invasion of wild/black mustard Brassica nigra; photo by carlbegge via Flickr
A California state legislator has proposed a bill to expand state efforts to counter invasive species. Should we support it – and others like it in other states?
The bill is Assembly Bill 2827 introduced by Assembly Member (and former Majority Leader) Eloise Reyes of the 50th Assembly District. She represents urban parts of southwestern San Bernardino County, including the cities of Rialto, Colton, and Fontana.
According to media reports, Reyes was prompted to act by the current outbreak of exotic fruit flies, which as of some months ago resulted in detections in 15 California counties.
The bill is much broader than agricultural pests, however. It would find and declare that it is a primary goal of the state to prevent the introduction, and suppress the spread, of invasive species within its borders. I applaud the language of the “findings” section:
(a) Invasive species have the potential to cause extensive damage to California’s natural and working landscapes, native species, agriculture, the public, and economy.
(b) Invasive species can threaten native flora and fauna, disrupt ecosystems, damage critical infrastructure, and result in further loss of biodiversity.
Paragraph (c) cites rising threats associated with increased movement of goods, international travel, and climate change — all said to create conditions that may enhance the survival, reproduction, and spread of these invasive species, posing additional threats to the state.
(d) It is in the best interest of the state to adopt a proactive and coordinated approach to prevent the introduction and spread of invasive species.
California sycamore attacked by invasive shot hole borer; photo by Beatriz Nobua-Behrmann
The bill calls for
The state agencies, in collaboration with relevant stakeholders, to develop and implement pertinent strategies to protect the state’s agriculture, environment, and natural resources.
The state to invest in research, outreach, and education programs to raise awareness and promote responsible practices among residents, industries, and visitors.
State agencies to coordinate efforts with federal, local, and tribal authorities.
However, the bill falls short when it comes to action. Having declared that countering bioinvasion is “a primary goal of the state”, and mandated the above efforts, the bill says only that the California Department of Food and Agriculture (which has responsibility for plant pests) is to allocate funds, if available, to implement and enforce this article. Under this provision, significant action is likely to depend on holding agencies accountable and providing increased funding.
removing coast live oak killed by goldspotted oak borer; photo by F.T. Campbell
Would this proposed legislation make a practical difference? I have often complained that CDFA has not taken action to protect the state’s wonderful flora. For example, CDFA does not regulate firewood to prevent movement of pests within the State. It has not regulated numerous invasive plants or several wood-boring insects. These include the goldspotted oak borer; the polyphagous and Kuroshio shothole borers; and the Mediterranean oak borer.
On the other hand, CDFA is quick to act against pests that might enter the state from elsewhere in the country, e.g., spongy moth (European or Asian), emerald ash borer and spotted lanternfly.
I hope Californians and the several non-governmental organizations focused on invasive species will lobby the legislature to adopt Assembly Bill 2827. I hope further that they will try to identify and secure a source of funds to support the mandated action by CDFA and other agencies responsible for managing the fauna, flora, and other taxa to which invasive species belong.
I applaud Ms. Reyes’ initiative. I hope legislators in other states will consider proposing similar bills.
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 bullfrog (Lithobates catesbeianus); photo by Will Brown via Wikimedia; one of invasive animals deliberately introduced to Europe in the past
In February 2024 the European Parliament approved legislation outlawing “ecocide” and providing sanctions for environmental crimes. Member states now have two years to enshrine its provisions in national law.
The new rules update the list of environmental crimes adopted in 2008 and enhance the sanctions. The goal is to ensure more effective enforcement. Listed among the offenses are:
the import and use of mercury and fluorinated greenhouse gases,
the import of invasive species,
the illegal depletion of water resources, and
pollution caused by ships.
This action followed an in-depth analysis of the failures of the previous EU environmental directive, first adopted in 2008 (Directive 2008/99/EC). The review found that:
The Directive had little effect on the ground.
Over the 10 years since its adoption few environmental crime cases were successfully investigated and sentenced.
Sanction levels were too low to dissuade violations.
There had been little systematic cross-border cooperation.
EU Member states were not enforcing the Directive’s provisions. They had provided insufficient resources to the task. They had not developed the needed specialized knowledge and public awareness. They were not sharing information or coordinating either among individual governments’ several agencies or with neighboring countries.
The review found that poor data hampered attempts by both the EU body and national policy-makers to evaluate the Directive’s efficacy.
The new Directive attempts to address these weaknesses. To me, the most important change is that complying with a permit no longer frees a company or its leadership from criminal liability. These individuals now have a “duty of care”. According to Antonius Manders, Dutch MEP from the Group of the European People’s Party (Christian Democrats), if new information shows that actions conducted under the permit are “causing irreversible damage to health and nature – you will have to stop.” This action reverses the previous EU environmental crime directive – and most member state laws. Until now, environmental crime could be punished only if it is unlawful; as long as an enterprise was complying with a permit, its actions would not be considered unlawful. Michael Faure, a professor of comparative and international environmental law at Maastricht University, calls this change revolutionary.
Lorton Prison; via Flickr
Another step was to make corporate leadership personally liable to penalties, including imprisonment. If a company’s actions cause substantial environmental harm, the CEOs and board members can face prison sentences of up to eight years. If the environmental harm results in the death of any person, the penalty can be increased to ten years.
Financial penalties were also raised. Each Member state sets the fines within certain parameters. Fines may be based on either a proportion of annual worldwide turnover (3 to 5%) or set at a fixed fine (up to 40 million euros). Companies might also be obliged to reinstate the damaged environment or compensate for the damage caused.Companies might also lose their licenses or access to public funding, or even be forced to close.
Proponents of making ecocide the fifth international crime at the International Criminal Court argue that the updated directive effectively criminalizes “ecocide”— defined as “unlawful or wanton acts committed with knowledge that there is a substantial likelihood of severe and either widespread or long-term damage to the environment being caused by those acts.”
Individual member states also decide whether the directive will apply to offences committed outside EU borders by EU companies.
Some members of the European Parliament advocate for an even stronger stance: creation of a public prosecutor at the European Union level. They hope that the Council of Europe will incorporate this idea during its ongoing revision of the Convention on the Protection of the Environment through Criminal Law. To me, this seems unlikely since the current text of the Convention, adopted by the Council in 1998, has never been ratified so it has not come into force.
The Council of Europe covers a wider geographic area than the European Union – 46 member states compared to 27. Members of the Council of Europe which are not in the EU include the United Kingdom, Norway, Switzerland, Bosnia-Hercegovina, Serbia, Kosovo, Albania; several mini-states, e.g., Monaco and San Remo; and countries in arguably neighboring regions, e.g., Armenia, Azerbaijan, Georgia, and Turkey.
While I rejoice that invasive species are included in the new Directive, I confess that I am uncertain about the extent to which this inclusion will advance efforts to prevent spread. The species under consideration would apparently have to be identified by some European body as “invasive” and its importation restricted. As we know, many of the most damaging species are not recognized as invasive before their introduction to a naïve environment. On the other side, the requirement that companies recognize new information and halt damaging actions – even when complying with a permit! – provides for needed flexibility.
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
U.S. Department of Agriculture headquarters; lets lobby these people! photo by Wikimedia
Twenty-three scientists based around the world published a Letter to the Editor titled “Overwhelming evidence galvanizes a global consensus on the need for action against Invasive Alien Species” It appears in the most recent edition of Biological Invasions (2024) 26:621–626.
The authors’ purpose is to draw attention to the release of a new assessment by the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services’ (IPBES).
The report was issued in September 2023. It is described as the most comprehensive global synthesis of the current knowledge on the bioinvasion process and the impacts of invasive alien species (952 pages!). Its preparation took nearly a decade. Most important, it represents the first consensus among governments and scientists worldwide on the magnitude and extent of the threats that bioinvasions pose to nature, people, and the economy.
The proposed solutions are astoundingly broad and ambitious: transformation of how governments and societies perform. I don’t disagree! However, we need interim steps – “bites of the elephant.” In my view, the report falls short on providing these.
Our challenge: join others in bringing this analysis to decision-makers’ attention. Can we pull out information that will help persuade U.S. decision-makers – governmental and non-governmental – that the threat is both urgent and solvable? How do we more effectively advocate for the aggressive, science-based action that we all know is needed?
(I hope that the fact that the report was prepared under the auspices of the Convention on Biodiversity, to which the U.S. is not a party, does not intensify the challenge for us.)
Why we need to restructure the behavior of governments and societies
Bioinvasions are facilitated by policies, decision-making structures, institutions, and technologies that are almost always focused on achieving other goals. Species transport and introduction are driven by policies aimed at promoting economic growth – especially trade. Later stages of invasions, i.e., establishment and some spread, are accelerated by certain uses of land and sea plus climate change. For example, activities that fragment habitats or cause widespread habitat disturbance provide ready places for bioinvasions. Rarely are those who gain by such policies held accountable for the harms they produce via bioinvasions.
To address these unintended consequences, the IPBES report calls for “integrated governance.” Its authors want coordination of all policies and agencies that touch on the indirect drivers, e.g., conservation; trade; economic development; transport; and human, animal, and plant health. Policy instruments need to reinforce – rather than conflict with — strategic invasive species management across sectors and scales. This involves international agreements, national regulations, all governmental sectors, as well as industry, the scientific community, and ordinary people – including local communities and Indigenous Peoples.
The report also calls for establishment of open and inter-operable information systems. This improved access to information is critical for setting priorities; evaluating and improving regulations’ effectiveness; and reducing costs by avoiding duplication of efforts.
Critically important information that is often unspoken:
Indirect causes underlying the usual list of human activities that directly promote bioinvasions are the rapid rise of human population and even more rapid rise in consumption and global trade.
Biosecurity measures at international borders have not kept pace with the growing volume, diversity, and geographic origins of goods in trade.
Continuation of current patterns is expected to result in one-third more invasive species globally by 2050. However, this is an underestimate because today’s harms reflect the consequences of past actions – often from decades ago. Drivers of invasions are expected to grow in both volume and impact.
We can prevent and control invasive alien species – but that success depends on the availability of adequate, sustained resources, plus capacity building; scientific cooperation and transfer of technology; appropriate biosecurity legislation and enforcement; and engaging the full range of stakeholders. These require political will.
A major impact of bioinvasion is increased biotic homogenization (loss of biological communities’ uniqueness). This concerns us because we are losing the biotic heterogeneity that provides insurance for the maintenance of ecosystem functioning in the face of ongoing global change.
The IPBES study asserts that successfully addressing bioinvasions can also strengthen the effectiveness of policies designed to respond to other drivers, especially programs addressing conservation of biological diversity, ensuring food security, sustaining economic growth, and slowing climate change. All these challenges interact. The authors affirm that evidence-based policy planning can reflect the interconnectedness of the drivers so that efforts to solve one problem do not exacerbate the magnitude of others and might even have multiple benefits.
