Florida’s bromeliads: who will protect them?

*Tillandsia utriculata*; photo courtesy of Teresa Cooper

Florida has 16 native species of bromeliads. Up to 15 invertebrates depend on bromeliads, especially the water that collects at the bases of the leaves. See Frank (1983) and Frank and Fish (2008) for more information about bromeliads’ ecological importance.

Eight species had been listed as threatened or endangered in the state (visit here) (Frank and Cave 2005) before arrival of the Mexican bromeliad weevil. The Florida Endangered Plant Advisory Council added two of its hosts – Tillandsia utriculata and Tillandsia fasciculata. The website now lists 12 species as threatened by the weevil.

Twelve species are believed to be vulnerable to the weevil. T. utriculata is at most immediate risk. Before arrival of the Mexican bromeliad weevil, some populations of this species were described by Teresa Cooper as “giant” and dense, containing very large mature plants and producing huge volumes of of seed.

One such population — in the Enchanted Forest Sanctuary in Brevard County – succumbed to the weevil quickly: 87% died in six months. At 27 months after arrival of the weevil, less than 3% of the original population remained. It is now rare to see large T. utriculata plants. Medium-sized plants put out inflorescences. The weevil persists at low levels, so is able to feed on and reproduce on medium and small plants. Several endemic wildlife species that inhabit the water pooled in T. utriculata’s base face global extinction (T.M. Cooper pers. comm. 2026).

damage to a bromeliad caused by the weevil; photo courtesy of Teresa Cooper

The first reason T. utriculata is at higher risk is that each plant can support many weevil larvae. Its leaves are softer (so more easily consumed) and its large stems contain lots of nutrients — for a bromeliad). More important is the plant’s very slow reproduction in contrast to the weevil’s rapid attack. T. utriculata plants do not produce seed until after they have grown for up to 20 years. Furthermore, an extremely small fraction of the ~ 10,000 seeds released by a mature plant make it to reproductive age. A plant that after 18 years begins growing an inflorescence — which then takes ~ a year to flower and produce seed – can be killed by a weevil within a few months. In addition, T. utriculata is monocarpic – it doesn’t produce offshoots. T.M. Cooper asserts that if the bromeliad weevil extirpates T. utriculata from Florida, it will not be because the weevil ate the very last plant, it will be because T. utriculata seed output has fallen below a sustainable level.

Teresa Cooper demonstrating the size of a *T. utricuclata* flower spike; photo courtesy of T. Cooper

A second host, Tillandsia fasciculata, had large and dense population in Loxahatchee National Wildlife Refuge (Palm Beach County) in 2002 to 2005. Almost every tree had multiple T. fasciculata plants; many were huge with multiple pups. When Teresa Cooper revisited about 10 years later, T. fasciculata plants were very sparse, large plants very rare; none was really large (T.M. Cooper pers. comm. 2026).

T. fasciculata is expected to decline more slowly than T. utriculata. First among several advantages is that its leaves are tougher and stems smaller, so they provide less nutrition to the weevil. Probably more important, T. fasciculata is polycarpic (it produces both seed and offshoots). A large T. fasciculata plant can sprout up to a dozen blooming rametes at a time. If the weevil kills one, other rametes will bloom and seed. At the same time, the plant might produce numerous up to 20 offshoots which come detatched from the “mother” plant and start a new individual. Still, as the weevil kills these rametes year after year the plant shrinks in size and eventually dies (T.M. Cooper pers. comm. 2026).

Teresa Cooper has documented damage on Tillandsia paucifolia, T. simulate, and T. variabilis (T.M. Cooper pers. comm. 2026). She has no funding to continue studying these species. I have been unable to find any more recent information about the status of the Tillandsia species. The Florida Department of Agriculture and Consumer Services does not have an active program addressing the weevil (Patrick Gordon, FDACS, pers. comm. June 2026).

There is considerably more information about another endangered bromeliad, Guzmania monostachia. This species has the broadest distribution of any species in the genus, stretching across northern South America, Central America, the Caribbean, as well as southern Florida. As the northernmost population, the Florida population might harbor unique genetics important for the species’ adaptation to climate change (Krupar et al. 2023).

Formerly more common, G. monostachia is now found in six fragmented and disjunct populations in five areas in Florida. The species has apparently been extirpated from four other sites by habitat loss, not depredations by the weevil. One set of habitats is in wetland sloughs on the peninsula’s west coast on the peninsula’s west coast. These forests comprise pond apple (Annona glabra) and Florida ash (Fraxinus caroliniana); Kupar et al. (2023) don’t mention whether this species is vulnerable to the emerald ash borer, which has not yet been detected in Florida. The largest of the bromeliad populations is in Fakahatchee Strand State Preserve, in Collier County in Southwest Florida. Fakahatchee Strand Preserve is Florida’ oldest and largest state park. The bromeliad’s population was previously estimated to exceed two million individuals. Their number has been halved by weevil attack. Also, since M. callizona prefers larger plants, the proportion of reproductively mature individuals had been reduced from roughly 50% to only 10–20% by 2021. T.M. Cooper reports (pers. comm. 2026) fears this species might also be extirpated eventually.

*Guzmania monostachia*; photo courtesy of Teresa Cooper

The adjacent Big Cypress National Preserve is home to two populations of G. monostachia; they are separated by ~2,900 km²so there is no interaction between them. The southern population consists of ~ 1000 individuals, the northern population comprises only~ 200 individuals.

Guzmania monostachia is in an even more precarious situation on the eastern side of the peninsula: tiny populations of two or three plants are found in three locations: Everglades National Park, Fuchs (formerly Sykes) Hammock Preserve, and Meissner Hammock Preserve.

The Mexican bromeliad weevil Metamasius callizona (Chevrolat) is native to southern Mexico, Guatamala, (Frank and Thomas 1994; Frank and Cave 2005) and Belize (Cooper, Cave and Frank 2023). Its presence in Florida was detected in a nursery in Ft. Lauderdale, Broward County, in 1989. Probably introduced on a shipment of ornamental bromeliads from Mexico. The nursery treated the infested plants, but the weevil had already established on native bromeliad species in the natural environment. By 1991, it was detected in four counties in southern Florida; by 1999, it was found in 12 more. It is now in most counties of peninsular Florida from Daytona (Volusa County) south to Miami-Dade (Patrick Gordon, Florida Department of Agriculture and Consumer Services, pers. comm. June 2026).

While chemical control is feasible in nurseries and display plantings, it can’t be used in natural areas, where the epiphytes are not accessible from the ground and stakeholders fear likely non-target effects.

Therefore, scientists initially focused on classical biocontrol. They made 16 expeditions to Mexico and Central and South America from 1992 to 2010 searching for natural enemies of the weevil. The chose parasitoid tachinid fly, Lixadmontia franki. A colony was established in the University of Florida’s quarantine laboratory and research on fly-weevil interactions was conducted for several years. Releases into the environment were begun under Federal and State permits in 2007. Although more than 3,100 flies were released, the species failed to establish. So this approach is no longer being pursued (Cooper, Cave, and Frank (2024); T.M. Cooper pers. comm. June 2026).

Meanwhile, in 2010, Dr. Frank and D. Giardina of the Florida Fish and Wildlife Conservation Commission observed that in one location in Belize the weevils had no apparent detrimental effect on bromeliads identified as Tillandsia utriculata (although there is some disagreement on this classification). Cooper, Cave, and Frank (2024) undertook a study to determine how the bromeliad could coexist with Metamasius callizona in Belize while being so vulnerable in Florida. They compared life cycle parameters (oviposition and pupation rates, egg hatch rate, adult emergence and size, and developmental time) of weevils from Florida and Belize. They also observed how populations of the weevil from Florida fared on three hosts: pineapple (Ananas comosus), T. utriculata collected from Florida, and T. utriculata collected from Belize. Finally, they quantified the hosts’ nutritional content using two measures (percent soluble solids and leaf toughness).

Their most important finding is that weevil larva from Florida could not develop past the third instar on leaves of the T. utriculata from Belize. The weevil larvae starved because the Belize bromeliad’s leaves had fewer nutrients and were tougher than the leaves from T. utriculata collected in Florida.

bromeliads seen in Big Cypress National Preserve in the early 1980s; photo by F.T. Campbell

Based on this finding, Cooper, Cave, and Frank (2024) recommend that authorities introduce T. utriculata plants from Belize into Florida’s forests and allow them to colonize and/or hybridize naturally with Florida’s remaining plants. An alternative would be to breed hybrid Tillandsia in the lab and release them into Florida’s forests. I note that the American Chestnut Foundation pursued a similar strategy for decades to develop American chestnuts (Castanea dentata) able to tolerate the chestnut blight fungus (Cryphonectria parasitica).

Sources

Cave, R.D. 1997. Admontia sp., a potential biological control agent of Metamasius callizona. J Brom Soc. 47:244-249.

Cave, R.D. 2008. Biological control of the MEXICO bromeliad weevil. Biocontrol News and Information 29(1):1N-2N.

Cooper T.M. 2006. Ecological and demographic trends and patterns of Metamasius callizona (Chevrolat), an invasive bromeliad-eating weevil, and FLORIDA’s native bromeliads [Master’s thesis]. [Gainesville (FLORIDA)]: University of FLORIDA. 69 p.

Cooper, T.M., R.D. Cave, and J.H. Frank. 2023. Potential bottom-up control of Metamasius callizona in Florida, USA. Entomologia Experimentales et Applicata. 2024. 172;4090421

Frank, JH. 1983. Bromeliad phytotelmata and their biota, especially mosquitoes. In: Frank J.H., Lounibos, L.P., editors. Phytotelmata: terrestrial plants as hosts for aquatic insect communities. Medford (NJ): Plexus. p. 101-128.

