Category: drivers of extinction

Updates on 1) hemlocks 2) shot hole borers/Fusarium & 3) beech leaf disease

symptoms of beech leaf disease; photo by Dr. Chagas de Freitas

Three webinars during April and May provided updates on efforts to address three non-native, tree-killing pests: hemlock woolly adelgid (HWA), link invasive shot hole borers (ISHB), link and beech leaf disease (BLD) link. I attended each and summarize here.  

  1. Hemlock conservation in North Carolina  – the NC Hemlock Restoration Initiative (HRI) see SaveHemlocksNC.org  

The webinar was recorded at Hope for the Hemlocks: HWA Management Approaches on Public and Private Lands in North Carolina.  You probably need to be a member of the Natural Areas Association to watch the archived version.

I was pleased to learn about the major effort under way in North Carolina, where eastern and Carolina hemlocks are extremely important components of multiple ecosystems. In 2013, the Commissioner of Agriculture decided to make protecting hemlocks a signature project. He wanted to ensure that three state agencies – the Forest Service, Wildlife Department, and State Parks – worked together to improve the efficacy of treating trees. (Treatments available at the time were expensive and time-consuming.)

HRI treatment at Conestee Falls; HRI photo

Thom Green described the result: North Carolina’s Hemlock Restoration Initiative (HRI). The initiative is administered by the Western North Carolina Communities – a non-governmental organization with strong connections to rural communities and a history of successful collaborative projects that support agriculture and forestry. It engages state agencies, local and county governments, local NGOs, and federal agencies and works on both public and private lands with the goal of ensuring that hemlocks can survive to maturity.

HRI staff work with local partners to identify priority hemlock conservation areas (HCAs). It then sends a “strike team” to guide the partners in treating as many trees as possible. (North Carolina allows non-licensed volunteers to apply pesticides under supervision; also, landowners can treat trees on their own property.) These collaborative projects can treat up to 1,000 trees per day.

The chemicals used are imidacloprid and, where poor tree health justifies emergency treatment, dinotefuran. These are usually applied as a soil drench because it is easier for people to transport the equipment into the woods. Bark spray is used in sensitive areas. They have found that imidacloprid provides five to seven years of protection. A new product, CoreTech, is even easier to transport and works much faster than imidacloprid, however, it costs more.

The HRI believes it is minimizing non-target impacts of the neonictenoid imidacloprid because:

  • hemlocks are pollinated by wind, not insects
    • hemlocks don’t exude resins that attract insects
    • pesticide applications are tightly targetted at the base of trunk, with 10-foot setbacks from water
    • long intervals between treatments (5 – 7 years) allow soil invertebrates to recover

The program has treated 100,000 trees between 2016 and 2021 on state and private lands. Now they are starting the second round of treatments for trees treated at the beginning of the program.

Treatment priorities are based primarily on the extent to which the trees are able to take up the chemical, evaluated by the percentage of the crown that is alive and the density of foliage. Since imidacloprid can take a year to reach the canopy of a mature tree, it is used only on trees with greater than half the crown rated as healthy. When trees have a lower status, dinotefuran is added (because it can reach the canopy within weeks).  Trees with less than 30% live crown are not treated.

The Initiative also supports biocontrol programs. It has assisted releases of Laricobius nigrinis (a beetle in the family Derodontidae) and helps volunteers monitor releases and survival. Dr. Green reports that L. nigrinis has spread almost throughout western North Carolina but that questions remain regarding its impact on tree health. He thinks biocontrol is not yet reliable as stand-alone tool; success will require a suite of predatory insects.

Forest Restoration Alliance potting hemlock seedlings; HRI photo

The HRI measures the success of various treatments (Hurray!). “Impact plots” are established at the start of treatment. Staff or volunteers return every three years to monitor all aspects of the health of a few designated trees – including untreated ones. So far, they have seen encouraging responses in crown density and new growth.

  • Invasive Shot Hole Borers (ISHB) in California

See www.ishb.org and video recordings of the meeting at:  

https://youtu.be/RyqJYyLkshk (Day 1); and https://youtu.be/kWmtcbjTczw (Day 2)

A host of scientists from California spent two full days describing research and management projects funded by specific state legislation – Assembly Bill (AB)-2470 on two invasive shot hole borers.

Adoption of this legislation resulted largely from lobbying by John Kabashima. Additional funding was provided by CalFire (the state’s forestry agency). The agency responsible for managing invasive species – California Department of Food and Agriculture (CDFA) had designated these organisms as not a threat to agriculture. So it did not fund many necessary activities.

The Problem and Where It Is

“Fusarium dieback” is the disease caused by this insect-pathogen complex. The insects involved are two ambrosia beetles in the Euwallacea genus – the polyphagous (E. whitfordiodendrus) and Kuroshio (E. Kuroshio) shot hole borers. link to DMFAccording to Dr. Bea Nabua-Behermann, Urban Forestry and Natural Resources Advisor with University of California Cooperative Extension (UCCE), other fungi are present on both beetle species but its matching Fusarium sp. is the principal associated fungus and is required for the beetle’s reproduction. These are Fusarium euwallaceae and F. kuroshium.

As of spring 2022, the beetle/fungus complex has spread as far north as Santa Barbara /Santa Clarita; and inland to San Bernardino and Riverside (see the map here). They are very widespread in Orange and San Diego counties. At least 65 tree species in southern California are reproductive hosts (globally, it is 77 species; see full list here). The preferred and most succeptible hosts are several species in the Acer, Parkinsonia, Platanus, Quercus, and Salix genera. Box elder (A. negundo) is so susceptible that it is considered a sentinel tree.

Because the beetles spend most of their life inside trees, their life cycle leaves few opportunities to combat them. Females (only) fly but tend to bore galleries on their natal tree. Several speakers on the webinar said management should focus on heavily infested “amplifier trees”. Much spread is human assisted since the beetles can survive in dead wood for months if it is damp enough for the fungus.  Possible vectors are green waste, firewood, and even large wood chips or mulch.

Management – from Trapping to Rapid Response to Restoration

Akiv Eskalen of University of California Davis discussed trapping and monitoring techniques to confirm presence of the insect and pathogen. Also, he talked about setting priorities for treating trees based on the presence of reproductive hosts, host value, infestation level, and whether the trees pose a safety hazard. The disease causes too little damage to some hosts to warrant management. He emphasized the importance of preventing spread. This requires close monitoring of infested trees to see whether beetles move to neighbors. Dr. Eskalen described a major and intensive monitoring and treatment program at Disneyland. The 600 acres of parks, hotels, and parking lots have ~16,000 trees belonging to 681 species.

Several speakers described on-going efforts in Orange County. Danny Hirchag (IPM manager for Orange County Parks) described how his agency is managing 60,000 acres of variable woodlands containing 42,000 trees, of which 55% are hosts of ISHB and their associated fungi. Of greatest concern are California sycamore and coast live oak in areas of heavy public use. The highest priority is protecting public safety; next is protecting historic trees (which can’t be replaced); third is minimizing impacts to ecosystem services. Orange County Parks is currently removing fewer than 50 trees each year. Hirchag noted the importance of collaborating in the research trials conducted by the University of California Cooperative Extension.

infested California sycamore; photo by Bea Nabua-Behermann

Maximiliano Regis and Rachel Burnap, of County of Los Angeles Department of Agricultural Commissioner/Weights and Measures, described Los Angeles County’s efforts more broadly. The challenge is clear: LA County has more than 160 parks. In 2021, they placed nearly 2,500 traps, mapped infected trees, carried out on-ground surveys to find amplifier trees, removed both amplifier and hazard trees (using funds provided by CalFire), and educated the public. Their efforts were guided by an early detection-rapid response (ED/RR) Plan (2019) developed by Rosi Dagit (see below). While London plane trees (Platanus x hispanica) and California sycamores (Platanus racemose) were initially most affected, now black locusts (Robinia pseudoacacia) and box elders (Acer negundo) are succumbing. [Note: both are widespread across North America.] The researchers are trying to determine why some areas are largely untouched, despite the presence of the same tree species. Regis and Burnap noted the increasing difficulty getting confirmation of the pathogen’s presence because laboratories are overwhelmed. They continue looking for funding sources.

Rosi Dagit, Senior Conservation Biologist, Resource Conservation District of the Santa Monica Mountains, described the creation of that ED/RR system for Los Angeles County as a whole, without regard for property lines. Participants established random study plots across the entire Santa Monica Mountains Natural Recreation Area (NRA), based on proximity to areas of particularly sensitive ecological concerns. The fact that the NRA’s forests are aging and that the risk of infestations is especially high in riparian forests helped persuade policy-makers to fund the effort. The accompanying rapid response plan informs everyone about what to do, who should do it, and who pays. This information incorporates agencies’ rules about what and where to plant. It also provides measures to evaluate whether the action was effective. It did take more than two years for the county to set staffing needs etc.

John Kabashima link discussed his criteria for replanting and ecosystem restoration following tree removal in the southern California region. He recommends prompt removal of amplifier trees – especially box elder and California sycamore. He relies on replanting guidance developed by UC-Irvine (which is on the website) – especially avoiding monocultures. Kabashima reiterated the importance of close monitoring to track beetle populations and responding quickly if they build up.

Economics of Urban Forests and Cities Most at Risk

Karen Jetter (an economist at the UC Agriculture Issues Center) has developed a model to compare the costs of an early detection program to the environmental and monetary costs of infestation by Fusarium disease.  She noted that early detection and monitoring programs are often hard to justify because — when they are successful — nothing changes! She found that averted or delayed costs (including tree removals, lost ecosystem services, lost landscape asset value [replanting value] and the cost to replant) always far exceeded the cost of monitoring programs. Unfortunately, a written report about this effort (Jetter, K., A. Hollander, B.E. Nobua-Behrmann, N. Love, S. Lynch, E. Teach, N. Van Dorne, J. Kabashima, and J. Thorne. 2022. Bioeconomic Modeling of Invasive Species Management in Urban Forests; Final Report)   appears to be available only through the University of California “collaborative tools” website dedicated to practitioners and stakeholders engaged on ISHB issues. If you are not a member of the list, contact me using the comment button and ask that I send it to you. Include your email address (the comment process makes determining emails difficult if not impossible.)

