drivers of extinction – Page 13 – Center for Invasive Species Prevention

Plant Invasions’ Impacts on Wildlife

spotted knapweed (*Centaurea maculosa*); photo by Alan Vernon via Wikipedia

Litt and Pearson (full citation at the end of the blog) are trying to improve scientists’ ability to predict the impact of biological invasions. Their goal is to predict which organisms will be winners, which losers, in the face of anthropogenic ecosystem change.

They focus on exotic plant invasions, because they are ubiquitous. Furthermore, plant invasions affect ecosystems by reassembling the plant community in ways that affect the niches used by native animals and hence the animals’ success under the new conditions. After determining the differences between the traits exhibited by invasive plants vs. the native plants they are displacing, scientists can then identify which native animals are most likely to be affected, as well as how and why they might respond to exotic plant invasion. [Note that Doug Tallamy is looking at similar issues.]

Litt and Pearson have developed a framework to assess how plants’ traits might affect associated wildlife. Applying the framework requires certain baseline information about the ecosystem in question.

This knowledge is applied in stepwise fashion:

1) Identify the fauna of interest and their linkage to the native plant community. This association might be food or habitat values such as shelter. Then the researcher determines the relevant plant traits of importance to that animal and approximates the strength of the animal’s dependence on these traits. Note that the focus is on plant traits relevant to the animal users, rather than specific plant species.

2) Determine overall importance of the plant traits for the area under study by (a) averaging dependence of a representative subsample of individuals to obtain a community-level value for each plant species or functional group and (b) quantifying the relative abundance of the plant functional group in the community (e.g., cover or biomass).

3) Plot the way the animal species’ abundance changes with resource abundance.

4) Understand how the invasive plants will alter the distributions of the native plants’ traits and potentially introducing novel traits that might alter the faunal community.

Litt and Pearson reviewed earlier studies to test how well this framework explained the responses of three groups of fauna to plant invasions in different ecosystems.

searching for spotted knapweed; photo by Oregon Department of Agriculture

Spiders in invaded grasslands

Intermountain grasslands of western Montana are heavily invaded; non-native plants already comprise 25–60% of average total plant cover.

One group of native spiders construct their irregular webs entirely within a single plant. A second group – orb weavers – suspend their larger webs from multiple plants. The former depend on the architectural complexity of individual plants; they can build larger webs in plant species possessing greater branching and/or longer branches of the flowering stalks. Orb spiders depend more on the complexity of the overall plant community.

Plant architecture is closely tied to the plant’s functional groups, that is, whether they are grasses or forbs.

These grasslands are generally dominated by perennial grasses. The irregular-web spiders can use grasses, but strongly favor forbs, particularly those with the most complex flowering structures. Orb weavers are generalists, incorporating multiple plant species; but they also tend to favor forbs, presumably because they are more robust.

Invasive plants in the Western Montana grasslands are of two types: an annual grass, cheatgrass (Bromus tectorum), and numerous perennial and annual forbs. Cheatgrass largely replaces the dominant native grasses with a similar architecture – although cheat is shorter. The exotic forbs, which can collectively invade at levels comparable to cheatgrass, tend to be taller and more complex structurally than the native forbs. Thus, invasion by exotic forbs strongly shifts the community-level distribution of the key trait toward greater structural complexity by replacing the dominant, but structurally simplistic, native grasses, and the more diminutive native forbs. These changes increased the abundance of both spider groups, but especially the specialist irregular web weavers. They find the new conditions meet their needs. Both spider groups appeared to expand their realized niches in response to invasion, i.e., they are able to use a broader range of plant architectures than was available in the native system.

Rodents in semi-desert grasslands invaded by Lehmann lovegrass

In the semi-desert grasslands of the American southwest, native grasses and forbs provide food and habitat for a variety of rodents. This vegetation influences which species of rodents are present in two ways: the size of the plants’ seeds and the density of vegetative cover. Litt and Steidl examined both. They divided the rodents into separate guilds based on diet and preferred vegetative cover. The two sets of guilds did not overlap for all species.