More Key Findings
Overall, 9% (3,500) of an estimated 37,000 alien species established in novel environments are invasive (those for which scientists have evidence of negative impacts). Proportions of invasives is high among many taxonomic groups: 22% of all 1,852 alien invertebrates; 14% of all 461 alien vertebrates; 11% of all 141 alien microbes; and 6% of all 1,061 alien plants. (The discussion of probable undercounts relates to aquatic systems and certain geographic regions. However, I believe these data are all undermined by gaps in studies.)
Invasive alien species – solely or in combination with other drivers – have contributed to 60% of recorded global extinctions. Invasive species are the only driver in 16% of global animal and plant extinctions. Some invasive species have broader impacts, affecting not just individual species but also communities or whole ecosystems. Sometimes these create complexoutcomes that push the system across a threshold beyond which ecosystem restoration is not possible. (No tree pests are listed among the examples.)
dead whitebark pine in Glacier National Park; photo by National Park Service
The benefits that some non-native – even invasive – species provide to some groups of people do not mitigate or undo their negative impacts broadly, including to the global commons. The report authors note that beneficiaries usually differ from those people or sectors that bear the costs. The authors cite many resulting inequities.
There are insufficient studies of, or data from, aquatic systems, and from Africa; Latin America and the Caribbean; and parts of Asia.
The number of alien species is rising globally at unprecedented and increasing rates. There are insufficient data specifically on invasive species, but they, too, are thought to be rising at similar rates.
Horticulure is a major pathway for introducing 46% of invasive alien plant species worldwide.
Regarding invasive species’ greater impact on islands,the IPBES report mentions brown tree snakes on Guam and black rats on the Galapagos Islands. It also notes that on more than a quarter of the world’s islands, the number of alien plants exceeds the total number of native ones. See my blogs on non-native plants on Hawai`i and Puerto Rico. In addition, I have posted several blogs regarding disease threats to rare bird species in Hawai`. The IPBES report does not mention these.
Where the Report Is Weak: Interim Steps
The report endorses adoption of regulated species (“black”) lists.
The report emphasizes risk analysis of species. Unfortunately IPBES’ analysis was completed before publication of the critique of risk analysis methods by Raffa et al. ( (2023) (see references). However, we must take the latter into consideration when deciding what to advocate as U.S. policy.
The report authors call for more countries to adopt national legislation or regulations specifically on preventing and controlling invasive species. (They note that 83% of countries lack such policies). They also list the many international agreements that touch on invasive species-relevant issues. However, Raffa et al. found that the number of such agreements to which a country is a party bears no relationship to the numbers of alien species detected at its border or established on its territory.
The challenge to risk assessment posed by multiple sources of uncertainty can be managed by recognizing, quantifying, and documenting the extent of that uncertainty.
Beech leaf disease – one of many non-native pests that were unknown before introduction to a naive ecosystem. Photo by Jennifer Koch, USDA Forest Service
I appreciate the report’s emphasis on the importance of public awareness and engagement, but I thought the discussion of effective campaigns lacked original ideas.
The report did not fulfill its own goal of fully exploring unappreciated impacts of policies in its discussion of habitat fragmentation. For example, the report notes that grazing by feral alien ungulates facilitates the spread of invasive alien plant species. However, it does not mention the similar impact by livestock grazing (Molvar, et al. 2024).
SOURCES
Molvar, E.M., R. Rosentreter, D. Mansfield, and G.M. Anderson. 2024. Cheat invasions: History, causes, consequences, and solutions. Hailey, Idaho: Western Watersheds Project, 128 pp.
Raffa, K.F., E.G. Brockerhoff, J-C. GRÉGOIRE, R.C. Hamelin, A.M. Liebhold, A. Santini, R.C. Venette, and M.J. Wingfield. 2023. Approaches to forecasting damage by invasive forest insects and pathogens: a cross-assessment. BioScience 85 Vol. 73 No. 2 (February 2023) https://academic.oup.com/bioscience
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
In the context of reading about forest succession (see previous blogs) I came upon a new publication by Akresh et al. (full citation at the end of this blog.) The article explores the impact of various silvicultural treatments on bird conservation in eastern North America. They note that forest managers are challenged to balance the opposing habitat needs of organisms that, on one hand, depend on structurally diverse old-growth forests and, on the other hand, those that inhabit more open areas or shrubs.
The authors conducted a meta-analysis of studies that examined birds’ responses to three silvicultural regimes: low-retention stands, shelterwoods, and high-retention stands. These terms were not defined in the article. According to Michigan State University extension, in shelterwood systems, all mature trees are harvested in a two- or three-stage process over several years. The other classes presumably reflect the proportion of trees remaining after the harvest.
Akresh et al. focussed on “community conservation scores,” not on protecting individual bird species. They followed the level of conservation concern for the two communities developed by the Partners-in-Flight program
Shrubland Birds
Akresh et al. note that a high proportion of open-canopy, shrubland bird species are declining range-wide; their habitat is already quite limited in eastern North America and continues to decline. Consequently, the Birds-in-Flight program gives them a high priority for conservation measures.
eastern bluebird – prefers open areas; photographer not named; Pickpic
The researchers found that clearcuts (presumably = low retention) and shelterwoods typically had the highest conservation scores because they provide habitat for the declining avian group, shrubland birds. More heavily harvested forests also support non-avian taxa such as pollinators and other arthropods, mammals, snakes, and vascular plants.
Forest Birds
Stands on which 40%–70% of tree were retained also have a high conservation score because they provide habitat for both shrubland and “mature forest” species. Only a few species, e.g., ovenbird and brown creeper, had lower densities in moderately harvested stands than in unharvested forests. The majority of “mature forest” species had relatively higher or equal densities in the sites on which 40%–70% of trees are retained than in unharvested stands. They suggest that several mature-forest bird species prefer the increased understory vegetation density found in these stands.
ovenbird; Wildreturn; Wikimedia Commons
Unharvested and lightly thinned stands, in which 70%–100% of trees remain, had the lowest conservation scores. The first explanation is that these forests don’t support shrubland bird species.
A second reason, Akresh et al. suggest, is that the second-growth forests now widespread in eastern North America are quite young (even if they have not been logged for at least 50 years). They are even-aged and lack the structural diversity of true old-growth forests. The authorsappear to place the greatest importance on the lack of dense understory vegetation, although otherkey elements of mature forests are also missing, e.g., large-diameter trees and snags, continuous canopy, and deep leaf litter. They concede that some bird species depend on forest characteristics that they did not examine. They did not provide examples of these other ecological attributes.
Akresh et al. note that their study concerns only bird species’ use of forests during the breeding season. Some species use other habitat types at other seasons. Furthermore, data were insufficient to analyze some species altogether. A more comprehensive analysis might have raised the conservation score of older forests. I would add that restoration of true old-growth forests depends on allowing some late-seral stands to continue aging.
Finally, fauna other than birds also depend on forest ecosystems and need to be considered when choosing management approaches. The authors mention salamanders and other amphibians, fungi, invertebrates, and lichens – some of which might be of conservation concern themselves.
Gaps in the forest: complicating factors
Akresh et al. mention deer browsing as an influence on understory conditions once, but do not explore this. I am surprised that they don’t expand this statement by a paragraph or two, given the role deer play in suppressing understory vegetation.
Nor do they mention possible impacts of invasions by non-native plants. As my earlier blogs have reported, plant invasions are common in many forested areas in eastern North America. These studies recommend great care in activities that open the forest canopy. Drs. Akresh and King have told me that they believe that forest managers in this region are well aware of invasive plant issues and already incorporate this concern into their management decisions. They referred me to two studies that indicate a very mixed picture of invasive plant impacts on birds (Labbe and King, 2020; Nelson et al, 2017. see full citations below).
multiflora rose – most common invasive plant on forest plots; photo by Famartin
Not Discussed: Insects as food sources
The studies analyzed by Akresh et al. explore levels of nesting success and bird species’ foraging on fruits of non-native shrubs. Others have focused on the reduced numbers of insects feeding on non-native plants; these insects are the principle food for many perching birds’ nestlings. Douglas Tallamy has documented lower numbers of a wide variety of birds which depend on the insect food supply.
SOURCES
Akresh, M.E., D.I. King, S.L. McInvale, J.L. Larkin, A.W. D’Amato. 2023. Effects of forest management on the conservation of bird communities in eastern North America: A meta-analysis. Ecosphere. 2023; 14:e4315. https://onlinelibrary.wiley.com/r/ecs2
Labbe, M.A. and D.I. King. 2020. Songbird Use of Native and Invasive Fruit in the Northeastern USA. Wildlife Society Bulletin. Volume 44, Issue 3. September 2020
Nelson, S.B, J.J. Coon, C.J. Duchardt, J.DL Fischer, A.J. Kranz, C.M. Parker, S.C. Schneider, T.M. Swartz, J.R. Miller. 2017. Patterns and mechanism of invasive plant impacts on North American birds: a systemic review. Biological Invasions. Volume 19, pp. 1547-1563.
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
Gettysburg battlefield; now under attack by emerald ash borer (see below)
Kathryn Miller and colleagues (full citation at end of blog) have published a study that examined the status and trends of forest regeneration in 39 National parks from Virginia to Maine. Four-fifths of the forest plots in the study are classified as mature or late successional – so at first glance the forests look healthy. However, the researchers made an alarming finding: in 27 of 39 parks, forest regeneration is failing – either imminently or probably. Acadia National Park is an exception; it is the only park in the study experiencing healthy regeneration. They warn that without intense, sustained – and expensive! – intervention, these forests are likely to be converted to other types of ecosystems. [I blogged recently about findings regarding regeneration in eastern forests: here and here and here and here.
The forests’ understories have too few seedlings and – especially – saplings to maintain themselves. Worse, in many cases the seedlings and saplings are not the same species as the mature trees that form the canopy. The saplings are shorter species that never reach the canopy. That is, species like pawpaw (Asimina triloba), American holly (Ilex opaca), American hornbeam (Carpinus caroliniana), and eastern redbud (Cercis canadensis) are regenerating, rather than the oaks (Quercus spp.), hickories (Carya spp.), maples (Acer spp.), and pines (Pinus spp.) that constitute the canopies of mature forests in these parks.
Miller and colleagues call these “regeneration mismatches.” Inabout half of the parks, these native canopy tree species make up less than half of current saplings and seedlings. This situation suggests the forests’ species composition will shift substantially, thereby undermining resilience in the face of other challenges, such as invasive plants and pests and climate change.
In many of these National parks, Miller and colleagues found abundant ash regeneration. For example, ash (Fraxinus spp.) constitute more than half of all seedlings in four parks (Johnstown Flood and Friendship Hill in Pennsylvania; Catoctin Mountain in Maryland; Manassas Battlefield in Virginia). Miller and colleagues consigned ash species to the “subcanopy class” because the emerald ash borer (EAB) has caused such high mortality of mature trees. They think regard it unlikely that current and future seedlings will ever reach full size. The devastating impact is most starkly illustrated in Gettysburg National Battlefield Park. Consistent deer management since 1996 has been rewarded: the Park ranks at the top for regeneration among the 39 parks. However, more than half of the seedlings and a quarter of the saplings are ashes. EAB has shifted the Park’s otherwise secure regeneration status into probable failure.