Shannon Lynch (UC Davis) developed a model to estimate vulnerability of urban areas based on phylogenetic structure (relationship between tree species), host abundance, and number of beetle generations/year (linked to temperature). She found that areas with less favorable host communities can become vulnerable if the climate becomes favorable. Where the host community is already favorable, climate not important.

She evaluated 170 California cities based on their tree inventories. The cities at highest risk were San Diego, Los Angeles, the San Francisco Bay area, and the Central Valley – e.g., Sacramento. For areas lacking tree inventories, she based her risk determination on the estimated number of generations of beetles per year – based on climate. This analysis posited a very high risk in the eastern half of southern California and the Central Valley. Participants all recognized the need to apply this model to cities in Arizona and Nevada.

Possible Management Strategies

Shannon Lynch (UC Davis) studied whether endophytes might be used to kill the Fusarium fungi. She reported finding 771 fungal strains and 657 bacterial strains in tree microbiomes. Some of the fungal isolates impeded growth of the Fusarium fungi in a petri dish. She began testing whether these fungi can be used to inoculate cuttings that are to be used for restoration. She also planned to test more endophytes, and more native plant species to explore creation of a multi-fungus cocktail.

Richard Stouthamer of UC Riverside is exploring possible biocontrol agents. Of three he has evaluated, the most promising is Phasmastichus sp., which is new to science. He is still trying to establish laboratory cultures so he can test its host specificity.

See bldresearch@lists.osu.edu

symptoms of beech leaf disease; photo by Dr. Chagas de Freitas

At this meeting, scientists described research aimed at improving basic understanding of beech leaf disease’s causal agents, its mechanisms of spread, etc.  Their findings are mostly preliminary.

These findings are of greatest importance now:

  • presence of the nematodes varies considerably across leaf surface – if one collects samples from the wrong site on leaf, one won’t detect nematode (Paulo Vieria, Agriculture Research Service)
    • developing predictive risk maps that combines temperature, humidity, elevation, soils (Ersan Selvi, Ohio State). So far, he has found that BLD is greater in humid areas – including under closed forest canopies. The USFS is funding studies aimed at incorporating disease severity in detection apps.
    • determining extent of nematode presence. Sharon Reed of Ontario has found nematode DNA in trap fluids throughout the Province. It is much more common at known disease sites. Reed is also studying the presence of arthropods on beech leaves and buds.

Longer term findings and questions

  • possible vectors:
    • nematode DNA has been detected from birds – although it is not clear whether the DNA came from bird  feces, feathers, or dust (DK Martin)
    • a few live nematodes have been extracted from the excrement of caterpillars that fed on infected leaves (Mihail Kantor, ARS)
    • nematode damage to leaves:
      • presence of the nematode in leaf buds before they open (Vieria and Joe Mowery, both ARS). The nematode can create considerable damage in leaf buds before they open. Nematodes are present as early as October of the preceding year.
      • damage to leaves by nematode (Mowery, ARS) Leaf epidermal cells are distorted, stomata blocked, chlorobasts are larger than normal, irregular shape
    • possible management tools
      • are there parasites that might attack the nematode? (Paulo Vieria, ARS)
      • experimental treatment of infested trees using phosphite (Kandor, ARS)
    • ecology: how do root microbiomes compare on infested and healthy trees? (Caleb Kime, Ohio State; and David Burke, Vice President for Science at Holden Arboretum)
infested European beech in Rhode Island; photo by Dr. Nathanial A. Mitkowski

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

or

www.fadingforests.org

Search for Asian giant hornet

Asian giant hornet (Vespa mandarinia); photo by University of Florida Dept. of Entomology

Washington State’s “Giant Hornet – Hornet Herald” for June asks people to help with detecting this pest by monitoring paper wasp nests (hornets attack them). Hornet visits last 5 – 10 minutes while the hornet removes paper wasp larvae.  How to help:

  • Locate paper wasp nests that you have access to and can monitor through October. Then log the nest locations using the form here
  • Visit the nests each week, observe them, and then log your nest activity on a different form – here. Please monitor the nests for at least 5 minutes during the day once per week, but you can check the nests for as long and as often as you would like.

If you would like guidance on how to become a citizen-science monitor or trapper of Asian giant hornets – or presumably other bioinvaders – go here

Meanwhile, Washington State Department of Agriculture entomologists are in South Korea testing several hornet attractants and studying hornet foraging behavior. The goal  is to improve Washington’s trapping and tracking techniques.

Of course, 2022 is only half over, but so far neither Washington nor British Columbia has confirmed any detections.

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

or

www.fadingforests.org

Boxwood Blight – Another Failure of the Global Phytosanitary System

boxwood garden at Gunston Hall – home of founding father George Mason; Virginia; photo by Roger 4336 via Wikimedia

Boxwood blight is a disease caused by a group of fungal pathogens. While boxwoods are horticultural plants in the U.S. – important ones! – they are keystone forest species in several regions of the tropics and subtropics.

The situation with boxwood blight is yet another example of a too-frequent pattern for plant pathogens. This pattern applies even to plant taxa that are important to the ornamental horticulture industry – not only plants that are important in natural ecosystems. [See other blogs posted here under the category “plants as pest vectors”, e.g., here. The boxwood blight pathogens:  

  • are of unknown origin;
  • have a wide range of known hosts; additional hosts probable;
  • have been introduced to many new sites over about 30 years;
  • have caused considerable economic, aesthetic, and ecological harm;
  • are a threat to centers of endemism;
  • have no known methods to treat plants in forests;
  • are spread by international plant trade;
  • complicate detection by having hosts that sometimes are asymptomatic; or symptoms can be suppressed by fungicides;
  • apparently few efforts to apply phytosanitary measures to prevent further spread.

Also typical: concerned scientists are trying to promote adoption of phytosanitary measures. This takes the form of a study by Barke, Coop and Hong (full citation at the end of the blog; unless otherwise stated, information in this blog is from this source). They use several models based largely on climatic factors to predict additional geographic areas where else boxwood blight might establish.

I think it is most unfortunate that the U.S. horticultural industry prefers to avoid federal regulation despite the significant costs to its members. Instead, it has advocated for a primarily voluntary response (see below). This undermines efforts to restructure regulatory programs to improve phytosanitary agencies’ management of pathogens. Since the U.S. is such a powerful player on this issue, reducing pressure on APHIS to find more effective measures has global implications. I recognize that preventing transmission of unknown and cryptic pathogens is an intrinsically difficult task. However, tackling this problem should be a top priority for people concerned about retaining healthy floral communities.

Specifics About Boxwood Blight

Boxwood blight is caused by two ascomycete fungi, Calonectria pseudonaviculata [synonym Cylindrocladium buxicola] and Calonectria henricotiae. Both can infect and blight boxwood foliage, resulting in rapid plant death. C. henricotiae is known from only five countries in Europe; C. pseudonaviculata is currently established in 24 countries in three geographic areas: Europe and western Asia; New Zealand; and North America (30 US states and British Columbia). The disease caused by C. pseudonaviculata could spread well beyond its currently invaded range in these regions.

range of Buxus sempervirens; via Wikimedia

Native plants in the family Buxaceae grow in tropical or subtropical areas around the world. Plants in the genera Buxus, Didymeles, Haptanthus, Pachysandra, Sarcococca, and Styloceras are found in some areas of western and southern Europe; Turkey and the Caucuses into Iran; several countries in southeast and east Asia (China, Japan, South Korea, Vietnam, Indonesia); coastal Australia; high elevation areas of Africa, including Madagascar; parts of South America (southern Brazil, Uruguay, northern Argentina, and southern Chile, and foothills of the Andes); parts of Central America and the Caribbean. Asia is home to about 40 species of Buxus, four species of Pachysandra, and 11 species of Sarcococca.  In the Andes region, all five species of Styloceras are endemic. Central America and the Caribbean are home to about 50 species of Buxus; there are 37 species endemic to Cuba! Madagascar has nine endemic Buxus species.

Many Buxus species occur in small and isolated distributions resulting from both natural causes (e.g., island endemism) and anthropogenic disturbances (including deforestation and invasions of by other non-native pests, such as the box tree moth Cydalima perspectalis in Europe and western Asia).

In native stands of Buxus sempervirens in Georgia and northern Iran, where C. pseudonaviculata was detected in 2010, the disease has caused rapid and intensive defoliation of boxwood plants of different ages. [See also Lehtijarvi, Dogmus-Lehtijarvi and Oskay. Boxwood Blight in Turkey: Impact on Natural Boxwood Populations and Management Challenges. Baltic Forestry 2017, vol. 23(1)] Infected plants are also vulnerable to attacks by secondary opportunistic pathogens that can lead to eventual death. Damage to these forests could lead to reductions in soil stability and subsequent declines in water quality and flood protection, changes in forest structure and composition, and declines in Buxus-associated biodiversity (at least 63 species of lichens, fungi, chromista and invertebrates might be obligate).

Barke, Coop and Hong expect excessive heat and seasonal dryness at one extreme and excessive cold at the other to limit areas in North America and Europe/central Asia where the disease can establish. Areas with oceanic rather than continental climates are probably more vulnerable. However, heat and aridity barriers could be overcome by artificial irrigation of horticultural plantings.

Indeed, the conditions favoring C. pseudonaviculata establishment – warm temperatures and high humidity or water on the leaves – are commonly found in production nurseries. Overhead irrigation exacerbates the risk. Production nurseries also have large numbers of host plants in close proximity – so it is easy for disease to spread (Douglas). 