In southern Arizona, the native plant community is dominated by several grass species and herbaceous forbs; most species produce relatively large seeds. Vegetative cover is generally low, but varies in a patchy fashion. The rodent communities in uninvaded native grasslands are dominated by seed-eaters that prefer sparse cover.

Invasion of these grasslands by Lehmann lovegrass (Eragrostis lehmanniana) results in increased vegetative cover but the grass produces very small seeds that probably provide little to no food for rodents. Another result is a decrease in overall abundance of arthropods. The new conditions favor different rodent species from those most common in uninvaded habitat.

Two more specialized seed-eating rodent species, which seek both lower cover and larger seeds, decreased in abundance. A rodent species which favors lower vegetative cover and feeds on larger invertebrates also declined. In contrast, abundance increased for two other rodent species that prefer more dense cover and are more opportunistic in their feeding. One species surprised the scientists: Dipodomys merriami increased in abundance, despite the fact that this species favors more open environments. Perhaps other functional traits or biotic interactions are important to this species? There was no apparent change in abundance for three other species, suggesting either a lack of statistical power (2 were less abundant) or that these rodents were able to persist through a balance of positive and negative changes in food and habitat characteristics.

Lucy’s warbler [nest in saguaro, not cottonwood); photo by Dominic Sherony

Warblers in Riparian Habitats in the Southwest

Riparian habitats in the same desert region have been aggressively invaded by the exotic shrub saltcedar (Tamarix spp.). Litt and Pearson consider the findings of Mahoney et al. of this invasion’s impact on two ecologically similar warbler species. One, the yellow warbler (Setophaga petechia), is very widely distributed across North America; it is considered a generalist. The other, Lucy’s warbler (Oreothlypis luciae), is endemic to a small region of the southwest United States and northern Mexico.

The two species have similar feeding behaviors but differ in their nesting requirements. The yellow warbler constructs open cup nests in the branches of shrubs and trees. Lucy’s warbler nests in cavities in larger trees excavated by others. Hence, these species were expected to respond similarly to changes in food resources and foraging habitat, but differ in their responses to changes in nesting substrate.

Native vegetation in the region consists primarily of willows and cottonwoods in the riparian corridors, with oak and mesquite woodlands in the adjacent uplands. Saltcedar invasion rapidly displaces the willows; it takes much longer to displace cottonwoods since are large and long-lived. Upland vegetation is uninvaded and unaffected. While saltcedar is structurally similar to native willows, its leaf architecture allows more light to penetrate in saltcedar stands. This can exacerbate heat stress on nestlings in these hot, arid environments, as well as expose the nestlings to nest predation. These effects are exacerbated by the presence of a biocontrol leaf beetle (Diorhabda spp.), which cause widespread defoliation of saltcedar during nesting season. Meantime, the cavity nests used by Lucy’s warbler are barely affected.

The study by Mahoney et al. showed that in low-invasion riparian sites, the two warblers occur at comparable abundances. When saltcedar invasion replaces willows, yellow warblers decline by ~50% while there is no apparent change in abundance of Lucy’s warblers.

Litt and Pearson point out that their framework is based on two key assumptions that establish the context for its efficacy.

The first is that bottom-up forces fuel ecological processes. Plants are key to making the sun’s energy available to consumer animals and – thence to predators. Consumers’ and predators’ top-down effects are secondary. The authors’ framework thus provides better predictions of community outcomes when systems are predominantly structured by bottom-up forces. As top-down forces increase or when invasive plants differentially affect multiple dimensions of the consumer niche space, it will be more challenging to track and predict outcomes, as our rodent example demonstrates.

The second assumption is that exotic plant invasions will most strongly influence bottom-up processes. Invasive plants displace native plants and their plant traits, thus directly affecting consumers by altering the quality and quantity of food and habitat resources. However, plant community changes caused by plant invasions can also affect predators directly and indirectly via several interactions. These changes in predators’ abundance and/or their per capita effects on prey might create feedbacks that can complicate interpreting and predicting invasion outcomes.

Litt and Pearson concluded that their approach is promising but has inherent limitations linked to the dynamic nature of ecological systems.