When regeneration fails: too many deer
Throughout the study region, the overwhelming reason regeneration fails is browsing by overabundant deer. The level of deer browse is considered “acceptable” in only four parks. Deer suppress the number of seedlings and saplings. They also skew species composition of native subcanopy species toward those less palatable. Miller and colleagues found that canopy tree density and cover and past human land use had minimal impacts on seedling and sapling numbers or species composition.
Overabundant deer also promote invasion and spread of non-native plants, which are the second most important factor impeding regeneration. Together, invasive plants and non-native earthworms are ecosystem engineers that negatively impact soil and cause cascades of biotic and abiotic impacts throughout forest ecosystems.
Many of the parks experiencing the most severe impacts of chronic deer browse also have the highest invasions by non-native plants. A natural process of regeneration occurs when the death or collapse of mature trees create gaps in the forest canopy. Where deer and invasive shrubs overlap, this process is often hijacked. Instead of nearby native tree species accelerating their growth toward the canopy, thickets of invasive shrubs crowd the space.
For this reason, Miller and colleagues recommend that park management prioritize treating invasive plants in canopy gaps of disturbed stands to avoid forest loss. They recommend deliberate creation of canopy gaps to promote resilience only for parks, or stands within parks, that have low deer and invasive plant abundance or the capacity to intensively manage invasive plants in gaps.
In most parks, non-native tree species are rare, less than 2% of total regeneration. In seven parks, though, non-native trees exceed ten percent of seedlings and/or saplings. In three parks, saplings of non-native trees are increasing. These are primarily tree-of-heaven (Ailanthus altissima) and Norway maple (Acer platanoides). In Saratoga National Historical Park, seedlings of common buckthorn (Rhamnus cathartica) are increasing.
Beech regeneration in Prince William Forest Park
Role of other pests
Miller and colleagues express fear that beech bark disease andbeech leaf disease might have effects similar to those of EAB, leading to a greater “regeneration debt” in parks where American beech (Fagus grandifolia) is the dominant regeneration component. They cite specifically Prince William Forest Park in northern Virginia, [25 mi2]Rock Creek Park in the District of Columbia, [2.7mi2] and Saratoga National Historical Park. [5.3 mi2] The authors also suggest that thickets of beech root sprouts formed in response to BBD can suppress regeneration of other native canopy species and so might need to be managed.
Miller and colleagues mention hemlock woolly adelgid (HWA), but provide very little information. They report that Saint-Gaudens National Historical Park in New Hampshire (the home and studio of sculptor Augustus Saint-Gaudens) is at particular risk because of growth of both beech and eastern hemlock (Tsuga canadensis). I know that Delaware Water Gap National Recreation Area [109m2] has experienced major losses of mature hemlocks. [Shenandoah National Park has also, but it was not included in the study.]
Hemlock Ravine, Delaware Water Gap National Recreation Area; photo by Nicholas T via Flickr
Miller and colleagues report that Acadia National Park is seeing recovery of red spruce (Picea rubens) from a major fire in 1947 and possibly also from acid rain. They do not mention the longer-term threat from the brown spruce longhorned beetle. Their focus is on forest dynamics largely unaffected by deer.
In the same way, the authors make no mention of the absence of dogwood trees, presumably because they had been eliminated by dogwood anthracnose decades ago. Nor do they mention vascular streak dieback of redbud; the causal agent still uncertain. [See Annie Self’s presentation to National Plant Board, August 2023.]
dead ash tree in Shenandoah National Park
One omission is large enough that it might affect the study’s findings. At mi2Shenandoah is the largest National Park in the region. It was not included in the study because the Park’s forest monitoring process is not compatible with those in other NPS units. All the other parks – including Acadia (562 mi) – are much smaller, protecting historic sites like Civil War battlefields.
RECOMMENDATIONS
Miller and colleagues recommend that deer management be initiated in parks classified as at imminent or probable regeneration failure, if such programs are not already under way. They warn that effective deer management requires sustained commitment. Studies of deer exclosures show that full forest recovery from chronic deer overabundance can take as long as 40–70 years.
The authors also recommend actions to open the subcanopy to facilitate growth of saplings belonging to desired species. They caution that deer predation must be controlled. Furthermore, either invasive plant cover must be low, or management must ensure that that the park has sufficient resources to sustain an invasive plant control program – especially if invasive plants are combined with abundant deer.
Parks experiencing compositional mismatches and that are dominated by oak–hickory forest types might also benefit from prescribed burning. Again, deer browse pressure must be minimized. In addition, regeneration of oaks and hickories must already be present.
In park forests dominated by species vulnerable to lethal pests, e.g., beech-, ash-, or hemlock-dominated forest stands, Miller and colleagues recommend considering planting alternative native canopy species and protecting those plantings from deer. Park managers should also consider thinning beech thickets formed after beech bark disease kills canopy trees.
Media coverage
The Washington, D.C., public radio station, WAMU, reported on this research on the air (broadcast December 20) and on its website. It is written by Jacob Fenston, with great photographs by Tyrone Turner. The story emphasized the link between deer and invasive plants – since regeneration in eastern deciduous forest happens by saplings taking advantage of gaps formed when mature trees die. The story quotes DC-area people on their efforts to contain vines. The Natural Resource Manager at Catoctin Mountain Park [8 mi2] describes that park’s longstanding deer control program. The story also mentions impacts of EAB and threat of BLD.
News – Funding for these parks to counter the threats!
Lead author Kathryn Miller has informed me that the Bipartisan Infrastructure Law and Inflation Reduction Act has provided the 39 parks involved in this study over $10 million to improve forest resilience largely through reduction of invasive plants and overabundant deer.
Of course, invasive species threats to National parks are not limited to the Northeast – nor are they new. I have raised this problem from the beginning. To see these blogs, on the “nivemnic” website, scroll down below the archives to the “categories”, then click on “national parks”.
SOURCE
Miller, K.M., S.J. Perles, J.P. Schmit, E.R. Matthews, M.R. Marshall. 2023. Overabundant deer and invasive plants drive widespread regeneration debt in eastern United States national parks. Ecological Applications. 2023;33:e2837. https://onlinelibrary.wiley.com/r/eap Open Access
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
whitebark pine in Glacier National Park killed by white pine blister rust; National Park Service photo
As I noted in the accompanying blog, the U.S. Department of Interior has also weighed in on how to mitigate climate change as part of the Nation’s response to COP24 of the UN Framework Convention on Climate Change.
Interior’s Nature-Based Solutions “Roadmap” (citation at the end of the blog) is 480 pages long! It includes lots of pictures and extensive lists of examples of various types of projects. The document reviews “nature-based” restoration techniques, the benefits they provide in various realms (ecosystem, economy, social values); and the challenges or barriers likely to be encountered. These analyses cover six types of ecosystems – coastal (further divided into five subgroups), forests, grasslands (two types), inland wetland habitats, riverine habitats (three subgroups), and built environments. The obvious emphasis on aquatic and semi-aquatic habitats reflects the Department’s responsibilities. The threat from invasive species is recognized in each case. Plus there are separate chapters discussing management/removal of invasive pests and pathogens, plants, and vertebrates in all types of ecosystems.
The document’s purpose is to provide Interior’s staff – and others who are interested – with reliable information on determining the conditions and goals under which “nature-based” strategies perform best, the benefits they are likely to provide, instructive examples, and additional resources. Much of the information is intended to help staff persuade skeptics that a “nature-based” approach can solve a climate-related problem, such as sea level rise, as well as, or better than, “grey” infrastructure. This includes discussion of: construction and maintenance costs, efficacy in solving a specific problem, and managing conflicts over land use. Also, it considers benefits to other realms, for example, protecting biodiversity and providing opportunities for recreation and mental and physical well-being.
I will focus on aspects dealing with forests. These occur in several chapters. Each chapter has a brief description of the climate and other services provided by that ecosystem type, followed by sections on ways forward (“Technical Approach”), factors affecting site suitability, tools and training resources, likely benefits and outcomes (economic and ecological), barriers and solutions, and examples of projects.
The forest chapter (Chapter 10) discusses forest conservation and restoration with an emphasis on improving forest health, including fuels management, reforestation, and addressing threats from native and non-native pests. One proposed solution is thinning. This measure is said to enhance tree health and promote invasive plants. The “Roadmap” does not recognize that experts consider thinning is helpful in managing native pests such as mountain pine beetle but not non-native pests.
I was startled to find another suggestion – to plant native tree species that are resistant to non-native pests to restore stands. The “Roadmap” refers readers to the National Park Service Resilient Forests Initiative for Region 1 [which reaches from Virginia to Maine]. The Initiative encourages collaboration among parks with similar issues; provides park-specific resource briefs for 39 parks in the Region; and offers management strategies for a host of problems. These include invasive species control, prescribed fire, deer management, silvicultural treatments, tree planting, and fencing. My confusion is that – as far as I know – there are no sources of trees resistant to the non-native pests plaguing forests of the Northeast, e.g., beech, butternut, chestnut, hemlocks, ash, and oaks.
test planting of pathogen-resistant whitebark pine seedlings in Glacier National Park; photo by Richard Sniezko
In the “Tools” section Chapter 10 lists forest restoration guides published by the U.S. Forest Service (USFS) and the International Union of Forest Research Organizations. The “Examples” section includes a few thinning projects.
Chapter 16 advises on enhancing urban forests, which provide many benefits. The chapter stresses the importance of ensuring that projects’ budgets can support protecting trees from such risks as flooding, fire, pests, disease, “invasive species” (presumably other than insects or pathogens), and climate change. The authors note that urban trees are often more susceptible to pests because of their proximity to human activities that facilitate pests’ spread. However, there is no mention that such pests spread to nearby natural forests. They warn against planting a single tree species. An issue noted but not discussed in detail is the use of non-native species in urban forests, some of which have already become invasive.
Three chapters discuss invasive species per se — insects and pathogens (Chap. 26), plants (Chap 27), and vertebrates (Chap. 28) Each chapter summaries invasion stages and stresses the importance of preventing new introductions, detecting them early, and responding rapidly. Most of the text deals with managing established populations – with the emphasis on applying integrated pest management (IPM). Each raises caveats about biological control agents possibly attacking non-target organisms. Again, the authors emphasize the necessity of ensuring availability of adequate resources to carry out the program.