I am reminded that the causal agent of sudden oak death, Phytophthora ramorum,  has been spread from production nurseries located in hot, dry areas that were considered unsuitable to the pathogen – because conditions inside the nursery were suitable.

wild Buxus on island of Corsica; photo by Sten Porse via Wikimedia

As I noted, the origin of C. pseudonaviculata is unknown. Barke, Coop and Hong think it is most likely in eastern Asia, which is thought to be the likely native region of box tree moth. However, they cannot rule out some other center of diversity for Buxaceae species e.g., the Caribbean or Madagascar.

Barke, Coop and Hong call for additional studies to

  1. Explore potential effects of climate change on establishment risk, especially higher latitude areas expected to see increasing humidity, precipitation, and rising temperatures.
  2. Determine ability of C. pseudonaviculata microsclerotia to survive higher temperatures, e.g. in parts of the U.S. Deep South that may have ideal growing conditions during cool seasons.
  3. Modify the CLIMEX model developed for this study to predict the potential distribution of C. henricotiae, a closely related but genetically distinct species with greater tolerance of higher temperatures.

They call for a strict phytosanitary protocol for risk mitigation of accidental intro, with effective surveillance for early detection, and development of a recovery plan.

Regulatory (non) Response

Boxwood blight was first detected in the United Kingdom in mid-1990s; then in New Zealand in 2002. Only then was the causal agent determined. It was first detected in the U.S. in October 2011 (in Connecticut). It was quickly determined to be established in the mid-Atlantic region. Apparently the British, other European countries, and APHIS all decided the pathogen was too widespread to regulate (Douglas).

The U.S. is relying on a voluntary program. The nursery industry, through its Horticultural Research Institute (HRI), and the National Plant Board developed guidance for best management practices – updated as recently as 2020. 

boxwood blight symptoms; Oregon State University; via Flickr

In contrast, APHIS has acted to regulate the boxwood tree moth, Cydalima perspectalis. The moth was first detected in North America near Toronto in 2018. U.S. nurseries in six states received infected plants in spring 2021. On May 26, 2021, APHIS prohibited importation of host plants from Canada, including boxwood (Buxus spp), Euonymus (Euonymus spp), and holly (Ilex spp).

In July 2021, the moth was detected in Niagara County, New York. It was thought that the moths had flown or been blown into the area from Canada.  New York adopted an intrastate quarantine of three counties (Erie, Niagara, and Orleans) in December 10, 2021. APHIS followed with an interstate quarantine on March 23, 2022.

SOURCES

Barke, B.S., L. Coop and C. Hong. 2022.  Potential Distribution of Invasive Boxwood Blight Pathogen (Calonectria pseudonaviculata) as Predicted by Process-Based and Correlative Models. Biology 2022, 11, 849. https://doi.org/10.3390/biology11060849 www.mdpi.com/journal/biology

Douglas, S.M. Fact sheet; Connecticut Agricultural Experiment Station https://portal.ct.gov/-/media/CAES/DOCUMENTS/Publications/Fact_Sheets/Plant_Pathology_and_Ecology/2020/Boxwood-Blight-(1).pdf?la=en&hash=A4C6AF39765F27FDDEB5B4DC3FD3B6F3

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

or

www.fadingforests.org

Invasions cost protected areas more than $22 billion in 35 years

Burmese python in Everglades National Park; photo by Bob Reed, US FWS

Scientists continue to apply data collected in an international database (InvaCost; see “methods” section of Cuthbert et al.; full citation at end of this blog) to estimate the economic costs associated with invasive alien species (IAS). These sources reported $22.24 billion in economic costs of bioinvasion in protected areas over the 35-year period 1975 – 2020. Because the data has significant gaps, no doubt invasions really cost much more.

Moodley et al. 2022 (full citation at end of this blog) attempt to apply these data to analyze economic costs in protected areas. As they note, protected areas are a pillar of global biodiversity conservation. So it is important to understand the extent to which bioinvasion threatens this purpose. 

Unfortunately, the data are still too scant to support any conclusions. Such distortions are acknowledged by Moodley et al. I will discuss the data gaps below a summary of the study’s findings.

The Details

Of the estimated $22.24 billion, only 4% were observed costs; 96% were “potential” costs (= extrapolated or predicted based on models). Both had generally increased in more recent years, especially “potential” costs after 1995. As is true in other analyses of InvaCost data, the great majority (73%) of observed costs covered management efforts rather than losses due to impacts. The 24% of total costs ascribed to losses, or damage, exceeded the authors’ expectation. They had thought that the minimal presence of human infrastructure inside protected areas would result in low records of “economic” damages.

The great majority (83%) of reported management costs were reactive, that is, undertaken after the invasion had occurred. In terrestrial environments, there were significantly higher bioinvasion costs inside protected areas than outside (although this varied by continent). However, when considering predicted or modelled costs, the importance was reversed: expected management costs represented only 5% while these “potential” damages were 94%.

Higher expenditures were reported in more developed countries – which have more resources to allocate and are better able to carry out research documenting both damage and effort. 

More than 80% of management costs were shouldered by governmental services and/or official organizations (e.g. conservation agencies, forest services, or associations). The “agriculture” and “public and social welfare” sectors sustained 60% of observed “damage” and 89% of “mixed damage and management” costs respectively. The “environmental” and “public and social welfare” sectors together accounted for 94% of all the “potential” costs (predicted based on models) generated by invasive species in protected areas; 99% of damage costs. With the partial exception of the agricultural sector, the economic sectors that contribute the most to movement to invasive species are spared from carrying the resulting costs.

Lord Howe Island, Australia; threatened by myrtle rust; photo by Robert Whyte, via Flickr

Invasive plants dominated by numbers of published reports – 64% of reports of observed costs, 79% of reports of “potential”. However, both actual and “potential” costs allotted to plant invasions were much lower than for vertebrates and invertebrates. Mammals and insects dominated observed animal costs.

It is often asserted that protected areas are less vulnerable to bioinvasion because of the relative absence of human activity. Moodley et al. suggest the contrary: that protected areas might be more vulnerable to bioinvasion because they often host a larger proportion of native, endemic and threatened species less adapted to anthropogenic disturbances. Of course, no place on Earth is free of anthropogenic influences; this was true even before climate change became an overriding threat. Plenty of U.S. National parks and wilderness areas have suffered invasion by species that are causing significant change (see, for example, here, here, and here).

Despite Best Efforts, Data are Scant and Skewed

Economic data on invasive species in protected areas were available for only a tiny proportion of these sites — 55 out of 266,561 protected areas.

As Moodley et al. state, their study was hampered by several data gaps:

  1. Taxonomic bias – plants are both more frequently studied and managed in protected areas, but their reported observed costs are substantially lower than those of either mammals or insects.
  2. The data relate to economic rather than ecological effects. The costliest species economically might not cause the greatest ecological harm.
  3. Geographical bias – studies are more plentiful in the Americas and Pacific Islands. However, studies from Europe, Africa and South America more often report observed costs. The South African attention to invasive species (see blogs here, here, and here), and economic importance of tourism to the Galápagos Islands exacerbate these data biases.
  4. Methodological bias – although reporting bioinvasion costs has steadily increased, it is still erratic and dominated by “potential” costs = predictions, models or simulations.

I note that, in addition, individual examples of high-cost invasive species are not representative. The highest costs reported pertained to one agricultural pest (mango beetle) and one human health threat (mosquitoes).

Great Smokey Mountains National Park; threatened by mammals (pigs), forest pests, worms, invasive plants … Photo by Domenico Convertini via Flickr

As these weaknesses demonstrate, a significant need remains for increased attention to the economic aspects of bioinvasion – especially since political leaders pay so much greater attention to economics than to other metrics. However, the reported costs – $22.24 billion over 35 years, and growing! – are sufficient in the view of Moodley et al. to support advocating investment of more resources in invasive species management in protected areas, including – or especially – it is not quite clear — preventative measures.

SOURCES

Cuthbert, R.N., C Diagne, E.J. Hudgins, A. Turbelin, D.A. Ahmed, C. Albert, T.W. Bodey, E. Briski, F. Essl, P.J. Haubrock, R.E. Gozlan, N. Kirichenko, M. Kourantidou, A.M. Kramer, F. Courchamp. 2022. Bioinvasion cost reveals insufficient proactive management worldwide. Science of The Total Environment Volume 819, 1 May, 2022, 153404

Moodley, D., E. Angulo, R.N. Cuthbert, B. Leung, A. Turbelin, A. Novoa, M. Kourantidou, G. Heringer, P.J. Haubrock, D. Renault, M. Robuchon, J. Fantle-Lepczyk, F. Courchamp, C. Diagne. 2022.  Surprisingly high economic costs of bioinvasions in protected areas. Biol Invasions. https://doi.org/10.1007/s10530-022-02732-7

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

or www.fadingforests.org

Two Teams with a New Take: Insect Losses Due to Invasive Plants

monarch butterfly on swamp milkweed; photo by Jim Hudgins, USFWS

I have been impressed recently by two groups of scientists who are trying to broaden understanding of the impacts of invasive plants by examining the interactions of those plants with insects. As they note, herbivorous insects are key players in terrestrial food webs; they transfer energy captured by plants through photosynthesis to other trophic levels. This importance has been recognized since Elton first established the basic premises of food webs (1927) [Burghardt et al.; full citation at end of blog] Arthropods comprise significant members of nearly every trophic level and are especially important as pollinators. If introduced plants cause changes to herbivore communities, there will probably be effects on predators, parasites, and other wildlife through multitrophic interactions [Lalk et al.; Tallamy, Narango and Mitchell].

[I briefly summarize the findings of a third group of scientists at the end of this blog. The third group looks at the interaction between agriculture – that is, planting of non-native plants! – and climate change.]