[Ecologists continue to evaluate the impacts of saltcedar eradication efforts on another bird species, the federally endangered southwestern willow flycatcher (Empidonax extimus trailii). See, for example, Goetz, A., I. Moffit and A.A. Sher. 2022. Recovery of a native tree following removal of an invasive competitor with implications for endangered bird habitat. Biological Invasions Vol. 24, pp. 2769-2793.]

SOURCE

Litt, A.R. and D.E. Pearson. 2022. A functional ecology framework for understanding and predicting animal responses to plant invasion. Biol Invasions https://doi.org/10.1007/s10530-022-02813-7

& Supporting Information [warblers in riparian ecosystems invaded by tamarisk]

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

www.fadingforests.org

Tree Planting – Warning from New Zealand

*Pinus radiata* plantation in New Zealand; photo by Jon Sullivan

As countries and conservation organizations ramp up tree planting as one solution to climate change, I worry that many of the plantings will use species not native to the region – with the risk of promoting more bioinvasions. My second fear is that inadequate attention will be paid to ensuring that the propagules thrive.

Warning from New Zealand

New Zealand has adopted a major afforestation initiative (“One Billion Trees”). This program is ostensibly governed by a policy of “right tree, right place, right purpose”. However, Bellingham et al. (2022) [full citation at end of blog] say the program will probably increase the already extensive area of radiata pine plantations and thus the likelihood of exacerbated invasion. They say the species’ potential invasiveness and its effects in natural ecosystems have not been considered.

Bellingham et al. set out to raise the alarm by evaluating the current status of radiata, or Monterrey, pine (Pinus radiata) in the country. They note that the species already occupies ~1.6 M ha; the species makes up 90% of the country’s planted forests. Despite the species having been detected as spreading outside plantations in 1904, it is generally thought not to have invaded widely.

The authors contend that, to the contrary, radiata pine has already invaded several grasslands and shrublands, including three classes of ecosystems that are naturally uncommon. These are geothermal ecosystems, gumlands (infertile soils that formerly supported forests dominated by the endemic and threatened kauri tree Agathis australis), and inland cliffs. Invasions by pines – including radiata pine – are also affecting primary succession on volcanic substrates, landslides on New Zealand’s steep, erosion-prone terrain, and coastal sand dunes. Finally, pine invasions are overtopping native Myrtaceae shrubs during secondary succession. Bellingham et al. describe the situation as a pervasive and ongoing invasion resulting primarily from spread from plantations to relatively nearby areas.

kauri; photo by Natalia Volna, iTravelNZ

The New Zealanders cite data from South America and South Africa on the damaging effects of invasions by various pine species, especially with respect to fire regimes.

Furthermore, their modelling indicates that up to 76% of New Zealand’s land area is climatically capable of supporting radiata pine — most of the country except areas above 1000 m in elevation or receiving more than 2000 mm of rainfall per year. That is, all but the center and west of the South Island. This model is based on current climate; a warmer/drier climate would probably increase the area suitable to radiata pine.

These invasions by radiata pine have probably been overlooked because the focus has been on montane grasslands (which are invaded by other species of North American conifers). [See below — surveys of knowledge of invasive plants’ impacts.]

Bellingham et al. recognize the economic importance of radiata pine. They believe that early detection of spread from plantations and rapid deployment of containment programs would be the most effective management strategy. They therefore recommend

1) taxing new plantations of non-indigenous conifers to offset the costs of managing invasions, and

2) regulating these plantations more strictly to protect vulnerable ecosystems.

They also note several areas where additional research on the species’ invasiveness, dispersal, and impacts is needed.

Survey of Awareness of Invasive Plants

A few months later a separate group of New Zealand scientists published a study examining tourists’ understanding of invasive plant impacts and willingness to support eradication programs (Lovelock et al.; full citation at end of the blog). One of the invasive plant groups included in the study are conifers introduced from North America and Europe. These conifers are invading montane grasslands, so they are not the specific topic of the earlier article. The other is a beautiful flowering plant, Russell lupine. These authors say that both plant groups have profound ecological, economic, and environmental impacts. However, the conifers and lupines are also highly visible at places valued by tourists. Lovelock et al. explored whether the plants’ familiarity – and beauty – might affect how people reacted to descriptions of their ecosystem impacts.