Chapter 26 addresses Invasive and Nuisance Insects and Pathogens. Examples listed include Asian longhorned beetle, emerald ash borer, hemlock woolly adelgid, spongy moth, Dutch elm disease, sudden oak death, laurel wilt, white pine blister rust, chestnut blight and butternut canker. (All these invaders are profiled under the “invasive species” tab here). The examples also include several native pests, e.g., mountain pine beetle, southern pine beetle, and several pathogens, including Swiss needlecast. I am confused by a statement that priorities for management should be based on pests’ traits; my understanding of the science is that other factors are more important in determining a pest’s impact. See, for example, Lovett et al. 2006.This chapter reiterates the impractical advice to plant trees resistant to the damaging pest. I also wonder at the following statement:
“The process of detection and prevention will need to continue over time to prevent reintroductions or reinvasions of nuisance or invasive pests and pathogens. In some cases, long-term management will be required to contain and prevent spread.” [p. 425] I believe long-term management will required in all cases!
The tools listed in the chapter include various DOI websites re: training and funding; the USDA website listing states’ plant diagnostic laboratories; a USDA IPM “road map”; The Nature Conservancy’s guidebook for assessing and managing invasive species in protected areas; the DOI Strategic Plan; and the University of Georgia’s Center for Invasive Species and Ecosystem Health.
Chapter 27 discusses invasive and nuisance plants. It starts by noting that an estimated 5,000 non-native plant species are stablished in the US. While not all are invasive, there is still potential for these plants to spread and cause harm. The authors state that controlling such plants reduces fire risk and lowers demand for water in arid areas.
The authors say early management is crucial to eradicate or control invasive plant species. Because plant invasions cross property lines, agencies must form partnerships with other agencies and private landowners. Because invasive and nuisance plant species are so widespread, managers must set priorities. The “Roadmap” suggests focusing on sites at the highest risk, e.g., heavily trafficked areas. Continued effort will be necessary to prevent reinvasions or reintroductions. However, long-term management and containment can be incredibly costly and labor-intensive.
lesser celandine invade bottomlands of Delaware Water Gap National Recreation Area
The “Roadmap” complains that many invasive and nuisance plant species are still offered for sale; in fact, that this is the primary pathway by which invasive plants enter the US, (While which we have known this for decades, it is encouraging to see a U.S. government report say: “Advocating for federal regulation and cohesive local policies for preventing invasive [plant] sales is essential to avoid disjointed state rulings.” – even if it does not specify which agencies should take the lead.
In the “Tools” section the chapter lists two USFS guides on managing invasive plants; two California Invasive Plant Council guides; the Interior Department’s 2021 Invasive Species Strategic Plan; EDDMapS (a University of Georgia site on which members of the public can report invasive species); and the TNC guidebook for Assessing and Managing Invasive Species in Protected Areas.
Chapter 28 addresses invasive & nuisance vertebrates (called “wildlife”). It notes that invasive animals are present in more than half of all US National parks. It briefly mentions the Lacey Act as providing legal power to curb the introduction and spread of these animals. It does not discuss strengths and weaknesses of this statute, both of which are substantial. This chapter repeats the odd wording from the pest and pathogen chapter – that in some cases long-term management will be required to contain and prevent spread of invasive species. I find it doubtful that short-term actions will be effective in virtually all cases.
Tools listed include Interior guides on IPM, funding sources, and protecting aquatic systems along with the Department of Interior’s 2021 Invasive Species Strategic Plan. Other tools include the USDA guide on IPM, EDDMapS, and the TNC guidebook.
Forests were also mentioned in the discussion of assisted migration of coastal wetlands to avoid drowning by rising seas (Chapter 1). The text notes that forests upland from coastal wetlands might be killed – either as a result of waterlogging as sea levels rise or as deliberate action to make room for the new marsh. Mortality in either case will reduce carbon sequestration. The authors also note the probability that invasive plants – shrubs in the woods, Phragmites on the edge of the wetland — will be present and have to be controlled.
SOURCES
Lovett, G.M, C.D. Canham, M.A. Arthur, K.C. Weathers, R.D. Fitzhugh. 2006. Foret Ecosystem Responses to Exotic Pests and pathogens in Eastern North America. BioScience Vol 56 No. 5 May 2006.
Warnell, K., S. Mason, A. Siegle, M. Merritt, & L. Olander. 2023. Department of the Interior Nature-Based Solutions Roadmap. NI R 23-06. Durham, NC: Nicholas Institute for Energy, Environment & Sustainability, Duke University. https://nicholasinstitute.duke.edu/publications/department-interior-nature-based-solutions-roadmap.
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
hardwood regeneration in northern Virginia forest; photo F.T. Campbell
I have posted several blogs recently about tree species’ regeneration. One blog found poor regeneration of many species throughout forests of the eastern United States. Regeneration is particularly poor in the Great Lakes region, western New York and Pennsylvania, along the Mid-Atlantic and New England coasts, and the coastal plain from southern South Carolina to eastern Texas.
A second blog focused on forest succession in New Hampshire. These findings, by Ducey et al., explicitly recognized the impact of non-native tree-killing insects and pathogens. A third article (Payne and Peet, 2023; full citation at the end of this blog) reports similar findings in North Carolina – and explicitly says that the same conditions are found in forests across the eastern United States.
The locations of neither in-depth study – New Hampshire or North Carolina – include those identified by Potter and Riitters (2022) as suffering particularly poor regeneration.
Payne and Peet find that forest succession in the Piedmont region of North Carolina is not proceeding as expected, based on earlier studies conducted in the same region. The differences are apparent at both the canopy and understory levels. Especially notable is the low recruitment of oaks (Quercus species) and hickories (Carya species) – the genera which previous studies indicated would be the climax taxa. One explanation is the disappearance since early in the 20th Century of fire as a driver of disturbance.
The understory communities are also novel, due largely to invasive species: dramatic loss of flowering dogwood (Cornus florida) killed by the non-native pathogen dogwood anthracnose (Discula destructiva), plus overcrowding of the shrub level by invasive plant species. Other drivers are probably suppression of growth of woody species caused by excessive deer herbivory, and overall accelerated shifts in successional trajectory due to hurricane damage.
flowering dogwood autumn display; F.T. Campbell
Forests in eastern North America in the 21st Century face several drivers of change that are either novel or greatly heightened. In addition to the disappearance of chronic fire, these are frequency and timing of hurricanes, feeding by herbivore populations, and introduction of non-native tree-killing pests and plants. Payne and Peet say scientists and managers need to consider these additional drivers – and their interactions! – when anticipating successional change.
Like Ducey et al. in New Hampshire, Payne and Peet used 80 years of data from 33 permanent plots established and 55 years of data from another 3 plots. Twenty-eight of the plots are transitioning from loblolly pine (Pinus taeda) to hardwood dominance; eight plots have been mixed-age hardwood stands since before the study plots were established.
In the North Carolina piedmont, the composition of canopy trees in plots evolving from pine compared to hardwood stands continue to be different 90–120 years after succession began. Canopy trees in upland and bottomland hardwood stands also differ. These differences reflect the relative species in the forest at the initiation of succession dynamics. Hurricanes – especially Hurricane Fran in 1996 – apparently accelerated succession in some plots by toppling the oldest pines. Despite the persistent differences, the species compositions of both canopy and subcanopy layers are trending toward increasing similarity.
deer-damaged red maple; photo by Eli Sagor via Flickr
The impact of deer browsing is complicated. Deer populations in the study area quadrupled after measurement began in 1980. Deer herbivory suppressed growth of all plant species when their stems were thin (3 – 10 cm DBH). However, after 1996 rapid growth of plants in openings caused by Hurricane Fran’s passage began to reverse the effects of deer browsing. Also, while deer browsing decreases regeneration, growth, and abundance of oak and hickory seedlings and saplings, it also decreases the abundance of other tree species that have – nevertheless – increased in abundance, e.g., red maple (A. rubrum) and black cherry(Prunus serotina).
Payne and Peet found that soil attributes (wetness, texture, organic matter and chemical components), as well as topographic position were minor factors in determining succession trajectories. Increased light availabilitydue to the new or exacerbated drivers of change (thinning of understory vegetation by disease and deer herbivory and opening of the canopy by hurricanes) overcame the influence of nutrients. At most, aunique soil condition might constraining the impacts of these disturbances. Furthermore, these soil-related conditions and other environmental variables change through time — and as a result so does the vegetation. Specifically, the conditions that once supported establishment of oaks and hickories apparently differ today. Payne and Peet conclude that other drivers might be continuing to impact these species’ maturation.
A partial exception is soil nitrogen, through its influence on mycorrhizal patterns. I review mycorrhizal patterns in the discussion of individual tree species, below.
How are Individual Tree Species Responding?
Oaks and hickories are not expanding as expected – either as canopy-sized trees or as seedlings / saplings in the understory. Payne and Peet agree that century-long suppression of low-intensity ground fires is probably the most significant factor in this compositional shift. This decline has been exacerbated by selective logging and deer herbivory. Hickories have established more widely, possibly because young stems have greater shade tolerance. Only plots located on sandy and acidic soils and plots with the greatest hurricane damage have moderate recruitment of oaks and hickories. Oaks and hickories on the poor soils might be aided by the types of ectomycorrhizal fungi that survive in acidic soils with relatively low nitrogen levels. In addition, these soils’ lower water retention probably impedes competition by more mesic, faster-growing, shade-tolerant species. However, even oaks and hickories that have established as seedlings or saplings only rarely progress to canopy dominance. Payne and Peet conclude that oaks might have lost competitive advantage in many of the undisturbed stands.
More mesophytic hardwoods, especially red maple(Acer rubrum), are becoming more numerous and larger – a trend seen throughout forests of the eastern United States. Damage from Hurricane Fran apparently accelerated this trend. However, red maple growth is significantly inhibited by competition from thicket-forming shrubs, especially in bottomland plots. The invasive non-native species thorny olive or oleasterElaeagnus pungensincreased dramatically following Hurricane Fran in 1996. The situation is likely to worsen: two other invasive species, Amur honeysuckle Lonicera maackii and privet Ligustrum japonicum were first detected in the Duke Forest plots in the 2013 survey.
[In New Hampshire, Ducey et al. detected an unexpected levelling off of red maple increases and decline in sugar maple (Acer saccharum); they were unable to determine a cause.]
beech-dominated understory in northern Virginia; F.T. Campbell
Another mesophytic hardwood – American beech(Fagus grandifolia) – has become very abundant in bottomland hardwood stands, especially in small-stem size classes in the understory. Beech prefers sandy soils and its ectomycorrhizal associations are apparently more tolerant of more acidic soils.
Payne and Peet mention – briefly and vaguely – uncertainty about the future of beech. The reference cited discusses the impact of beech bark disease (BBD) in the northeast. Range maps indicate that BBD is well established in the southern Appalachians along the North Carolina/Tennessee border; it has apparently not spread as far east as the study area. There is no mention of beech leaf disease (BLD), which is the primary threat to seedlings and saplings. BLD is currently known to be in northern Virginia. It is unknown whether the disease has any climatic or other barrier that would prevent its moving farther south.