Frank, J.H. 1996. Bromeliad biota: history of Metamasius callizona FLORIDA [online]. Gainesville (FLORIDA): University of FLORIDA [cited 2010 Feb 1]. Available from https://www.entnemdept.uFlorida.edu/frank/bromeliadbiota/wvbrom6.htm

Frank, J.H. and R.D. Cave. 2005. Metamasius callizona is destroying FLORIDA’s native bromeliads. In: Hoddle MS, editor. USDA Forest Service Publication FHTET-2005-08. Vol 1. Second International Symposium on Biological

Control of Arthropods; 2005 Sep 12-16; Davos, Switzerland. Washington D.C.: USDA Forest Service. p. 91-101.

Frank, J.H. and D. Fish. 2008. Potential biodiversity loss in Florida bromeliad phytotelmata due to Metamasius callizona (Coleoptera: Dryophthoridae), an invasive species. Florida Entomol. 91(1):1-8.

Frank, J.H. and M.C. Thomas. 1994. Metamasius callizona (Chevrolat) (Coleoptera:Curculionidae), an immigrant pest, destroys bromeliads in FLORIDA. Can Entomol.126(1):673-682.

Krupar, S., A.A. Naranjo, G. Godden, N. Cellinese. The Fate of Guzmania monostachia in Florida Rests with Humans. Diversity 2023, 15, 525. https://doi.org/10.3390/d15040525

Potter, K.M., Escanferla, M.E., Jetton, R.M., Man, G., Crane, B.S., Prioritizing the conservation needs of US tree species: Evaluating vulnerability to forest insect and disease threats, Global Ecology and Conservation (2019), doi: https://doi.org/10.1016/

Salas, J. and J.H. Frank. 2001. Development of Metamasius callizona (Coleoptera:Curculionidae) on pineapple stems. Florida Entomol. 84(1):123-126.

Wood, D.M. and R.D. Cave. 2006. Description of a new genus and species of weevil parasitoid from Honduras (Diptera: Tachinidae). Florida Entomol. 89(2):239-24.

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

https://fadingforests.org

Plant invasions grow everywhere

invasion of Chinese privet (*Ligustrum sinense*)

A decade ago I posted a blog reporting that 39% of forests surveyed under the Forest Inventory and Analysis (FIA) system were invaded by one or more invasive plants (Oswald et al. 2015). By regions, Hawai`i had the highest invasion intensity – 70%. The second highest density was in the eastern forests – 46%. Forests in the West ranked third, with 11% of plots containing at least one of the monitored invasive plant species. Finally, forests in Alaska and the Intermountain regions both had 6% of plots invaded.

I rejoice that US Forest Service scientists have continued to analyze their data on plant invasions. Analysis of the most recent data shows alarming increases in invasions everywhere since 2015. However, the scientists could not determine a nation-wide percentage because many areas in the West had not yet been surveyed anew. They did determine that the number of inventory plots containing invasive plant species rose in 58.9% of surveyed counties. Furthermore, in 73.2% of the counties the plots experienced an increase in species richness of invading plant species. While increases were observed in all regions, they were greater in the East than in the West — and in the USFS Southern region compared to the Northern region. Specifically, the proportion of forest plots in the East (USFS Southern and Northern regions) invaded has risen from 46% to 52.8%. In the Rocky Mountains they rose from 6% to 11%. In Hawai`i plots having invasive plants grew from 70% to 83.2%. Surveys in the Pacific Coast states have not yet been completed so this region is not included in the analysis (Potter et al. 2026). It is not clear to me how the current boundaries of the western regions – which are based on Bailey’s ecosystem boundaries relate to the 2015 boundaries, which were based on USFS official regions. Hawai`i is clearly the same.

Porter et al. (2026) concluded that in the forests of the East plant invasions are so extensive that elimination of their impacts is practically impossible. Their spread to new areas is unhindered now and, I would add, is likely to remain so without heroic counter measures.

Forests in the East have a greater mean richness of invasive plant species than do western forests. In particular, there is a profusion of shrubs and vines as well as trees. The West has a greater diversity of invasive forbs. The diversity of invasive grasses is high in both regions.

kudzu (*Pueraria montana*) spreading from edge into forest in Virginia; photo by F.T. Campbell

Potter at al. (2026) worry that the apparently lower level of plant invasions in the West might be an artifact of a higher proportion of plant species being at an earlier stage of invasion. That is, the species have not yet established sufficiently widely to be classified as invasive.

thicket of guava (*Psidium cattleianum* ) replacing `ohi`a killed by ROD; Hawai`i Island; photo by F.T. Campbell

Of course, the situation in Hawai`i is much worse. Another, more detailed, discussion of invasive plant species in Hawai`i pointed out that relying on data reflecting canopy-level trees obscures the real picture. While “only” 29% of large trees across the Islands are non-native, about two-thirds of saplings and seedlings are. Potter et al. (2023) expected that plant succession will result in non-native tree species taking over the canopy. This likelihood exists regardless of the impact of rapid ‘ohi’a death since ‘ohi’a lehua (Metrosideros polymorpha) is not reproducing even when seed sources are plentiful and people remove invasive forbs and grasses Potter et al. (2023).

The nation-wide analysis of Potter et al. (2026) does not include forests on U.S. Caribbean islands, i.e., Puerto Rico and the Virgin Islands. See here for a description of this situation. In summary, 33 of 57 (58%) of non-native tree species tallied by FIA surveyors are actual or potential high-impact bioinvaders. Furthermore, 21 (38%) of the non-native species occurred on at least 2% of the FIA plots – far above the seven species fitting this description in the continental U.S.

As these sources, and those with a broader perspective, demonstrate that we should not ignore invasions of our forests by non-native plants. These species erode forest productivity and provision of the full range of ecosystem services, hinder shifting (?) forest uses, and degrade biodiversity and habitat.

These invasions also impose extensive financial costs from lost or damaged resources (Potter et al. ( 2022). Potter et al. (2026) note that these negative outcomes depend on interactions between the traits of the non-native plants and the biomes being invaded. These impacts are greatly exacerbated in Hawai`i because more than 95% of native species on the Islands are endemic. This includes 67% of the large trees still present in the forests. As Potter et al. (2023) point out, extirpation of any of these species is a global loss.

ʻōhiʻa lehua (*Metrosideros polymorpha*); photo by F.T. Campbell

Data issues

Potter et al. (2026) note that in the Northern region only about 20% of plots were surveyed for invasive plants. They state that these difference in sampling intensity does not affect statistical analyses across broad scales.

The regional lists of invasive plants were developed by experts. They include those species thought at the time to be most damaging. Of course, there are other non-native plant species that might be present – and some might prove to be invasive over time (Potter et al. 2026). I have been unable to determine whether the regional lists are updated periodically. Because of this structure of the FIA system, these surveys can assess only spread of already-established species. It is not suitable for early detection of new species entering the forest.

For all these reasons, the analyses in Porter et al. (2026) probably underestimate the total abundance of non-native plant species in U.S. forests. Indeed, the time lag between introduction or even identification of invasive species and their eventual ecological and economic impact obscures their full impact. This ever-increasing invasion debt probably contributes to decisions not to implement effective countermeasures.

Recommendations

How do we set priorities for responding to nearly unmanageable situations? We sharpen our focus on the most damaging pathways of introduction, the most vulnerable regions, and the most at-risk species.

The high-risk pathways are imports of plants for planting and wood – including but not limited to crates, pallets, and other forms of packaging.

Vulnerable regions start with the Hawaiian Islands, Puerto Rico, and the Virgin Islands; and include many biodiversity-rich areas on the continent. We should enhance monitoring of these vulnerable regions by federal, state, and tribal agencies, conservation organizations, citizen scientists, and others. Surveys must report all non-native plant present, not just those already known to be invasive. These data will improve detection of new species and better inform us about factors affecting species’ spread.

Also, I support Potter et al.’s (2026) emphasis on the wildland-urban interface as an area of high human-environment conflict.These include, but are not limited to, plant invasions. The authors point out that we need new policy, management, and scientific tools to address threats in these vulnerable and too-often ignored social and ecological zones.

This increase in available information must be paired with management of the factors that facilitate invasion. Some of these are associated with ecosystems. But the key target must be plant species being brought into the region by people for various purposes. This is often for ornamental horticulture.

lesser celandine (*Ficaria verna*) dominating herb layer in a Virginia forest; photo by F.T. Campbell

We must ask state legislatures and Congress to empower regulatory agencies – e.g., their state departments of agriculture and USDA’s Animal and Plant Health Inspection Service – to be far more more assertive and pro-active. For example, they must give higher priority to the full range of ecological and economic impacts of invading plants, not just damage to agriculture.

Evans et al. (2024) urged prioritizing for state regulation those species in the ornamental trade that are projected to remain or become abundant under evolving climate conditions. Beaury et al. (2023) called for regulating the nursery trade at the national level – reflecting the scope of sales.