One approach to studying this issue, taken by Douglas Tallamy of the University of Delaware and colleagues, is to look at the response of herbivorous insects to NIS woody plants fairly generally. They integrate their studies with growing concern about the global decline in insect populations and diversity. They note that scientists have focused on light pollution, development, industrial agriculture, and pesticides as causes of these declines. They decry the lack of attention to disruption of specialized evolutionary relationships between insect herbivores and their native host plants due to widespread domination by non-indigenous plants [Richard, Tallamy and Mitchell].

In their studies, Tallamy and colleagues consider not just invasive plants, but also non-native plants deliberately planted as crops or ornamentals, or in forestry. They point out that such introduced plants have completely transformed the composition of plant communities in both natural and human-dominated ecosystems around the globe. At least 25% of the world’s planted forests are composed of tree species not native to their locale. At least one-sixth of the globe is highly vulnerable to plant invasions, including biodiversity hotspots [Richard, Tallamy and Mitchell].

A different approach, taken by Lalk and colleagues, is more closely linked to concern about impacts of the plants themselves. They have chosen to pursue knowledge about relationships between individual species of invasive woody plants and the full range of arthropod feeding guilds – pollinators, herbivores, twig and stem borers, leaf litter and soil organisms. In so doing, they decry the general absence of data.

Both teams agree that:

  • Invasive plants are altering ecosystems across broad swaths of North America and the impacts are insufficiently understood.
  • The invasive plant problem will get worse because non-native species continue to be imported and planted. (Reminder: the Tallamy team considers impacts of deliberate planting as well as bioinvasion.)
  • Plant-insect interactions are the foundation of food webs, so changes to them will have repercussions throughout ecosystems.

Tallamy team

Non-native plants have replaced native plant communities to a greater or lesser extent in every North American biome – including anthropogenic landscapes [Burghardt]. The first trophic level in suburban and urban ecosystems throughout the U.S. is dominated by plant species that evolved in Southeast Asia, Europe, and South America [Tallamy and Shropshire]. Abundant non-native plants not only dominate plant biomass; they also reduce native plant taxonomic, functional and phylogenetic diversity and heterogeneity. Thus, they indirectly alter the abundance of native insects  [Burghardt; Richard, Tallamy and Mitchell].

I think these articles might actually underestimate the extent of these impacts. While Richard, Tallamy and Mitchell report that more than 3,300 species of non-native plants are established in continental U.S., years ago Rod Randall said that more than 9,700 non-native plant species were naturalized in the U.S. (probably includes Hawai` i.   The Tallamy team cites USDA Forest Service data showing 9% of forests in the southeast are invaded by just 33 common invasive plant species [Richard, Tallamy and Mitchell], I have cited that and other sources showing even greater extents of plant invasion in the east and here; other regions and here. 

The Tallamy team has conducted several field experiments that demonstrate that the presence of non-native plants suppress numbers and diversity of native lepidopteran caterpillars. These non-native woody plants have not replaced the ecological functions of the native plants that used to support insect populations. This is true whether or not the non-native plants are deliberately planted or are invading various ecosystems on their own. [Richard, Tallamy and Mitchell]. (Of course, they expect plant invasions to grow; they note that some of the many ornamental species that are not yet invasive will become so.)

The result is disruption of the ecological services delivered by native plant communities, including the focus of their studies: plants’ most fundamental contribution to ecosystem function: generation of food for other organisms [Burghardt].

They note that plants’ relationship to insects differs depending on the insects’ feeding guilds — folivores, wood eaters, detritivores, pollinators, frugivores, and seed-eaters; and among herbivores with different mouthparts — chewing or sucking; and as host plant specialists or generalists. They decry studies that fail to recognize these differences [Tallamy, Narango, and Mitchell].

The Tallamy team explores why insect populations decline among non-native plants. That is,  

1) Do insects directly requiring plant resources have lower fitness when using non-native plants; do they not recognize them as viable host plants; or do they avoid them altogether? 

2) Are reductions in numbers of specialist herbivores mitigated by generalists? A decade of research shows that both specialists and generalists decline.

The team’s studies focus on lepidopteran larvae (caterpillars). Insect herbivores are both the largest taxon of primary consumers and extremely important in transferring energy captured by plants through photosynthesis to other trophic levels [Burghardt]. In addition, insects with chewing mouthparts are typically more susceptible to defensive secondary metabolites contained in leaves than are insects with sucking mouthparts that tap into poorly defended xylem or phloem fluids [Tallamy, Narango and Mitchell].

A study by Burghardt et al. found that 75% of all lepidopteran species and 93% of specialist species were found exclusively on native plant species. Non-native plants that were in the same genus as a native plant often supports a lepidopteran community that is a similar but depauperate subset of the community found on its native congener. In fact, the insect abundance and species richness supported by non-native congeners of native species was reduced by 68%.

A meta-analysis of 76 studies by other scientists found that, with few exceptions, caterpillars had higher survival and were larger when reared on native host plants. Plant communities invaded by non-native species had significantly fewer Lepidoptera and less species richness. In three of eight cases examined, non-native plants functioned as ecological traps, inducing females to lay eggs on plants that did not support successful larval development. Richard, Tallamy and Mitchell cite as an example the target of many conservation efforts, monarch butterflies (Danaus plexxipus), which fail to reproduce when they use nonnative swallowworts (Vincetoxicum species.) instead of related milkweeds (Asclepias species.).

Tallamy and Shropshire ranked 1,385 plant genera that occur in the mid-Atlantic region by their ability to support lepidopteran species richness. They found that introduced ornamentals are not the ecological equivalents of native ornamentals. This means that solar energy harnessed by introduced plants is largely unavailable to native specialist insect herbivores.

Tallamy, Narango, and Mitchell describe the following patterns:

1) Insects with chewing mouthparts are typically more susceptible to defensive secondary metabolites contained in leaves than are insects with sucking mouthparts that tap into poorly defended xylem or phloem fluids. As a result, sucking insects find novel non-indigenous plants to be acceptable hosts more often. However, there are more than 4.5 times as many chewing (mandibulate) insect herbivores than sucking (haustellate) species. It follows that the largest guild of insect herbivores is also the most vulnerable to non-native plants as well as being the most valuable to insectivores.

native azalea Rhododendron periclymenoides; photo by F.T. Campbell

2) Woody native species, on average, support more species of phytophagous insects than herbaceous species.

3) Although insects are more likely to accept non-native congeners or con-familial species as novel hosts, non-native congeners still reduced insect abundance and species richness by 68%.

4) Host plant specialists are less likely to develop on evolutionarily novel non-indigenous plants than are insects with a broader diet. There are far more specialist species than generalists, so generalists will not prevent serious declines in species richness and abundance when native plants are replaced by non-indigenous plants. In addition, non-native plants cause significant reductions in species richness and abundance even of generalists. In fact, generalists are often locally specialized on particular plant lineages and thus may function more like specialists than expected.

5) Any reduction in the abundance and diversity of insect herbivores will probably cause a concomitant reduction in the insect predators and parasitoids of those herbivores – although few studies have attempted to measure this impact beyond spiders, which are abundant generalists. The vast majority of parasitoids are highly specialized on particular host lineages.

6) Studies comparing native to non-native plants must avoid using native species that support very few phytophagous insects as their baseline, e.g., in the mid-Atlantic region tulip poplar trees (Liriodendron tulipifera) and Yellowwood (Cladrastus kentuckea).

7) Insects that feed on well-defended living tissues such as leaves, buds, and seeds are less likely to be able to include non-native plants in their diets than are insects that develop on undefended tissues like wood, fruits, and nectar. Although this hypothesis has never been formally tested, they note the ease with which introduced wood borers – emerald ash borer, Asian longhorned beetle, polyphagous and Kuroshio shot-hole borers, redbay ambrosia beetle, Sirex woodwasp (all described in profiles posted here — have become established in the US.

palamedes swallowtail Papilio palamedes; photo by Vincent P. Lucas; this butterfly depends on redbay, a tree decimated by laurel wilt disease vectored by the redbay ambrosia beetle

Lalk and Colleagues

As noted, Lalk and colleagues have a different frame; they focus on individual introduced plant species rather than starting from insects. They also limit their study to invasive plants. The authors say there is considerable knowledge about interactions between invasive herbaceous plants and arthropod communities, but less re: complex interactions between invasive woody plants and arthropod communities, including mutualists (e.g., pollinators), herbivores, twig- and stem-borers, leaf-litter and soil-dwelling arthropods, and other arthropod groups.

They ask why this knowledge gap persists when invasive shrubs and trees are so widespread and causing considerable ecological damage. They suggest the answer is that woody invaders rarely encroach on high-value agricultural systems and some are perceived as contributing ecosystem services, including supporting some pollinators and wildlife.

Lalk and colleagues seek to jump-start additional research by summarizing what is currently known about invasive woody plants’ interactions with insects. They found sufficient data about 11 species – although even these data are minimal. They note that all have been cultivated and sold in the U.S. for more than 100 years. All but one (mimosa) are listed as a noxious weed by at least one state; two states (Rhode Island and Georgia) don’t have a noxious weed list. None of the 11 is listed under the federal noxious weed statute.

Ailanthus altissima

Illustrations of how minimal the existing information is:

  • Tree-of-heaven (Ailanthus altissima) is noted to be supporting expanded populations of the Ailanthus webworm moth (Atteva aurea), which is native to Central America; and to be the principal reproductive host for SLF (Lycorma delicatua)  https://www.dontmovefirewood.org/pest_pathogen/spotted-lanternfly-html/
  • Chinese tallow (Triadica sebifera) is thought to benefit both native generalist bee species and non-indigenous European honeybees (Apis mellifera).
  • Chinese privet (Ligustrum sinense) appears to suppress populations of butterflies, bees, and beetles.

Lalk and colleagues then review what is known about interactions between individual invasive plant species in various feeding guilds. They point out that existing data on these relationships are scarce and sometimes contradictory.

