information about impacts in northwest Michigan are from Engelken and McCullough (2020). Full citations for both are at the end of the blog.
All study areas had high ash densities before EAB’s arrival. One study (Engelken and McCullough 2020) found ash densities high in the immediate riparian areas (in one case, a strip reaching 100 meters from the streambank) but scattered in surrounding forests.
In all these study areas, populations of mature (reproductive age) ash crashed within 10-15 years after EAB invasion:
In northeast Michigan, EAB carrying capacity was reduced by 94% – 99%; total ash basal area was reduced by 87 – 97.7% (Siegert, Engelken, McCullough. 2021);
In southern Michigan, more than 85% of the basal area of green (F. pennsylvanica) and black ash (F. nigra) had been killed by 2020. An estimated 96% of the overstory ash phloem area had died, thus radically reducing EAB carrying capacity (Engelken, Benbow and McCullough 2020);
In northwest Michigan, more than 95% of the overstory ash have been killed. (Engelken and McCullough 2020).
The worst impact has been on black ash– which plays such an important ecological role in riparian areas and wetlands and has enormous importance in Native American cultures. In all these study areas, there is no stump sprouting by black ash (Siegert, Engelken, McCullough. 2021; Engelken, Benbow and McCullough 2020; Engelken and McCullough 2020). In three watersheds of northwest Michigan where black ash constituted up to a quarter of the overstory species before the EAB invasion, scientists found no black ash recruits, only eight saplings, and a single seedling.
Green ash (F. pennsylvanica) has survived in much higher numbers – so far. However, this species’ ability to grow into reproductive size is still uncertain. In northwest Michigan, green ash saplings are abundant in canopy gaps created by EAB-caused mortality of mature ash. These saplings had established before the EAB invasion so some call them the “orphaned cohort”. However, there are few seedlings of any woody plant species in these gaps because sedges form such dense mats.
Green ash reproduction faces many challenges before persistence of the species can be considered assured. First, populations of EAB – now reduced by the lack of mature ash to support them – might resurge when young ash grow to larger sizes. It is not yet clear the extent to which introduced biocontrol agents and native predators, e.g., woodpeckers, will protect these trees as they grow to reproductive size. Here, again, green ash has an advantage over black ash. While green ash produce seed at a relatively young age, black ash don’t produce seed until they reach 30–40 years. Even then, they produce seeds only sporadically, with intervals of five or more years.
A second challenge is the lack of seed sources – at least until and unless young trees are able to reach reproductive size.
A third challenge is competition for resources from other plants. The canopy gaps eliminate competition for light for the taller plants, i.e., the existing ash saplings. However, the sapling cohort is not supported by a seedling cohort. There are very few seedlings of all woody plant species (including invasive species!). Seed germination is suppressed by the dense mats of wetland-adapted sedges and possibly the higher water tables (which resulted from reduced evapotranspiration following mortality of the mature trees).
Competition for resources is also a factor in the forests outside the immediate riparian zone. There, ash seedlings sprout, but shade created by lateral ingrowth suppresses their growth. In southern Michigan, Engelken, Benbow and McCullough (2020) note that the forests are apparently transitioning from red oak dominated forests to red maple and black cherry dominated forests. This transition is apparently intensified by forest mesophication resulting from reduced fire frequency, decreasing light availability in forest understories and increasing soil moisture content.
Fourth, while stump sprouting of green ash was noted in southern Michigan, in the northwestern forests all the sprouts died. I have already noted the absence of stump sprouting by black ash at all sites.
Beaver & Green Ash in Northern Virginia
photos of beaver feeding on ash saplings in northern Virginia; photos by F.T. Campbell
In spring 2022 I noticed along one stream in northern Virginia that beavers had cut down green ash saplings; McCullough and Siegert report that this does not appear to be a problem in their study areas.
By December 2022, the beaver-cut trees tried to recover: see the sprouts from a stump [below]. (I think deer or rabbits ate the tips of the sprouts.)
The beavers also continued feeding on the ash — the tree photographed in the spring when it was half-chewed through has now been felled and its branches removed [see below].
Ecosystem Impacts, Especially on Streams
Across much of the upper Midwest, massive ash mortality is causing widespread changes in forest systems.
Riparian forests, i.e., areas adjacent to waterways where periodic inundation occurs, are functionally linked to the aquatic systems. Loss of such a significant proportion of the overstory changes the transfer of energy to adjacent waterways that takes the form of inputs of nutrients from leaf litter and coarse woody debris. Intact forests also stabilize stream banks and maintain channel depth by preventing erosion. Forests moderate temperature of the water. Finally, forests with “coarse woody debris” increase habitat structure. These impacts might be especially important along first order streams, (defined as perennial streams that have no permanently flowing tributaries). These streams are too small to buffer the impacts of major tree loss. The scientists say they are uncertain whether these changes continue to affect larger streams downstream.
Unshaded streams have higher water temperatures that can affect populations of fish, in particular salmonids, by delaying migration, reducing egg viability and increasing egg mortality. Higher temperatures can also alter primary productivity of aquatic algae, potentially increasing eutrophication (Engelken and McCullough 2020).
The scientists expect increasing abundance of coarse woody debris in the forests and streams of northwest Michigan as the 75% of dead ash that are still standing fall. Such debris provides nutrients and habitat for an array of plants and animals, thereby influencing the abundance, activity and species compositions of several ground dwelling insects and seedling establishment. In streams, coarse woody debris provides complex habitat and refuges. It also retains organic matter. Recreationists do find that debris impedes boating.
Loss of ash specifically
As described by Engelken, Benbow and McCullough (2020), and in my earlier blog, ash leaf litter – particularly black ash leaf litter – is highly nutritious. Ash leaf litter has efficient turnover rates and contributes important soil nutrients such as nitrogen, organic carbon and exchangeable cations. Invertebrate communities in headwater streams feed largely on coarse organic material such as leaf litter (Engelken and McCullough 2020). Consequently, loss of the annual influx of ash leaf litter will likely have adverse effects on nutrient availability in riparian forests and adjacent streams.
SOURCES
Engelken, P.J., M.E. Benbow, D.G. McCullough. 2020. Legacy effects of emerald ash borer on riparian forest vegetation and structure. Forest Ecology and Management 457 (2020) 117684
Engelken, P.J. and D.G. McCullough. 2020. Riparian Forest Conditions Along Three Northern Michigan Rivers Following Emerald Ash Borer Invasion. Canadian Journal of Forest Research.