SOURCES

Beaury, E.M., J.M. Allen, A.E. Evans, M.E. Fertakos, W.G. Pfadenhauer, B.A. Bradley. 2023. Horticulture could facilitate invasive plant range infilling and range expansion with climate change. BioScience 2023 0 1-8 https://doi.org/10.1093/biosci/biad069

Evans, A.E., C.S. Jarnevich, E.M. Beaury, P.S. Engelstad, N.B. Teich, J.M. LaRoe, B.A. Bradley. 2024. Shifting hotspots: Climate change projected to drive contractions and expansions of invasive plant abundance habitats. Diversity and Distributions 2024;30:4154

Potter, K.M., C. Giardina, R.F. Hughes, S. Cordell, O. Kuegler, A. Koch, E. Yuen. 2023. How invaded are Hawaiian forests? Non-native understory tree dominance signals potential canopy replacement. Landsc Ecol 2023 https://doi.org/10.1007/s10980-023-01662-6

Potter, K.M., B.V. Iannone III, K.H. Riitters, Q. Guo, K. Pandit, C.M. Oswalt. 2026. US Forests are Increasingly Invaded by Problematic Non-Native Plants. Forest Ecology and Management 599 (2026) 123281

Potter K.M., K.H. Riitters, and Q Guo. 2022. Non-native tree regeneration indicates regional and national risks from current invasions. Frontiers in Forests & Global Change Front. For. Glob. Change 5:966407. doi: 10.3389/ffgc.2022.966407

Potter, K.M., K.H. Riitters, B.V. Iannone, III, Q. Guo and S. Fei. 2024. Forest plant invasions in eastern US: evidence of invasion debt in the wildland‑urban interface. Landsc Ecol (2024) 39:207 https://doi.org/10.1007/s10980-024-01985-y

Posted by Faith Campbell

https://fadingforests.org

Invasive plants threaten integrity of eastern U.S. forests

garlic mustard (*Alliaria petiolata*); photo by Katja Schulz via Wikimedia

I welcome a recent series of studies documenting the extent of plant invasions in forests of the eastern United States and the socio-economic conditions that contribute to a state of affairs increasingly recognized as a crisis. I wish, however, that the authors had devoted more attention to the role of deliberate planting of non-native species and the resulting propagule pressure.

I summarize here findings of two studies written by largely the same scientists and relying on the same underlying data: surveys of forest plots conducted under the Forest Inventory and Analysis (FIA) program. In this blog, if focus on the extent of invasive plant presence in the forests of the eastern United States. In an accompanying blog I will summarize the status of plant invasions in forests nation-wide.

As I have noted in earlier blogs, link a decade ago one or more invasive plant species had already invaded 46% of FIA plots in the eastern U.S. (Oswald et al. 2015). This situation has worsened. Updated data show that 52.8% of these plots contain invasive plants. In the USFS Southern Region, invasive plants have been documented on 55.3 million ha. In the Northern Region, they are found on 36.9 million ha. (Only ~20% of FIA plots in the Northern Region were surveyed for invasive plants.) In some counties of the 37 states constituting these two USFS regions, 80% of inventoried forest plots contain invasive plants. Areas with lower levels of invasion are found in parts of New England, the Great Lakes states, southern Appalachians, southeastern coastal plain, and western Texas and Oklahoma (Potter et al. 2026). Spread of these bioinvaders is largely unchecked – either throughout the East or “just” in the South. In any case, the extent and intensity of these invasions are so great that their complete removal – or elimination of their impacts – is “practically impossible” (Potter et al., 2024; Potter et al. 2026). [It is not clear whether the scientists mean “nearly” or “in practice”. Or that this difference is important.]

[In comparison, in the West less than 30% of FIA plots are invaded, on average. In Hawai`i, more than70% are (Potter et al. 2026).]

The scientists analyzing the FIA data warn that the extent and impact of plant invasions in eastern forests is undoubtedly worse than these data indicate. The records include only some of the non-native plant species present — those considered to be the worst invaders at the time regional lists were compiled – apparently in the first years of the 21^st Century (Potter et al. 2026).

Japanese honeysuckle (*Lonicera japonica*) photo by Chuck Bargeron

The scientists emphasize the role of disturbance in promoting plant invasions. They cite various studies as well as the FIA data to document that forest edges facilitate non-native plant establishment and spread into forests. They stress various aspects of suburban development, including roads and other transportation corridors. It follows that invasion rates are highest in the “wildland-urban interface (WUI).” [The wildlife-urban interface is the zone of transition between unoccupied land and human development; the zone where structures meet or intermix with undeveloped land and its vegetation.] They worry that the WUI is growing faster than any other land use type in the country – and especially rapidly in the East. As a result, the scientists expect more and worse invasions in the future (Potter et al., 2024 and Potter et al. 2026).

I appreciate that they highlight the uniqueness of WUI ecosystems. Housing development in the WUI has numerous effects on natural ecosystems, including habitat modification and fragmentation followed by diffusion of the direct and indirect effects of anthropogenic activities into neighboring ecosystems at different scales. As regards specifically non-native plants, this transmission occurs through a combination of (1) human-driven disturbances to native ecosystems that promote plant invasion and (2) providing a source of non-native plant propagules in their yards and gardens. These plants can then spread into and establish in nearby ecosystems (in this case, forests). [I note that tree-killing arthropods and pathogens also can be introduced in the WUI.] (Scroll below “Archives” to “Categories”, click on “forest pests” and “wood packaging”.)

They also found that plant invasions are more strongly related to older, than more recent, land-cover changes. Survey plots that have been located in the WUI since 1990 or earlier had on average 2.6% more invasive plant cover and 0.33 more invasive plant species than those that were classified as being in the WUI in 2000 or 2010. Their explanation is that the WUI forests experienced decreased spatial integrity, increased forest-developed area edges, and falling proportions of forest in the surrounding landscapes. In addition, the human population in the vicinity might have grown. All these factors that would increase forest fragmentation and the plots’ susceptibility to invasion.

The other side of the coin is propagule pressure. Both Potter et al (2024) and Potter et al. (2026) note that the flora of residential landscapes – rural as well as suburban – is typically dominated by non-native plant species. Still, I think these studies downplay the impact of this ubiquity of non-native plants in all anthropogenic landscapes.

In discussing the higher invasion rates found in survey plots located in WUIs dating from the 1990s they made no mention of human activities that promote plant invasions. There are several. Plants growing in those older yards had one or two more decades to flower – and for their fruits and seeds to be transported into the forest by birds, wind, or water. Residents might have decided to beautify their neighborhood by planting shrubs or flowers in the woods. Maybe they succumbed to the temptation to dump yard waste in the woods – thinking it would be absorbed by “nature”. Since plant invasions take time to unfold, these additional years of human-mediated exposure are highly relevant. Another factor is that people who choose to live in wooded surroundings probably choose horticultural plants that thrive under such conditions – exactly those best able to establish beyond the property line.

Another opportunity to discuss these factors came from the discovery that plant invasion rates are higher in association with “interface” rather than “intermix” WUI forests. [“WUI interface forests” are those where settled areas abut wildlands. In “WUI intermix forests” the structures are scattered.] They speculate about reasons. Potter, et al. (2024) mention that invasions originating from older housing developments have had more time to establish (or at least to be detected) given the well-known lag associated with plant invasions.

I wish they had focused more on the probable difference in suburban development across time. While I was growing up in expanding suburbs in the 1950s, I observed that the earlier housing developments were either built on land that had been cleared to support agriculture or the builders cleared the forest to make construction easier and cheaper. More recently, wealthier buyers have sought residences on more wooded sites – so creating an “intermix” WUI. Potter et al. (2024) speculate that locations in the “interface” WUI are closer to high-density urbanization so have higher exposure to non-native plants. They do not discuss whether the “interface” WUIs are older, thus giving associated plantings longer years to proceed through the stages of bioinvasion.

burning bush (*Euonymus alatus*) invading a forest in Virginia; photo by F.T. Campbell

The Role of Deliberate Planting?

I recognize that these authors analyzed mountains of data. However, I wish they had incorporated the findings of numerous scientists who have analyzed the role of deliberate planting – especially ornamental horticulture – in facilitating introduction and spread of invasive plants. (Scroll below “Archives” to “Categories” and click on “invasive plants”. Also See Reichard and White 2001 and Mack 2000).

As I hope USFS scientists are aware, recent studies confirm the continuing role of ornamental horticulture in plant invasions. Kinlock et al. (2025) blog 440 found that more than 1,600 plant species sold by nursery and seed catalogs over 200 years had “naturalized” somewhere in the continental 48 states. They do not discuss what proportion of these species are truly damaging invaders. Fertakos and Bradley (2024) found that species were likely to establish if they were introduced to as few as eight locations. Beaury et al. (2024) found that half of 89 plant species recognized as invasive are sold in the same locations where they are invasive. Another 25 species are sold by one or more nurseries located in an area that is currently unsuitable for those species, but that will become more suitable for invasion as temperatures warm.

Potter et al. (2026) acknowledge that the ornamental plant trade is likely to continue introducing new plant species into U.S. forests. However, they recommend only updating the lists of invasive plants to be included in future surveys. Apparently these lists have not been updated since 2004.

Potter et al. (2024) go farther, urging efforts to encourage homeowners to plant more native and environmentally friendly private landscapes. They note that such advocacy is complicated by the fact that non-native – even invasive – species provide valued ecosystem and cultural services.

I add that the nursery industry and their customers enjoy enormous lobbying clout.

Many associations – native plant societies, regional or state invasive plant councils, etc. – are pursuing this approach. To research these efforts, visit the websites for the state native plant societies and the Southeast Exotic Pest Plant Council, Mid-Atlantic Invasive Plant Council, and Midwest Invasive Plant Network. These voluntary efforts have yielded some success. But they have not resulted in adequate protection for our ecosystems. Dr. Douglas Tallamy points out that even non-invasive, non-native plants disrupt food webs.

The insufficient attention to the role of the plant trade in articles intended to be comprehensive has crucially important impacts. As both Potter, et al. (2024) and Potter et al. (2026) affirm, determining which factors are most important in facilitating plant invasions of eastern American forests is the necessary foundation for identifying and implementing the most efficient and effective counter measures.