Another bottomland indicator taxon that is also increasing in abundance is ash (Fraxinus species). Along with sweetgum (Liquidambar styraciflua), tulip poplar (Liriodendron tulipifera) and black cherry Prunus serotina, ash density and basal area increased dramatically in plots heavily damaged by Hurricane Fran. Payne and Peet expect most ash trees to be killed by emerald ash borer (Agrilus planipennis) by 2022. The beetle was detected in the study area in 2015.
ash killed by EAB on Potomac lowlands; F.T. Campbell
Flowering dogwood (Cornus florida)was one of the most abundant understory species throughout the study area until the late 1980s. The species has declined by more than 80% since then due to the non-native disease dogwood anthracnose (Discula destructiva). No other species has experienced as precipitous a decline. There is now almost no regeneration in most upland sites.
A second species almost eradicated from the study area by a non-native pathogen is American elm (Ulmus americana). Its basal area in 2013 was 5% of peak levels in the 1950s. Most of this loss occurred by the 1960s, shortly after arrived of Dutch elm disease (DED) in North Carolina. A congeneric species, slippery elm U. alata, is reported to beabundant; it is somewhat resistant to DED. There is no mention of the zig-zag sawfly (Aproceros leucopoda) which has been detected in North Carolina, a few counties away from the study area. The foliage-feeding insect’s long-term impact on elm species is not yet understood.
Payne and Peet note that the study area has twice experienced loss of important components due to specialist non-native pathogens: elms and dogwoods. A third similar event looms: ash [The article does not discuss prospects for biological control.] A fourth is less certain: beech. [This numbering assumes that American chestnut and eastern hemlock were not significant components of forests in the study area.] In their view, these events demonstrate the drastic impacts such non-native organisms can have, especially when the host species is highly abundant or otherwise dominant in a specific community. The resulting shifts in community dynamics and modifications to light and water availability due to such losses, can be dramatic and long-lasting, even resulting in novel successional trajectories.
Members of the 23rd Civil Engineer Squadron/23rd Wing chainsaw a tree lying across a street in the NCO housing area- damage to piedmont North Carolina by Hurricane Fran. Photo courtesy of U.S. National Archives.
Payne and Peet also emphasize the impact of large, episodic disturbances (in their case, hurricanes). These can have widespread and long-lasting impacts on plant community dynamics. Hurricanes’ frequency, intensity, and timing relative to successional stage are key in determining their impacts on successional trajectories. E.g., strong storms that felled the even-aged pine canopy accelerated succession toward more mixed hardwoods. These changes affect biomass, diversity, competitive dynamics, and invasion by invasive plant species, especially in sites with advantageous soil conditions.
Scientists must also evaluate interactions (both reinforcing and antagonistic) between these drivers. For example, in this study deer herbivory and damage from episodic storms had opposite effects on the density of stems in the understory and therefore the future dynamics of forested stands. Hurricane aftereffects frequently accelerated existing or developing trends resulting from various other drivers (e.g., loss of dogwood to anthracnose disease). [While Ducey et al. also detected lasting impacts from hurricane damage in New Hampshire, these effects did not include changes in tree species composition.] Broader regional and global drivers of change, especially those associated with climate change and nitrogen deposition, interact with these many indicators in novel ways based on their own local loadings.
The Nature Conservancy focuses on fire
The Nature Conservancy magazine for Winter 2023 carries an article describing the organization’s experimental efforts to promote oak succession in the Piedmont forests of North Carolina. Greg Cooper, TNC’s forest ecologist in North Carolina, describes retaining dominance by oaks and hickories – rather than maples and poplars – as vital to protecting the region’s faunal diversity and minimizing impacts from climate change. He says this is because oaks use a quarter of the water of maples and poplars.
Cooper links oaks’ failure to reproduce on fire suppression. TNC kills midstory maples and poplars through hack and squirt methods. This allows more light to penetrate the forest and foster oak seedling recruitment. Then they apply controlled fire. “We currently have 700 acres of [controlled-] burn plots, some of which have been burned twice, some of which have been burned once, [and already] we’re getting more light and an immediate flush of herbaceous diversity. We’re getting a lot more berry species, more wildflowers.” TNC is monitoring plots that have been burned, with and without the pre-burn herbicide treatments, and those that have not been burned. They hope to have results in five to ten years that will indicate whether they are achieving the desired improvement in oak regeneration. If so, they also hope is that in future prescribed burns will be sufficient.
Cooper adds that through the Fire Learning Network and a 23-person fire crew they carry out similar work not just on TNC properties, but also federal and state properties.
SOURCES
Ducey, M.J, O.L., Yamasaki, M. Belair, E.P., Leak, W.B. 2023. Eight decades of compositional change in a managed northern hardwood landscape. Forest Ecosystems 10 (2023) 100121
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
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
garlic mustard – a widespread invasive plant in forests of the Northeast and Midwest; photo by Chris Evans, University of Illinois; via Bugwood
This blog summarizes important new research on invasive plant species. You’ll find a USFS update on the spread of invasive plants in regional forests. The second paper used a new funding source to assess the impacts of removing invasive shrub honeysuckles on forest canopies – and to begin restoration. The third paper uses Google searches to study news coverage of plant invaders (spoiler alert: it’s poor). I welcome the attention!
A. Update on Invasive Plants in the Northeast: 44 Species, in 24 States, on Forested Land
The USDA Forest Service Northern Research Station (NRS) continues to publish lots of studies of invasive plants in the forests of the Northeast and Midwest. Kurtz (see USDA citation at end of blog) summarizes data on the extent and intensity of plant invasions collected by the USFS Forest Inventory and Analysis (FIA) program. Since 2012, scientists have compiled data on 44 invasive species on forested land. Crews recorded these plants’ presence on 6,361 plots in 2014 and 4,244 in 2019. [The species surveyed are listed in Table 2 of the publication.]
1. Presence —
Overall, the number of forested plots on which one or more of the 44 monitored invasive species occurs rose from 52% in 2014 to 55% in 2019. Eighteen of the 24 states in the region experienced an increase in the percentage of plots with invasive plants. The number of invasive plants per plot also increased.
The plots with the greatest number of these invasive plants are Ohio (98%), Indiana (97%), North Dakota (93%), and Illinois and Iowa (92% each). The North Dakota data bear a high uncertainty because only 15 plots were surveyed in 2019. The states with the lowest number of invaded plots are New Hampshire (16%) and Vermont (22%). South Dakota also ranks near the bottom with 38% of plots invaded.
A different picture emerges when considering the number of invasive plants per plot. States ranking highest on this criterion were Pennsylvania (13 species on at least one plot) and Illinois (11 species on at least one plot). (See map in USDA publication from References)Plots with five to eight species appear in a band from the Ohio/West Virginia border, across western Maryland and western and eastern Pennsylvania, and into western New York and western Massachusetts. Note the apparent abse nce of invasive plants in the Adirondacks!
2. Species
Of the 44 species monitored, 41 species were observed on plots during these surveys. Two of the three species not found — punktree/Melaleuca and Chinese tallow tree (Triadica sebifera) – grow in the deep South so I am not surprised by their absence. The third – Bohemian knotweed (Polygonum xbohemicum) is invasive in colder climates, e.g., Washington State. Multiflora rose (Rosa multiflora) is the most common species – again, not surprising since it was long planted deliberately to provide food for wildlife. Three species were observed in one but not both inventories: Chinaberry (Melia azedarach) was only found in 2014; saltcedar (Tamarix spp.) and European swallow-wort (Cynanchum rossicum) were found only in 2019.
Amur honeysuckle; photo by pverdunk via Flickr
Most of the 44 plant species increased their presence as measured by the proportion of plots on which they were observed (see Table 2). Among these, Amur honeysuckle (Lonicera maackii) increased by 5.38%; garlic mustard (Alliaria petiolata) by 3.21%; Japanese stiltgrass (Microstegium vimineum) by 2.95%; and bush honeysuckles (Lonicera species.) by 2.77%. Twelve species decreased – all by less than one percent.
3. One of the Worst: Amur Honeysuckle
Amur honeysuckle is one of most actively spreading invasive plant in the entire region – especially in urban forests. The proportion of plots invaded by this species rose from half a percent to six percent between 2014 and 2019. Amur honeysuckle dominates due to competitive growth, allelopathy (producing a biochemical that impedes the germination, growth, and survival of other plants), and its ability to resprout after cutting. The presence of honeysuckle negatively affects native plant communities. It suppresses plant recruitment, homogenizes forest communities, and alters ecosystem processes. It also shapes the canopy structure by affecting the growth and composition of overstory trees, as well as the amount of leaf material (Fotis et al.).
The riparian areas which Amur honeysuckle often invades provide numerous ecosystem services. These include filtering nutrients, preventing soil erosion, filtering sediment from runoff, offering shade and wildlife habitat, and reducing the likelihood of floods to croplands and downstream communities (Fotis et al.).
B. Managing Honeysuckle, Restoring Riparian Forests
Forest managers in central Ohio, near Columbus, have begun a significant restoration experiment to improve the resilience and function of disturbed riparian forests. Fotis and colleagues took advantage of this to track and characterize the immediate, short-term, and long-term impacts of removing Amur honeysuckle on forest canopies. They used new technology: portable canopy imaging, detection, and ranging (LiDAR).
Fotis et al. found that
Honeysuckle presence had a stronger influence on tree species diversity than on the size or number of trees.
Removing honeysuckle from areas where its abundance is high and native tree density is low promoted native tree growth (e.g., the height of tallest trees) and increases in the tree canopy’s structural complexity for up to 10 years.
Honeysuckle removal, followed by treating honeysuckle stumps with herbicides to prevent resprouting, is key for establishing a healthy restored riparian forest.
Planting a diverse suite of native species to fill different ecological niches helps create more resilient forest systems.
Forest recovery began within two years of honeysuckle removal.
At some sites, the researchers planted a variety of woody plants, including understory shrubs and mid- and full-canopy trees, after removing the honeysuckle. The article did not discuss the results.
giant hogweed; photo by NY State DEC fia Flickr
C. Poor Media Coverage of Invasive Plants Undermines Management Efforts
Woodworth et al. (full citation at end of blog) studied how media coverage of invasive plants interacts with low public interest, which the authors claim hampers management efforts and efficacy.
Woodworth et al. note that strong public awareness of urgent environmental issues is linked to the development of new public policy. However, public awareness of invasive species is low despite their causing high monetary costs and significant damage to the environment and human health.
The authors recognize the public generally has a low level of “plant awareness”. They argue that some plants are “charismatic” in one way or another, e.g., some form widespread and conspicuous monocultures; some are attractive and sold in the ornamental trade; and some are harmful to people by producing allergens or bearing thorns/spines/prickles. Such plants often attract more attention.
They asked four questions: Is public interest in these invasive plant species driven by (1) their abundance? (2) their traits? (3) the quantity and sentiment of news articles written about them? Finally, (4) How do these factors combine to drive interest – or lack thereof! – in invasive plant species?