Siegert, N.W., P.J. Engelken, D.G. McCullough. 2021 Changes in demography and carrying capacity of green ash and black ash ten years after emerald ash borer invasion of two ash-dominant forests. Forest Ecology and Management Vol 494, August 2021
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
Hakalau Forest, Hawai“i; nearly 90% of Hawaiian flora is unique to the Islands
A recent article by Yang et al. 2021 (full citation at the end of this blog) seeks to determine the extent to which introduced plants reduce the uniqueness of regional floras. They analyzed data from 658 regions covering about 65.7% of the Earth’s ice-free land surface and about 62.3% of the planet’s known plant species.
They found strong homogenization of plant species’ taxonomic and phylogenetic diversity results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.
There are two aspects of a region’s floral uniqueness. One is the number of species that it shares with other regions. This is taxonomic uniqueness. The other is the distinctiveness of the evolutionary history of the region. When several species are endemic to a region’s flora, and lack close relatives in other regions, that equals phylogenetic uniqueness.
The effect of a species introduction differs depending on which of these aspects one focuses on. Thus, naturalization of a species closely related to native species (e.g., a congeneric species) will have less impact on the phylogenetic floristic uniqueness of the region than naturalization by a distantly related species. Taxonomic uniqueness, however, will be affected to the same degree, irrespective of the phylogenetic distance between the naturalized and native species.
Yang et al. found strong homogenization of plant diversity. They found that species introductions increased the taxonomic similarity in 90.7% of all regional pairs and phylogenetic similarity in 77.2% of all region pairs. Most homogenization results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.
This loss of regional biotic uniqueness or distinctiveness changes biotic interactions and species assemblages. These, in turn, have ecological and evolutionary consequences at larger scales and higher levels.
The degree of homogenization between regions’ floras depends on three factors:
1) The distance between the donor and recipient regions. Since nearby regions share more species, an introduction from a more distant origin is more likely to be a novel species and so contribute to homogenization of “donor” and “receiving” floras.
2) Climatic similarity, especially temperature. A plant species introduced from a climatically similar but geographically distant place is more likely to establish than a species from a different climatic zone. As a result, the recipient area’s flora is changed to more closely resemble the flora of the donor region with which it shares climatic conditions – regardless of the distance between them.
3) The level of exchange of goods and people between two regions. The higher the rate of exchange between two regions, the greater the chance that a species will be introduced and become established. Yang et al. used the existence of current or past administrative relationships (e.g., colonial relationship) between two regions as a proxy for intensity of trade and transport between donor and recipient regions. They found that floras of regions with current or past administrative links have taxonomically become more similar to each other than the floras of regions with no such links.
flora of the Cape Floral Kingdom – South Africa; photo from Michael Wingfield
Establishment of introduced species can increase floristic similarity of the donor and recipient regions (= floristic homogenization) when the species is native to one of the two regions and naturalizes in the other, or when it is not native to both regions and naturalizes in both. On the other hand, a species introduction can decrease the floristic similarity of the two regions (i.e., enhance floristic differentiation) when the species is not native to both regions but naturalized in only one.
Homogenization hotspots differed slightly depending on whether one focused on taxonomic or phylogenetic aspects.
The regions with the greatest average increase in taxonomic similarity with other regions due to naturalized alien species were New Zealand, portions of Australia, and many oceanic islands. The Australasian situation probably reflects its long biogeographic isolation from other parts of the globe and its highly unique native flora. As a result, nearly all non-native plants introduced to Australasia strongly increase levels of its floristic similarity to the rest of the world. Oceanic islands have species-poor floras with large proportions of unique endemics. They have also received high numbers of naturalized alien plants.
Hotspots of phylogenetic homogenization on continents are the same as those for taxonomic homogenization, but this is not true for islands. Yang et al. think this is because islands’ native floras were established by natural colonization from nearby continental floras so – despite subsequent speciation – they retain their phylogenetic relationship to the donor areas’ floras.
Yang et al. concede that they lacked high-quality data on native and naturalized alien species lists for a third of Earth’s ice-free terrestrial surface, especially Africa, Eastern Europe, and tropical Asia. They believe, however, that data from these regions are unlikely to change the overall finding. (Scientists are beginning to compile lists of forest pests in Africa). link to blog
Yang et al. note that introduction and naturalization of alien species are likely to increase in the future, thusaccelerating floristic homogenization. The ecological, evolutionary and socioeconomic consequences are largely unknown.They call for stronger biosecurity regulations of trade and transport and other measures to protect native vegetation.
SOURCE
Yang, Q., P. Weigelt, T.S. Fristoe, Z. Zhang, H. Kreft, A. Stein, H. Seebens, W. Dawson, F. Essl, C. König, B. Lenzner, J. Pergl, R. Pouteau, P. Pyšek, M. Winter, A.L. Ebel, N. Fuentes, E.L.H. Giehl, J. Kartesz, P. Krestov, T. Kukk, M. Nishino, A. Kupriyanov, J.L. Villaseñor, J.J. Wieringa, A. Zeddam, E. Zykova and M. van Kleunen. 2021. The global loss of floristic uniqueness. NATURE COMMUNICATIONS (2021) 12:7290. https://doi.org/10.1038/s41467-021-27603-y
Posted by Faith Campbell
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 report here on recent developments on breeding resistant trees. These include both promising results from decades-long efforts and also a promising start to addressing a new challenge.
These programs have benefited from major commitments by the USDA Forest Service. I hope they encourage similar commitments for other priority species – such as those named by the CAPTURE program.
Port-Orford cedar – ready to be planted in the forest!
Scientists who have been working for decades to breed seedlings of Port-Orford cedar (POC) trees resistant to the root rot caused by Phytophthora lateralis https://www.dontmovefirewood.org/pest_pathogen/port-orford-cedar-root-disease-html/now say that they have seedlings ready for planting in the forest. They made this case in a webinar in late February. It can be viewed here. The full webinar runs somewhat over two hours.
The scientist who led early studies of POC and the root disease, Don Zobel, Professor Emeritus, Oregon State University, described the ecological requirements that should guide planting programs. POC produces high-calcium litter. It grows from the sea coast to 1950 meters elevation, on sand dunes, fens, soils with hardpans; mafic & ultramafic rocks (serptentines) and fertile soils on some sedimentary rocks. POC is less shade tolerant than western hemlock but more fire tolerant. It can form a secondary canopy under Douglas-fir and supercede other conifers when fire occurs repeatedly. The tree needs surface water, e.g., seepages and stream sides; but the water must be flowing, not stagnant. Seedlings are especially vulnerable to drying during winter.