These scientists are employees of the U.S. Department of Agriculture. If departmental leadership interpret their studies as justifying inaction on regulating plant sales, USDA’s regulatory agencies will not respond. And we will continue failing to curtail introduction and spread of damaging plant invasions.

I agree with the authors on the need for enhanced monitoring and management of WUI zones in the East to detect new species or new locations of invasion and the need to develop better tools for these purposes. However, I ask all stakeholders to follow Evans et al. (2024), who urge prioritizing for state regulation those species in the ornamental trade that are projected to remain or become abundant under evolving climate conditions. Or, more aggressively, follow Beaury et al. (2023)’s call for regulating the nursery trade in a manner consistent with the scope of the horticultural trade at the national level. That would require legislation, since the Federal Noxious Weed Act does not currently address long-established, widespread species. Beaury et al. (2023) also note that existing state restrictions are outdated, tend to include only a few weeds that plague agriculture rather than those that invade natural systems, and are irregularly enforced.

orchids in Everglades National Park; photo by F.T. Campbell

I conclude by agreeing with the scientists that managing the disturbance component of plant invasions points to protecting particularly forests of high conservation value. They suggest adoption of land-use planning rules aimed at this goal. However, as they point out, such action will be extremely unlikely given the magnitude of predicted land-use changes in the country and powerful demographic factors driving them. I would add other barriers: the lobbying clout of the real estate industry and homeowners plus the local nature of zoning decisions.

SOURCES

Fertakos, M.E. and B.A. Bradley. 2024. Propagule pressure from historic U.S. plant sales explains establishment but not invasion. Ecology Letters 2024;27:e14494 doi: 10.1111/ele.14494

Kinlock, N.L., D.W. Adams, W. Dawson, F. Essl, J. Kartesz, H. Kreft, M. Nishino, Jan Pergl, P. Pyšek, P. Weigelt and M. van Kleunen. 2025. Naturalization of ornamental plants in the United States depends on cultivation and historical land cover context. Ecography 2025: e07748 doi: 10.1002/ecog.07748

Oswalt, C.M., S. Fei, Q. Guo, B.V. Iannone III, S.N. Oswalt, B.C. Pijanowski, K.M. Potter. 2016. A subcontinental view of forest plant invasions. NeoBiota. 24:49-54 http://www.srs.fs.usda.gov/pubs/48489

Potter, K.M., K.H. Riitters, B.V. Iannone III, Q. Guo and S. Fei. 2024. Forest plant invasions in the eastern United States: evidence of invasion debt in the wildland‑urban interface. Landsc Ecol (2024) 39:207 https://doi.org/10.1007/s10980-024-01985-y

Posted by Faith Campbell

https://fadingforests.org

Will we act to minimize sales of invasive plants?

*Wisteria floribunda*; photo by Jack Stane via Wikimedia

For decades, it has been clear that deliberate introduction of plant species for cultivation plays a central role in the early stages of bioinvasion by plants (and associated insects, plant pathogens, earthworms … even vertebrates. Viz. coqui frogs in Hawai`i.)

Repeatedly over the two plus decades since Sarah Reichard demonstrated this role of ornamental horticulture (see Reichard and White 2001 and Mack 2000), new studies have provided corroborative details. Publications during the past two years show the risks we are still accepting in the United States. Will we act to protect our environment?

Fertakos and Bradley (2024) found that species were likely to establish if they were introduced to as few as eight locations. However, introduction history was not a strong predictor of an established species’ ultimate invasive success. They suggest that other characteristics, like plant traits and local-scale processes (e.g., interspecific interactions), may better predict whether a plant becomes invasive.

Kinlock et al. (2025) also found that plant species that were cultivated longer or were sold by more catalogs were more likely to have “naturalized”. This conclusion was based on analysis of the behavior of nearly 4,000 species sold in nursery and seed catalogs in the continental United States over 200 years. Nearly 41% of these species naturalized somewhere in the “lower 48” states. Unfortunately, they do not discuss what proportion of these species are truly damaging invaders.

Neither Fertakos and Bradley (2024) nor Kinlock et al. (2025) mention the concept of a lag between a species’ establishment and recognized symptoms of invasiveness. Has this concept been repudiated?

Evans et al. (2024) were focused on analyzing which regions of the eastern United States are likely to suffer the worst plant invasions under climate change. In this context, they worry that people will assist non-native plant species’ movement to newly suitable habitats. Evans et al. urge prioritizing for state regulation species in the ornamental trade that are projected to remain or become abundant under the new climate conditions. They say we Americans are poorly prepared to take this action, however, because plant sales are so poorly regulated and only 10% of land managers in eastern North America monitor for new invasive taxa. They say this is because the managing agencies lack of funding and personnel. After 2025’s losses of programs, appropriations, grants, and staff, this deficit is probably worse – not just for federal agencies but also the many state, local, and volunteer programs that have been supported by federal funding.

Beaury et al. (2024) investigated whether plant species recognized as invasive are sold in the same locations as where they are invasive. They found that half of the 89 species named as invasive were sold by a nursery within 21km of an observed record of invasion. The authors say that data gaps mean that these findings underestimate the number of species sold near locations of documented invasions. They warn that at least 25 species are sold by one or more nurseries located in an area that is currently unsuitable for those species, but that will become more suitable for invasion as temperatures warm. Like Evans et al. (2024), they urge proactive regulation to limit these species’ spread.

burning bush *Euonymus*, Japanese honeysuckle, & English ivy invading a bottomland hardwood site in Fairfax County, Virgina; photo by F.T. Campbell

U.S. Regulatory response is completely inadequate

Beaury et al. (2023) call for regulating the nursery trade in a manner consistent with the scope of the horticultural trade – sales by both e-commerce and brick and mortar stores go to customers far outside a specific state’s jurisdiction. Despite the interstate nature of the trade, sales of horticultural plants are regulated primarily by state governments. Even when a state does restrict the sale of a specified list of invasive plants, the regulations are outdated, tend to include only a few weeds that plague agriculture rather than those that invade natural systems, or are irregularly enforced. The result is a checkerboard of places where a species may legally be offered for sale next to places where that sale is prohibited. Finally, the regulations are reactive; they rarely include plants in anticipation of their spread to new areas. Beaury et al. (2023) call this as a missed opportunity to reduce the likelihood of ornamental escapes.

Evans et al. (2024) also note that online plant sales are relatively unregulated, and state regulations are inconsistent.

Under the Constitution, the appropriate entity for regulating interstate commerce is the federal government. The U.S. Department of Agriculture’s Animal and Plant Health Inspection Service is responsible for populating and managing the federal noxious weed list. Unfortunately, APHIS lists only those taxa that qualify as quarantine pests under the definition of that term in the International Plant Protection Convention (IPPC) Glossary of Phytosanitary Terms. This means that the taxon is either not yet present in the United States or, if present, is not widely distributed and is being officially controlled. Under these criteria, the federal noxious weed list is required to exclude nearly all the invasive plant species sold by the nursery trade.

To counter this enormous regulatory failure, many associations – native plant societies, regional or state invasive plant councils, etc. – publish their own lists of invasive plants. They often encourage their members and the public to either avoid planting these species voluntarily or to plant predominantly native plants. Also, these stakeholders urge nurseries to halt sales of invasive species voluntarily. Dr. Douglas Tallamy points out that even non-invasive, non-native plants disrupt food webs.

These voluntary efforts have yielded some success. But they have not resulted in adequate protection for our ecosystems.

Will Americans choose to invigorate the regulatory system? At a minimum, can we urge neighboring states to adopt a regional approach? More difficult, but also more effective, would be to persuade Congress to strengthen APHIS’ invasive plant regulations to outlaw interstate sales of at least those species documented to be invasive.

*Cortadera selloana*; picture by Alex Borland via PublicDomainPictures.net

Do you have other suggestions?

SOURCES

Fertakos, M.E. and B.A. Bradley. 2024. Propagule pressure from historic U.S. plant sales explains establishment but not invasion. Public? doi: 10.1111/ele.14494.

Fridley, J.D., P.J. Bellingham, D. Closset-Kopp, C.C. Daehler, M.S. Dechoum, P.H. Martin, H.T. Murphy, J. Rojas- Sandoval, D. Tng. 2025. A general hypothesis of forest invasions by woody plants based on whole-plant carbon economics.

Posted by Faith Campbell

https://fadingforests.org

New Shothole Borer in California — Alert! & Opportunity to Advise Whether the State or County Should Lead the Response

several *Euwallaceae* species; E. interjectus is 2nd from the top. Photo from Gomez et al. 2018; ZooKeys 768 19-68

In December 2024, California officials announced detection of a third species of invasive shothole borer beetle in the state. This invasion was found in Santa Cruz County in October 2024. The beetle has been identified as Euwallacea interjectus; the associated fungus is Fusarium floridanum. Like other non-native shothole borers in the same genus already known to be in California, Euwallacea interjectus is native to Southeast Asia.

So far, the infestation extends across at least 75 acres (CDFA proposal). It is affecting primarily box elders (Acer negundo). Other tree species have also been attacked: California sycamore (Platanus racemose), coast live oak (Quercus agrifolia), arroyo willow (Salix lasiolepis), red willow (Salix laevigata), and black cottonwood (Populus balsamifera ssp. trichocarpa). [See the CDFA risk assessment referred to below]. While it is too early to know precisely, E. interjectus is expected to pose a risk of tree dieback in urban, wildland and agricultural landscapes similar to that already caused by its relatives — the Polyphagous shot hole borer (Euwallacea fornicatus s.s. [PSHB]) and Kuroshio shot hole borer (Euwallacea kuroshio [(KSHB)].