Visitors from elsewhere in New Zealand were more aware of invasive plants’ impacts and more willing to support eradication programs for these species specifically. Asian visitors had lower awareness and willingness to support eradication of the invasives than tourists from the United Kingdom, Europe, or North America. This pattern remained after the tourists were informed about the plants’ ecological impacts. All groups were less willing to support eradication of the attractive Russell lupine than the conifers.

Conifers invading montane grasslands are perhaps the most publicized invasive plants in New Zealand [as noted above]. Lovelock et al. report that New Zealand authorities have spent an estimated $NZ166 million to eradicate non-native conifers over large tracts of land on the South Island. Still, only about half the New Zealand visitors surveyed were aware of the ecological problems caused by wild conifers.

Russell lupine (Lupinus × russellii) is invading braided river systems, modifying river flows, reducing nesting site availability for several endangered birds, and provides cover for invasive predators. While initially planted in gardens, the lupines were soon being deliberately spread along the roads to ‘beautify’ the landscape. Foreign tourists often specifically seek river valley invaded by the lupine because pictures of the floral display appear in both official tourism promotional material & tourist-related social media. It is not surprising, then, that even among New Zealanders, only a third were aware of the lupines’ environmental impacts.

The oldest participants (those over 60) had the lowest acceptance of wild conifers. Participants 50–59 years old were most aware of ecological problems caused by wild conifers. Participants 30–39 years old showed the highest acceptance of wild conifers and lowest awareness of ecological issues.

Female participants showed a higher preference for the landscape with wild conifers (45.90%) than males (36.89%). Female participants were also half as aware of ecological problems (25.62% v. 46.12% among male participants).

Nearly all survey participants (96.1%) preferred the landscape with flowering lupine; only 19.4% were aware of associated ecological problems. New Zealand domestic visitors were more aware. After the impacts of lupines were explained, half decided to support eradication. However, the same proportion of all survey participants (42.5%) still wanted to see lupines in the landscape.

Once again, participants older than 50 were more aware of ecological problems arising from lupine invasions. Both men and women greatly preferred the landscape with Russell lupins.

While the authors do not explore the ramifications of the finding that younger people are less aware of invasive species impacts, I think they bode ill for future protection of the country’s unique flora and fauna. They did note that respondents had a high level of acceptance overall for these species on the New Zealand landscapes.

While the study supported use of simple environmental messaging to influence attitudes about invasive species, also showed that need to consider such social attributes as nationality and ethnicity. So Lovelock et al. call for investigation of how and why place of origin and ethnicity are important in shaping attitudes towards invasives. Conveying conservation messages will be more difficult because tourist materials often contain photographs of the lupines. Much of this information comes from informal media such as social media, which are beyond the control of invasive species managers.

SOURCES

Bellingham, P.J., E.A. Arnst, B.D. Clarkson, T.R. Etherington, L.J. Forester, W.B. Shaw, R. Sprague, S.K. Wiser, and D.A. Peltzer. 2022. The right tree in the right place? A major economic tree species poses major ecological threats. Biol Invasions Vol.: (0123456789) https://doi.org/10.1007/s10530-022-02892-6

Lovelock B., Y. Ji, A. Carr, and C-J. Blye. 2022. Should tourists care more about invasive species? International and domestic visitors’ perceptions of invasive plants and their control in New Zealand. Biological Invasions (2022) 24:3905–3918 https://doi.org/10.1007/s10530-022-02890-8

Posted by Faith Campbell

www.fadingforests.org

Invasive Species Costs Point to Inadequate Effort – especially Prevention

EAB-killed ash tree falls before it can be taken down; photo courtesy of former Ann Arbor mayor John Hieftje

Concerned by growing impacts of bioinvasion and inadequate responses by national governments worldwide and by international bodies, a group of experts have attempted to determine how much invasive species are costing. They’ve built the global database – InvaCost. See Daigne et al. 2020 here.

Several studies have been based on these data. In two earlier blogs, I summarized two of these articles, e.g., Cuthbert et al. on bioinvasion costs, generally, and Moodley et al. on invasive species costs in protected areas, specifically. Here, I look at two additional studies. Ahmed et al. focusses on the “worst” 100 invasives affecting conservation — as determined by the International Union of Conservation and Nature (IUCN). The second, by Turbelin et al., examines pathways of introduction. Full citations of all sources appear at the end of this blog.