1. Searching for Answers
Woodworth et al. analyzed data on Google searches (in Google Trends) for 209 plant species, 2010 – 2020. The searches revealed whether members of the public sought information about invasive plants, whether news media covered invasive plant issues, whether such coverage had a positive or negative slant, and whether it affected public awareness and attitudes toward invasive plants.
They found that public search interest was highest for the species that are most abundant at both national and state levels. Plant abundance was the second strongest predictor of Google search interest. Also, the more widespread the invader, the more articles were published and the more negative the terms used to describe it. For some species, high search interest was limited to the locations where the plant occurs. An example is garlic mustard (Alliaria petiolata). For other species, there was high search interest across the US although the plant occurs only in some localities. This was true, for example, for giant hog-weed (Heracleum mantegazzianum).
In a related finding, plant species posing a risk to human health ranked high in Google searches. These include giant hogweed (which causes serious skin rashes), allergen-producing grasses, plants that harbor ticks, and those that contribute to fires in the West like cheatgrass (Bromus tectorum). Regarding ticks, they cite multiflora rose but not barberry (Berberis spp). These results suggest to the authors that public interest in invasive plants is motivated primarily by the likelihood of encountering these species, with direct consequences for health and well-being. These concerns can be amplified by the media. The authors suggest that articles spelling out health risks of invasive plants might increase public support for wider management efforts.
cheatgrass invasion of BLM lands in SW Idaho; photo by Thane Tuoson via Flickr
The greater the number of articles published about a species, the more frequently readers utilized Google to search for information. Woodworth et al. suggest that more attention by the media translates to more public interest. They caution, however, that there are two other possible explanations. First, more searches might drive science journalists [I add: or newspaper editors!] to write more articles on species already known to be of interest to the public. Or, search and media interest might be unrelated to each other, but driven by the same external factors, such as a recent fire. In other words, journalists and the general public might be interested in the same aspects of invasive plants.
2. Which Invaders Got Attention
The media and public focus on only some invasive plants. Media articles discussed only 175 (84%) of the 209 species included in the study. More than 50% of news articles were written about only 10 species (5% of all the species); 80% on just the top 25 species. Public interest was even narrower. No one searched for information on 60 of the species (29%) over the 10-year period.
The authors – and I – are distressed that invasive plants that are sold as ornamentals were both written about and searched for less than invasive plants that are not so marketed. Worse, articles about the ornamental species had a more positive tone than articles about non-marketed invasives.
English ivy (Hedera helix) – an invasive plant widely used in horticulture; Washington State Weed Control Board
Finally, they found that species that form monocultures did not garner more media attention, more negative coverage, or search interest. Their example is Japanese stiltgrass (Microstegium vimineum), which they describe as being a problematic and prolific invader of eastern forest understories. There are exceptions to this finding: common reed Phragmites and kudzu. I note that these species are very noticeable, and thus “charismatic.” The former is a large plant and masses along roadsides and in open habitats. Kudzu has been notorious for decades as the “vine that ate the South”.
Woodworth et al. concluded that the media’s narrow focus on “notorious” invasive plant species, when combined with the lower and more positive coverage of ornamental introductions, could send mixed messages and weaken public awareness of their threats. The authors believe, however, that there is ample opportunity to improve messaging and increase public awareness. This requires more media coverage and a greater focus on invasive plants’ negative impacts. One potentially “sticky” message is about “loss of control.”
SOURCES
Fotis, A., Flower, C.E.; Atkins, J.W. Pinchot, C.C., Rodewald, A.D., Matthews, S. 2022. The short-term and long-term effects of honeysuckle removal on canopy structure and implications for urban forest management. Forest Ecology and Management. 517(6): 120251. 10 p. https://doi.org/10.1016/j.foreco.2022.120251 .
USDA Forest Service Northern Research Station Rooted in Research ISSUE 18 | SEPTEMBER 2023
Kurtz, C.M. 2023. An assessment of invasive plant species in northern U.S. forests. Res. Note NRS-311. http://doi.org/10.2737/NRS-RN-311
Woodworth, E. A. Tian, K. Blair, J. Pullen, J.S. Lefcheck and J.D. Parker. 2023. Media myopia distorts public interest in US invasive plants. Biol Invasions (2023) 25:3193–3205 https://doi.org/10.1007/s10530-023-03101-8
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
tanoak trees in southern Oregon killed by sudden oak death; photo by Oregon Department of Forestry; this pathogen is not mentioned by USFS RPA report
In August the USDA Forest Service published the agency’s 2020 assessment of the future of America’s forests under the auspices of the Resources Planning Act. [See United States Department of Agriculture Forest Service Future of America’s Forests and Rangelands, full citation at the end of the blog.] To my amazement, this report is the first in the series (which are published every ten years) to address disturbance agents, specifically invasive species. In 2023! Worse, I think its coverage of the threat does not reflect the true state of affairs – as documented by Forest Service scientists among others.
This is most unfortunate because policy-makers presumably rely on this report when considering which threats to focus on.
Here I discuss some of the USFS RPA report and what other authors say about the same topics.
The RPA Report’s Principle Foci: Extent of the Forest and Carbon Sequestration
The USFS RPA report informs us that America’s forested area will probably decrease 1- 2% over the next 50 years (from 635.3 million acres to between 619 and 627 million acres), due largely to conversion to other uses. This decline in extent, plus trees’ aging and increases in disturbance will result in a slow-down in carbon sequestration by forests. In fact, if demand for wood products is high, or land conversion to other uses proceeds apace, U.S. forest ecosystems are projected to become a net source of atmospheric CO2 by 2070.
Eastern forests sequester the majority of U.S. forest carbon stocks. These forests are expected to continue aging – thereby increasing their carbon storage. Yet we know that these forests have suffered the greatest impact from non-native pests.
I don’t understand why the USFS RPA report does not explicitly address the implications of non-native pests. In 2019, Songlin Fei and three USFS research scientists did address this topic. Fei et al. estimated that tree mortality due to the 15 most damaging introduced pest species have resulted in releases of an additional 5.53 terragrams of carbon per year. Fei and colleagues conceded this is probably an underestimate. They say that annual levels of biomass loss are virtually certain to increase because current pests are still spreading to new host ranges (as demonstrated by detection of the emerald ash borer in Oregon). Also, infestations in already-invaded ranges will intensify, and additional pests will be introduced (for example, beech leaf disease).
I see this importance of eastern forests in sequestering carbon as one more reason to expand efforts to protect them from new pest introductions, and the spread of those already in the country, etc.
A second issue is the role of non-native tree species in supporting the structure and ecological functions of forests. Ariel Lugo and colleagues report that 18.8 million acres (7.6 million ha, or 2.8% of the forest area in the continental U.S.) is occupied by non-native tree species. (I know of no overall estimate for all invasive plants.) They found that non-native tree species constitute 12–23% (!) of the basal area of those forest stands in which they occur.
Norway maple (Acer platanoides); one of the most widespread invasive species in the East. Photo by Hermann Falkner via Flickr
Lugo and colleagues confine their analysis of ecosystem impacts to carbon sequestration. They found that the contribution of non-native trees to carbon storage is not significant at the national level. In the forests of the continental states (lower 48 states), these trees provide 10% of the total carbon storage in the forest plots where they occur. (While Lugo and colleagues state that the proportion of live tree biomass made up of non-native tree species varies greatly among ecological subregions, they do not provide examples of areas on the continent where their biomass – and contribution to carbon storage — is greater than this average.) In contrast, on Hawai`i, non-native tree species provide an estimated 29% of live tree carbon storage. On Puerto Rico, they provide an even higher proportion: 36%.
Brazilian pepper (Schinus terebinthifolius) – widespread invasive in Hawai`i and Florida; early stage invasive in Puerto Rico. Photo by Javier Alexandro via Flickr
In the future, non-native trees will play an even bigger role. Since tree invasions on the continent are expanding at ~500,000 acres (202,343 ha) per year, it is not surprising that non-native species’ saplings provide 19% of the total carbon storage for that size of trees in the lower 48 states (Lugo et al.).
Forming a More Complete Picture: Biodiversity, Disturbance, and Combining Data.
The USFS RPA report has a chapter on biodiversity. However, the chapter does not discuss historic or future diversity of tree species within biomes, nor the genetic diversity within tree species.
Treatment of Invasive Species
The USFS 2020 RPA report is the first to include a chapter on disturbance, including invasive species. I applaud its inclusion while wondering why they have included it only now? Why is the coverage so minimal? I think these lapses undercut the report’s purpose. The RPA is supposed to inform decision-makers and stakeholders about the status, trends, and projected future of renewable natural resources and related economic sectors for which USFS has management responsibilities. These include: forests, forest products, rangelands, water, biological diversity, and outdoor recreation. The report also has not met its claim to “capitalize on” areas where the USFS has research capacity. One excuse might be that several important publications have appeared after the cut-off date for the assessment (2020). Still, the report’s authors cite some of the evaluations that were in preparation as of 2020, e.g., Poland et al.
I suggest also that it would be helpful to integrate data from other agencies, especially the invasive species database compiled by the U.S. Geological Survey, into the RPA. For example, the USGS lists just over 4,000 non-native plant species in the continental U.S. (defined as the lower 48 plus Alaska). On Hawai`i, the USGS lists 530 non-native plant species as widespread. Caveat: many of the species included in these lists probably coexist with the native plants and make up minor components of the plant community.
Specifically: Invading Plants
The USFS RPA report gives much more attention to invasive plants than non-native insects and pathogens. The report relies on the findings of Oswalt et al., who based their data on forested plots sampled by the Forest Inventory and Analysis (FIA) program. (The RPA also reports on invasive plants detected on rangelands, primarily grasslands.) Oswalt et al. found that 39% of FIA plots nationwide contained at least one plant species that the FIA protocol considers to be invasive and monitors. The highest intensity of plant invasions is in Hawai`i – 70% of the plots are invaded. The second-greatest intensity is in the eastern forests: 46%. However, the map showing which plots were inventoried for invasive plants makes clear how incomplete these data are – a situation I had not realized previously.
I appreciate that the USFS RPA report mentions that propagule pressure is an important factor in plant invasions. This aspect has often been left out in past analyses. I also appreciate the statement that international trade in plants for ornamental horticulture will probably lead to additional introductions in the future. Third, I concur with the report’s conclusions that once forest land is invaded, it is unlikely to become un-invaded. Invasive plant management in forests often results in one non-native species being replaced by another. In sum, the report envisions a future in which plant invasion rates are likely to increase on forest land.
If you wish to learn more about invasive plant presence and impacts, see the discussion of invasive plants in Poland et al., my blogs based on the work by Doug Tallamy, and several other of my blogs compiled under the category “invasive plants” on this website.
I believe all sources expect that the area invaded by non-native plant species, and the intensity of existing invasions, will increase in the future.