The Santa Cruz County Department of Agriculture and University of California Cooperative Extension Service are coordinating with the California Department of Food and Agriculture (CDFA) to monitor and respond to the infestation. Research is being conducted by the University of California to evaluate the full range of potential tree species that may be affected by the beetle.

CDFA is seeking input on whether to designate Euwallacea interjectus as a category “B” pest. Under this category, response to the pest would be carried out by the counties at their own discretion, not by the state. You can advise CDFA’s on this decision until 17 February. Go here.

In its proposal, CDFA notes that several tree hosts of the beetle grow throughout California. The analysis gave a risk ranking of “High (3)” in four categories: climate/host interaction, host range, dispersal and reproduction, and ecosystem-level impacts. The economic risk rank is “Medium (2)” because it might attack only stressed trees – although CDFA concedes that drought stress is common in California. The overall determination is that the consequences of Euwallacea interjectus’ introduction to California is “High (14)”. Still, CDFA proposes to leave response to this introduction up to affected counties.

Santa Cruz County is outside the areas identified by a model developed by Lynch et al. (full citation below) as being at high risk of establishment of the Euwallacea-Fusarium complex, based on analysis of sites where Euwallacea fornicatus and E. kuroshio are already established. Nearby areas are ranked at high risk; these include drier areas in the San Francisco Bay region.

There are at least three four beetles in the Euwallacea fornicatus species complex. Several look almost identical to one another; the only reliable way to tell them apart is by looking at their DNA. However, E. interjectus is substantially larger than E. fornicatus and E. kuroshio, the two already-established shothole borers causing damage in southern California.

Various members of the Euwallacea fornicatus species complex have invaded countries around the world and other parts of the United States. While many of these introductions occurred decades ago – e.g., Hawai`i, Florida, possibly Israel, there appears to have been a spurt of introductions around or after 2000. The PSHB was first detected in California in 2003; the KSHB in 2013. As of 2022, disease caused by these two complexes had spread throughout Orange, San Diego, Los Angeles, Riverside, San Bernardino and Ventura counties. Outbreaks have been detected as far north as Santa Barbara /Santa Clarita. The KSHB had “jumped” to more distant locations in San Luis Obispo and Santa Clara counties. So far, the two later detections apparently do not represent established populations. In November 2023, the PSHB beetle–pathogen complex was confirmed killing hundreds of trees in riparian forests in San Jose, in the San Francisco Bay region. Two host trees – California sycamore and valley oaks – are important in the urban forest canopy of the region

NOTE: the invasive shot hole borers and their associated fungi attacking trees in California are completely unrelated to the laurel wilt complex killing trees in the Lauraceae family in eastern States. This complex involves an ambrosia beetle Xyleborus glabratus and associated fungus Harringtonia (formerly Raffaelea) lauricola.

SOURCES

California Department of Food and Agriculture, California Pest Rating Proposal Euwallaceae interjectus (Blanford): Boxelder ambrosia beetle https://blogs.cdfa.ca.gov/Section3162/wp-content/uploads/2025/01/Euwallacea-interjectus.pdf

Comments due by February 17, 2025.

Lynch, S.C., E. Reyes-Gonzalez, E.L. Bossard, K.S. Alarcon, N.L.R. Love, A.D. Hollander, B.E. Nobua-Behrmann & G.S. Gilbert. 2024. A phylogenetic epidemiology approach to predicting the establishment of multi-host plant pests Communications Biology

Posted by Faith Campbell

www.fadingforests.org

Too Many Deer; Too Few Forest Seedlings & Wildflowers

white flowered trillium (Trillium grandiflorum); via PICRYL One of the “charismatic wildflowers” mentioned by Blossey and colleagues

Bernd Blossey, Darragh Hare, and Don Waller have published a plea that America’s federal government take the lead in formulating a new national program on managing deer. Otherwise, they fear that deer populations will not be reduced to ecologically sustainable levels. I find their argument convincing and well-sourced. I agree that Americans need to figure out how to address this threat. (The full citation is at the end of this blog).

First, Blossey and colleagues describe the damage caused by overabundant deer:

severe declines in populations of many native forest herbs and shrubs, probably including disappearing wildflowers;
their replacement by non-native species that are less palatable;
poor regeneration of many canopy hardwood species;
decreased forest resilience, lowering forests’ ability to adapt to stressors, especially climate change;
decreased ability of forests to deliver benefits that are of increasing value to many people;
increased prevalence of wildlife and human diseases associated with the spread and size of growing tick populations; and
people – and deer — killed vehicle accidents on roads.

The widespread impacts of white-tailed deer (Odocoileus virginianus) in forests of the East are well-documented (see my previous blogs for a few examples; scroll below the “Archives” to find “Categories”). Blossey and colleagues note examples of similar impacts in the West, attributed to elk (Cervus elaphus) and black-tailed and mule deer (Odocoileus hemionus).

The authors review the decimation of deer populations in earlier centuries and the efforts of state wildlife agencies to rebuild their populations during the 20^th Century. The problem, in their view, is that federal and — especially — state wildlife agencies have retained their traditional focus on managing wildlife for recreational hunters. However, recreational hunters make up a small and shrinking proportion of all Americans. Many more people now engage in “non-consumptive” enjoyment of wildlife.

lack of regeneration in Rock Creek Park, Washington D.C.; photo by Sam Sheline, NatureServe, via Flickr

State agencies’ narrow focus might partly arise from fragmented authorities. Agencies other than wildlife departments are responsible for addressing some repercussions of overabundant deer. These include threats to human health, loss of agricultural crops.

For several reasons, Blossey and colleagues call for federal leadership. They think that only a national strategy can address, in a holistic way, the interrelated deer, human health, forest, and biodiversity crises. The strategy’s goal should be to protect species that are in decline because of over-browsing by deer and to avoid further declines in environmental and human health.

The authors reason that states are tied to traditional constituencies. Also, they have difficulty acting across jurisdictional boundaries. Second, few state wildlife agencies have authority to protect plant and invertebrate species. Yet these are the taxa most directly affected by overabundant deer. Blossey and colleagues point out that, of the ~1,300 species listed under the federal Endangered Species Act, 942 are plants and 287 are invertebrates.

They point out that deer also suffer the effects of overpopulation. Chronic wasting disease is spreading. It causes a slow, agonizing death of affected animals. Another 2.1 million deer are killed each year in vehicle crashes. [According to the World Animal Foundation, the current number of deer killed in traffic crashes is 1.8 million — 300,000 fewer.) Again, these deaths are often gruesome. Finally, the principal population “control” now is death by starvation in winter. This, too, is cruel.

Blossey and colleagues say that return of large predators, even where feasible, will not result in sufficient reduction in deer populations. Nor will encouragement of greater hunting pressure on does.

They note that the federal government owns nearly 30% of the United States’ terrestrial surface area. Management is divided among many agencies – National Park Service, Fish and Wildlife Service, Bureau of Land Management, USDA Forest Service, Department of Defense, and many smaller agencies. Management approaches vary. However, it would be possible to bring them into agreement – although, in some cases, this would require new legislation.

Another issue requires resolution: federal agencies’ authority to manage wildlife on federal land.. The states have repeatedly claimed constitutional and legal authority to manage (vertebrate) wildlife on the federal lands within their borders. This assertion was countered years ago by Nie et al. (2017):

‘Federal land management agencies have an obligation, not just the discretion, to manage and conserve fish and wildlife on federal lands. … [M]ost states have not addressed the conservation obligations inherent in trust management; rather, states wish to use the notion of sovereign ownership as … a source of unilateral power but not of public responsibility. Furthermore, the states’ trust responsibilities for wildlife are subordinate to the federal government’s statutory and trust obligations over federal lands and their integral resources.’

Blossey and colleagues assert that managing wildlife (typically defined as mammals, birds, and fish) is much broader than establishing hunting seasons or methods. Furthermore, the concept of “public trust resources” means resources should be managed for all citizens, not just the fewer than 10% of US residents who hunt. A growing proportion of society expects this management to support healthy and diverse environments.

The authors stress that reducing deer overpopulations is necessary to meet numerous policy goals. These include fulfilling obligations under international treaties related to climate change, invasive species, and threatened species; restoring and conserving the nation’s forests to provide habitat; and adopting “nature-based” climate adaptations, such as carbon sequestration. They express the hope that recent presidential mandates to better quantify and value natural assets will increase awareness of the harm caused by deer overpopulation. Their proposed national strategy would develop goals and metrics to match specific environmental and human health outcomes.

Of course, management of deer must extend beyond federal property lines. This will require cooperation among states, Tribes, and private landowners.

The paper proposes the North American Waterfowl Management Plan as a model. Under this scheme the US Fish and Wildlife Service works with states, tribal governments, Mexico, and Canada to ensure accurate information on waterfowl populations a to calculate harvest levels. States implement their assigned quotas through their own regulations. Waterfowl hunters purchase Duck Stamps to fund the monitoring efforts. This program has worked well for most species covered by the program. Waterfowl are one of the few bird groups that have not declined dramatically.

Reducing deer populations will probably require lethal control. Studies indicate that at least 60% of does must be removed from a population to reduce herd sizes over time. Other means have been attempted at regional or larger landscape levels, such as sterilization, fertility control. These methods have failed even when paired with recreational hunting. Lethal approaches will probably distress many people. However, Blossey, Hare, and Waller believe the program would be supported if it is understood to be undertaken with the goal of improving the health of both humans and also the environment.