It is clear from all of these papers that the authors (and I!) are frustrated by the laxity with which virtually all governments respond to bioinvasions. Thus more robust actions are needed. The authors and I also agree that data on economic costs influence political decision-makers more than ecological concerns. However, InvaCost – while the best source in existence — is not yet comprehensive enough to generate the thoroughly-documented economic data about specific aspects of bioinvasion that would be most useful in supporting proposed strategies.

Scientists working with InvaCost recognize that the data are patchy. At the top level, these data demonstrate high losses and management costs imposed by bioinvasion. The global total – including both realized damage and management costs – is estimated at about $1.5 trillion since 1960. In fact, these overall costs are probably substantially underestimates (Cathbert et al.). [For a summary of data gaps, go to the end of the blog.] Furthermore, they recognize that species imposing the highest economic costs might not cause the greatest ecological harm (Moodley et al).

citrus longhorned beetle exit hole in bonsai tree; USDA APHIS photo

Comparing estimated management costs to estimated damage, the authors conclude that countries invest too little in bioinvasion management efforts and — furthermore — that expenditures are squandered on the wrong “end” of bioinvasion – after introduction and even establishment, rather than in preventive efforts or rapid response upon initial detection of an invader. While I think this is true, these findings might be skewed by the fact that fewer than a third of countries reporting invasive species costs included data on specifically preventive actions. Cuthbert et al. notes that failing to try to prevent introductions imposes an avoidable burden on resource management agencies. Ahmed et al. developed a model they hope will overcome the perverse incentives that lead decision-makers to either do nothing or delay.

Why Decision-Makers Delay

Citing the InvaCost data, the participating experts reiterate the long-standing call for prioritizing investments at the earliest possible invasion stage. Ahmed et al. found that this was the most effective practice even when costs accrue slowly. They ask, then, why decision-makers often delay initiating management. I welcome this attention because we need to find ways to rectify this situation.

They conclude, first, that invasive species threats compete for resources with other threats to agriculture and natural systems. Second, Cuthbert et al. and Ahmed et al. both note that decision-makers find it difficult to justify expenditures before impacts are obvious and/or stakeholders demand action. By that time, of course, management of invasions are extremely difficult and expensive – if possible at all. I appreciate the wording in Ahmed et al.: bioinvasion costs can be deceitfully slow to accrue, so policy makers don’t appreciate the urgency of taking action.

Cuthbert et al. also note that impacts are often imposed on other sectors, or in different regions, than those focused on by the decision-makers. Stakeholders’ perceptions of whether an introduced species is causing a “detrimental” impact also vary. Finally, when efficient proactive management succeeds – prevents any impact – it paradoxically undermines evidence of the value of this action!

Ahmed et al. point out that in many cases, biosecurity measures and other proactive approaches are even more cost effective when several species are managed simultaneously. They cite as examples airport quarantine and interception programs; Check Clean Dry campaigns encouraging boaters to avoid moving mussels and weeds; ballast water treatment systems; and transport legislation e.g., the international standard for wood packaging (ISPM#15) [I have often discussed the weaknesses in ISPM#15 implementation; go to “wood packaging” under “Categories” (below the archive list)].

pallet “graveyard”; photo by Anand Prasad

Pathways of Species’ Introduction

Tuberlin et al. focus on pathways of introduction, which they say influence the numbers of invaders, the frequency of their arrival, and the geography of their eventual distribution. This study found sufficient data to analyze arrival pathways of 478 species – just 0.03% of the ~14,000 species in the full database. They found that intentional pathways – especially what they categorized as “Escape” – were responsible for the largest number of invasive species (>40% of total). On the other hand, the two unintentional pathways called “Stowaway” and “Contaminant” introduced the species causing the highest economic costs.