They believe this is because interactions vary depending on phylogenetic relationships, trophic guild, and behavior (e.g., specialized v. generalist pollinator). Arthropods can be “passengers” of a plant invasion. That is, they can be affected by that invasion, with follow-on effects to other arthropods in the community. Also, arthropods can be “drivers” of invasion, increasing the success of the invasive plants.

They then summarize the available information on various interactions. For example, they note that introduced plants can compete with native plants in attracting pollinators, causing cascading effects. Or they can increase pollination services to native plants by attracting additional pollinators.

They note that herbivore pressure on invasive plants can have important impacts on growth, spread, and placement within food webs. They note that these cases support the “enemy release hypothesis”, although they think there are probably additional driving mechanisms.

Lalk and colleagues note that most native twig- and stem-borers (Coleoptera: Buprestidae, Curculionidae, Cerambycidae; Hymenoptera: Siricidae) are not considered primary pests but that some of our most damaging insect species are wood borers (see above).

Some of these borers are decomposers; in that role, they are critical in nutrient cycling.

Arthropods in leaf litter and soil also serve important roles in the decomposition and cycling of nutrients, which affects soil biota, pH, soil nutrients, and soil moisture. They act as a trophic base in many ecosystems. Lalk and colleagues suggest these arthropod communities probably change with plant species due to differences in leaf phytochemistry. They cite one study that found litter community composition differed significantly between litter beneath tree-of-heaven, honeysuckle (Lonicera maackii), and buckthorn (Rhamnus cathartica) compared to litter underneath surrounding native trees.

Recommendations

Both the Tallamy and Lalk teams call for ending widespread planting of non-native plants. Lalk and colleagues discuss briefly the roles of

  • The nursery industry (including retailers); they produce what sells.
  • Scientists and educators have not sufficiently informed home and land owners about which species are invasive or about native alternatives.
  • Private citizens buy and plant what their neighbors have, what they consider aesthetically pleasing, or what is being promoted.
  • States have not prohibited sale of most invasive woody plants. Regulatory actions are not a straightforward matter; they require considerable time, supporting information, and compromise.

Tallamy team calls for restoration ecologists in the eastern U.S. to consider the number of Lepidopterans hosted by a plant species when deciding what to plant. For example, oaks (Quercus), willows (Salix), native cherries (Prunus)and birches (Betula) host orders of magnitude more lepidopteran species in the mid-Atlantic region than tulip poplar.(Those lepidopteran in turn support breeding birds and other insectivorous organisms.) [Tallamy & Shropshire]

Lalk and colleagues focused on identifying several key knowledge gaps:

  • How invasive woody plants affect biodiversity and ecosystem functioning
  • How they themselves function in different habitats.
  • Do non-native plants drive shifts in insect community composition, and if so, what is that shift, and how does it affect other trophic levels?
  • How do IAS woody plants affect pollinators?

The authors do not minimize the difficulty of separating such possible plant impacts from other factors, including climate change and urbanization.

Global Perspective

oil palm plantation in Malaysia; © CEphoto, Uwe Aranas

Outhwaite et al. (full citation at end of this blog) note that past studies have shown that insect biodiversity changes are driven primarily by land-use change (which is another way of saying planting of non-native species – as Dr. Tallamy and colleagues describe it) and increasingly by climate change. They south to examine whether these drivers interact. They found that the combination of climate warming and intensive agriculture is associated with reductions of almost 50% in the abundance and 27% in the number of species within insect assemblages relative to levels in less-disturbed habitats with lower rates of historical climate warming. These patterns were particularly clear in the tropics (perhaps partially because of the longer history of intensive agriculture in temperate zones). They found that high availability of nearby natural habitat (that is, native plants) can mitigate these reductions — but only in low-intensity agricultural systems.

Outhwaite et al. reiterate the importance of insect species in ecosystem functioning, citing pollination, pest control, soil quality regulation & decomposition. To prevent loss of these important ecosystem services, they call for strong efforts to mitigate climate change and implementation of land-management strategies that increase the availability of natural habitats.

SOURCES

Burghardt, K. T., D. W. Tallamy, C. Philips, and K. J. Shropshire. 2010. Non-native plants reduce abundance, richness, and host specialization in lepidopteran communities. Ecosphere 1(5):art11. doi:10.1890/ES10-00032.

Lalk, S. J. Hartshorn, and D.R. Coyle. 2021. IAS Woody Plants and Their Effects on Arthropods in the US: Challenges and Opportunities. Annals of the Entomological Society of America, 114(2), 2021, 192–205 doi: 10.1093/aesa/saaa054

Outhwaite, C.L., P. McCann, and T. Newbold. 2022.  Agriculture and climate change are shaping insect biodiversity worldwide. Nature 605 97-192 (2022)  https://www.nature.com/articles/s41586-022-04644-x

Richard, M. D.W. Tallamy and A.B. Mitchell. 2019. Introduced plants reduce species interactions. Biol Invasions

Tallamy, D.W., D.L. Narango and A.B. Mitchell. 2020. Ecological Entomology (2020), DOI: 10.1111/een.12973 Do Non-native plants contribute to insect declines?

Tallamy, D.W. and K.J. Shropshire. 2009. Ranking Lepidopteran Use of Native Versus Introduced Plants Conservation Biology, Volume 23, No. 4, 941–947 2009 Society for Conservation Biology DOI: 10.1111/j.1523-1739.2009.01202.x

Posted by Faith Campbell

We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.

For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm

Plants Depend on Animals – and They are Disappearing

black berry eating hawthorn berries; photo by Paul D. Vitucci

Articles by Evan Fricke and colleagues remind us to look more broadly at bioinvasion to consider the impact on ecosystem function and evolution. They focus on animal interactions with plants in the shared environment, especially animals’ role as seed dispersers.

The authors also remind us that natural barriers explain why there are different species in different areas and thus how evolution and speciation follow different paths in different places. Think of Galapagos finches evolving in isolation from a few ancestors that somehow made it over the ocean from mainland South America.

These points are made in two recent articles.

In the first, Fricke and Svenning 2020 (full citation at end of this blog) note that about half of all plant species depend on animals to disperse their seeds. Animal seed dispersal is influenced by several drivers of global change, including local or generalized extinction (= defaunation); bioinvasion; and habitat fragmentation. The decline of large vertebrates has a particularly important role in these interactions.

Their study focused on fleshy-fruited plants that are dispersed by animals. (The study does not include nuts, e.g., acorns, which are presumably subject to some of the same pressures.) They expect evolution of the affected plants and animals to proceed differently as a result of the new partnerships, but they did not study any such interactions.

Their study covered animal seed-dispersal interactions with plants at 410 locations. The data encompassed 24,455 unique animal-plant pairs involving 1,631 animal and 3,208 plant species. Three quarters of the animals were birds; most of the rest were mammals, primarily bats and primates. Only 1% were in other animal groups – lizards, tortoises, or fish.

fruit bats on Luzon, Philippines; photo by Francesco Vernonesi; Flickr.com

They found that introduced plants and animals are twice as likely as native species to interact with introduced partners. The resulting interactions are likely to amplify biotic homogenization in future ecosystems. Already, introduced species have largely replaced missing native frugivore species in some places. In fact, mutualisms in which either or both the plant and animal is an introduced species are now about seven times higher than decades ago.

These mutual-benefit interactions of introduced species are even more prevalent in areas where human modification of the environment is greater. The proportion of introduced species and of novel interactions caused by introduced plant or animal species was higher for oceanic island systems than for continental bioregions. This finding adds a new dimension to the already recognized heightened susceptibility of remote islands to invasion and their loss of native species. Continental bioregions’ networks typically had few introduced animals and a greater prevalence of intro plants than animals.

Fricke and colleagues think plant-frugivore networks are likely to increasingly favor a relatively few introduced generalists over many native species, reducing the uniqueness of future biotas. The result might be to reduce resilience of terrestrial ecosystems by, first, allowing perturbations to propagate more quickly; and, second, by exposing disparate ecosystems to similar drivers. They called for giving higher priority to managing increasing ecological homogenization.

In the second article, Fricke, Ordonez, Rogers, and Svenning (2022) note that climate change requires many plant species to shift their populations hundreds of meters to tens of kilometers per year to track their climatic niche. Earth is also experiencing the formation of novel communities as species introductions and shifting ranges result in co-occurrence of species that do not share co-evolutionary history. They conclude that the novel mutualistic interaction networks will influence whether certain plant species persist and spread.

These authors examined four scenarios to assess how current long-distance dispersal has been affected by past defaunation and invasion and how it is threatened by species endangerment. These scenarios are as follows:

1st scenario (current scenario) = natural and introduced ranges of extant species today.

2nd scenario (natural scenario) = mammal and bird ranges as they would be if unaffected by extinctions, range contractions, or introductions.

3rd scenario (extinction scenario) = those bird and mammal species listed as vulnerable or endangered by the IUCN go extinct.

4th scenario (extirpation of introduced species scenario) = introduced species are extirpated.

Fricke and colleagues estimate that extinction of at least local populations of seed-dispersing mammals and birds has already reduced the capacity of plants to track climate change by 60% globally. The effect is strongest in temperate regions and regions with little topographic complexity. Two examples are eastern North America and Europe. These regions face a double threat: rapid climate change and loss of large mammals that provided long-distance dispersal.

The extinction scenario is most evident in Southeast Asia and Madagascar. The remaining animal seed dispersers are already threatened or endangered. Fricke and colleagues project that future loss of vulnerable and endangered species from their current ranges would result in a further reduction of 15% in the capacity of plants to track climate change.