[I posted a separate blog about other trees native to this region, including serpentine soils, here.]
One purpose of the webinar was to encourage owners and managers of lands within POC’s historic range (see the map under Dr. Zobel’s presentation) to begin planting the species in appropriate sites. With this in mind, Dr. Zobel emphasized criteria for selecting sites:
Climates in coastal areas of the range are less likely to change under climate change
Quartenary marine terraces are the best geologic type; Lookingglass and Roseburg geologic types are also acceptable
Availability of water during summer, e.g., streamside and seepage areas. Try planting beneath alder. However, avoid interior valley stream corridors if the soils are not ultramafic. And avoid stagnant water.
a POC tree in a bog next to the endemic pitcher plant of southern Oregon, Darlingtoniacalifornica; photo by Richard Sniezko
Dr. Zobel also says one should plant pathogen-resistant genotypes and pay attention to local genetic varieties (which have largely been determined).
Dr. Richard Sniezko of the USFS Dorena Genetic Resource Center described the Center’s 30-year effort to find and exploit resistance to the pathogen. Funding has come from the USFS Forest Health Protection program, other parts of the USFS, and the Bureau of Land Management (BLM). The goal all along has been to produce seedlings for restoration to the forest – meaning not just resistant to the pathogen but also adapted to various local conditions. The program can now provide resistant seedlings in large quantities for planting by landowners and public land managers.
Dr. Sniezko emphasizes that success depends on engagement of four sets of people: research by university scientists; application of that research and development of propagule growing methods by the Dorena Center; support from USFS leaders to continue the program; involvement of land managers who choose to plant the resistant seedlings.
USFS and BLM staff described efforts to determine where POC grows on land under their management, the status of disease in those areas, and efforts to slow the spread of the disease, especially along roadsides and as result of timber or engineering projects. Some of this sanitation work has been funded by USFS Forest Health Protection program — not the National Forest System.
Richard Sniezko stated that the seedlings’ quantitative disease resistance means that some seedlings will die. He expects 40-50% survival of seedlings from many of the breeding zones. This is well above the level of resistance in un-improved populations.
Both BLM and the Rogue-River-Siskiyou National Forest have planted tens of thousands of resistant seedlings in recent years and plan to continue. Funding provided by COVID-19 legislation might allow increased effort. [See Dr. Sniezko’s presentation on the webinar for photos from some plantings.]
POC seedlings at Dorena; photo by Richard Sniezko
Norma Kline of the Oregon State University extension program has distributed more than 10,000 seedlings to small/non-industrial landowners. Many of the recipients shared seedlings with neighbors or are coordinating their planting over a large area. They were motivated primarily by conservation concerns. Her monitoring showed that the POC seedlings survived but did not thrive under dense tanoak canopy. They did well in competition with grass in areas near the coast where there was more moisture. They also did well under Douglasfir as long as there was dappled sunlight.
The non-governmental organization American Forests is likely to participate actively in the planting effort.
In an email to me, Dr. Sniezko asks that people who have planted POC outside its native range inform him where the tree(s) is/are thriving. This information would enhance scientists’ understanding of the species’ environmental tolerances.
Posted by Faith Campbell
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
‘Ohi‘a (Metrosideros polymorpha) is the most abundant native forest tree in Hawai`i and of enormous ecological, cultural, and economic importance. Five species of endemic Metrosideros are recognized on the Hawaiian islands. Only one — M. polymorpha — is found throughout the state. Eight varieties are recognized. These varieties inhabit different environments and have adapted to selective pressures characteristic of these locations. There are at least five other species in the Metrosideros genus, each endemic to one or a few nearby islands. Blaine et al. (2022) [full citation at end of this blog] provide a helpful summary of the tree’s ecological importance and its apparently on-going speciation.
‘Ohi‘a provides habitat for endemic birds, insects, and plants, many of which are endangered. Thus, conservation of this species — and all Hawaiian Metrosideros – is vital for the conservation of countless other taxa. In addition, high elevation ʻōhiʻa forests protect vitally important watersheds across the state. For Native Hawaiians, ʻōhiʻa is a physical manifestation of multiple Hawaiian deities so is the subject of many proverbs, chants, stories, and a foundation of scared hula. Finally, the tree is beautiful!
Native Hawaiian forests face multiple threats — invasive animals and plants, wildfire, and land-use changes. Due to such threats, natural ʻōhiʻa regeneration is largely absent in most lower-elevation forests. In this case, competition with invasive species and the presence of diseases such as ʻōhiʻa rust (Austropuccinia psidii) are probably the specific causes. Multiple government and non-governmental entities have made substantial effort to mitigate these threats.
ROD-infected ʻōhiʻa; photo by J.B. Friday
The disease Rapid ‘Ohi‘a Death (ROD) is an unprecedented threat to this species and the forests it constitutes. The disease is caused by two newly described fungal pathogens: Ceratocystis lukuohia and C. huliohia. The disease caused by C. lukuohia is more severe. To date it has been detected on the two islands farthest apart in the chain — Hawai`i (the Big Island) and Kaua‘i. C. huliohia causes a canker disease that kills trees more slowly. It is more widespread, found on Maui and O‘ahu in addition to Hawai`i and Kaua‘i. Blaine et al. (2022) and the profile here describe the two diseases’ epidemiologies, progression, impacts, and challenges.
Because of the clear threat to Hawaiian ecosystems, ecosystem services, and cultural assets, considerable effort has put into delimitation and research on possible mitigation actions since ROD was discovered in 2010. The first strategic plan covered the period 2017–2019. It focused on expanded efforts to map outbreaks, research on the epidemiology of the pathogens, and most-promising management practices. The second strategic plan covers 2020–2024. It provides for continued surveillance and improvement of these technologies; expanding outreach and public engagement; research on possible vectors of the pathogens; collection and preservation of seeds for research and future restoration; and comprehensive evaluation and development of disease resistance in ʻōhiʻa.
Soon after the causal agents were clarified, the USDA Agriculture Research Service (ARS) began screening for disease resistance. By 2016, ARS had demonstrated that five individuals from two varieties of M. polymorpha had survived inoculation by the more virulent pathogen, C. lukuohia. Their survival raised hopes that natural resistance might be present in wild populations of at least some varieties. However, more comprehensive screening of trees from throughout the species’ range is needed to provide an accurate baseline on the frequency, level, and distribution of genetic resistance to both pathogens. The goal is to produce material resistant to both pathogens that can be used to preserve the ecology, culture, and biotic communities that are dependent on this tree species.