Tuberlin et al. therefore emphasize the importance of managing these unintentional pathways. Also, climate change and emerging shipping technologies will increase potential invaders’ survivability during transit. Management strategies thus must be adapted to countering these additive trends. They suggest specifically:

eDNA detection techniques;
Stricter enforcement of ISPM#15 and exploring use of recyclable plastic pallets (e.g., IKEA’s OptiLedge); [see my blog re: plastic pallets, here]
Application of fouling-resistant paints to ship hulls;
Prompt adoption of international agreements addressing pathways (they cite the Ballast Water Management Treaty as entered into force only in 2017 — 13 years after adoption);
Ensuring ‘pest free status’ (per ISPM#10) before allowing export of goods—especially goods in the “Agriculture”, “Horticulture”, and “Ornamental” trades; and
Increasing training of interception staff at ports.

What InvaCost Data say re: Taxa of greatest concern to me

Two-thirds of reported expenditures are spent on terrestrial species (Cuthbert et al.). Insects as a Class constitute the highest number of species introduced as ‘Contaminants’ (n = 74) and ‘Stowaways’ (n = 43). They also impose the highest costs among species using these pathways. 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.). Indeed, one of 10 species for which reported spending on post-invasion management is highest is the infamous Asian longhorned beetle (Tuberlin et al.)

ALB pupa in wood packaging; Pennsylvania Dept. of Natural Resources via Bugwood

Mammals and plants are often introduced deliberately – either as intentional releases or as escapes. Plant invasions are reported as numerous but impose lower costs.

Tuberlin et al. state that intentional releases and escapes should in theory be more straightforward to monitor and control, so less costly. They propose two theories: 1) Eradication campaigns are more likely to succeed for plants introduced for cultivation and subsequently escaped, than for plants introduced through unintentional pathways in semi-natural environments. 2) Species introduced unintentionally may be able to spread undetected for longer; they expect that better measures already exist to control invasions by deliberate introductions. I question both. Their theories ignore that constituencies probably like the introduced plants … and the near absence of attention to the possible need to control their spread. This is odd because elsewhere they recognize conflicts over whether to control or eradicate “charismatic” species.

Geographies of greatest concern to me

North America reported spending 54% of the total expenditure in InvaCost. Oceania spent 30%. The remaining regions each spent less than $5 billion. (Cuthbert et al.)
North America funded preventative actions most generously than other regions. Cuthbert suggests this was because David Pimentel published an early estimate of invasive species costs. I doubt it. The Lacey Act was adopted in 1905. USDA APHIS was formed in 1972 – based on predecessor agencies — because officials recognized the damage by non-native pests to agriculture. APHIS began addressing natural area pests with discovery of the Asian longhorned beetle in 1996. Of course, most of APHIS’ budget is still allocated to agricultural pests. I conclude that North America’s lead in this area has not resulted in adequate prevention programs.

Oregon ash swamp before attack by EAB (photo by Wyatt Williams, Oregon Dept. of Forestry)

Equity Issues

Tuberlin et al and Moodley et al. address equity issues of who causes introductions vs. who is impacted. This is long overdue.

More than 80% of bioinvasion management costs in protected areas fell on governmental services and/or official organizations (e.g. conservation agencies, forest services, or associations). With the partial exception of the agricultural sector, the economic sectors that contribute the most to movement of invasive species are spared from carrying the resulting costs (Moodley et al.)
A lack of willingness to invest might represent a moral problem when the invader’s impacts are incurred by regions, sectors, or generations other than those that on whom management action falls (Ahmed et al.)
People are perhaps more inclined to spend money to mitigate impacts that cause economic losses than those that damage ecosystems (Tuberlin et al.)

Data deficiencies

Only 41% of countries (83 out of 204) reported management costs; of those, only 24 reported costs specifically associated with pre-invasion (prevention) efforts (Cuthbert et al.).
Reliable economic cost estimates were available for only 60% of the “worst” invasive species (Cuthbert et al.)
Only 55 out of 266,561 protected areas reported losses or management costs (Moodley et al.).
Information on pathways of introduction was available for only three species out of 10,000 (Turbelin et al).
Taxonomic and geographic biases in reporting skew examples and possibly conclusions (Cuthbert et al.).