The contrary situation is found on islands which have few native mammals. Introduced species are now important long-distance seed dispersers. In some cases, the introduced animals are dispersing invasive plant seeds, e.g., on Hawai`i feral hogs are spreading the invasive plant strawberry guava (Psidium cattleianum).

strawberry guava on Maui; photo by Forest and Kim Starr

People’s actions have resulted in ecoregions disproportionately losing the species that provide long-distance seed dispersal function, i.e., large mammals. In other words, human activities have caused not only rapid climate change—requiring broad-scale range shifts by plants—but also defaunation of the birds and mammals needed by plants to do so. Habitat fragmentation and other land-use changes will likely amplify existing constraints on plant range shifts.

Fricke and colleagues say their findings emphasize the importance of not only promoting habitat connectivity to maximize the functional potential of current seed dispersers but also restoring biotic connectivity through the recovery of large-bodied animals to increase the resilience of vegetation communities under climate change.

SOURCES

Fricke, E. C., & Svenning, J. C. (2020). Accelerating homogenization of the global plant–frugivore meta-network. Nature, 585(7823), 74-78. https://www.nature.com/articles/s41586-020-2640-y

Fricke, E. C., Ordonez, A., Rogers, H. S., & Svenning, J. C. (2022). The effects of defaunation on plants’ capacity to track climate change. Science, 375(6577), 210-214. https://www.science.org/doi/full/10.1126/science.abk3510

Posted by Faith Campbell

We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.

For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm

West Coast Responding to EAB

nearly pure stand of Oregon ash in Ankeny National Wildlife Refuge, Oregon; photo by Wyatt Williams, Oregon Department of Forestry

While Michiganders document the impacts of the emerald ash borer (EAB) there, conservationists on the West Coast are jump-starting efforts to save their regional species, Oregon ash (Fraxinus latifolia). Earlier field tests in the Midwest showed that EAB will attack Oregon ash (press release) – something West Coast state would like to counter as early and effectively as possible.

Oregon ash is a wide-ranging species, occurring from California to Washington and possibly into British Columbia. The species has not been studied extensively (it is not a timber species!), but it is clearly an imponearlrtant component of riparian forests. In wetter parts of the Willamette Valley, ash is the predominant tree species. See the photo of the riparian forest in the Ankeny National Wildlife Refuge; this forest is nearly 100% Oregon ash (ODA/ODF EAB Response Plan).

As is true in the Midwest, ash provides important food and habitat resources along creeks and rivers where seasonally high water-tables can exclude nearly all other tree species. Standing and fallen dead ash biomass can alter soil chemistry and affect rates of decomposition, nutrient, and water cycling, i.e., nutrient resource availability for the remaining trees. Gaps in tree canopy can increase soil erosion, stormwater runoff and elevated stream temperatures. In dense stands of Oregon ash, understory vegetation is often sparse, consisting primarily of sedges. The authors of the Response Plan anticipate invasion by non-native plants into canopy gaps caused by the loss of ash trees as a result of an EAB invasion. In Michigan, though, it is the sedges that dominate these gaps.

The Oregon Department of Forestry, the state Department of Agriculture, and other entities have actively participated in “don’t move firewood” campaigns for at least a decade. The Departments of Forestry and Agriculture also led a team that prepared the EAB Response Plan in 2018 (full citation at the end of this blog). It lays out in considerable detail the roles of both government agencies and non-governmental stakeholders. Oregon’s quarantine is broad, covering all insects not on an approved list (Williams, pers. comm.)

California has inspected incoming firewood for years. In April 2021 – after APHIS terminated the federal quarantine on EAB — California Department of Food and Agriculture established a state quarantine on the beetle and articles that could transport it into the state. In doing so, CDFA noted that commercially grown olive trees might also be at risk to EAB.

Washington State operates a statewide trapping program for invasive insects. There has also been significant attention to non-native insect threats to urban forests. These have included a study in 2016 led by the Washington Invasive Species Council (WISC). It involved a partnership of WISC with the Washington Department of Natural Resources Urban and Community Forestry Program as well as and statewide stakeholder meetings [Bush, pers. comm.].

Of these various state-wide initiatives, the institutions in Oregon appear to be most pro-active. The Tualatin Soil and Water Conservation District provided $10,000 to fund some of the genetics work and testing for EAB resistance. Other funding came from the USDA Forest Service Forest Health Protection unit of State and Private Forestry (not from USFS’ Research Program). As described by USFS geneticist Richard Sneizko in an article in the publication TreeLine (full citation at end of blog), participants hope to find at least some level of genetic resistance to EAB. Any such resistance might be deployed in several ways: 1) promoting reproduction by resistant trees to enhance their numbers before EAB gets to Oregon; 2) using seeds from resistant trees for restoration of natural areas; or 3) cross-breeding resistant trees to build genetically diverse stocks of resistant trees for future restoration.

Participants think it is vitally important to work from seeds collected over much of the range of Oregon ash – first, to search for probably very rare resistant trees; and second, to preserve the full diversity of the tree species’ genome so that restored ash will be adapted to the wide variety of conditions in which ash grow.

Participants in this effort include the forest genetics/tree improvement community – specifically, the USDA Forest Service Dorena Genetic Resource Center (located in Cottage Grove, Oregon) and Washington State University at Puyallup Research & Extension Center. Also engaged is the public gardens community, specifically the Huntington Botanical Gardens in San Marino, Los Angeles County.  The garden is collecting seed of Oregon and other western ashes from California and Washington State.

The first step in assessing resistance is collecting seed from ash trees across the range of Oregon ash. This began in 2019. Carried out by, inter alia, some USFS and Interior’s Bureau of Land Management units, Oregon State University, citizen scientists [Sniezko] and the Oregon Department of Forestry [press release & Sniezko pers. comm.] Also, some seeds were collected in Washington State in 2020. Additional collections in Oregon are scheduled for 2022.

The collected seeds have been evaluated for vitality and stored by the USFS Dorena Center and at the USFS National Seed Lab (Macon, GA).

Oregon ash planting at Dorena; photo by Emily Boes

The USFS Dorena Center and Washington State University have begun germinating and growing some of the seedlings for various tests of possible resistance. There is concern that the 2021 drought might have killed some of the seedlings in Oregon; those in Washington are not affected. The initial seedlings are mostly from Oregon but there is space to add additional families from a wider geographical area. Experimenters plan to collect data annually on bud break, yearly growth, and any diseases or pests that develop on the trees. (Chastagner pers. comm.)

The next step is systematic testing whether some of the ash show genetic resistance to EAB. Richard Sneizko has sent seedlings of 17 ash families to USFS colleague Dr. Jennifer Koch. She operates a breeding facility in northern Ohio where they can be tested for resistance. Testing is expected to begin this year. [Tree Line]

The Dorena Center is also helping a researcher at Penn State University, Dr. Jill Hamilton, to set up a landscape genomics project. She will evaluate the genetic variability in the species by using leaf samples from about 20 trees from many populations across the Oregon ash’s range (California to British Columbia).  This potentially includes a collection from the Dorena population of ash in late Spring 2022. [Sniezko]

These various ash plantings can also be “sentinel” plantings to assist in early detection of newly arriving EAB. [Tree Line]

SOURCES

Bush J. Executive Coordinator | Washington Invasive Species Council

ODF and ODA Emerald Ash Borer Readiness and Response Plan. 2018. 

Click to access EAB+Plan+2018.pdf

ODF press release Feb 24, 2022

Treeline Newsletter May 13, 2021. Richard Sniezko. Is There a Future for Oregon Ash? Forest Genetics to the Rescue? Genetic & Emerald Ash Borer Resistance Projects https://www.nnrg.org/wp-content/uploads/2022/02/Treeline_newsletter-June-2021.pdf

The newsletter is issued by Bonneville Environmental Foundation for a consortium of conservation agencies

Sniezko pers comm Feb 2022  22-2/24

A video explaining the campaign to save Oregon ash is at https://youtu.be/uZmfLrxEA7g or https://youtu.be/S8y-XK285S8

Posted by Faith Campbell

We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.

For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm

In Michigan: Devastating News for Black Ash; Merely Bad News for Green Ash

results of EAB infestation; photo by Nate Siegert, USFS

A series of studies by Patrick J. Engelken, M. Eric Benbow, Deborah G. McCullough, Nate Siegert, Randall Kolka, Melissa Youngquist and others examine the status of ash (Fraxinus spp.) in the aftermath of the emerald ash borer (EAB) invasion. Initial studies documented the crash of biomass supporting EAB numbers when the large ash trees died (Siegert, Engelken, McCullough. 2021; full citation at end of blog.) More recent studies have focused on bogs and forests in the riparian areas where ash were especially numerous and arguably ecologically most important. I posted a blog about black ash bogs earlier.

I will focus here on the studies in riparian areas of southern and northwest Michigan. Information about impacts in forests of southern Michigan are from Engelken, Benbow and McCullough (2020); information about impacts in northwest Michigan are from Engelken and McCullough (2020). Full citations for both are at the end of the blog.

All study areas had high ash densities before EAB’s arrival. One study (Engelken and McCullough 2020) found ash densities high in the immediate riparian areas (in one case, a strip reaching 100 meters from the streambank) but scattered in surrounding forests.

In all these study areas, populations of mature (reproductive age) ash crashed within 10-15 years after EAB invasion:

  • In northeast Michigan, EAB carrying capacity was reduced by 94% – 99%; total ash basal area was reduced by 87 – 97.7% (Siegert, Engelken, McCullough. 2021);
  • In southern Michigan, more than 85% of the basal area of green (F. pennsylvanica) and black ash (F. nigra) had been killed by 2020. An estimated 96% of the overstory ash phloem area had died, thus radically reducing EAB carrying capacity (Engelken, Benbow and McCullough 2020);
  • In northwest Michigan, more than 95% of the overstory ash have been killed. (Engelken and McCullough 2020).