To carry the expanded effort forward, in 2018 a collaborative partnership of state, federal, and non-profit groups was formed. Participants in the ‘Ohi‘a Disease Resistance Program (‘ODRP) include: the Akaka Foundation for Tropical Forests; USDA’s Forest Service and Agriculture Research Service; the state’s Division of Forestry and Wildlife and Agriculture Research Center; programs of the University of Hawai‘i at Manoa and at Hilo; Purdue and Arizona State universities; the Tropical Hardwood Tree Improvement and Regeneration Center; and Kalehua Seed Conservation Consulting.
Blaine et al. (2022) have now outlined a framework to guide the overall effort to identify and develop ROD resistance in M. polymorpha and, possibly, all Hawaiian Metrosideros species. The framework calls for the following activities:
(1) evaluating and operationalizing methods for inoculation-based screening and greenhouse-based production of test plants; and
(2) short-term greenhouse screenings of seedlings and rooted cuttings sampled from native Metrosideros throughout Hawai’i.
Once these tasks have been achieved, the effort is expected to expand to address:
(3) establishing field trials to validate the short-term greenhouse assays and monitor durability and stability of resistance;
drivers of susceptibility and resistance to characterize the durability and stability of genetic resistance to ROD;
(5) developing remote sensing and molecular methods to rapidly detect ROD-resistant individuals;
(6) if necessary, conducting breeding to increase the efficacy of resistance and improve durability of ROD resistance; and
(7) supporting already established and ongoing Metrosideros conservation, including state-wide seed collection and banking, with information on not only genotypes resistant to ROD but also production of ROD-resistant seed.
Blaine et al. (2022) outline how to proceed on each step, and describe the challenges that must be overcome. Challenges range from building growing and screening capacity to handle the thousands of plants required, to developing the remote sensing tools to identify diseased trees in the forest, to identifying sites for seed orchards. Actions by ‘ODRP will focus on Stage II screening in the field to examine the durability of resistance under the wide variety of ecological conditions in which ʻōhiʻa grows and in the presence of a potentially evolving pathogen. Resistance studies must expand beyond M. polymorpha varieties from only one island (the Big Island) to include the other Hawaiian Metrosideros taxa.
Once ROD-resistant M. polymorpha trees are discovered and groundwork has been laid to satisfy initial needs for resistant tree seedlings for forest restoration, scientists can begin research into the genetic basis of ROD resistance. This knowledge will assist breeding efforts which might be necessary if resistance to one of the pathogens does not confer resistance to the other, since the goal is to provide seedlings that are resistant to both.
Blaine et al. (2022) note that the state and others continue efforts to address other aspects of ROD management. These include
1) controlling the spread of the pathogen through local quarantines on movement of infected material and increased public education on bio-sanitation for forest users;
2) testing repellants to reduce beetle attack on infected trees and subsequent frass production.
3) reducing wounding of trees by fencing more pristine forests and removing feral ungulates
SOURCE
Blaine C. Luiz, Christian P. Giardina, Lisa M. Keith, Douglass F. Jacobs, Richard A. Sniezko, Marc A. Hughes, James B. Friday, Philip Cannon, Robert Hauff, Kainana Francisco, Marian M. Chau, Nicklos Dudley, Aileen Yeh, Gregory Asner, Roberta E. Martin, Ryan Perroy, Brian J. Tucker, Ale.alani Evangelista, Veronica Fernandez, Chloe Martins-Keli.iho.omalu, Kirie Santos, Rebekah Ohara. 2022. A framework for establishing a rapid ‘Ohia death resistance program. New Forests. https://doi.org/10.1007/s11056-021-09896-5
See also the video at https://www.bigislandvideonews.com/2019/06/16/video-to-save-ohia-a-genetic-resistance-program-will-be-built/
Posted by Faith Campbell
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
The Invasive Species Prevention and Forest Restoration Act (H.R. 1389) is before Congress. It is co-sponsored by Reps. Peter Welch [VT], Ann Kuster and Chris Pappas [NH], Chellie Pingree [ME], Elise Stefanik and Antonio Delgado [NY], Brian Fitzpatrick [PA], Mike Thompson [CA], Deborah Ross [NC].
Ask your Member of Congress/Representative to co-sponsor this bill. Ask your Senators to sponsor a companion bill.
In summary, this bill will:
Expand USDA APHIS’ access to emergency funds to eradicate or contain newly detected pest outbreaks.
Establish a pair of grant programs to support strategies aimed at restoring tree species decimated by non-native plant pests or noxious weeds. Such strategies include biological control of pests and enhancement of a tree host’s pest resistance.
One grant program supports research to explore and develop these strategies.
The second program support application of resistance breeding and other measures to restore forest tree species. Funded programs must incorporate a majority of the following components: collection and conservation of native tree genetic material; production of sufficient numbers of propagules; preparation of planting sites in the species’ former habitat; planting and post-planting maintenance.
Mandate a study to identify actions to overcome the shortfall of mission, leadership, and prioritization; identify agencies’ expertise and resources; improve coordination among agencies and with partners; and develop national strategies for saving tree species.
Organizations eligible for these grants include federal agencies; state cooperative institutions; colleges or universities offering a degree in the study of food, forestry, and agricultural sciences; and nonprofit entities with non-profit status per §501(c)(3) of the Internal Revenue Code.
Endorsements: Vermont Woodlands Association, American Forest Foundation, The Association of Consulting Foresters (ACF), Audubon Vermont, Center for Invasive Species Prevention, Ecological Society of America, Entomological Society of America, Maine Woodland Owners Association, Massachusetts Forest Alliance, National Association of State Foresters (NASF), National Woodland Owners Association (NWOA), The Nature Conservancy (TNC) Vermont, New Hampshire Timberland Owners Association, North American Invasive Species Management Association (NAISMA), Pennsylvania Forestry Association, Reduce Risk from Invasive Species Coalition, The Society of American Foresters (SAF), and a broad group of university professors and scientists.
Legislative Point of Contact: Alex Piper, Legislative Assistant, office of Rep. Welch. Contact me – providing your email! – if you wish me to send you Alex’ contact information. [The “contact” form does not provide your email and I will not reply in a public way.]
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
American beech (Fagus grandifolia) is a widespread and beautiful tree of the eastern deciduous forest. Its range reaches from Nova Scotia to eastern Wisconsin, then south to Mississippi and Louisiana and east to mid-Georgia. It is an important food source for 40 wildlife species, particularly in the northern parts of its range where few other species produce hard mast. (See Lovett et al. 2006.)