The USFS RPA links these invasions to expansion of the “wildland-urban interface” (“WUI”). These areas increased rapidly before 2010. At that time, they occupied 14% of forest land. The report published in 2023 did not assess their future expansion over the period 2020 to 2070. However, it did project increased fragmentation in many regions, especially in the RPA Western and Southeastern regions. Since “fragmentation” is very similar to wildland-urban interfaces, the report seems implicitly to project more widespread plant invasions in the future.
plant invasions facilitated by fragmentation; northern Virginia; photo by F.T. Campbell
Specifically: Insects and Pathogens
The USFS RPA report on insects and pathogens is brief and contains puzzling errors and gaps. It says that the tree canopy area affected by both native and non-native mortality-causing agents has been consistently large over the three most recent five-year FIA assessment periods. It notes that individual insects or diseases have extirpated entire tree species or genera and fundamentally altered forests across broad regions. Examples cited are chestnut blight and emerald ash borer.
The USFS RPA report warns that pest-related mortality might be underreported in the South, masked by more intense management cycles and higher rates of tree growth and decay. On the other hand, the report asserts that pest-related mortality is probably overrepresented in the Northern Region in the 2002 – 2006 period because surveyors drew polygons to encompass large areas affected by EAB and balsam woolly adelgid (Adelges piceae) infestations. The latter puzzles me; I think it is probably an error, and should have referred to hemlock woolly adegid, A. tsugae. Documented mortality has generally been much more widespread from insects than diseases, e.g., bark beetles, including several native ones, across all regions and over time, especially in the West – where the most significant morality agents are several native beetles. The USFS RPA report mentions that the Northern Region has been particularly affected by non-native pests, including EAB, HWA, BWA, beech bark disease, and oak wilt. It mentions that Hawai`i has also suffered substantial impacts from rapid ʻōhiʻa death.
Defoliating insects have affected relatively consistent area over time. This area usually equaled or exceeded the area affected by the mortality agents. Principal non-native defoliators in the Northern Region have been the spongy moth (Lymantria dispar); larch casebearer (Coleophora laricella); and winter moth (Operophtera brumata). In the South they list the spongy moth.
More disturbing to me is the USFS RPA report’s conclusion that the future impact of forest insects is highly uncertain. The authorsblame the complexity of interactions among changing climate, those changes’ effects on insect and tree species’ distributions, and overall forest health. Also, they name uncertainty about which new non-native species will be introduced to the United States. I appreciate the report’s avoidance of blanket statements regarding the effects of climate change. However, other studies – e.g., Poland et al. – have incorporated these complexities while still offering conclusions about a number of currently established non-native pests. Finally, I am particularly dismayed that the USFS RPA does not provide analysis of any forest pathogens beyond the single mention of a few.
I am confused as to why the USFS RPA report makes no mention of Project CAPTURE (Conservation Assessment and Prioritization of Forest Trees Under Risk of Extirpation). This is a multi-partner effort to prioritize U.S. tree species for conservation actions based on invasive pests’ threats and the trees’ ability to adapt to them. Several USFS units participated, including the Southern Research Station, the Eastern Forest Environmental Threat Assessment Center, and the Forest Health Protection program. The findings were published in 2019. See here. Lead scientist Kevin Potter was one of the authors of the RPA’s chapter on disturbance.
redbay (Persea borbonia) trees in Georgia killed by laurel wilt; photo by Scott Cameron. Redbay is ranked by Project CAPTURE as 5th most severely at risk due to a non-native pest
“Project CAPTURE” provided useful summaries of non-native pests’ impacts, including the facts that
54% of the tree species on the continent are infested by one or more non-native insect or pathogen;
nearly 70% of the host/agent combinations involve angiosperm (broadleaf) species, 30% gymnosperms (e.g., conifers). When considering only non-native pests, pests attacking angiosperms had greater average severity.
Disease impacts are more severe, on average, than insect pests. Wood-borers are more damaging than other types of insect pests.
Non-native agents have, on average, considerably more severe impacts than native pests.
Project CAPTURE also ranked priority tree species based on the threat from non-native pests (Potter et al., 2019). Tree families at the highest risk to non-native pests are: a) Fagaceae (oaks, tanoaks, chestnuts, beech), b) Sapindaceae (soapberry family; includes maples, Aesculus (buckeye, horsechestnut); c) in some cases, Pinaceae (pines); d) Salicaceae (willows, poplars, aspens); e) Ulmaceae (elms) and f) Oleaceae (includes Fraxinus). I believe this information should have been included in the Resources Planning Act report in order to insure that decision-makers consider these threats in guiding USFS programs.
I also wish the USFS RPA had at least prominently referred readers to Poland et al. Among that study’s key points are:
Invasive (non-native) insects and diseases can reduce productivity of desired species, interactions at other trophic levels, and watershed hydrology. They also impose enormously high management costs.
Some non-native pests potentially threaten the survival of entire tree genera, not just individual species, e.g., emerald ash borer and Dutch elm disease. I add white pine blister rust and laurel wilt.
Emerald ash borer and hemlock woolly adelgid are listed as among the most significant threats to forests in the Eastern US.
White pine blister rust and hemlock woolly adelgid are described as so profoundly affecting ecosystem structure and function as to cause an irreversible change of ecological state.
Restoration of severely impacted forests requires first, controlling the non-native pest, then identifying and enriching – through selection and breeding – levels of genetic resistance in native populations of the impacted host tree. Programs of varying length and success target five-needle pines killed by Cronartium ribicola; Port-Orford cedar killed by the oomycete Phytophthora lateralis; chestnut blight; Dutch elm disease; butternut canker (causal agent Ophiognomonia clavigignenti juglandacearum), emerald ash borer; and hemlock woolly adelgid.
Climate change will almost certainly lead to changes in the distribution of invasive species, as their populations respond to increased variability and longer-term changes in temperature, moisture, and biotic interactions. Predicting how particular species will respond is difficult but essential to developing effective prevention, control, and restoration strategies.
Poland et al. summarizes major bioinvaders in several regions. Each region except Hawai`i (!!) includes tree-killing insects or pathogens.
It is easier to understand the RPA report’s not mentioning priority-setting efforts by two other entities, the Morton Arboretumand International Union for the Conservation of Nature (IUCN). These studies were published in 2021 and their lead entities were not the Forest Service – although the USFS helped to fund the U.S. portion of the studies.
The Morton Arboretum led in the analysis of U.S. tree species. It published studies evaluating the status of tree species belonging to nine genera, considering all threats. The Morton study ranked as of conservation concern one third of native pine species; 31% of native oak species; significant proportion of species in the Lauraceae. The report on American beech — the only North American species in the genus Fagus – made no mention of beech leaf disease – despite it being a major concern in Ohio – only two states away from the location of the Morton Arboretum near Chicago.
valley oak (Quercus lobata) in Alameda Co, California; photo by Belinda Lo via Flickr
Most of the species listed by the Morton Arboretum are of conservation concern because of their small populations and restricted ranges. The report’s coverage of native pests is inconsistent, spotty, and sometimes focuses on odd examples.
Tree Species’ Regeneration
Too late for consideration by the authors of the USFS RPA report come new studies by Potter and Riitters that evaluate species at risk due to poor regeneration. This effort evaluated 280 forest tree species native to the continental United States – two-thirds of the species evaluated in the Kevin Potter’s earlier analysis of pest impacts.
The results of Potter and Riitters 2023 only partially matched those of the IUCN/Morton studies. The Morton study did not mention three genera with the highest proportions of poorly reproducing species according to Potter and Riitters: Platanus,Nyssa, and Juniperus. Potter, Morton, and the IUCN largely agree on the proportion of Pinus species at risk. Potter et al. 2023 found about 11% of oak species to be reproducing poorly, while Morton designated a third of 91 oak species to be of conservation concern.
I believe Potter and Riitters and the Morton study agree that the Southeast and California are geographic hot spots of tree species at risk.
Potter and Riiters found that several species with wide distributions might be at risk because they are reproducing at inadequate rates. Three of these exhibit poor reproduction across their full range: Populus deltoids (eastern cottonwood), Platanus occidentalis (American sycamore), and ponderosa pine(Pinus ponderosa). Four more species are reported to exhibit poor reproduction rates in all seed zones in which they grow (the difference from the former group is not explained). These are two Juniperus,Pinus pungens, and Quercus lobata. As I point out in my earlier blog, valley oak is also under attack by the Mediterranean oak borer.
SOURCES
Fei, S., R.S. Morin, C.M. Oswalt, and A.M. 2019. Biomass losses resulting from insect and disease invasions in United States forests. Proceedings of the National Academy of Sciences. Vol. 116, No. 35. August 27, 2019.
Lugo, A.E., J.E. Smith, K.M. Potter, H. Marcano Vega, and C.M. Kurtz. 2022. The Contribution of Nonnative Tree Species to the Structure and Composition of Forests in the Conterminous United States in Comparison with Tropical Islands in the Pacific and Caribbean. USDA USFS General Technical Report IITF-54
Poland, T.M., T. Patel-Weynand, D.M. Finch, C.F. Miniat, D.C. Hayes, V.M. Lopez, eds. 2021. Invasive Species in Forests and Rangelands of the United States: A Comprehensive Science Synthesis for the United States Forest Sector. Springer Verlag. Available gratis at https://link.springer.com/book/10.1007/978-3-030-45367-1
Potter, K.M., M.E. Escanferla, R.M. Jetton, G. Man, and B.S. Crane. 2019. Prioritizing the conservation needs of United States tree species: Evaluating vulnerability to forest insect and disease threats. Global Ecology and Conservation.
Potter, K.M. and Riitters, K. 2023. A National Multi-Scale Assessment of Regeneration Deficit as an Indicator of Potential Risk of Forest Genetic Variation Loss. Forests 2022, 13, 19. https://doi.org/10.3390/f13010019
United States Department of Agriculture Forest Service. 2023. Future of America’s Forests and Rangelands: The Forest Service 2020 Resource Planning Act Assessment. GTR-WO-102 July 2023
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 Eastern deciduous forest is large and important ecologically. The forest is important for biological diversity: it shelters many endangered species, especially plants, molluscs and fish, mammals, and reptiles. In addition, the majority of forest carbon stocks in the U.S. are those of theeastern states.
But the Eastern deciduous forest is also under many anthropogenic stresses – including high numbers of non-native insects and pathogens, Liebhold map high numbers of invasive plants, blog invasive earthworms, blog browsing by overabundant deer, and timber extraction. In the southern portions of the forest, human populations are expanding, resulting in landscape fragmentation (USDA FS 2023b RPA, full reference at end of this blog).
map showing number of non-native pests in each county, as of ~2010
Agency and academic scientists in the USDA Forest Service Eastern Region (Maine to Minnesota; Delaware to West Virginia, then north of the Ohio River to Missouri) are trying to understand how long-term, continuous stressors, like deer browsing and invasive plants and earthworms, – interact with short-term gap-forming events. They call the long-term stressors “press” disturbances to distinguish them from the short-term “pulse” disturbances (Reed, Bronson, et al.; full citation at end of this blog). Understanding the processes by which forests recover from disturbance is increasingly important. Climate change is expected to raise the frequency and intensity of catastrophic natural disturbances (Spicer and Reed, Royo et al.).
The scientists emphasize that the impacts of these stressors – and effective solutions — vary depending on context.
Invasive Earthworms
USDA APHIS is responsible for regulating introduction of new species. For earthworms, APHIS’ principal concern is clearly the possibility that imported worms or soil might transport pathogens. However, the agency’s website does mention worms’ ability to disrupt the soil and possibly cause undesirable impacts on plant growth and diversity. At the 2023 National Plant Board meeting in early August 2023, Gregg Goodman, Senior Agriculturalist in APHIS PPQ NPB website for agenda? discussed issues that he considers when evaluating whether to grant permits for importing earthworms. APHIS allows imports to be used for fish bait. Dr. Goodman explained that APHIS surveyed fishermen to determine where they dump unused bait. He found no damage to plants along streams, etc. where they are dumped. A state plant health official from a northern state and I objected that the ecosystem damage caused by earthworms is well documented and we doubted that dumping of bait is not a pathway for introducing worms into natural areas.
Reed, Bronson et al. found lower earthworm biomass and density in both deer exclosures and canopy gaps. They hypothesize that the new plant growth associated with canopy gaps attracts deer, resulting in increased browse pressure. That browse pressure then affects the plant community, succession and forest structure. The changed plant community affects soil properties that then affect soil-dwelling fauna like earthworms. They believe the higher worm densities in closed-canopy sites might be the result of nutrient-rich tree leaf litter which provides both shelter and food. Another factor might be lack of recent soil disturbances in closed canopy sites.
While they say need more research is needed on the complex, combined effects of earthworms and deer, Reed, Bronson et al. still suggest that reducing deer populations or – where that is not possible – creating gaps might help manage earthworm invasions.
Deer Interactions
The long-term, chronic effect of excessive deer herbivory are well documented. See the many presentations at the recent Northern Hardwood research forum (USDA FS 2023b Proceedings). Most studies show that deer browsing overwhelms other disturbances, such as fire and canopy gaps that typically promote seedling diversity. However, recent results refine our understanding.
Samuel P. Reed and colleagues (Reed, Royo et al.) found that on the Allegheny Plateau of western Pennsylvania high deer densities at the time of stand initiation resulted in long-term reduced tree species diversity, density, and basal area. These responses were still detectable nearly four decades later. Stands are dominated by the unpalatable black cherry (Prunus serotina). The reduced stand density and the cherries’ narrower crowns lead to less above-ground biomass and reductions in above-ground carbon stocks. These scientists recommend that managers reduce deer populations to prevent changes in forest structure with probably long-term and important ramifications for many ecosystem functions.
Prunus serotina; photo by Awinch1001 via Flickr
Hovena et al. concurred with the importance of reducing deer densities, but suggested focussing on wet sites where, in their study, deer browsing had its greatest effects. On drier sites deer browsing had no effect on the diversity of woody plant seedlings.
Spicer et al. seek particularly to maintain a heterogeneous landscape to allow coexistence of both early- and late-successional species. In the Eastern Deciduous Forest biome, herbs, shrubs, and vines comprise 93% of the species richness of vascular plants
These authors found that the impact of deer browsing diverged depending on vegetation management actions. In wind-throw gaps where the plant community was retained, deer caused a 14% decline in shrub cover. In contrast, when scientists removed the extant vegetation at the beginning of recovery, deer exclusion caused a 67% increase in shrub cover. The authors speculate that vegetation removal stimulated abundant blackberry (Rubus species) regrowth. Where they had access (in gaps lacking exclosures), deer heavily browsed young Rubus stalks that sprouted after the competing vegetation was cut down. However, when the pre-established vegetation was not removed, older Rubus thickets might have protected other herbs and shrubs from browsing. Spicer et al. did not observe any major shifts in browse-tolerant species in deer-exclusion plots.
Invasive Shrubs
Hovena et al. found that in drier forest plots, the presence of non-native shrubs reduced native seedling abundance, richness, and diversity. Instead there were more seedlings of introduced species, including Lonicera maackii, L. morrowii, Ligustrum sp., and Rosa multiflora. They are concerned that replacement by invasive honeysuckles might be particularly strong in gaps resulting from death of ash trees caused by emerald ash borer. Woodlands could become dominated introduced shrubs, reducing diversity. Consequently, they recommend removing non-native shrubs in drier forests to promote seedling numbers and diversity.
In contrast, in wetter forests basal area of non-native shrubs did not affect introduced seedling abundance. However, the shrubs’ size did promote greater proportions of Lonicera maackii and Ligustrum seedlings. They suggest this might be the outcome of either abundant seed sources or allelopathic properties of some invasive shrubs e.g., L. maackii. In such sites, seedling diversity is already limited to plants that tolerate waterlogging. A hopeful note is that one native shrub, Lindera benzoin, seems able to prevent establishment of L. maackii.
Lonicera maackii; photo by pverdonk via Flickr
Hovena et al. do worry that death of ash trees might lead to declining transpiration rates, raising water tables, and further reducing seedling species richness and diversity.
Impact of Salvage Logging and Vegetation Removal
Spicer et al. studied how anthropogenic stressors affect succession. These scientists took advantage of tornado-caused gaps to compare interactions with deer browsing, salvage logging, and mechanical removal of the understory.
Contrary to expectations, none of these anthropogenic disturbances delayed community recovery or reduced diversity in comparison to the natural disturbance (tornado blowdown). Instead, adding either salvage logging or mechanical removal of understory vegetation substantially enhanced herbaceous species richness and shrub cover.
However, each major plant growth form responded differently. First, none of the manipulations affected species diversity or abundance of tree seedlings and saplings. Second, salvage logging in the wind-throw gaps increased species richness of herbs by 30%. Shrub abundance was doubled and cover almost tripled, but species richness did not change. Third, removing competing understory vegetation caused an increase of 23% in mean herbaceous cover. I have already discussed the impact of excluding deer.
Spicer et al. greet these increases in species richness with enthusiasm; they recommend managing to create a patchwork of combined natural and anthropogenic disturbances to promote plant diversity. However, I have some questions about which species are being promoted.
This study identified a total of 264 vascular plant species: 40 trees, 190 herbs, 15 shrubs, 17 vines, and 2 of unknown growth form. Only about half of these, 123 species, grew in portions of the mature forest not affected by either the tornado or one of the anthropogenic manipulations.
Gaps contained more plant species – as is to be expected. Natural blowdown areas where no manipulation was carried out had 49 more species than the undisturbed forest community (172 species). Blowdown sites subjected to salvage logging added another 53 species for a total of 225 species, or 102 more than the undisturbed reference forest.
A total of 17 species occurred only once in the authors’ data [= unique species]. Eight of these species grew only in the undisturbed forest. Two grew only in the tornado-impacted plots. Spicer et al. do not elaborate on whether these species are officially rare in that part of Pennsylvania – although it seems they might be. I wish Spicer et al. had addressed whether these possibly rare species might be affected by the forest management they recommend, i.e., intentionally creating a patchwork of various disturbances. An additional seven unique species were found in plots that had been subjected to an anthropogenic disturbance — either salvage logging or removal of remnant vegetation.
nodding trillium (Trillium cernuum); imperiled by restricted range or low populations; photo by Jason Ryndock, Pennsylvania Natural Heritage Program
In tornado-disturbed sites, one native species associated with areas where vegetation was left intact is one of the gorgeous wildflowers of eastern deciduous forests: a Trillium (species not indicated). The one native plant associated with plots from which vegetation was removed was a grass (unspecified).
Spicer et al. report that the proportion of the flora composed of non-native species was very similar between the salvage-logged area (7%) and the undisturbed reference forest (5%). Half of the non-native plant species (3.5% or 9 species) are listed as invasive in Pennsylvania (the article does not list them).
Spicer et al. say these non-native species are relatively uncommon and that they pose a minimal threat. They do concede that the invasive thorny shrub barberry (Berberis thunbergii) was more common in disturbed than intact areas. [I saw plenty of barberry along forest edges in Cook State Forest, which is only 100 miles away from the study site.] I think Spicer and others are too blasé since invasive plant populations can build up quickly when seed sources are present.
Spicer et al. raise two caveats. First, their results regarding the beneficial effects of salvage logging and vegetation manipulation probably will not apply to situations in which vast areas are logged.
More pertinent to us, they warn that their results would also not apply to forest areas in which propagules have been drastically depleted. This can result from previous human land-use or repeated catastrophic disturbances, such as canopy fires. Nor would their results apply to forests that are more threatened by invasive species. They note that a widespread and dense understory of multiple non-native species can create invasional meltdowns, resulting in a lasting depauperate state. This is especially the case when invaders at higher trophic levels, such as earthworms, are part of the mix.
Other Lessons
Reed, Bronson et al. conclude that forest canopies’ responses to disturbance are too variable to be measured by a single method. Evaluating proposals for management will require multiple measures. The overwhelming recommendation of presenters at the recent northern hardwoods research symposium (USDA FS 2023a Proceedings) was to adapt more flexible management strategies to promote forest sustainability and species diversity.
Hovena et al.’s principal finding is that interactions among site wetness, non-native shrubs and the total basal area of trees in the stand had the largest impacts on the species composition of seedlings. In Ohio, site wetness and chronic stressors like deer and introduced shrubs are acting together to shift seedling communities towards fewer native species. Of these three long-term “press” stresses, the interaction between introduced shrubs and soil wetness overshadowed even the impact of deer herbivory on seedling species richness and abundance. Surprisingly, site-specific characteristics – e.g., wetness, canopy tree competition, deer herbivory and introduced shrubs – were more influential than ash mortality in shaping woody seedling communities.
SOURCES
Hoven, B.M., K.S. Knight, V.E. Peters, D.L. Gorchov. 2022. Woody seedling community responses to deer herbivory, introduced shrubs, and ash mortality depend on canopy competition and site wetness. Forest Ecology and Management 523 (2022) 120488
Reed, S.P., D.R. Bronson, J.A. Forrester, L.M. Prudent, A.M. Yang, A.M. Yantes, P.B. Reich, and L.E. Frelich. 2023. Linked disturbance in the temperate forest: Earthworms, deer, and canopy gaps. Ecology. 2023;104:e4040. https://onlinelibrary.wiley.com/r/ecy
Reed, S.P, A.A. Royo, A.T. Fotis, K.S. Knight, C.E. Flower, and P.S. Curtis. 2022. The long-term impacts of deer herbivory in determining temperate forest stand and canopy structural complexity. Journal of Applied Ecology. 2022; 59:812-821
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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