In the end, Blossey, Hare, and Waller say they are not willing to leave the killing to cars, disease, and starvation. They emphasize our moral responsibility to protect humans and the many other species that rely on diverse ecosystems. Our policies and choices created the problem, so we must try to correct it.

SOURCES

Blossey. B., D. Hare, and D.M. Waller, 2024. Where have all the flowers gone? A call for federal leadership in deer management in the US. Front. Conserv. Sci. 5:1382132. doi: 10.3389/fcosc.2024.1382132

Nie, M., C. Barns, J. Haber, J. Joly, K. Pitt and S. Zellmer. 2017. Fish and Wildlife Management on Federal Lands: Debunking State Supremacy. Environmental Law, Vol. 47, no. 4 (2017).

Posted by Faith Campbell

www.fadingforests.org

Invasive Tree Species in the U.S. Caribbean: New Attention!

African Tulip Tree (*Spathodea campanulata*) on Puerto Rico; photo by Joe Schlabotnik via Flickr

While it is widely accepted that tropical island ecosystems are especially vulnerable to invasions, there has been little attention to terrestrial bioinvaders in the Caribbean; there has been more attention to marine bioinvaders such as lionfish. I am glad that is starting to change. Here I review a new study by Potter et al. (full citation at end of this blog), supplemented by information from other recent studies, especially Poland et al.

Potter et al. used USFS Forest Inventory and Analysis (FIA) survey data to examine regeneration rates by non-native tree species introduced to the continental United States, Hawai`i, and Puerto Rico. I rejoice that they have included these tropical islands, often left out of studies. They are part of the United States and are centers of plant endemism!

Potter et al. sought to learn which individual non-indigenous tree species are regenerating sufficiently to raise concern that they will cause significant ecological and economic damage in the future. That is, those they consider highly invasive. They defined such species as those for which at least 75% of stems of that species detected by FIA surveys are in their small tree categories – saplings or seedlings. They concluded that these species are successfully reproducing after reaching the canopy so they might be more likely to alter forest ecosystem functions and services. They labelled species exhibiting 60 – 75% of stems in the “small” categories as moderately invasive.

The authors recognize that many factors might affect tree species’ regeneration success, especially at the stand level. They assert that successful reproduction reflects a suite of factors such as propagule pressure, time since invasion, and ability of a species to adapt to different environments.

As I reported in an earlier blog, link 17% of the total flora of the islands of the Caribbean archipelago – including but not limited to Puerto Rico – are not native (Potter et al.). In Puerto Rico, two-thirds of forests comprise novel tree assemblages. The FIA records the presence of 57 non-native tree species on Puerto Rico. Potter et al. identified 17 non-native tree species as highly invasive, 16 as potentially highly invasive, and two as moderately invasive. That is, 33 of 57 nonnative tree species, or 58% of those species tallied by FIA surveyors, are actual or potential high-impact bioinvaders. While on the continent only seven non-native tree species occurred on at least 2% of FIA plots across the ecoregions in which they were inventoried, on Puerto Rico 21 species occurred on at least 2% of the FIA plots (38%). They could not assess the invasiveness of the eight species that occurred only as small stems on a couple of survey plots. These species might be in the early stages of widespread invasion, or they might never be able to reproduce & spread.

The high invasion density probably reflects Puerto Rico’s small size (5,325 mi² / 1,379,000 ha); 500 years of exposure to colonial settlement and global trade; and wide-scale abandonment of agricultural land since the middle of the 20^th Century

Naming the invaders

The most widespread and common of the highly invasive non-native tree species are river tamarind (Leucaena leucocephala), on 12.6% of 294 forested plots; algarroba (Prosopis pallida) on 10.9%; and African tuliptree (Spathodea campanulata)on 6.1%. Potter et al. attribute the prevalence of some species largely to land-use history, i.e., reforestation of formerly agricultural lands. In addition, some of the moderately to highly invasive species currently provide timber and non-timber forest products, including S. campanulata, L. leucocephala, Syzgium jambos (rose apple) and Mangifera indica (mango).

Potter et al. contrast the threat posed by Spathodea campanulata with that posed by Syzgium jambo. The latteris shade tolerant and can form dense, monotypic stands under closed canopies. Because it can reproduce under its own canopy, it might be able to remain indefinitely in forests unless it is managed. In contrast S. campanulata commonly colonizes abandoned pastures. Since it is shade intolerant, it might decline in the future as other species overtop it. Meanwhile, they suggest, S. campanulata might provide habitat appropriate for the colonization of native tree species.

Second-growth forest in Caribbean National Forest “El Yunque”

Poland et al. say the threat from Syzgium jambos might be reduced by the accidentally introduced rust fungus Puccinia psidii (= Austropuccinia psidii), which has been killing rose apple in Puerto Rico. In Hawai`i, the same fungus has devastated rose apple in wetter areas.

Potter et al. note that stands dominated by L. leucocephala and Prosopis pallida in the island’s dry forests are sometimes arrested by chronic disturbance – presumably fire. However, they do not report whether other species – native or introduced – tend to replace these two after disturbance. The authors also say that areas with highly eroded soils might persist in a degraded state without trees. The prospect of longlasting bare soil or trashy scrub is certainly is alarming.

Potter et al. warn that the FIA’s sampling protocol is not designed to detect species that are early in the invasion process. However, they do advise targetting eradication or control efforts on the eight species that occurred only as small stems on a couple of survey plots. While their invasiveness cannot yet be determined, these species might be more easily managed because presumably few trees have yet reached reproductive age. They single out Schinus terebinthifolius (Brazilian pepper), since it is already recognized as moderately invasive in Hawai`i. I add that this species is seriously invasive in nearby peninsular Florida and here! APHIS recently approved release of a biocontrol insect in Florida targetting Brazilian pepper. It might easily reach nearby Puerto Rico or other islands in the Caribbean. I am not aware of native plant species in the Caribbean region that might be damaged by the biocontrol agent. However, two native Hawaiian shrubs might be harmed if/when this thrips reaches the Hawaiian Islands. Contact me for specifics, or read the accompanying blog about Potter et al. findings in Hawai`i.

Poland et al. looked at the full taxonomic range of possible bioinvaders in forest and grassland ecosystems. The Caribbean islands receive very brief coverage in the chapter on the Southeast (see Regional Summary Appendices). This chapter contains a statement that I consider unfortunate: “Introduction of species has enriched the flora and fauna of Puerto Rico and the Virgin Islands.” The chapter’s authors assert that many of the naturalized species are restoring forest conditions on formerly agricultural lands. They say that these islands’ experience demonstrates that introduced and native species can cohabitate and complement one another. I ask – but in what kind of forest? These forests, are novel communities that bear little relationship to pre-colonial biodiversity of the islands. Was not this chapter the right place to note that loss? Forests are more than CO₂sinks.

I also regret that the chapter does not mention that the Continental United States can be the source of potentially invasive species (see several examples below).

Mealybug-infested cactus at Cabo Rojo National Wildlife Refuge, Puerto Rico. Photo by Yorelyz Rodríguez-Reyes

The chapter does concede that some introduced species are causing ecological damage now. See Table A8.1. Some of these troublesome introduced species are insects:

the South American Harrisia cactus mealybug (Hypogeococcus pungens) is killing columnar cacti in the islands’ dry forests. The chapter discusses impacts on several cactus species and control efforts, especially the search for biocontrol agents.
the agave snout weevil (Scyphophorus acupunctatus), native to the U.S. Southwest and Mexico , is threatening the endemic and endangered century plant (Agave eggersiana) in St. Croix & Puerto Rico.
Tabebuia thrips (Holopothrips tabebuia) is of unknown origin. It is widespread around mainland Puerto Rico. Its impacts so far are primarily esthetic, but it does apparently feed on both native and introduced tree species in the Tabebuia and Crescentia genera.

The Caribbean discussion also devotes welcome attention to belowground invaders, i.e., earthworms. At least one species has been found in relatively undisturbed cloud forests, so it is apparently widespread. Little is known about its impact; more generally, introduced earthworms can increase soil carbon dioxide (CO₂) emissions as through speeded-up litter decomposition and soil respiration.

A factsheet issued by the British forestry research arm DEFRA reports that the pine tortoise scale Toumeyella parvicornis has caused the death of 95% of the native Caicos pine (Pinus caribaea var. bahamensis) forests in the Turks and Caicos Islands (a UK Overseas Territory). The scale is native to North America. It has recently been introduced to Italy as well as to Puerto Rico, and the Turks and Caicos Islands.

SOURCES

Lugo, A.E., J.E. Smith, K.M. Potter, H. Marcano Vega, C.M. Kurtz. 2022. The Contribution of Non-native Tree Species to the Structure & Composition of Forests in the Conterminous United States in Comparison with Tropical Islands in the Pacific & Caribbean. USFS International Institute of Tropical Forestry General Technical Report IITF-54.

Poland, T.M., Patel-Weynand, T., Finch, D., Miniat, C. F., and Lopez, V. (Eds) (2019), Invasive Species in Forests and Grasslands of the United States: A Comprehensive Science Synthesis for the United States Forest Sector. Especially the Appendix on the Southeast and Caribbean. Springer Verlag. Available gratis at https://link.springer.com/book/10.1007/978-3-030-45367-1

Potter K.M., Riitters, K.H. & Guo. Q. 2022. Non-nativetree regeneration indicates regional & national risks from current invasions. Frontiers in Forests & Global Change Front. For. Glob. Change 5:966407. doi: 10.3389/ffgc.2022.966407

Posted by Faith Campbell

www.fadingforests.org

West Coast Steps Up Efforts to Protect Ash

Oregon-ash dominated swamp in the Ankeny National Wildlife Refuge, Willamette Valley, Oregon; photo by Wyatt Williams, Oregon Department of Forestry

In April 2022 I blogged about efforts on the West Coast to prepare for arrival of the emerald ash borer (EAB).

That blog focused on Oregon ash (Fraxinus latifolia), which is an important component of riparian forests. I alerted you to the availability of ODA/ODF EAB 2018 Response Plan.

I also mentioned Oregon’s active participation in “don’t move firewood” campaigns.

California has long inspected incoming firewood. In 2021 it establishment of a state quarantine in response to APHIS ending the federal quarantine. Washington State operates a statewide trapping program for invasive insects but does not regulate firewood.

Contributions from the Tualatin Soil and Water Conservation District enabled the USDA Forest Service Dorena Genetic Resource Center to begin testing Oregon ash for resistance to EAB and related genetics work. Other funding came from the USFS Forest Health Protection program.

EAB has now been detected in Oregon — in the Willamette Valley! (See photo above, by Wyatt Williams) Concerned stakeholders have established a new newsletter to keep people informed and promote cooperative efforts.

The newsletter is “Ash across the West”.

The first issue of the newsletter provides the following information:

there are eight ash species in the West; all are vulnerable to the emerald ash borer (EAB)

Single-leaf ash (Fraxinus anomala) CA, NV, AZ, UT, NM, CO, WY

Fragrant ash (Fraxinus cuspidata) NV, AZ, NM, UT

Calif ash (Fraxinus dipetala) CA, NV, AZ, UT

Fresnillo (Fraxinus gooddingii) AZ

Gregg’s ash (Fraxinus greggii) AZ

OR ash (Fraxinus latifolia) WA, OR, CA

Chihuahuan ash (Fraxinus papillosa) AZ, NM, TX

Velvet ash (Fraxinus velutina) CA, NV, AZ, UT, NM, TX

EAB Risk Map for OR: based upon known occurrences of ash & corresponding human activities associated with known pathways of EAB introduction and establishment.
2022 status of the two field trials
- the Dorena Genetic Resource Center (DGRC): planted 600 seedlings from 27 families; 85% survival in 2022; controlling competing vegetation
- Washington State University Puyallup Research Center: planted seedlings from 26 of these families; 95% survival rate. Possible complication from a foliar disease.
Seedlings from 17 Oregon ash families (including 14 of those in the DGRC field trial) sent to Dr. Jennifer Koch (USFS) in Ohio) for EAB resistance/susceptibility testing.
Seed collections began in 2019; interrupted by COVID-19 in 2020 but resumed in 2021 and continue in 2022. Several consortia are involved in Oregon and Washington. In California and the other states, The Huntington Botanical Gardens will lead the collecting effort. Funding is from USFS Forest Health Protection. Seeds are stored for gene conservation; some are used for the field trials in Oregon and Washington and the initial EAB-resistance studies going on in Ohio.
Penn State Ash Genomic Project: Dr. Jill Hamilton is trying to create a ‘genomic passport’ for Oregon ash populations for use in establishing genotype-environment associations to inform seed transfer guidelines. If you would like to help Dr. Hamilton collect leaves for sampling, contact: Dr. Jill Hamilton at jvh6349@psu.edu

To help with seed collection, ash monitoring, documenting the importance of ash to various communities, and other activities; or to get on the mailing list for the newsletter, contact Richard Sniezko at Richard.sniezko@usda.gov

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

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

Forest Pests: What’s Improved, What’s Still to Do

sassafras – vulnerable to the rapidly spreading laurel wilt disease; photo by F.T. Campbell

In summer 2019 I posted several blogs summarizing my analysis of forest pest issues after 30 years’ engagement. I reported the continuing introductions of tree-killing insects and pathogens; their relentless spread and exacerbated impacts. I noted the continued low priority given these issues in agencies tasked with preventing and solving these problems. Also, Congress provides not only insufficiently protective policies but also way too little funding. I decried the impediments created by several Administrations; anti-regulatory ideology and USDA’s emphasis on “collaborating” with “clients” rather than imposing requirements.

In my blogs, I called for renewed effort to find more effective strategies – as I had earlier advocated in my “Fading Forests” reports (link provided at the end of this blog), previous blogs, and Lovett et al. 2016

Areas of Progress

Now two years have passed. I see five areas of progress – which give me some hope.

1) Important Activities Are Better Funded than I had realized

a) The US Forest Service is putting significant effort into breeding trees resistant to the relevant pests, more than I had realized. Examples include elms and several conifer species in the West – here and here.

b) USDA has provided at least $110 million since FY2009 to fund forest pest research, control, and outreach under the auspices of the Plant Pest and Disease Disaster Prevention Program (§10201 of the Farm Bill). This total does not include additional funding for the spotted lanternfly. Funded projects, inter alia: explored biocontrol agents for Asian longhorned beetle and emerald ash borer; supported research at NORS-DUC on sudden oak death; monitored and managed red palm weevil and coconut rhinoceros beetle; and detected Asian defoliators. Clearly, many of these projects have increased scientific understanding and promoted public compliance and assistance in pest detection and management.

This section of the Farm Bill also provided $3.9 million to counter cactus pests – $2.7 million over 10 years targetting the Cactoblastis moth & here and $1.2 million over four years targetting the Harissia cactus mealybug and here.

flat-padded *Opuntia* cactus – vulnerable to the *Cactoblastis* moth; National Park Service photo

2) Additional publications have documented pests’ impacts – although I remain doubtful that they have increased decision-makers’ willingness to prioritize forest pests. Among these publications are the huge overview of invasive species published last spring (Poland et al.) and the regional overview of pests and invasive plants in the West (Barrett et al.).

3) There have been new efforts to improve prediction of various pests’ probable virulence (see recent blogs and here.

4) Attention is growing to the importance of protecting forest health as a vital tool in combatting climate change — see Fei et al., Quirion et al., and IUCN. We will have to wait to see whether this approach will succeed in raising the priority given to non-native pests by decision-makers and influential stakeholders.

5) Some politicians are responding to forest pest crises – In the US House, Peter Welch (D-VT) is the lead sponsor of H.R. 1389. He has been joined – so far – by eight cosponsors — Rep. Kuster (D-NH), Pappas (D-NH), Stefanik (R-NY), Fitzpatrick (R-PA), Thompson (D-CA), Ross (D-NC), Pingree (D-ME), and Delgado (D-NY). This bill would fund research into, and application of, host resistance! Also, it would make APHIS’ access to emergency funds easier. Furthermore, it calls for a study of ways to raise forest pests’ priority – thus partially responding to the proposal by me and others (Bonello et al. 2020; full reference at end of blog) to create federal Centers for Forest Pest Control and Prevention.

This year the Congress will begin work on the next Farm Bill – might these ideas be incorporated into that legislation?

What Else Must Be Done

My work is guided by three premises:

1) Robust federal leadership is crucial:

The Constitution gives primacy to federal agencies in managing imports and interstate trade.
Only a consistent approach can protect trees (and other plants) from non-native pests that spread across state lines.
Federal agencies have more resources than state agencies individually or in likely collective efforts – even after decades of budget and staffing cuts.

2) Success depends on a continuing, long-term effort founded on institutional and financial commitments commensurate with the scale of the threat. This requires stable funding; guidance by research and expert staff; and engagement by non-governmental players and stakeholders. Unfortunately, as I discuss below, funding has been neither adequate nor stable.

3) Programs’ effectiveness needs to be measured. Measurement must focus on outcomes, not just effort (see National Environmental Coalition on Invasive Species’ vision document).

Preventing New Introductions – Challenges and Solutions

We cannot prevent damaging new introductions without addressing two specific challenges.

1) Wood packaging continues to pose a threat despite past international and national efforts. As documented in my recent blogs, the numbers of shipping containers – presumably with wood packaging – are rising. Since 2010, CBP has detected nearly 33,000 shipments in violation of ISPM#15. The numbers of violations are down in the most recent years. However, a high proportion of pest-infested wood continues to bear the ISPM#15 mark. So, ISPM#15 is not as effective as it needs to be.

We at CISP hope that by mid-2022, a new analysis of the current proportion of wood packaging harboring pests will be available. Plus there are at least two collaborative efforts aimed at increasing industry efforts to find solutions – The Nature Conservancy with the National Wooden Pallet and Container Association; and the Cary Institute with an informal consortium of importers using wooden dunnage.

2) Imports of living plants (“plants for planting”) are less well studied so the situation is difficult to assess. However, we know this is a pathway that has often spread pests into and within the US. There have been significant declines in overall numbers of incoming shipments, but available information doesn’t tell us which types of plants – woody vs. herbaceous, plant vs. tissue culture, etc. – have decreased.

APHIS said, in a report to Congress (reference at end of blog), that introductions have been curbed – but neither that report nor other data shows me that is true.

Scientists are making efforts to improve risk assessments by reducing the number of organisms for which no information is available on their probable impacts (the “unknown unknowns”).

Solving Issues of Prevention

While I have repeatedly proposed radical revisions to the international phytosanitary agreements (WTO SPS & IPPC) that preclude prevention of unknown unknowns (see Fading Forests II and blog), I have also endorsed measures aimed at achieving incremental improvements in preventing introductions, curtailing spread, and promoting recovery of the affected host species.

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

The more radical suggestions focus on: 1) revising the US Plant Protection Act to give higher priority to preventing pests introductions than to facilitating free trade (FF II Chapter 3); 2) APHIS explicitly stating that its goal is to achieve a specific, high level of protection (FF II Chapter 3); 3) APHIS using its authority under the NAPPRA program to prohibit imports of all plants belonging to the 150 genera of “woody” plants that North America shares with Europe or Asia; 4) APHIS prohibiting use of packaging made from solid wood by countries and exporters that have a record of frequent violations of ISPM#15 in the 16 years since its implementation.

Another action leading to stronger programs would be for APHIS to facilitate outside analysis of its programs and policies to ensure the agency is applying the most effective strategies (Lovett et al. 2016). The pending Haack report is an encouraging example.

I have also suggested that APHIS broaden its risk assessments so that they cover wider categories of risk, such as all pests that might be associated with bare-root woody plants from a particular region. Such an approach could speed up analyses of the many pathways of introduction and prompt their regulation.

Also, APHIS could use certain existing programs more aggressively. I have in mind pre-clearance partnerships and Critical Control Point integrated pest management programs. APHIS should also clarify the extent to which these programs are being applied to the shipments most likely to transport pests that threaten our mainland forests, i.e. imports of woody plants belonging to genera from temperate climates. APHIS should promote more sentinel plant programs. Regarding wood packaging, APHIS could follow the lead of CBP by penalizing importers for each shipment containing noncompliant SWPM.

Getting APHIS to prioritize pest prevention over free trade in general, or in current trade agreements, is a heavy lift. At the very least, the agency should ensure that the U.S. prioritize invasive species prevention in negotiations with trading partners and in developing international trade-related agreements. I borrow here from the recent report on Canadian invasive species efforts. (I complained about APHIS’ failure to even raise invasive species issues during negotiation of a recent agricultural trade agreement with China.)

Solving Issues of Spreading Pests

The absence of an effective system to prevent a pest’s spread within the U.S. is the most glaring gap in the so-called federal “safeguarding system”. Yet this gap is rarely discussed by anyone – officials or stakeholders. APHIS quarantines are the best answer – although they are not always as efficacious as needed – witness the spread of EAB and persistence of nursery outbreaks of the SOD pathogen.

APHIS and the states continue to avoid establishing official programs targetting bioinvaders expected to be difficult to control or that don’t affect agricultural interests. Example include laurel wilt, and two boring beetles in southern California – goldspotted oak borer, Kuroshio shot hole borer and polyphagous shot hole borer and their associated fungi.

One step toward limiting pests’ spread would come from strengthening APHIS’ mandate in legislation, as suggested above. A second, complementary action would be for states to adopt quarantines and regulations more aggressively. For this to happen, APHIS would need to revise its policies on the “special needs exemption” [7 U.S.C. 7756]. Then states could adopt more stringent regulations to prevent introduction of APHIS-designated quarantine pests (Fading Forests III Chapt 3).

Finally, APHIS should not drop regulating difficult-to-control species – e.g., EAB. There are repercussions.

APHIS’ dropping EAB has not only reduced efforts to prevent the beetle’s spread to vulnerable parts of the West. It has also left states to come up with a coherent approach to regulating firewood; they are struggling to do so.

Considering interstate movement of pests via the nursery trade, the Systems Approach to Nursery Certification (SANC) program) is voluntary and was never intended to include all nurseries. Twenty-five nurseries were listed on the program’s website as of March 2020. It is not clear how many nurseries are participating now. The program ended its “pilot” phase and “went live” in January 2021. Furthermore, the program has been more than 20 years in development, so it cannot be considered a rapid response to a pressing problem.

Solving Issues of Recovery and Restoration via Resistance Breeding

I endorse the findings of two USFS scientists, Sniezko and Koch citations. They have documented the success of breeding programs when they are supported by expert staff and reliable funding, and have access to appropriate facilities. The principle example of such a facility is the Dorena Genetic Resource Center in Oregon. Regional consortia, e.g., Great Lakes Basin Forest Health Collaborative and Whitebark Pine Ecosystem Foundation are trying to overcome gaps in the system. I applaud the growing engagement of stakeholders, academic experts, and consortia. Questions remain, though, about how to ensure that these programs’ approaches and results are integrated into government programs.

resistant and vulnerable ash seedlings; photo courtesy of Jennifer Koch, USFS

In Bonello et al., I and others call for initiating resistance breeding programs early in an invasion. Often other management approaches, e.g., targetting the damaging pest or manipulating the environment, will not succeed. Therefore the most promising point of intervention is often with by breeding new or better resistance in the host. This proposal differs slightly from my suggestion in the “30 years – solutions” blog, when I suggested that USFS convene a workshop to develop consensus on breeding program’s priorities and structure early after a pest’s introduction.

Funding Shortfalls

I have complained regularly in my publications (Fading Forests reports) and blogs about inadequate funding for APHIS Plant Protection program and USFS Forest Health Protection and Research programs. Clearly the USDA Plant Pest and Disease Management and Disaster Program has supported much useful work. However, its short-term grants cannot substitute for stable, long-term funding. In recent years, APHIS has held back $14 – $15 million each year from this program to respond to plant health emergencies. (See APHIS program reports for FYs 20 and 21.) This decision might be the best solution we are likely to get to resolve APHIS’ need for emergency funds. If we think it is, we might drop §2 of H.R. 1389.

Expanding Engagement of Stakeholders

Americans expect a broad set of actors to protect our forests. However, these groups have not pressed decision-makers to fix the widely acknowledged problems: inadequate resources – especially for long-term solutions — and weak and tardy phytosanitary measures. Employees of federal and state agencies understand these issues but are restricted from outright advocacy. Where are the professional and scientific associations, representatives of the wood products industry, forest landowners, environmental NGOs and their funders, plus urban tree advocates – who could each play an important role? The Entomological Society’s new “Challenge” is a welcome development and one that others could copy.

SOURCES

Bonello, P., Campbell, F.T., Cipollini, D., Conrad, A.O., Farinas, C., Gandhi, K.J.K., Hain, F.P., Parry, D., Showalter, D.N, Villari, C. and Wallin, K.F. (2020) Invasive Tree Pests Devastate Ecosystems—A Proposed New Response Framework. Front. For. Glob. Change 3:2. doi: 10.3389/ffgc.2020.00002

Green, S., D.E.L. Cooke, M. Dunn, L. Barwell, B. Purse, D.S. Chapman, G. Valatin, A. Schlenzig, J. Barbrook, T. Pettitt, C. Price, A. Pérez-Sierra, D. Frederickson-Matika, L. Pritchard, P. Thorpe, P.J.A. Cock, E. Randall, B. Keillor and M. Marzano. 2021. PHYTO-THREATS: Addressing Threats to UK Forests and Woodlands from Phytophthora; Identifying Risks of Spread in Trade and Methods for Mitigation. Forests 2021, 12, 1617 https://doi.org/10.3390/f12121617ý

Krishnankutty, S., H. Nadel, A.M. Taylor, M.C. Wiemann, Y. Wu, S.W. Lingafelter, S.W. Myers, and A.M. Ray. 2020. Identification of Tree Genera Used in the Construction of Solid Wood-Packaging Materials That Arrived at U.S. Ports Infested With Live Wood-Boring Insects. Journal of Economic Entomology 2020, 1 – 12

Liebhold, A.M., E.G. Brockerhoff, L.J. Garrett, J.L. Parke, and K.O. Britton. 2012. Live plant imports: the major pathway for forest insect and pathogen invasions of the US. Front. Ecol. Environ. 2012; 10(3):135-143

Lovett, G.M., M. Weiss, A.M. Liebhold, T.P. Holmes, B. Leung, K.F. Lambert, D.A. Orwig, F.T. Campbell, J. Rosenthal, D.G. McCullough, R. Wildova, M.P. Ayres, C.D. Canham, D.R. Foster, SL. Ladeau, and T. Weldy. 2016. NIS forest insects and pathogens in the US: Impacts and policy options. Ecological Applications, 26(5), 2016, pp. 1437–1455

Mech, A.M., K.A. Thomas, T.D. Marsico, D.A. Herms, C.R. Allen, M.P. Ayres, K.J. K. Gandhi, J. Gurevitch, N.P. Havill, R.A. Hufbauer, A.M. Liebhold, K.F. Raffa, A.N. Schulz, D.R. Uden, & P.C. Tobin. 2019. Evolutionary history predicts high-impact invasions by herbivorous insects. Ecol Evol. 2019 Nov; 9(21): 12216–12230.

Poland, T.M., Patel-Weynand, T., Finch, D., Miniat, C. F., and Lopez, V. (Eds) (2019), Invasive Spp in Forests and Grasslands of the US: A Comprehensive Science Synthesis for the US Forest Sector. Springer Verlag. (in press).

Roy, B.A., H.M Alexander, J. Davidson, F.T Campbell, J.J Burdon, R. Sniezko, and C. Brasier. 2014. Increasing forest loss worldwide from invasive pests requires new trade regulations. Front Ecol Environ 2014; 12(8): 457–465

Schulz, A.N., A.M. Mech, M.P. Ayres, K. J. K. Gandhi, N.P. Havill, D.A. Herms, A.M. Hoover, R.A. Hufbauer, A.M. Liebhold, T.D. Marsico, K.F. Raffa, P.C. Tobin, D.R. Uden, K.A. Thomas. 2021. Predicting non-native insect impact: focusing on the trees to see the forest. Biological Invasions.

United States Department of Agriculture Animal and Plant Health Inspection Service. Report on the Arrival in the US of Forest Pests Through Restrictions on the Importation of Certain Plants for Planting. https://www.caryinstitute.org/sites/default/files/public/downloads/usda_forest_pest_report_2021.pdf

Posted by Faith Campbell