Threats
Unfortunately American beech is under threat from three non-native organisms or complexes: 1) beech bark disease, 2) beech leaf disease, and 3) beech leaf mining weevil. A fourth pest, a previously unknown – and still unnamed bark beetle in the genus Agrilus – has been detected in New York City on European beech trees. It is not yet known whether it will attack American beech and, if so, whether it will also cause serious damage (Michael Bohne, USFS, pers. comm.)
symptoms of beech bark disease; photo by Linda Haugen, USFS; via Bugwood
Beech bark disease (BBD) results from the interaction of the introduced European beech scale insect (Cryptococcus fagisuga) and several fungi in the Neonectria genus – some of which are also introduced. The resulting disease has been killing American beech trees since the beginning of the 20th Century. It has spread from Nova Scotia to much of the tree’s range. It has dramatically altered the composition and structure of stands containing beech.
symptoms of beech leaf disease; photo provided by Jennifer Koch, USFS
Beech leaf disease (BLD) was initially detected in 2012, near Cleveland. As of December, 2021, it has spread due east across New York, Pennsylvania, and New Jersey to the Atlantic, then up the coast through Connecticut and eastern Massachusetts, with a separate outbreak in central Maine. The disease is apparently associated with a nematode, Litylenchus crenatae ssp. mccanni, although additional pathogens, like bacteria, might also play a role. The origin of the North American population of the nematode is unknown; it is a related but separate subspecies from a Japanese nematode (Reed et al. 2022).
American beech defoliated by leaf mining weevil; photo courtesy of Jon Sweeney, CFS
Beech leaf mining weevil (Orchestes fagi) is, so far, limited to Nova Scotia. However, it is expected that the weevil will continue spreading throughout the range of American beech through both natural dispersal and human-assisted movement. Repeated defoliation by the weevil might increases mortality rates in forests that are surviving in the “aftermath” stage of BBD (Sweeney et al. 2020).
A new study (Reed et al. 2022) concludes that, despite being detected only 10 years ago, BLD has already become pervasive in forests surrounding Lake Erie in the U.S. and Ontario. While somewhat more prevalent in U.S. states on the eastern side of the Great Lakes (on 54% of trees) than in Ontario (on 46% of trees), BLD is spreading rapidly and affecting every canopy layer. Mortality is highest in seedlings and saplings; understory saplings die within 2 – 5 years. The occasional mortality of overstory trees occurs within seven years of [observed] infection. Defoliation and mortality of saplings allow more light to pass through to the understory; this is expected to alter plant communities on the forest floor.
Beech scale is more widespread in Ontario (found on 60% of trees) than in the U.S. (38% of trees). This is not surprising since the scale was detected in Ontario in 1960, 24 years before it was detected in portions of Ohio, New York and Pennsylvania included in the study (in 1984). Throughout this region, beech scale is disproportionately affecting overstory trees.
Only 4% of trees throughout the study area are infected with Neonectria cankers. In other words, full-scale beech bark disease is not yet widespread and is spreading surprisingly slowly. Scientists do not understand this phenomenon.
These findings are based on a network of monitoring plots a network of monitoring plots set up in 2019 set up in areas surrounding the Great Lakes. They comprise 34 plots at 17 locations in southwest Ontario and 30 plots at 25 locations in Ohio, Pennsylvania, and New York. In total the plots hold 646 live American beech trees — 412 saplings; 85 in the intermediate/suppressed (subcanopy) category; and 149 in the dominant/codominant (canopy) class.
Forest composition is similar throughout the study area. The most common species in association with American beech are sugar and red maples (Acer spp.), and white and green ash (Fraxinus spp.). Other tree species present include eastern hemlock (Tsuga canadensis), white pine (Pinus strobus), oaks (Quercus spp.), and birches (Betula spp.). Study plots had few invasive plants – although the invasive species present are well-documented to invade forests.
Ontario disease assessment
In Ontario, BLD was identified in 25 of the 34 plots. It was present on 171 saplings, 53 intermediate trees, and 70 dominant trees. Both prevalence and severity were greatest on intermediate trees. Beech scale was present at all 34 plots. While scales were found on trees of all sizes, they were almost two times more prevalent and were more severe on mature trees than saplings. Neonectria cankers were detected at 34 plots. Neonectria was rare but most severe on dominant trees. Fewer than one third of saplings and one-sixth of mature trees were pest free.
U.S. disease assessment
BLD was present in 17 of the 30 plots. It was found on 75 saplings, 30 intermediate trees, and 38 dominant trees. Saplings and dominant trees had similar levels of disease; intermediate trees had significantly less. However, BLD severity was twice as high on saplings compared to mature trees. BLD was present on more than half of the seedlings assessed – 46 out of 82. Beech scale was present in 20 of the 30 plots. It was significantly less common and severe on saplings than on mature trees. Neonectria cankers were present in only 4 of 30 plots. Canker prevalence and severity did not differ significantly among size classes.
Distribution and Effects of Beech Scale and BBD
While beech scale attack facilitates invasion by the Neonectria fungi, the disease – BBD complex – had the most limited distribution of the three pests in this study. It was found on only ~4% of beech trees throughout the study area. The disease was first reported there in the early 2000s. Although no one knows why, it has spread more slowly there than in areas to the east (Reed et al. 2022).
As is the case with beech scale, BBD disproportionately affects large diameter trees. Typically, BBD kills more than half of mature beech within 10 years of its arrival. Dying trees produce prolific root sprouts resulting in dense beech sapling understories that impede regeneration of less shade-tolerant tree species. The persistence of thickets of disease-vulnerable small beech perpetuates the disease. BBD is the only forest disease in North America that can inadvertently intensify itself by increasing densities of its host while suppressing other species.
Beech Forest Community Change in Response to Combined Impacts of BBD and BLD
It is unclear how forests will change as beech die. Some expect saplings of species already present — red maple, white ash, and, especially sugar maple — to exploit the canopy gaps. Of course, white and green ash are under attack by the emerald ash borer; DMF their ability to reach the canopy will depend on the success of biocontrol agents.
However, if BBD or BLD resistant beech survive or if BLD fails to persist, future forests might instead consist of beech thickets that would prevent all but the most shade tolerant species from establishing. Heavy deer predation on maple seedlings and saplings might also play a role. A third possibility is that morbidity from BBD and BLD might lead to uneven-aged conditions that allow younger trees — perhaps even shade intolerant species e.g., oaks — to establish.