The worst impact has been on black ash– which plays such an important ecological role in riparian areas and wetlands and has enormous importance in Native American cultures. In all these study areas, there is no stump sprouting by black ash (Siegert, Engelken, McCullough. 2021; Engelken, Benbow and McCullough 2020; Engelken and McCullough 2020). In three watersheds of northwest Michigan where black ash constituted up to a quarter of the overstory species before the EAB invasion, scientists found no black ash recruits, only eight saplings, and a single seedling.

Green ash (F. pennsylvanica) has survived in much higher numbers – so far. However, this species’ ability to grow into reproductive size is still uncertain. In northwest Michigan, green ash saplings are abundant in canopy gaps created by EAB-caused mortality of mature ash. These saplings had established before the EAB invasion so some call them the “orphaned cohort”.  However, there are few seedlings of any woody plant species in these gaps because sedges form such dense mats.

Green ash reproduction faces many challenges before persistence of the species can be considered assured.  First, populations of EAB – now reduced by the lack of mature ash to support them – might resurge when young ash grow to larger sizes. It is not yet clear the extent to which introduced biocontrol agents and native predators, e.g., woodpeckers, will protect these trees as they grow to reproductive size. Here, again, green ash has an advantage over black ash. While green ash produce seed at a relatively young age, black ash don’t produce seed until they reach 30–40 years. Even then, they produce seeds only sporadically, with intervals of five or more years.

A second challenge is the lack of seed sources – at least until and unless young trees are able to reach reproductive size.

A third challenge is competition for resources from other plants. The canopy gaps eliminate competition for light for the taller plants, i.e., the existing ash saplings. However, the sapling cohort is not supported by a seedling cohort. There are very few seedlings of all woody plant species (including invasive species!). Seed germination is suppressed by the dense mats of wetland-adapted sedges and possibly the higher water tables (which resulted from reduced evapotranspiration following mortality of the mature trees).

Competition for resources is also a factor in the forests outside the immediate riparian zone. There, ash seedlings sprout, but shade created by lateral ingrowth suppresses their growth. In southern Michigan, Engelken, Benbow and McCullough (2020) note that the forests are apparently transitioning from red oak dominated forests to red maple and black cherry dominated forests. This transition is apparently intensified by forest mesophication resulting from reduced fire frequency, decreasing light availability in forest understories and increasing soil moisture content.

Fourth, while stump sprouting of green ash was noted in southern Michigan, in the northwestern forests all the sprouts died. I have already noted the absence of stump sprouting by black ash at all sites.

Beaver & Green Ash in Northern Virginia

photos of beaver feeding on ash saplings in northern Virginia; photos by F.T. Campbell

In spring 2022 I noticed along one stream in northern Virginia that beavers had cut down green ash saplings; McCullough and Siegert report that this does not appear to be a problem in their study areas.

By December 2022, the beaver-cut trees tried to recover: see the sprouts from a stump [below]. (I think deer or rabbits ate the tips of the sprouts.)

The beavers also continued feeding on the ash — the tree photographed in the spring when it was half-chewed through has now been felled and its branches removed [see below].

Ecosystem Impacts, Especially on Streams

Across much of the upper Midwest, massive ash mortality is causing widespread changes in forest systems.

Riparian forests, i.e., areas adjacent to waterways where periodic inundation occurs, are functionally linked to the aquatic systems. Loss of such a significant proportion of the overstory changes the transfer of energy to adjacent waterways that takes the form of inputs of nutrients from leaf litter and coarse woody debris. Intact forests also stabilize stream banks and maintain channel depth by preventing erosion. Forests moderate temperature of the water. Finally, forests with “coarse woody debris” increase habitat structure. These impacts might be especially important along first order streams, (defined as perennial streams that have no permanently flowing tributaries). These streams are too small to buffer the impacts of major tree loss. The scientists say they are uncertain whether these changes continue to affect larger streams downstream.

Unshaded streams have higher water temperatures that can affect populations of fish, in particular salmonids, by delaying migration, reducing egg viability and increasing egg mortality. Higher temperatures can also alter primary productivity of aquatic algae, potentially increasing eutrophication (Engelken and McCullough 2020).

The scientists expect increasing abundance of coarse woody debris in the forests and streams of northwest Michigan as the 75% of dead ash that are still standing fall. Such debris provides nutrients and habitat for an array of plants and animals, thereby influencing the abundance, activity and species compositions of several ground dwelling insects and seedling establishment. In streams, coarse woody debris provides complex habitat and refuges. It also retains organic matter. Recreationists do find that debris impedes boating.

Loss of ash specifically 

As described by Engelken, Benbow and McCullough (2020), and in my earlier blog, ash leaf litter – particularly black ash leaf litter – is highly nutritious. Ash leaf litter has efficient turnover rates and contributes important soil nutrients such as nitrogen, organic carbon and exchangeable cations. Invertebrate communities in headwater streams feed largely on coarse organic material such as leaf litter (Engelken and McCullough 2020). Consequently, loss of the annual influx of ash leaf litter will likely have adverse effects on nutrient availability in riparian forests and adjacent streams.

SOURCES

Engelken, P.J., M.E. Benbow, D.G. McCullough. 2020. Legacy effects of emerald ash borer on riparian forest vegetation and structure.  Forest Ecology and Management 457 (2020) 117684

Engelken, P.J. and D.G. McCullough. 2020. Riparian Forest Conditions Along Three Northern Michigan Rivers Following Emerald Ash Borer Invasion. Canadian Journal of Forest Research.

Siegert, N.W., P.J. Engelken, D.G. McCullough. 2021 Changes in demography and carrying capacity of green ash and black ash ten years after emerald ash borer invasion of two ash-dominant forests. Forest Ecology and Management Vol 494, August 2021

Posted by Faith Campbell

We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.

For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm

What Do Invasive Species Cost?

brown tree snake Boiga irregularis; via Wikimedia; one of the species on which the most money is spent on preventive efforts

In recent years a group of scientists have attempted to determine how much invasive species are costing worldwide. See Daigne et al. 2020 here.

Some of these scientists have now gone further in evaluating these data. Cuthbert et al. (2022) [full citation at end of blog] see management of steadily increasing numbers of invasive, alien species as a major societal challenge for the 21st Century. They undertook their study of invasive species-related costs and expenditures because rising numbers and impacts of bioinvasions are placing growing pressure on the management of ecological and economic systems and they expect this burden to continue to rise (citing Seebens et al., 2021; full citation at end of blog).

They relied on a database of economic costs (InvaCost; see “methods” section of Cuthbert et al.) It is the best there is but Cuthbert et al. note several gaps:

  • Only 83 countries reported management costs; of those, only 24 reported costs specifically associated with pre-invasion (prevention) efforts.
  • Data comparing regional costs do not incorporate consideration of varying purchasing power of the reporting countries’ currencies.  
  • Data available are patchy so global management costs are probably substantially underestimated. For example, forest insects and pathogens account for less than 1% of the records in the InvaCost database, but constitute 25% of total annual costs ($43.4 billion) (Williams et al., in prep.) .

Still, their findings fit widespread expectations.  

These data point to a total cost associated with invasive species – including both realized damage and management costs – of about $1.5 trillion since 1960.  North America and Oceania spent by far the greatest amount of all global money countering bioinvasions. North America spent 54% of the total expenditure of $95.3 billion; Oceania spent 30%. The remaining regions each spent less than $5 billion.

Cuthbert et al. set out to compare management expenditures to losses/damage; to compare management expenditures pre-invasion (prevention) to post-invasion (control); and to determine potential savings if management had been more timely.

Economic Data Show Global Efforts Could Be – But Aren’t — Cost-Effective

The authors conclude that countries are making insufficient investments in invasive species management — particularly preventive management. This failure is demonstrated by the fact thatreported management expenditures ($95.3 billion) are only 8% of total damage costs from invasions ($1.13 trillion). While both cost or losses and management expenditures have risen over time, even in recent decades, losses were more than ten times larger than reported management expenditures. This discrepancy was true across all regions except the Antarctic-Subantarctic. The discrepancy was especially noteworthy in Asia, where damages were 77-times higher than management expenditures.

Furthermore, only a tiny fraction of overall management spending goes to prevention. Of the $95.3 billion in total spending on management, only $2.8 billion – less than 3%  – has been spent on pre-invasion management. Again, this pattern is true for all geographic regions except the Antarctic-Subantarctic. The divergence is greatest in Africa, where post-introduction control is funded at more than 1400 times preventive efforts. It is also significant for Asia and South America.

Even in North America – where preventative actions were most generously funded – post-introduction management is funded at 16 times that of prevention.

Cuthbert et al. worry particularly about the low level of funding for prevention in the Global South. They note that these conservation managers operate under severe budgetary constraints. At least some of the bioinvasion-caused losses suffered by resources under their stewardship could have been avoided if the invaders’ introduction and establishment had been successfully prevented.

While in the body of the article Cuthbert et al. seem uncertain about why funding for preventive actions is so low, in their conclusions they offer a convincing (to me) explanation. They note that people are intrinsically inclined to react when impact becomes apparent. It is therefore difficult to motivate proactive investment when impacts are seemingly absent in the short-term, incurred by other sectors, or in different regions, and when other demands on limited funds may seem more pressing. Plus efficient proactive management will prevent any impact, paradoxically undermining evidence of the value of this action!

Aedes aegypti mosquito; one of the species on which the most money is spent for post-introduction control; photo by James Gathany; via Flickr

Delay Costs Money

The reports contained in the InvaCost database indicate that management is delayed an average of 11 years after damage was first been reported. Cuthbert et al. estimate that these delays have caused an additional cost of about $1.2 trillion worldwide. Each $1 of management was estimated to reduce damage by $53.5 in this study. This finding, they argue, supports the value of timely invasive species management.

They point out that the Supplementary Materials contain many examples of bioinvasions that entail large and sustained late-stage expenditures that would have been avoided had management interventions begun earlier.