SOURCES

Ahmed, D.A., E.J. Hudgins, R.N. Cuthbert, .M. Kourantidou, C. Diagne, P.J. Haubrock, B. Leung, C. Liu, B. Leroy, S. Petrovskii, A. Beidas, F. Courchamp. 2022. Managing biological invasions: the cost of inaction. Biol Invasions (2022) 24:1927–1946 https://doi.org/10.1007/s10530-022-02755-0

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

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

Turbelin, A.J., C. Diagne, E.J. Hudgins, D. Moodley, M. Kourantidou, A. Novoa, P.J. Haubrock, C. Bernery, R.E. Gozlan, R.A. Francis, F. Courchamp. 2022. Introduction pathways of economically costly invasive alien species. Biol Invasions (2022) 24:2061–2079 https://doi.org/10.1007/s10530-022-02796-5

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

www.fadingforests.org

Urgent Update on Beech Leaf Disease

banding symptoms of beech leaf disease; photo by Dr. Chagas de Freitas, Ohio State

Experts on beech leaf disease (BLD) hold conference calls every two months. I reported on the May meeting earlier in July. The July conference call of the experts emphasized not only the alarming spread of the disease but also the dilemmas frustrating efforts to slow its spread and protect beech.

Jerry Carlson, chief of forest health protection for the New York Department of Environmental Conservation called beech leaf disease “the next chestnut blight”.

Yet forestry, plant health, and conservation entities have been slow to support research needed to develop protective measures.

As was noted by participants, 10 years after scientists from Lake MetroParks (in Cleveland) first detected disease symptoms, scientists still are unsure about all aspects of BLD and how it spreads. Experts agree that the nematode (Litylenchus crenatae ssp mccannii) must be present to initiate the disease. Other possible factors, especially fungi in the genus Colletotrichum, appear to play important roles in causing the symptoms.

The lack of clarity about the causal agent means that USDA APHIS has not recognized the disease as a priority species for tracking. APHIS has provided some funds. However, scientists seeking to obtain funding through the Plant Pest and Disease Management and Disaster Prevention Program [laid out in the Plant Protection Act’s Section 7721] can’t get traction. Other funding sources also don’t quite fit. For example, the National Science Foundation funds basic, hypothesis-driven, research – not the more applied science needed to address BLD. Some participants wondered whether funding might be sought from wildlife-oriented sources, since beech are so important in providing hard mast, den and nest sites, etc., for a variety of wildlife.

Participants discussed ways to raise awareness – and alarm – in order to build a broader coalition. This effort should include Europe. Although the disease has not yet been detected in Europe, the native beech is vulnerable.

European beech in Rhode Island infected by BLD; photo by Dr. Nathaniel A. Mitkowski, University of Rhode Island

Data indicate that the disease is now significantly more widespread than was known last year. That is, BLD is more widespread from New York to Maine, with New Hampshire reporting its first detection. To the west, BLD has been detected in Michigan and in a forest fragment in western Ohio (near Toledo). Disease severity has also intensified. Of course, the disease is present at least a year before it is detected because leaf symptoms appear in the spring following infection. Therefore its presence is probably wider.

map of BLD presence as of early July 2022 (some states have not yet reported); note the many counties in fuschia – 2022 detections

While mortality of mature beech is still rarely documented, this might be related to difficulties determining the cause of mortality during standard forest health surveys. Participants discussed how to rectify this situation.

Meanwhile, concern is rising – as reflected in Dr. Carlson’s statement.

You can help by asking your state and national officials and conservation organizations to support applied research aimed at clarifying how the disease spreads, what ecological conditions are conducive to disease, improved detection and prediction tools, and possible containment strategies. Let’s overcome the roadblocks impeding protection of this magnificent and ecologically vital tree species.

Is this not worth protecting?

Posted by Faith Campbell

www.fadingforests.org

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.

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.

Beech Leaf Disease (BLD) – Ongoing Research by the BLD Research Group

See bldresearch@lists.osu.edu

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

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

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

Explore potential effects of climate change on establishment risk, especially higher latitude areas expected to see increasing humidity, precipitation, and rising temperatures.
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.
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

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:

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.
The data relate to economic rather than ecological effects. The costliest species economically might not cause the greatest ecological harm.
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.
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

Posted by Faith Campbell

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.

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

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:

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

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

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

4^th 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