Invasive plants also have the potential to fill gaps left by declining beech. While maple-beech forests often have sparse understories due to low understory light levels, pest-caused canopy gaps are expected to increase the abundance of invaders, especially in small woodlots and forests near urban areas. Several shade-tolerant invasive shrubs are already present in low numbers: Japanese barberry (Berberis thunbergii), tatarian honeysuckle (Lonicera tatarica), multiflora rose (Rosa multiflora), and buckthorn (Frangula sp.). Reed et al. (2022) note that these species, plus privet and autumn olive, can take advantage of small canopy gaps, especially when soils are disturbed, e.g., by active intervention to counteract the loss of beech.
Precautionary Research and Management
Reed et al. (2022) call for enhanced monitoring of beech forests focused on
the timing of BLD presence relative to tree age and size – which might affect competitiveness of sprouting beech in the understory; and
compositional and structural change in forests with BLD or to which it is likely to spread
They also recommend abandoning the management approach for BBD currently recommended by foresters. It calls for removing scale-susceptible beech so that resistant genotypes increase in prevalence. In forests with both BBD and BLD, they conclude, management of natural regeneration is unlikely to succeed because BLD will kill sprouts and saplings that might be resistant to scale. They recommend instead active management of the forest to promote mast-producing, shade intolerant species, such as oaks and hickories.
They also recommend increased support for resistance-breeding programs. Such programs already target BBD, based on the estimated 1% of American beech that show some resistance. Now those programs need to incorporate BLD resistance. (Reed et al. note that small numbers of beech show few or no BLD symptoms so might possess resistance or tolerance.)
grafted beech for resistance breeding; photo by Rachel Kappler, then USFS (now Great Lakes Basin Initiative & Holden Arboretum)
Unfortunately, the Canadian beech breeding program’s future funding is highly uncertain. To counter this threat, in part, Reed et al. (2022) suggest cryopreserving beech embryos from Canada to develop a beech conservation collection that would be available for a more robust, future Canadian breeding program. The USFS is trying to develop methods to screen trees for resistance to BLD, specifically to the nematode (J. Koch, USFS, pers. comm.)
Another approach would actively manage beech stands in which potentially BLD-resistant beech grow to help these trees reach the canopy and reproduce. In the absence of management, any BLD-resistant beech seedlings might be overtopped by faster growing, shade-intolerant species – especially if the gaps promote soil drying or sun scald.
Finally, breeding programs need to factor in the beech leaf mining weevil, DMF which — as I noted in the beginning — is spreading across Nova Scotia and could spread to the rest of the native range of beech (Sweeney et al., 2020).
The Department of Agriculture has created a website on the Department’s plant-breeding efforts. It includes a subwebsite on USFS efforts. However, I did not find much useful information there.
SOURCES
Lovett, G.M., C.D. Canham, M.A. Arthur, K.C. Weathers, and R.D. Fitzhugh. 2006. Forest Ecosystem Responses to Exotic Pests and Pathogens in Eastern North America. BioScience Vol. 56 No. 5 May 2006)
Reed, S.F., D. Volk, D.K.H. Martin, C.E. Hausman, T. Macy, T. Tomon, S. Cousins. 2022. The distribution of beech leaf disease and the causal agents of beech bark disease (Cryptoccocus fagisuga, Neonectria faginata, N. ditissima) in forests surrounding Lake Erie and future implications Forest Ecology and Management 503 (2022) 119753
Sweeney J.D., Hughes, C., Zhang, H., Hillier, N.K., Morrison, A. and Johns R. (2020) Impact of the Invasive Beech Leaf-Mining Weevil, Orchestes fagi, on American Beech in Nova Scotia, Canada. Frontiers in Forests and Global Change | www.frontiersin.org 1 April 2020 | Volume 3 | Article 46
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 often blog about the forest pest situation in the UK because its scientists provide lots of easily accessible information. This blog has new information on Britain and – I am pleased to add – on Ireland!
European ash trees alongside the Malham Cove path in Yorkshire; KA via geograph
UK Overview
British woodlands cover just 13% of total land area (just over one-third of the European average of 37%). Their value is increasingly recognized—especially their role in combating climate change through carbon sequestration, flood mitigation and urban cooling. Realization of these benefits is driving new policy to increase woodland cover. In 2019–2020, 13,700 ha of new woodland was created in the United Kingdom (Green et al. 2021). The U.K. government has pledged to plant 30,000 ha of broadleaf and coniferous woodland every year as part of its climate change mitigation strategy (Donald et al. 2021). One example, the ‘northern forest’ scheme, involves planting 50 million native trees over 25 years (Green et al. 2021).
Risks Associated with Conservation Plantings or Translocations
On-going conservation planting efforts plus these ambitious new plans prompted Donald et al. (2021) to assess risk that pathogens might be introduced into the environment as a result of putting native plants in natural habitats. The focus of their study is the planting of common juniper (Juniperus communis)in habitats throughout England, Scotland, and Wales.
Juniper killed by Phytophthora austrocedri; photo from British Forest Research
Juniper is one of three native conifers in the UK. It has been in decline for decades, which conservationists hope to reverse. Now, though, juniper populations are experiencing significant mortality from disease. In 2012, the causal agent was determined to be the non-native pathogen Phytophthora austrocedri. The presence of a single genotype in ~60 geographically separate locations across Scotland and England lends support to idea that the pathogen is being introduced to these sites through some human mechanism. Phytophthora austrocedri has not [yet] been detected in Northern Ireland.
British and European nurseries have contained many Phytophthora pathogens. Hence Donald et al. (2021) sought to determine whether the pathogen is being introduced through use of nursery stock in these well-intentioned plantings. (I have blogged about a similar problem in California restoration plantings.)
The authors found that 19% of P. austrocedri detections are within 2 km of a known planting. The more frequently junipers were planted at a site, and the more cuttings planted during each planting effort, the higher the likelihood that nearby junipers would be infested by P. austrocedri. They conclude that transplanting material is a significant risk pathway for the introduction of disease. The key factor appeared to be the origin of the material. A higher percentage of stock at sites with P. austrocedri outbreaks for which data were available had been raised in a central location by the organization doing the planting or obtained from commercial nurseries. No P. austrocedri was detected in Wales. There, unlike in Scotland or England, the majority of plants were sourced from a commercial nursery that only grew juniper collected from Welsh populations and did not trade with other retailers.
Planting juniper has risen rapidly since the mid-1990s. The highest percentage of planting events co-occurring with disease outbreaks were conducted in 2000–2009. It is likely that there is a time lag between planting and disease detectability. If so, the even larger planting effort since 2010 probably will produce many more P. austrocedri outbreaks that will become visible in the future. And that might not be the end. Planting guidelines have been revised based on pathogen detection. However, the entities doing the planting have not changed their approach, especially regarding site selection.
Donald et al. (2021) also found serious data gaps in these programs beyond the health of propagules. They found:
1) very incomplete knowledge of which organizations are doing the planting;
2) poor attention to traceability of source material; and
3) very little follow-up to check the success of planting projects.
The authors concluded that planting projects have had mixed success in restoring juniper populations. They called for changes in planting strategy to reduce the risks of pathogen introduction.
They also note that efforts to slow the spread of P. austrocedri – for which there is no treatment – are more expensive and less likely to succeed than measures aimed at ensuring that nurseries are free of Phytophthoras. California native plant nurseries have shown that nurseries can maintain Phytophthora-free stock.
Risk of Nursery-spread Pathogens & Willingness of UK Nurseries to Adopt BMPs
Great Britain has experienced an accelerating series of Phytophthora outbreaks and disease epidemics affecting British trees. Introductions detected just since early 2000s include P. ramorum, P. kernoviae, P. lateralis, P. austrocedri and P. pseudosyringae. In all the above cases, imported planting material either is confirmed or strongly implicated as the likely route of intro (Green et al. 2021).
To address this threat – and with massive planting projects proposed – in 2016 the British forest research entity initiated the multidisciplinary ”Phyto-threats” project. Its goal was to understand the drivers of rising Phytophthora infestations and opportunities for mitigating them. The project:
(i) examined Phytophthora distribution and diversity in different nursery management systems;
(ii) assessed the social and economic feasibility of a nursery accreditation programs to curb the risk; and
(iii) identified Phytophthora risks by modelling introduction, establishment and spread of species in relation to biological characteristics, environmental factors and trade flows.
The assessment of Phytophthora presence in nurseries involved collecting 3,624 water and root samples from 163 host genera growing in plant nurseries across the U.K. over a three-year period. Sampling was not random but targetted to facilities thought to harbor Phytophthora. About half of the samples tested positive. They identified 63 species of Phytophthora. Among the most commonly detected species are several that are considered pathogenic — P. cinnamomi, P. cryptogea/pseudocryptogea, P. syringae, P. cactorum, P. cambivora, P. plurivora and P. nicotianae. P. ramorum was found in 12 samples; P. lateralis and P. austrocedri were each found in 10 samples. Several Phytophthora species are potential new records for the U.K. (i.e., P. castanetorum, P. palmivora, P. pseudotsugae,P. tentaculata,P. terminalis, P. uliginosa).
They also saw evidence for Phytophthora root infections in newly arrived plants imported from the European Union.
Their finding raised question about whether Phytophthora can be transported in peat-free potting media, that is, coconut fiber or coir.
The widespread presence and the diversity of Phytophthora found in nurseries was linked to high-risk management practices. These included: careless disposal of culled plants, the near presence of trees along nursery boundaries, and, especially, open water sources. [These factors are essentially identical to infection-facilitating factors found by researchers in California, Oregon, and Washington State. See advisory issued by Oregon State University Extension.]
The project also assessed the feasibility of nursery accreditation programs. The authors consulted widely with nursery owners and customers and conducted a cost-benefit analysis. Regarding nursery practices, owners claimed they were already addressing issues related to water storage in enclosed tanks, clean/covered storage of growing media, installation of drains or free-draining gravel beds, raised benches, and tool disinfestation stations. Therefore the new analysis focused on seven other topics: water testing for pathogens; water treatment s; quarantine holding areas for imported plants; composting or incineration of culled plants; boot and vehicle washing stations; and purchase from only trusted or accredited UK suppliers.
The study found that nurseries would support an accreditation program. However, their support required that costs not be “prohibitive”, actions required not be “unreasonable”, the scheme provide a safety net; and that measures exist to deter non-compliance. Nursery staff wanted to see evidence of consumer demand – a willingness to drive farther to buy “clean” plants, or to pay higher prices for them. The cost-benefit analysis reached a worrying conclusion: nurseries would benefit financially from introducing best practices only when the program would prevent introduction of a wider range of pests and pathogens, not only Phytophthoras. Green et al. (2021) note that the overall net benefit to society from nurseries adopting best practices would be much more substantial. That is, healthy trees are important in meeting carbon sequestration goals. They did not explore whether society should subsidize nurseries’ participation in BMP accreditation programs.
Ireland and Northern Ireland
The island of Ireland (Ireland and Northern Ireland) is thought to have fewer plant pests than other European countries due to its island status and because of its national and international phytosanitary regulations. O’Hanlon et al. (2022) do not mention another possible factor: the likelihood that import volumes to Ireland were probably much lower until the recent vitalization of the Republic’s economy.
O’Hanlon et al. (2022) sought to establish baseline information so scientists can track changes as trade increases and the climate changes. Their search of the literature and unpublished sources identified 396 forest pests on the island, including 11 bacteria, 20 oomycetes, 150 fungi and 215 arthropods. They believe these figures are all probably underestimates. At least 44 of the pests or pathogens are probably non-native to Ireland. (Determining original ranges is difficult, especially for pathogens.)
The Republic of Ireland is one of the least forested countries in Europe. Forests cover ~ 11% of the land area. In Northern Ireland, it is even less: ~ 8%. These forests are predominantly plantations of exotic species. In the Republic, Picea sitchensis makes up 51% of the forest area, Pinus contorta another 10%. Other exotic species planted are Picea abies (4%) and Larix kaempferi. In Northern Ireland, ~ 62% of the forest area is composed of conifer mixtures. Planting of P. sitchensis has accelerated recently, probably as a result of removal of ash and larch because of their vulnerability to pests already established on the island.
Sitka spruce plantation in U.K. Adam Ward, Geograph.org.uk
O’Hanlon et al. (2022) note the great vulnerability of these monocultures to pests. They found 51 pests native to Ireland that are associated with non-indigenous tree genera. They are also concerned about pests introduced from other parts of Europe. For example, green spruce aphid (Elatobium abietinum, native to Central and Eastern Europe) is already attacking Sitka spruce. A second pest of spruce, Ips typographus, which is native to much of Europe but not the British/Irish isles, is not yet established on the island. Northern Ireland imports bark and wood from Europe for processing. Ips typographus has been associated with at least one such shipment.
Non-native forest pests and pathogens also threaten tree species native to Ireland. These include:
Dutch elm disease caused by fungi from the genus Ophiostoma vectored by bark beetles of the genus Scolytus. The second outbreak, caused by the more aggressive pathogen 0. novo-ulmi, was detected in Britain in 1965 and in Ireland in 1977. It caused considerable mortality of elms in Northern Ireland throughout 1970s.
Phytophthora ramorumwas recognized as a threat to forests in Europe only in 2010, when extensive mortality of Japanese larch was detected in Britain. The Republic of Ireland has only the EU1 lineage of the species. Northern Ireland has both the EU1 and EU2 lineages – the former only in nurseries.
Phytophthora disease of alder (caused by several Phytophthora species) was confirmed in Ireland in 2001. However, symptoms of the disease were noted as far back as 1995. It is likely that there are many other Phytophthora species present but not yet recorded.
Ash dieback disease (causal agent Hymenoscyphus fraxineus) on European ash (Fraxinus excelsior) has spread across Europe from Poland beginning in the 1990s. It was confirmed on the Irish island in 2012. Authorities made significant attempts to eradicate the disease, but were not successful. It is now recorded in every county in both Northern Ireland and Ireland. Damage to the economy, environment, and society are expected to be large. The Irish government had helped plant more than 13,000 ha of ash between 1992 and 2012. An estimated 2.9 million ash trees are in Northern Irish hedgerows. British scientists say more than 1,000 fauna species are associated with ash trees.
A second pest on ash — ash sawfly (Tomostethus nigritus) — was detected in Northern Ireland in 2016; it has defoliated hundreds of trees in Belfast.
In recent years, forest pest incursions have increased at a relatively steady rate, comparable to other countries, including Britain. In the 1970s, 26 species were reported; in the 1980s, 27; in the 1990s, 16; in the 2000s, 37; between 2010 and 2017, 28. See the graph in Fig. 2
There is a strong link between pest and pathogen findings in Britain and Ireland. O’Hanlon et al. (2021) list 16 insects and pathogens detected in Britain after 1960 which were later detected in Ireland. The list includes H. fraxineus, 0. novo-ulmi, Phytophthora ramorum, and Phytophthora lateralis. The average delay was 10 years. The authors note that the two islands share similar ecological conditions and hosts, are nearby, plus there is substantial travel and exchange of goods between them. For example, in 2018 an estimated 30,000 metric tonnes of conifer roundwood was sent from Scotland to Northern Ireland for processing.
There are very limited physical checks on plants or plant products moving between Ireland and Northern Ireland. The exception is conifer wood that is not bark-free. European Union regulations require that such shipments be accompanied by a plant passport that certifies that the wood has been inspected by a professional operator authorized by the NPPO of the exporting country. What rules will apply now, after BREXIT, remains unclear. Because of concerns about re-igniting sectarian conflict, most political figures want the border on the island to be almost invisible.
The Europhyt database for the period February 2006 – November 2016 documented interception of numerous high-risk pests at the British and Irish borders, including Anoplophora chinensis and A. glabripennis; I. typographus; Monochamus alternatus; H. fraxineus; and P. ramorum O’Hanlon et al. (2021). believe many more go undetected. O’Hanlon et al. (2021) report specifically on detections on commodities from China, especially on wood packaging. One detection on imported plants of interest to me is that of Discula destructiva(dogwood anthracnose). The article does not mention the origin of the shipment. The native British dogwood, Cornus sanguinea, would presumably be vulnerable to this Asian fungus, which has already caused widespread mortality of woodland dogwoods in North America.
Cornus sanguinea; photo by Hans Hillewaert
In addition to reviewing the current situation, O’Hanlon et al. (2021) note pertinent facts about current policy and future science. First, while the two political units on the island have a history of plant pathology expertise, there has recently been a reduction in the number of practicing forest pathologists, mycologists and entomologists. (I and others have complained about the same deterioration in expertise in the United States.)
Second, they describe the years of delay before official recognition that the pathogen Gremmeniella abietina was present in Northern Ireland. This delay resulted from officials refused to accept data from molecular detection tools.
O’Hanlon et al. (2021) add their voice to others criticizing the international phytosanitary system (they cite six major publications: Brasier 2008; Liebhold el. al. 2012; Santini et al. 2012; Eschen et al. 2015; Jung et al. 2016; Meurisse et al. 2019). The failures are (i) visual inspections can miss asymptomatic infections, (ii) limited resources mean only a small proportion of commodities can be inspected, (iii) allowing the use of fungicides masks disease symptoms on plants, (iv) list-based regulations don’t address undescribed organisms and (v) countries vary in how aggressively they carry out the required phytosanitary procedures. O’Hanlon et al. (2021) conclude that “Until these issues are addressed it is likely further increases in the numbers of non-native pests and pathogens of trees will increase.”
The authors note that Eschen et al. (2018) suggested that risk analysis should focus on the commodity (commodity risk assessment) rather than on an individual pest. I have made a similar suggestion, although less clearly worded.
Finally, O’Hanlon et al. (2021) note that climate change is expected to increase the island’s vulnerability to tree-killing pests and pathogens due to fewer frost days, more rain in winter, increased chance of drought in summer, increased average annual temperatures, and more frequent weather extremes. These changes are likely to affect the amount of damage caused by both native and introduced pests organisms. Range shifts in both pests and pathogens and their natural enemies; physiological or behavioral responses in the pests; phenological changes in the hosts; and increased stress on the trees will combine to affect damage.
SOURCES
Donald, F.; Purse, B.V.; Green, S. 2021. Investigating the Role of Restoration Plantings in Introducing Disease—A Case Study Using Phytophthora [UK] Forests 2021, 12, 764
Green, S., D.E.L. Cooke, M. Dunn, L. Barwell, B. Purse, D.S. Chapman, G. Valatin, A. Schlenzig, J. Barbrook, T. Pettitt, C. Price, A. Pérez-Sierra, D. Frederickson-Matika, L. Pritchard, P. Thorpe, P.J.A. Cock, E. Randall, B. Keillor and M. Marzano. 2021. PHYTO-THREATS: Addressing Threats to UK Forests and Woodlands from Phytophthora; Identifying Risks of Spread in Trade and Methods for Mitigation. Forests 2021, 12, 1617 https://doi.org/10.3390/f12121617ý
O’Hanlon, R., Ryan, C., Choiseul, J., Murchie, A.K. and Williams, C. D. 2021 Catalogue of P&P of trees on the island of Ireland. Biology and Enviro
Proceedings of the Royal Irish Academy 2021. Vol. 121, No. 1.12-45 DOI: 10.3318/ BIOE.2021.02
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