Although Cuthbert et al. are not as clear as I would wish, they seem to recognize also that stakeholders’ varying perceptions of whether an introduced species is causing a detrimental “impact” might also complicate reporting – not just whether any management action is taken

Cuthbert et al. are encouraged by two recent trends: growing investments in preventative actions and research, and shrinking delays in initiating management. However, these hopeful trends are unequal among the geographic regions.

Which Taxonomic Groups Get the Most Money?

About 42% of management costs ($39.9 billion) were spent on diverse or unspecified taxonomic groups. Of the costs that were taxonomically defined, 58% ($32.1 billion) was spent on invertebrates [see above re: forest pests]; 27% ($14.8 billion) on plants; 12% ($6.7 billion) on vertebrates; and 3% ($1.8 billion) on “other” taxa, i.e. fungi, chromists, and pathogens. For all of these defined taxonomic groups, post-invasion management dominated over pre-invasion management.  

When considering the invaded habitats, 69% of overall management spending was on terrestrial species ($66.1 billion); 7% on semi-aquatic species ($6.7 billion); 2% on aquatic species ($2.0 billion); the remainder was “diverse/unspecified”. For pre-invasion management (prevention programs), terrestrial species were still highest ($840.4 million). However, a relatively large share of investments was allocated to aquatic invaders ($624.2 million).

Considering costs attributed to individual species, the top 10 targetted for preventive efforts were four insects, three mammals, two reptiles, and one alga. Top expenditures for post-invasion investments went to eight insects [including Asian longhorned beetle], one mammal, and one bird.

Asian longhorned beetle

Just two of the costliest species were in both categories: insects red imported fire ant(Solenopsis invicta) and Mediterranean fruitfly (Ceratitis capitate). None of the species with the highest pre-invasion investment was among the top 10 costliest invaders in terms of damages. However, note the lack of data on fungi, chromists, and pathogens. (I wrote about this problem in an earlier blog.)

Discussion and Recommendations

Cuthbert et al. conclude that damage costs and post-invasion spending are probably growing substantially faster than pre-invasion investment. Therefore, they call for a stronger commitment to enhancing biosecurity and for more reliance on regional efforts rather than ones by individual countries. Their examples of opportunities come from Europe.

Drawing parallels to climate action, the authors also call for greater emphasis on during decision-making to act collectively and proactively to solve a growing global and inter-generational problem.

Cuthbert et al. focus many of their recommendations on improving reporting. One point I found particularly interesting: given the uneven and rapidly changing nature of invasive species data, they think it likely that future invasions could involve a new suite of geographic origins, pathways or vectors, taxonomic groups, and habitats. These could require different management approaches than those in use today.

As regards data and reporting, Cuthbert et al. recommend:

1) reducing bias in cost data by increasing funding for reporting of underreported taxa and regions;

2) addressing ambiguities in data by adopting a harmonized framework for reporting expenditures. For example, agriculture and public health officials refer to “pest species” without differentiating introduced from native species. (An earlier blog also discussed the challenge arising from  these fields’ different purposes and cultures.)

3) urging colleagues to try harder to collect and integrate cost information, especially across sectors;

4) urging countries to report separately costs and expenditures associated with different categories, i.e., prevention separately from post-invasion management; damage separately from management efforts; and.

5) creating a formal repository for information about the efficacy of management expenditures.

While the InvaCost database is incomplete (a result of poor accounting by the countries, not lack of effort by the compilers!), analysis of these data points to some obvious ways to improve global efforts to contain bioinvasion. I hope countries will adjust their efforts based on these findings.

SOURCE

Cuthbert, R.N., C. Diagne, E.J. Hudgins, A. Turbelin, D.A. Ahmed, C. Albert, T.W. Bodey, E. Briski, F. Essl, P. J. Haubrock, R.E. Gozlan, N. Kirichenko, M. Kourantidou, A.M. Kramer, F. Courchamp. 2022. Bioinvasion costs reveal insufficient proactive management worldwide. Science of The Total Environment Volume 819, 1 May 2022, 153404

Seebens, H. S. Bacher, T.M. Blackburn, C. Capinha, W. Dawson, S. Dullinger, P. Genovesi, P.E. Hulme, M.van Kleunen, I. Kühn, J.M. Jeschke, B. Lenzner, A.M. Liebhold, Z. Pattison, J. Perg, P. Pyšek, M. Winter, F. Essl. 2021. Projecting the continental accumulation of alien species through to 2050. Glob Change Biol. 2021;27:970-982.

Williams, G.M., M.D. Ginzel, Z. Ma, D.C. Adams, F.T. Campbell, G.M. Lovett, M. Belén Pildain, K.F. Raffa, K.J.K. Gandhi, A. Santini, R.A. Sniezko, M.J. Wingfield, and P. Bonello 2022. The Global Forest Health Crisis: A Public Good Social Dilemma in Need of International Collective Action. submitted

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

Global Loss of Floristic Uniqueness

Hakalau Forest, Hawai“i; nearly 90% of Hawaiian flora is unique to the Islands

A recent article by Yang et al. 2021 (full citation at the end of this blog) seeks to determine the extent to which introduced plants reduce the uniqueness of regional floras. They analyzed data from 658 regions covering about 65.7% of the Earth’s ice-free land surface and about 62.3% of the planet’s known plant species.

They found strong homogenization of plant species’ taxonomic and phylogenetic diversity results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.

There are two aspects of a region’s floral uniqueness. One is the number of species that it shares with other regions. This is taxonomic uniqueness. The other is the distinctiveness of the evolutionary history of the region. When several species are endemic to a region’s flora, and lack close relatives in other regions, that equals phylogenetic uniqueness.

The effect of a species introduction differs depending on which of these aspects one focuses on. Thus, naturalization of a species closely related to native species (e.g., a congeneric species) will have less impact on the phylogenetic floristic uniqueness of the region than naturalization by a distantly related species. Taxonomic uniqueness, however, will be affected to the same degree, irrespective of the phylogenetic distance between the naturalized and native species.

Yang et al. found strong homogenization of plant diversity. They found that species introductions increased the taxonomic similarity in 90.7% of all regional pairs and phylogenetic similarity in 77.2% of all region pairs. Most homogenization results from introductions of plant species to ecosystems beyond their native range. Homogenization caused by regional extinctions of native floral species occurs much less frequently.

This loss of regional biotic uniqueness or distinctiveness changes biotic interactions and species assemblages. These, in turn, have ecological and evolutionary consequences at larger scales and higher levels.

The degree of homogenization between regions’ floras depends on three factors:

1) The distance between the donor and recipient regions. Since nearby regions share more species, an introduction from a more distant origin is more likely to be a novel species and so contribute to homogenization of “donor” and “receiving” floras.

2) Climatic similarity, especially temperature. A plant species introduced from a climatically similar but geographically distant place is more likely to establish than a species from a different climatic zone. As a result, the recipient area’s flora is changed to more closely resemble the flora of the donor region with which it shares climatic conditions – regardless of the distance between them.

3) The level of exchange of goods and people between two regions. The higher the rate of exchange between two regions, the greater the chance that a species will be introduced and become established. Yang et al. used the existence of current or past administrative relationships (e.g., colonial relationship) between two regions as a proxy for intensity of trade and transport between donor and recipient regions. They found that floras of regions with current or past administrative links have taxonomically become more similar to each other than the floras of regions with no such links.

flora of the Cape Floral Kingdom – South Africa; photo from Michael Wingfield

Establishment of introduced species can increase floristic similarity of the donor and recipient regions (= floristic homogenization) when the species is native to one of the two regions and naturalizes in the other, or when it is not native to both regions and naturalizes in both. On the other hand, a species introduction can decrease the floristic similarity of the two regions (i.e., enhance floristic differentiation) when the species is not native to both regions but naturalized in only one.  

Homogenization hotspots differed slightly depending on whether one focused on taxonomic or phylogenetic aspects.

The regions with the greatest average increase in taxonomic similarity with other regions due to naturalized alien species were New Zealand, portions of Australia, and many oceanic islands. The Australasian situation probably reflects its long biogeographic isolation from other parts of the globe and its highly unique native flora. As a result, nearly all non-native plants introduced to Australasia strongly increase levels of its floristic similarity to the rest of the world. Oceanic islands have species-poor floras with large proportions of unique endemics. They have also received high numbers of naturalized alien plants.

Hotspots of phylogenetic homogenization on continents are the same as those for taxonomic homogenization, but this is not true for islands. Yang et al. think this is because islands’ native floras were established by natural colonization from nearby continental floras so – despite subsequent speciation – they retain their phylogenetic relationship to the donor areas’ floras.  

Yang et al. concede that they lacked high-quality data on native and naturalized alien species lists for a third of Earth’s ice-free terrestrial surface, especially Africa, Eastern Europe, and tropical Asia. They believe, however, that data from these regions are unlikely to change the overall finding.  (Scientists are beginning to compile lists of forest pests in Africa). link to blog

Yang et al. note that introduction and naturalization of alien species are likely to increase in the future, thusaccelerating floristic homogenization. The ecological, evolutionary and socioeconomic consequences are largely unknown.They call for stronger biosecurity regulations of trade and transport and other measures to protect native vegetation.

SOURCE

Yang, Q., P. Weigelt, T.S. Fristoe, Z. Zhang, H. Kreft, A. Stein, H. Seebens, W. Dawson, F. Essl, C. König, B. Lenzner, J. Pergl, R. Pouteau, P. Pyšek, M. Winter, A.L. Ebel, N. Fuentes, E.L.H. Giehl, J. Kartesz, P. Krestov, T. Kukk, M. Nishino, A. Kupriyanov, J.L. Villaseñor, J.J. Wieringa, A. Zeddam, E. Zykova  and M. van Kleunen. 2021. The global loss of floristic uniqueness. NATURE COMMUNICATIONS (2021) 12:7290. https://doi.org/10.1038/s41467-021-27603-y

Posted by Faith Campbell

We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.

For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm