The Eastern deciduous forest is large and important ecologically. The forest is important for biological diversity: it shelters many endangered species, especially plants, molluscs and fish, mammals, and reptiles. In addition, the majority of forest carbon stocks in the U.S. are those of theeastern states.
But the Eastern deciduous forest is also under many anthropogenic stresses – including high numbers of non-native insects and pathogens, Liebhold map high numbers of invasive plants, blog invasive earthworms, blog browsing by overabundant deer, and timber extraction. In the southern portions of the forest, human populations are expanding, resulting in landscape fragmentation (USDA FS 2023b RPA, full reference at end of this blog).
map showing number of non-native pests in each county, as of ~2010
Agency and academic scientists in the USDA Forest Service Eastern Region (Maine to Minnesota; Delaware to West Virginia, then north of the Ohio River to Missouri) are trying to understand how long-term, continuous stressors, like deer browsing and invasive plants and earthworms, – interact with short-term gap-forming events. They call the long-term stressors “press” disturbances to distinguish them from the short-term “pulse” disturbances (Reed, Bronson, et al.; full citation at end of this blog). Understanding the processes by which forests recover from disturbance is increasingly important. Climate change is expected to raise the frequency and intensity of catastrophic natural disturbances (Spicer and Reed, Royo et al.).
The scientists emphasize that the impacts of these stressors – and effective solutions — vary depending on context.
Invasive Earthworms
USDA APHIS is responsible for regulating introduction of new species. For earthworms, APHIS’ principal concern is clearly the possibility that imported worms or soil might transport pathogens. However, the agency’s website does mention worms’ ability to disrupt the soil and possibly cause undesirable impacts on plant growth and diversity. At the 2023 National Plant Board meeting in early August 2023, Gregg Goodman, Senior Agriculturalist in APHIS PPQ NPB website for agenda? discussed issues that he considers when evaluating whether to grant permits for importing earthworms. APHIS allows imports to be used for fish bait. Dr. Goodman explained that APHIS surveyed fishermen to determine where they dump unused bait. He found no damage to plants along streams, etc. where they are dumped. A state plant health official from a northern state and I objected that the ecosystem damage caused by earthworms is well documented and we doubted that dumping of bait is not a pathway for introducing worms into natural areas.
Reed, Bronson et al. found lower earthworm biomass and density in both deer exclosures and canopy gaps. They hypothesize that the new plant growth associated with canopy gaps attracts deer, resulting in increased browse pressure. That browse pressure then affects the plant community, succession and forest structure. The changed plant community affects soil properties that then affect soil-dwelling fauna like earthworms. They believe the higher worm densities in closed-canopy sites might be the result of nutrient-rich tree leaf litter which provides both shelter and food. Another factor might be lack of recent soil disturbances in closed canopy sites.
While they say need more research is needed on the complex, combined effects of earthworms and deer, Reed, Bronson et al. still suggest that reducing deer populations or – where that is not possible – creating gaps might help manage earthworm invasions.
Deer Interactions
The long-term, chronic effect of excessive deer herbivory are well documented. See the many presentations at the recent Northern Hardwood research forum (USDA FS 2023b Proceedings). Most studies show that deer browsing overwhelms other disturbances, such as fire and canopy gaps that typically promote seedling diversity. However, recent results refine our understanding.
Samuel P. Reed and colleagues (Reed, Royo et al.) found that on the Allegheny Plateau of western Pennsylvania high deer densities at the time of stand initiation resulted in long-term reduced tree species diversity, density, and basal area. These responses were still detectable nearly four decades later. Stands are dominated by the unpalatable black cherry (Prunus serotina). The reduced stand density and the cherries’ narrower crowns lead to less above-ground biomass and reductions in above-ground carbon stocks. These scientists recommend that managers reduce deer populations to prevent changes in forest structure with probably long-term and important ramifications for many ecosystem functions.
Prunus serotina; photo by Awinch1001 via Flickr
Hovena et al. concurred with the importance of reducing deer densities, but suggested focussing on wet sites where, in their study, deer browsing had its greatest effects. On drier sites deer browsing had no effect on the diversity of woody plant seedlings.
Spicer et al. seek particularly to maintain a heterogeneous landscape to allow coexistence of both early- and late-successional species. In the Eastern Deciduous Forest biome, herbs, shrubs, and vines comprise 93% of the species richness of vascular plants
These authors found that the impact of deer browsing diverged depending on vegetation management actions. In wind-throw gaps where the plant community was retained, deer caused a 14% decline in shrub cover. In contrast, when scientists removed the extant vegetation at the beginning of recovery, deer exclusion caused a 67% increase in shrub cover. The authors speculate that vegetation removal stimulated abundant blackberry (Rubus species) regrowth. Where they had access (in gaps lacking exclosures), deer heavily browsed young Rubus stalks that sprouted after the competing vegetation was cut down. However, when the pre-established vegetation was not removed, older Rubus thickets might have protected other herbs and shrubs from browsing. Spicer et al. did not observe any major shifts in browse-tolerant species in deer-exclusion plots.
Invasive Shrubs
Hovena et al. found that in drier forest plots, the presence of non-native shrubs reduced native seedling abundance, richness, and diversity. Instead there were more seedlings of introduced species, including Lonicera maackii, L. morrowii, Ligustrum sp., and Rosa multiflora. They are concerned that replacement by invasive honeysuckles might be particularly strong in gaps resulting from death of ash trees caused by emerald ash borer. Woodlands could become dominated introduced shrubs, reducing diversity. Consequently, they recommend removing non-native shrubs in drier forests to promote seedling numbers and diversity.
In contrast, in wetter forests basal area of non-native shrubs did not affect introduced seedling abundance. However, the shrubs’ size did promote greater proportions of Lonicera maackii and Ligustrum seedlings. They suggest this might be the outcome of either abundant seed sources or allelopathic properties of some invasive shrubs e.g., L. maackii. In such sites, seedling diversity is already limited to plants that tolerate waterlogging. A hopeful note is that one native shrub, Lindera benzoin, seems able to prevent establishment of L. maackii.
Lonicera maackii; photo by pverdonk via Flickr
Hovena et al. do worry that death of ash trees might lead to declining transpiration rates, raising water tables, and further reducing seedling species richness and diversity.
Impact of Salvage Logging and Vegetation Removal
Spicer et al. studied how anthropogenic stressors affect succession. These scientists took advantage of tornado-caused gaps to compare interactions with deer browsing, salvage logging, and mechanical removal of the understory.
Contrary to expectations, none of these anthropogenic disturbances delayed community recovery or reduced diversity in comparison to the natural disturbance (tornado blowdown). Instead, adding either salvage logging or mechanical removal of understory vegetation substantially enhanced herbaceous species richness and shrub cover.
However, each major plant growth form responded differently. First, none of the manipulations affected species diversity or abundance of tree seedlings and saplings. Second, salvage logging in the wind-throw gaps increased species richness of herbs by 30%. Shrub abundance was doubled and cover almost tripled, but species richness did not change. Third, removing competing understory vegetation caused an increase of 23% in mean herbaceous cover. I have already discussed the impact of excluding deer.
Spicer et al. greet these increases in species richness with enthusiasm; they recommend managing to create a patchwork of combined natural and anthropogenic disturbances to promote plant diversity. However, I have some questions about which species are being promoted.
This study identified a total of 264 vascular plant species: 40 trees, 190 herbs, 15 shrubs, 17 vines, and 2 of unknown growth form. Only about half of these, 123 species, grew in portions of the mature forest not affected by either the tornado or one of the anthropogenic manipulations.
Gaps contained more plant species – as is to be expected. Natural blowdown areas where no manipulation was carried out had 49 more species than the undisturbed forest community (172 species). Blowdown sites subjected to salvage logging added another 53 species for a total of 225 species, or 102 more than the undisturbed reference forest.
A total of 17 species occurred only once in the authors’ data [= unique species]. Eight of these species grew only in the undisturbed forest. Two grew only in the tornado-impacted plots. Spicer et al. do not elaborate on whether these species are officially rare in that part of Pennsylvania – although it seems they might be. I wish Spicer et al. had addressed whether these possibly rare species might be affected by the forest management they recommend, i.e., intentionally creating a patchwork of various disturbances. An additional seven unique species were found in plots that had been subjected to an anthropogenic disturbance — either salvage logging or removal of remnant vegetation.
nodding trillium (Trillium cernuum); imperiled by restricted range or low populations; photo by Jason Ryndock, Pennsylvania Natural Heritage Program
In tornado-disturbed sites, one native species associated with areas where vegetation was left intact is one of the gorgeous wildflowers of eastern deciduous forests: a Trillium (species not indicated). The one native plant associated with plots from which vegetation was removed was a grass (unspecified).
Spicer et al. report that the proportion of the flora composed of non-native species was very similar between the salvage-logged area (7%) and the undisturbed reference forest (5%). Half of the non-native plant species (3.5% or 9 species) are listed as invasive in Pennsylvania (the article does not list them).
Spicer et al. say these non-native species are relatively uncommon and that they pose a minimal threat. They do concede that the invasive thorny shrub barberry (Berberis thunbergii) was more common in disturbed than intact areas. [I saw plenty of barberry along forest edges in Cook State Forest, which is only 100 miles away from the study site.] I think Spicer and others are too blasé since invasive plant populations can build up quickly when seed sources are present.
Spicer et al. raise two caveats. First, their results regarding the beneficial effects of salvage logging and vegetation manipulation probably will not apply to situations in which vast areas are logged.
More pertinent to us, they warn that their results would also not apply to forest areas in which propagules have been drastically depleted. This can result from previous human land-use or repeated catastrophic disturbances, such as canopy fires. Nor would their results apply to forests that are more threatened by invasive species. They note that a widespread and dense understory of multiple non-native species can create invasional meltdowns, resulting in a lasting depauperate state. This is especially the case when invaders at higher trophic levels, such as earthworms, are part of the mix.
Other Lessons
Reed, Bronson et al. conclude that forest canopies’ responses to disturbance are too variable to be measured by a single method. Evaluating proposals for management will require multiple measures. The overwhelming recommendation of presenters at the recent northern hardwoods research symposium (USDA FS 2023a Proceedings) was to adapt more flexible management strategies to promote forest sustainability and species diversity.
Hovena et al.’s principal finding is that interactions among site wetness, non-native shrubs and the total basal area of trees in the stand had the largest impacts on the species composition of seedlings. In Ohio, site wetness and chronic stressors like deer and introduced shrubs are acting together to shift seedling communities towards fewer native species. Of these three long-term “press” stresses, the interaction between introduced shrubs and soil wetness overshadowed even the impact of deer herbivory on seedling species richness and abundance. Surprisingly, site-specific characteristics – e.g., wetness, canopy tree competition, deer herbivory and introduced shrubs – were more influential than ash mortality in shaping woody seedling communities.
SOURCES
Hoven, B.M., K.S. Knight, V.E. Peters, D.L. Gorchov. 2022. Woody seedling community responses to deer herbivory, introduced shrubs, and ash mortality depend on canopy competition and site wetness. Forest Ecology and Management 523 (2022) 120488
Reed, S.P., D.R. Bronson, J.A. Forrester, L.M. Prudent, A.M. Yang, A.M. Yantes, P.B. Reich, and L.E. Frelich. 2023. Linked disturbance in the temperate forest: Earthworms, deer, and canopy gaps. Ecology. 2023;104:e4040. https://onlinelibrary.wiley.com/r/ecy
Reed, S.P, A.A. Royo, A.T. Fotis, K.S. Knight, C.E. Flower, and P.S. Curtis. 2022. The long-term impacts of deer herbivory in determining temperate forest stand and canopy structural complexity. Journal of Applied Ecology. 2022; 59:812-821
Spicer, M.E., A.A. Royo, J.W. Wenzel, and W.P. Carson. 2023. Understory plant growth forms respond independently to combined natural and anthropogenic disturbances. Forest Ecology and Management 543 (2023) 12077
United States Department of Agriculture. Forest Service. 2023a. Proceedings of the First Biennial Northern Hardwood Conference 2021: Bridging Science and Management for the Future. Northern Research Station General Technical Report NRS-P-211 May 2023
United States Department of Agriculture. Forest Service. 2023b. Future of America’s Forests and Rangelands. Forest Service 2020 Resources Planning Act Assessment. GTR-WO-102. July 2023 https://www.fs.usda.gov/research/treesearch/66413
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
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 20th 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 CO2 sinks.
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
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
At CISP, our hearts go out to all those affected by the terrible August fires on Maui. May the departed rest in peace. May the living find comfort and all that is needed for recovery.
Fire and Invasive Grasses
A fire in non-native grasses on Maui in 2009; photo by Forrest and Kim Starr
Major U.S. and international media continue to detail the fires’ devastation, especially in Lahaina. As time has passed, more news has highlighted the role that the widespread presence of introduced, fire-prone grasses played in the rapid growth and spread of Maui’s fires.
For example, The Washington Post devoted seven paragraphs in one story to the issue of grasses. The story quotes several experts: Alison Nugent, an associate atmospheric scientist at the University of Hawaii’s Water Resources Research Center; Jeff Masters, a meteorologist for Yale Climate Connections; and Clay Trauernicht, a fire researcher at the University of Hawaii.
These and others have been widely quoted in the many recent articles. I am glad that they – and the media – are making clear that climate change is not the sole factor causing damaging wildfires. It is clear that Maui’s recent weather patterns – including the high-velocity winds and drought – have been within the range of normal climate patterns. Fluctuations in the Pacific’s weather have also been normal, especially under the influence of the current El Niño.
The dangers caused by Hawai’i’s fire-prone grasses are also clear – and have been for years. Experts have identified policy weaknesses at the county and state level. Also, they have specified changes to land management that could better prevent or mitigate wildfires. There has been far too little action.
On the other hand, there are hopeful signs.
endangered ‘akikiki photo by Carter Atkinson, USGS
The Hawai’i Wildfire Management Organization, a nonprofit, is educating and engaging communities state-wide. Elizabeth Pickett, a Co-Executive Director, presented an overview of wildfire at the Hawai’i Invasive Species Awareness Month in February 2023. The Big Island Invasive Species Committee has successfully eradicated two species of pampas grass on Hawai’i Island – after 13 years’ work. A native species has been planted where pampas formerly grew.
Another Postarticle reported on efforts by staff and fire departments to protect the Maui Bird Conservation Center, which houses critically endangered Hawaiian birds found nowhere else on Earth, including some currently extinct in the wild. As I have blogged previously, the palila, kiwikiu, ‘akikiki, ‘alalā [Hawaiian crow; extinct in the wild] and other birds are dying from avian malaria, carried by nonnative mosquitoes. The Center on Maui and another on the Big Island are run by the San Diego Zoo Wildlife Alliance. Conservationists have completed field trials of a proposed mosquito suppression process for Maui and are seeking public comments for a similar program on Kaua’i. These programs represent groundbreaking and long-awaited progress on countering a principal threat to the survival of Hawai`i’s unique avifauna.Loss of the Center and its birds would have devastated post-suppression efforts to rebuild and restore bird populations in the wild.
The Post carried a second story about the effort to protect Hawai`i’s endangered birds – a full page of print, even longer – with many photos, on the web. The article mentions the “Birds, Not Mosquitoes” program and varying views about it. I rejoice that the dire situation for the Islands’ biodiversity is getting attention in the Nation’s capital. Again, see my earlier blog.
Plant Invasions in Hawaiian Forests
A team of scientists from the USDA Forest Service and Natural Resources Conservation Service, plus the Hawaii Division of Forestry and Wildlife, has carried out a new assessment of the extent of invasive plant species in forests on the Hawaiian Islands (Potter et al. 2023; full citation at end of blog).
The results of their analysis are – in their words – “sobering”. They portend “a more dire future for Hawai`i`s native forests.”
First, regarding the recent fires, Potter et al. found significantly higher cover by invasive grasses on Forest and Inventory Analysis (FIA) plots on Hawai‘i and Maui than on O‘ahu, Kaua‘i, and Lana‘i. Grass invasions were particularly high on the eastern coast of Maui – near Lahaina. Even so, the authors say their study’s methods resulted in a gross underestimate of areas invaded by fire-prone grasses. That is, most of Hawai’i’s xerophytic dry forests were converted to grasslands before the FIA program began. Therefore these grasslands are not included in FIA surveys.
Psidium cattleyanum; photo by Forrest and Kim Starr
The extent of current invasions in wetter forests is already significant – but trends point to an even more worrying future.
Naturalized non-native plant taxa constitute half of the Hawaiian flora.
56% of Hawaii’s 553,000 ha of forest land contained non-native tree species; about 39% of these forest lands are dominated by non-native tree species. Invasive plant species of particular concern were found in the understory of 27% of surveyed forest plots.
Across all islands, six of the ten most abundant species are non-native: Psidium cattleyanum,Schinus terebinthifolius, Leucaena leucocepahala, Ardisia elliptica, Psidium guajava, and Acacia confusa.
While less than one-third (29%) of large trees across the Islands are non-native, this proportion increases to about two-thirds of saplings (63%) and seedlings (66%). Potter et al. focus on the likelihood that plant succession will result in transformation of these forests’ canopies from native tree species to non-native species.
75% of forests in lower-elevation areas of all islands are already dominated by non-native tree species. “Only” 31% of higher-elevation forests are so dominated. These montane forests have been viewed as refugia for native species, but all are invaded to some extent – and likely to become more degraded.
Potter et al. say the high elevation forests might be more resistant to domination by non-natives. Such a result would be counter to well-documented experience, though. Even the authors report that the montane rainforests and mesophytic forests of O‘ahu and Kaua‘i are heavily invaded by non-native tree species. Such species constitute 86% or more of large trees, saplings, and seedlings in mesophytic forests; 45% of large trees and 66% of seedlings in their montane rainforests.
The most abundant tree species in Hawai`i is the invasive species Psidium cattleyanum (strawberry guava). It was recorded on 88, or37%, of 238 FIA plots. There are nearly twice as many P. cattleyanum saplings as Hawai`i’s most widespread native species, ‘ohi’a lehua (Metrosideros polymorpha).
Widescale replacement of native trees by non-native species is likely. Several factors favor these changes: 1) tree disease – rapid ‘ohi’a death has had drastic impacts on ‘ohi’a populations on several islands; 2) invasions by forbs and grasses; 3) soil damage and other disturbances caused by invasive ungulates; and 4) climate change. If succession conforms to these trends, non-native tree species could eventually constitute 75% or more of the forest tree stems and basal area on all islands and across forest types and elevations.
Loss of Hawai’i’s native tree species would be disastrous for biodiversity at the global level. More than 95% of native Hawaiian tree species are endemic, occurring nowhere else in the world.
The authors analyzed plant presence data from 238 FIA plots. Plots spanned the state’s various climates, soils, elevations, gradients, ownership, and management. However, access issues precluded inclusion of forests from several islands: Moloka‘i, Kaho’olawe, and Ni‘ihau. I know that Moloka‘i, at least, has a protected forest reserve (a Nature Conservancy property) at the island’s highest elevations.
Protecting Native Trees
Federal, state, and private landowners have carried out numerous actions to protect native forests. These efforts might be having some success. For example, forests on public lands, in conservation reserves, or in areas fenced to exclude ungulates were less impacted by non-native plants than unfenced plots, on average. However, the authors could not determine how much of this difference was the result of management or because protections were established in forests with the lowest presence of IAS species. Fencing did not prevent invasions by forbs and grasses – possibly because they are so widespread that seed sources are everywhere.
Hawaii’s two National parks (Hawai`i Volcanoes and Haleakala) have made major efforts to control invasive plants. Hawai`i Volcanoes, on the Big Island, began its efforts in the 1980s; Haleakala (on Maui) more recently. This might be one explanation for the fact that a smaller proportion of the forests on these two islands have been invaded. These efforts have not fully protected the parks, however. Low elevation native rainforests now have a high presence of non-native shrubs. Such forests on Hawai`i Island also have significant invasions by non-native woody vines, forbs and grasses.
More discouraging, intensive efforts have not returned lowland wet forest stands to a native-dominated state. Native tree species are not regenerating—even where there is plentiful seed from native canopy trees and managers have repeatedly removed competing non-native understory plants.
Potter et al. conclude that other approaches will be needed. They suggest deliberate planting of native and non-invasive non-native species or creation of small artificial gaps that might facilitate recovery of native tree species. In montane forests on Hawai`i and Maui, where native tree seedlings account for more than 70% of all tree seedlings, they propose enhancing early detection/rapid response efforts targetting invasive forbs. This would include both National parks.Certainly Haleakala National Park has this priority in mind. It launched a serious effort to try to eradicate Miconia calvescens when this tree first was detected.
Lloyd Loope, much-mourned scientist with US Geological Survey, attacking Miconia on Maui
Potter et al. note the challenge of managing remnant xerophytic dry forests, where natural regeneration of native plants has been strongly limited by invasive grasses; loss of native pollinators and seed dispersers; and the increasing frequency and intensity of droughts. They note that expanded management efforts must be implemented for decades, or longer, to be successful.
Native Trees at Risk to Nonnative Insects
Beyond the scope of the Potter et al. study is the fact that at least two dry forest endemic trees have faced their own threats from non-native insects.
The Erythrina gall wasp, Quadrastichus erythrinae, appeared in Hawai`i in 2005; it originates in east Africa. It attacks the endemic tree, wiliwili, Erythrina sandwicensis. I believe a biocontrol agent, Eurytoma erythrinae, first released in 2008, has effectively protected the wiliwili tree, lessening this threat.
The Myoporum thrips, Klambothrips myopori, from Tasmania, was detected on the Big Island in 2009. It threatens a second native tree. Naio, (Myoporum sandwicense), grows in dry forests, lowlands, upland shrublands, and mesic and wet forest habitats from sea level to 3000 m. The loss of this species would be both a signifcant loss of native biodiversity and a structural loss to native forest habitats. The thrips continues to spread; a decade after the first detection, it was found on the leeward (dry) side of Hawai`i Island with rising levels of infestation and tree dieback.
Rhus sandwicensis on Maui; photo by Forrest and Kim Starr
Two native shrubs, Hawaiian sumac Rhus sandwicensis and Dodonea viscosa, might be at risk from a biocontrol agent in the future. APHIS has approved a biocontrol for the highly invasive Brazilian pepper, Schinus terebinthifolia. Brazilian pepper is the second-most abundant non-native tree species in the State. It was found on 28 of 238 (12%) FIA plots. However, the APHIS-approved biocontrol agent is a thrips—Pseudophilothrips ichini. It is known to attack both of these two native Hawaiian shrubs. The APHIS approval allowed release of the thrips only on the mainland US. However, many insects have been introduced unintentionally from the mainland to Hawai`i. Furthermore, Hawaiian authorities were reported to be considering deliberate introduction of P. ichini to control peppertree on the Islands.
In Conclusion
In conclusion, Potter et al. found that most Hawaiian forests are now hybrid communities of native and non-native species; indeed, a large fraction are novel forests dominated by non-native trees. Business-as-usual management will probably mean that the hybrid forests – and probably those in which the canopy is currently dominated by native species—will follow successional trajectories to novel, non-native- dominated woodlands. This likelihood results in a more dire future for native plants in Hawaiian forests than has been previously described.
Potter at al. hope that their findings can guide research and conservation on other islands, especially those in the Pacific. However, Pacific islands already have the most naturalized species globally for their size—despite what was originally considered their protective geographic isolation.
SOURCE
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 https://doi.org/10.1007/s10980-023-01662-6
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
Ponderosa pine, Coconico National Forest; photograph by Brady Smith, USFS
Have you noticed, as I have, a spurt of interest in conservation of trees? I can rejoice that more people now focus on this!!!!
I have blogged previously about international and national efforts to determine not only native species deserving conservation priority – by the Morton Arboretum and IUCN but also species most threatened by non-native pests. I have also reported on growing attention to breeding tree resistance to non-native pests.
Some scientists are now focusing on species’ regeneration as a way to understand the probable future of both native and introduced species. I hope that scientists will integrate these new data with existing information on the impacts of invasive non-tree plants and tree-killing introduced pests. We need such a comprehensive picture. That will be a challenge!
Also, I hope attempts to set conservation priorities will influence decisions by governmental and non-governmental funders – and those who influence them! So far, I see little evidence that these key players are paying attention. Some Forest Service scientists and academics are pushing for expanded resistance-breeding efforts. Others are writing sophisticated analyses of non-native pests’ ecosystem impacts. But is the USDA leadership supporting stronger pest-prevention measures? Or funding for research on restoration of species? Are conservation NGOs addressing introduced forest pests?
Here, I summarize new work by Kevin Potter and his colleagues, published in two papers (full references at the end of this blog). After reading my summary, I’d like to know: What do you think? Do you agree with the focus on individual species’ regeneration to set conservation and control priorities? Do you agree with the priority species and geographic regions they suggest?? How should we resolve inconsistencies compared to the priorities suggested by the IUCN and Morton Arboretum? If you do agree, how would you suggest we move forward? If not, what approach do you think would be more useful?
A New Approach to Evaluating Species at Risk
Potter and Riitters (2022) point out that a species’ successful regeneration is key to its population’s future genetic diversity. That, in turn, determines the organisms’ ability to adapt to environmental stress and change. The latter includes, but is not limited to, climate change. Because trees are immobile and long-lived, their populations probably require substantially more genetic variation than those of other kinds of plants.
Potter and colleagues (both articles) used FIA survey data to examine regeneration rates by both tree species native to the continental United States (= CONUS) and non-native tree species introduced to CONUS, Hawai`i, or Puerto Rico. I rejoice that they have included these tropical islands, which are part of the United States and are centers of plant endemism. (Two other blogs provide details on their findings in Hawai`i and Puerto Rico.
Native Trees at Risk: Focus on Poor Regeneration
For CONUS, Potter and Riitters (2022) asked whether 280 native forest tree species are regenerating at sustainable levels, both across their full ranges and in regional portions of their ranges, defined by provisional seed zones (an area within which plant materials are assumed to be adapted). Tree species for which FIA surveys placed 75% of the stems in the sapling or seedling classes are determined to be regenerating at sustainable levels. Tree species exhibiting lower proportions of their stems in these “small tree” classes are said to be failing to regenerate adequately.
Potter and Riitters (2022) found that 46 of the 280 native tree species (16.4%) might be at risk of losing important levels of genetic variation (see the list of species in Table 2 of the article). These included high proportions of species evaluated in the following genera: two of three Platanus species; two of four Nyssa species; about 40% of Juniperus and Pinus; and five of 46 Quercus species (10.9%).
[Many areas of the eastern forest, especially in the Mid-Atlantic region, are reported by Stout, Hille, and Royo (2023) to be have insufficient advance regeneration to replace canopy trees.]
Some species appear to be headed toward outright extinction, not only loss of genetic diversity. These include four relatively rare species in California: Pinus muricata, Platanus racemosa, Pseudotsuga macrocarpa, and Sequioadendron giganteum.No seedlings or saplings are recorded on the plots on which they occurred. I note that Platanus racemosa in southern California is being attacked and killed by the Fusarium dieback vectored by the polygamous and Kuroshio shot hole borers.
Platanus racemosa riddled by invasive shot hole borer; photo by Beatriz Nobua-Behrmann, University of California Cooperative Extension
I find it alarming that a few of the possibly at-risk species have extremely wide distributions. These are Populus deltoides (eastern cottonwood), Platanus occidentalis (American sycamore), and ponderosa pine (Pinus ponderosa). Another group of species are classified as at potential risk in all their seed zones: Juniperus californica, Juniperus osteosperma, Pinus pungens, and Quercus lobata (valley oak). I note that valley oak is also under attack by the recently introduced Mediterranean oak borer. Its vulnerability is exacerbated by its relatively small range.
Potter and Riitters (2022) found distinct geographic hot spots: 15 at-risk species occur primarily in the Southeast and 14 species are in California; both represent nearly a third of the at-risk species.
In general, high rates of regeneration failure are seen in the West. Nine at-risk species (19.6% of the 46) grow in the Southwest, eight in Texas (17.4%), and four in the Rocky Mountains (8.7%). However, the Northeast and Midwest are not immune. Seven species from the former and six from the latter are also regenerating poorly. Considering pines alone, seven of 14 at-risk speciesare in the West and five in the Southeast.
Seed Zones: a Proxy for Local Genotypes
As I noted at the beginning, Potter and Riitters (2022) used USDA Forest Service provisional seed zones as a proxy for areas in which a species is presumably locally adapted. In addition to the 46 species considered failing to regenerate adequately throughout their entire ranges, Potter and Riitters (2022) determined that another 39 species are at potential risk of losing locally adapted genotypes. That is, their regeneration levels fell below the threshold in at least half of the seed zones in which they occurred. These potentially at-risk species are in the same taxonomic groups: 13 pines (33.3% of the 39 species in the category), six junipers (15.3%), and three oaks (7.7 %). These, too are concentrated in the Southeast and California: 40% are in the former — including both bald-cypress species — and 30.8% are in California. Another seven species (17.9% of the 39) are in Texas. The Midwest is home to seven species, the Northeast and Southwest each has five species (12.8%), and the Rocky Mountain region has three species (7.7%).
Bald-cypress; photo by Kej605 via WikiMedia
The seed zones with the largest numbers of species regenerating poorly are in the East, specifically the central Great Lakes region, western New York and Pennsylvania, along the Mid-Atlantic and New England coasts, and the coastal plain from southern South Carolina to eastern Texas. Potter and Riitters (2022) say these areas have such high numbers of at-risk species because they are home to so many tree species. I note [although Potter and Riitters (2022) do not] that these regions have also experienced severe levels of tree mortality due to the emerald ash borer (mature and young trees), beech leaf disease (primarily young trees), and laurel wilt disease (sub-canopy trees).
A different geographic pattern appears when considering the proportion — rather than the number — of species facing deficits in regeneration. In several Western regions, 60 – 100% of the tree species fell below the study’s threshold of 75% of recorded stems being in the sapling or seedling sizes. These seed zones are found particularly in parts of California, the Southwest, the Great Basin, and the Pacific Northwest. In none of the seed zones in the East are more than 50% of tree species in the category of potentially losing genetic variation. The implication is that while more species might be lost from parts of the East, the loss of fewer species in some Western seed zones could result in larger impacts on the composition, structure, and function of forest ecosystems there.
Potter and Riitters (2022) say that their approach has limitations because it relies on an assumption that a lack of smaller (i.e., younger) trees is an indication that a species has inadequate regeneration across all or part of its distribution and thus is vulnerable to losing genetic variation. They are not able to quantify directly the genetic variation within most forest tree species. In addition, the choice of 75% or fewer of all trees being seedlings or saplings threshold as the threshold is arbitrary. They believe these decisions are defensible.
Potter and Riitters (2022) hope that indicators of forest sustainability such as this can bridge the gap between scientists, forest managers, policy makers, and other stakeholders.
Further, the authors hope that this approach will help prioritize species most in need of: 1) monitoring for genetic diversity, 2) in situ conservation, and 3) ex situ propagule collections. In a future blog I will compare the species highlighted by Potter and Riitters (2022) to the earlier priority list developed by the IUCN and Morton Arboretum. Finally, the focus on regeneration levels could help scientists design representative sampling protocols for range-wide ex situ propagule collections for genetic diversity studies using molecular markers.
Applying This Analysis to Invasions by Non-native Trees
In a second study, Potter, Riitters, and Guo (full citation at end of this blog) flipped the focus: they used the same approach to quantify the degree of invasion by non-native trees in the U.S. I’ve blogged about this study, in general, here. Also see my separate blogs for its welcome application to Hawai`i and Puerto Rico.
Again, Potter, Riitters, and Guo hope their approach will assist in the crucial, difficult task of distinguishing between high-impact and less threatening non-native species. They warn, however, that the FIA survey procotol does not suit the needs of an early detection system.
Differentiating Invasive Tree Species’ Impacts
Potter, Riitters, and Guo note that thousands of non-native tree species have been planted around world to provide an extensive list of ecosystem services. Globally, 400 tree species have been recognized as naturalized (= consistently reproducing) or invasive (= spreading) in areas outside their native ranges. Contrary to some expectations, even relatively undisturbed forests are affected by invasive plants. In the continental United States, many fewer invasive plant species are trees than other forms/habits – shrubs, forbs, gramminoids. On the tropical islands, a much higher proportion of invasive plants are trees.
Lugo et al. (2022; full citation at end of this blog) find non-native tree species occupy a tiny fraction of the forest area of the continental United States [= CONUS], i.e., only 2.8% of the area, and only 0.4% of all tree species recorded in the FIA plots. However, these non-native tree species are widespread. They are found in 61% of forested ecosections in CONUS. Also, they are becoming more common in invaded sites. [Ecosections are divisions within 37 ecological provinces in the hierarchical framework developed by Cleland et al. (2007). There are 190 ecosections in U.S. forest biomes.]
Potter, Riitters, and Guo categorized those non-native tree species with at least 75% of stems detected by FIA surveys to be in sapling or seedling size as highly invasive. In other words, these species are successfully reproducing after reaching the canopy. So they might be more likely to alter forest functions and ecosystem services than those reproducing less robustly. They classified as species with 60 – 75% of recorded stems in these “small tree” categories as “moderately invasive.”
Potter, Riitters, and Guo suggest that control might more productively target the moderately invasive species in geographic regions where they have spread less so far – so presumably fewer seed-bearing mature specimens are present. They list as examples Picea abies, Pinus sylvestris, and Paulownia tomentosa.
In CONUS, FIA protocols specify reporting of 30 non-indigenous tree species.
Acer platanoides
Ailanthus altissima
Albizia julibrissin
Alnus glutinosa
Castanea mollissima
Casuarina lepidophloia
Cinnamomum camphora
Citrus sp.
Elaeagnus angustifolia
Eucalyptus globulus
Eucalyptus grandis
Ginko biloba
Melaleuca quinquenervia
Melia azedarach
Morus alba
Paulownia tomentosa
Picea abies
Pinus nigra
Pinus sylvestris
Populus alba
Prunus avium
Prunus persica
Salix alba
Salix sepulcralis
Sorbus aucuparia
Tamarix spp
Triadica sebifera
Ulmus pumila
Vernicia fordii
About half of these –16 species – qualified under the Potter, Riitters, and Guo criteria as highly invasive: Acer platanoides, Ailanthus altissima,Albizia julibrissin, Cinnamomum camphora, Elaegnus angustifolia, Melia azedarach, Melaleuca quinquenervia, Morus alba, Picea abies, Pinus nigra, Prunus avium, Salix alba, Salix sepulcralis, Triadica sebifera, Ulmus pumila, Vernicia fordii. An additional four taxa are ranked as potentially highly invasive: Tamarix; Eucalyptus grandis and E. globulus, Populus alba.
ring-billed gulls eating berries of Chinese tallowtree (Triadica sebifera); photo by TexasEagle via Flickr
I ask : Do YOU agree that these taxa are the most important to be tracking as potentially invasive in forests of the continental United States?
Potter, Riitters, and Guo distinguish between the most “common” and the most “widespread” invasive tree species – although they do not define the differences. Some of the most “common” or “widespread” species are not a surprise: Ailanthus altissima, Triadica sebifera (syn. Sapium sebiferum), and Acer platanoides. Ailanthus is categorized as highly invasive in 39 of 44 ecoregions in which it occurs. It is also notoriously difficult to manage. Triadica sebifera is classified as highly invasive in every one of the 20 ecoregions in which it occurs. It produces prolific seed crops that are widely dispersed by birds and water. It can invade both disturbed and undisturbed habitats. Some of the common or widespread species do surprise me: Ulmus pumila, Morus alba and Picea abies.
Most of the non-native tree species occur on only 2% of plots in the ecoregions in which they occur. However, some highly invasive trees exceed this level:
Triadica sebifera is detected on 8.6% of plots on average across 20 ecoregions;
Ulmus pumila is detected on 3.7% of plots across 39 ecoregions;
Elaeagnus angustifolia is detected on 3.3% of plots in 13 ecoregions;
Melaleuca quinquenervia is detected on 2.7% of plots in 4 ecoregions.
A. altissima is detected on only 2% of plots in the 44 ecoregions. This is surprising to me. I see it everywhere in the Mid-Atlantic – and elsewhere!
[In USFS Region 9 (24 states in the Northeast and Midwest), FIA surveys in 2019 detected Ailanthus on only 3% of plots, Norway maple and Siberian elm each on only 1% of plots (Kurz 2023).]
Eastern U.S. forests are invaded at rates several times those in Western forests, both as a proportion of plots that are invaded and the diversity of plant growth forms. The probability of invasion is highest in Eastern forests that are relatively productive and located in fragmented landscapes that contain developed or agricultural land. Non-native invasive trees are most prevalent along the Gulf Coast and in Mid-Atlantic and Midwestern States. Highly invasive non-native trees are most diverse in the ecoregions of the Mid-Atlantic and Southeast. I note that these regions also rank high in numbers of native tree species determined by Potter et al.’s other study to be reproducing an unsustainable levels.
The study found that non-native trees are almost entirely absent from the Rocky Mountain States and Alaska. However, I have seen Ailanthus in riparian areas of Utah, Arizona, and New Mexico. While few non-native tree species are recorded from ecoregions along the Pacific Coast, those areas are heavily invaded by other types of plants. Lugo et al. say those shrubs and forbs are not interfering with forest regeneration. Do YOU agree?
BLM & USFS botanists removing Spanish broom from Rogue River Canyon; photo by Stacy Johnson, BLM
On tropical islands included in the study – Hawai`i and Puerto Rico – the situation is very different. Together, these islands’ tree canopy covers less than 0.5% that of the area in the lower 48. Hawai`i is recognized as a global hotspot of non-native species richness. Naturalized non-native plant taxa constitute about half of the Hawaiian flora. The US Forest Service tracks twice as many non-native tree species in Hawai`i (62) than over the entire continental U.S. plus Alaska.
Of these 62 species, Potter, Riitters, and Guo identified 26 tree species as either highly or moderately invasive, either already or potentially highly invasive, three as moderately invasive, seven as potentially moderately invasive. In general, the richness of non-native tree species is higher in lower-elevation ecoregions, especially the lowland/leeward dry and mesic forests on O’ahu and lowland wet and mesic forests of the Big Island. [The article makes a brief reference to the probable role of rapid ʻōhiʻa death opening the canopy of the mesic and wet forests, thereby facilitating plant invasions.]Most Hawaiian ecoregions, especially those on O’ahu and Hawai’i Island, had higher non-native tree species richness than even the most highly invaded ecoregions in the lower 48 states. Parts of O’ahu & Maui had the most non-native tree species classified as highly invasive.
The Caribbean archipelago – including but not limited to Puerto Rico – has a lower proportion of non-native plant species than Hawai’i — 17% of plant species are not native. However, their presence is even higher: two-thirds of Puerto Rico’s forests comprise novel tree assemblages. This is probably because Puerto Rico has half the land area of the Hawaiian archipelago and has been part of global trade networks for 500 years instead of 200. Potter and colleagues identified 17 non-native tree species as highly invasive, 16 as potentially highly invasive, and two as moderately invasive.
On the continent only seven of 30 non-native tree species occurr on at least 2% of FIA plots across the ecoregions in which they are inventoried. Hawai’i is stunningly different: 56 of 62 species occurr on at least 2% of plots across ecoregions on average; 24 species are present on at least 10% of plots on average. One species, Psidium cattleyanum, is present on nearly half of surveyed plots across 13 ecoregions! In Puerto Rico, 21 species occurred on at least 2% of the FIA plots.
Acacia confusa – highly invasive in dry forests of Hawai`i; photo by Forrest and Kim Starr
Potter, Riitters, and Guo could not assess the invasiveness of several species that occurred only as small stems in a couple of plots. There are 11 such species on Hawai`i, eight on Puerto Rico. These species might be in the early stages of widespread invasion, or they might never be able to reproduce and spread. Despite the uncertainty, the authors suggest that eradication or control efforts targetting these species might be more cost-effective since presumably few trees have reached reproductive age yet. In Puerto Rico, they single out Schinus terebinthifolius, since it is already recognized as moderately invasive in Hawai`i [I add – seriously invasive in nearby Florida!]. However, they also emphasize the threat from one of the widespread species, Syzgium jambos, because it is a shade-tolerant species that can form dense, monotypic stands under closed canopies
I have posted separate blogs providing more details on the invasive tree species in Hawai`i and Puerto Rico.
Limits of the FIA Dataset
As in the study of native species regeneration, Potter, Riitters, and Guo specify limits arising from use of the FIA dataset. Two seem particularly pertinent to evaluation of the situation on the tropical islands.
First, they cataloged only those non-native tree species chosen by the FIA program administrators to track in the three major regions. Again, I ask YOU whether you agree with the species being recorded. Should others species be included? Should some of these species be dropped?
Second, the survey protocol does not differentiate between sites with significantly different status and history. For example, non-native trees growing on abandoned agricultural sites are counted the same way as those growing in presumably old-growth forests. They conclude that including such sites might explain the records of Eucalyptus and pine species in surveys on the islands.
Finally, as noted in the other study, the program incorporates plots that contain at least 10% canopy cover by live trees or had such cover in the past. The inventory has not included urban parks – although in recent years an urban inventory protocol has been developed.
I remind you that Potter, Riitters, and Guo warned that the FIA inventory is not designed to detect newly introduced species that are early in the invasion process.
SOURCES
Kurtz, C.M. 2023. An assessment of invasive plant species in northern U.S. forests. Res. Note NRS-311. http://doi.org/10.2737/NRS-RN-311
Lugo, A.E., J.E. Smith, K.M. Potter, H. Marcano Vega, and C.M. Kurtz. 2022. The Contribution of NIS Tree Species to the Structure and Composition of Forests in the Conterminous United States in Comparison with Tropical Islands in the Pacific & Caribbean. USDA USFS General Technical Report IITF-54.
Potter, K.M and Riitters, K. 2022. A National Multi-Scale Assessment of Regeneration Deficit as an Indicator of Potential Risk of Forest Genetic Variation Loss. Forests 2022, 13, 19. https://doi.org/10.3390/f13010019.
Potter K.M., Riitters, K.H. and Guo, Q. 2022. Non-native tree regeneration indicates regional and national risks from current invasions. Frontiers in Forests and Global Change doi: 10.3389/ffgc.2022.966407
Stout, S.L., A.T. Hille, and A.A. Royo. 2023. Science-Management Collaboration is Essential to Address Current & Future Forestry Challenges. IN United States Department of Agriculture. Forest Service. 2023. Proceedings of the First Biennial Northern Hardwood Conference 2021: Bridging Science and Management for the Future. Northern Research Station General Technical Report NRS-P-211 May 2023
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
I have blogged many times about the risk of pest introductions on imports of live plants [= “plants for planting” in USDA’s terms]. Last October I reviewed 14-year old data indicating that nearly 70% of 455 damaging tree pests introduced to the continental U.S. had probably been introduced via plant imports. These included 95% of sap feeding and 89% of foliage feeding insects and about half of the pathogens. The approach rate of pests on imported plants was apparently 12% (Liebhold et al. 2012) — more than 100 times higher than the 0.1% approach rate found by Haack et al. (2014) for wood packaging.
First, those analyses focus almost exclusively on insects (MacLachlan et al. 2022 focused on a single insect order, the Hemiptera!), despite the many pathogens probably introduced by the plant trade in recent decades. Examples I cited included several Phytophthoras, rapid ohia death, beech leaf disease, and boxwood blight. There have been repeated detections of the Ralstonia solanacearum Race 3 biovar 2.
SOD- infected rhododendrons; photo by Jennifer Parke, Oregon State University
Second, most studies analyzing the pest risk associated with plant imports use port inspection data – which are not reliable indicators of the pest approach rate – as explained by Liebhold et al. 2012 and Haack et al. 2014 (as it pertains to wood packaging).
Third, many of the studies are based on data from a decade or longer in the past. This means the studies do not address whether APHIS’ recent changes in its approach – including adoption of NAPPRA – have resulted in reduced introductions.
A complication is that, since insects are difficult to detect, those associated with the high volumes of plants imported in recent years might not be detected for years or decades after their introduction.
I have called for APHIS to update the Liebhold et al. 2012 study to determine the approach rate for all types of organisms that threaten North American tree species. Any such study should include trees on Hawai`i, Guam, Puerto Rico, and other U.S possessions and territories. These islands are nearly always excluded from analyses of imported pests. I concede that there are probably scientific and data-management challenges but these islands are immensely important from a biodiversity point of view, and they are parts of the United States!
eastern hemlocks killed by hemlock woolly adelgid; Linville Gorge; photo by Steven Norman, USFS
MacLachlan et al. (2022) estimated that new establishments – of insects in the order Hemiptera – per unit of additional plant imports have shrunk substantially. They attribute this decline to a combination of increased imports and the presence of a growing number of insect species introduced in the past. They found that introductions to the Asian Palearctic and Neotropic regions have been reduced by depletion of species pools. Other factors are thought to explain the substantial decline in establishment likelihood for the other regions. However, lag times in detecting insect introductions complicate this assessment.
However, despite that significant decrease in risk per unit of imports, MacLachlan et al. (2022) found that the number of establishments has remained relatively constant over the past century because of substantial increases in overall import levels and diversification of the origins of imports across regions, which exposed the U.S. to new source species pools.
MacLachlan et al. (2022) suggested that APHIS should target biosecurity resources to the specific commodity-country pairs associated with a higher relative risk of introducing additional insect species.
Recent studies are taking a welcome new stance: looking at links between introductions of non-native plant and insect species. I first raised this approach a year ago. Studies by teams led by Doug Tallany and Sara Lalk [Lalk et al.; articles by Tallamy] agree that:
Non-native plants – some of which are invasive – are altering ecosystems across broad swaths of North America and the impacts are insufficiently understood.
The invasive plant problem will get worse because non-native species continue to be imported, planted … and to invade.
Plant-insect interactions are the foundation of food webs – they transfer energy captured by plants through photosynthesis to other trophic levels, plus play a major role as pollinators. Consequently, changes to a region’s flora will have repercussions throughout ecosystems.
Dr. Tallamy studies the response of herbivorous insects to non-native woody plants – not just invasive plants, but also non-native plants deliberately planted as crops or ornamentals, or in forestry. Introduced plants have completely transformed the composition of plant communities in both natural and human-dominated ecosystems world-wide. The impacts can be significant: Burghardt et al. found that 75% of North American lepidopteran species and 93% of specialist species were found exclusively on native plant species.
monarch butterfly on milkweed; photograph by Jim Hudgins, USFWS
Lalk and colleagues studied the relationships between individual species of invasive woody plants and the full range of arthropod feeding guilds – pollinators, herbivores, twig and stem borers, leaf litter and soil organisms. They decry the absence of data on the complex interactions between invasive woody plants and arthropod communities at a time when invasive shrubs and trees are so widespread and causing considerable ecological damage. (See the blog for their specific research recommendations.)
Nor is the impact of non-native plants on insect fauna limited to North America. Outhwaite et al. found that the combination of climate warming and intensive agriculture is associated with reductions of almost 50% in the abundance and 27% in the number of species within insect assemblages relative to levels in less-disturbed habitats with lower rates of historical climate warming. These patterns were particularly clear in the tropics (perhaps partially because of the longer history of intensive agriculture in temperate zones). They found that high availability of nearby natural habitat (that is, native plants) can mitigate these reductions — but only in low-intensity agricultural systems.
Recognizing that plant diversity drives global patterns of insect invasion, Liebhold et al. (2023) compared various factors associated with numbers of invasive insect species in 44 land areas.They determined that the numbers of established non-native insect species are primarily driven by diversity of plants – both native and non-indigenous. Other factors, e.g., land area, latitude, climate, and insularity, strongly affect plant diversity; thus they influence insect diversity as a secondary impact. When I blogged about this study, I noted that the article appeared more than four years earlier, but has apparently had little influence on either policy formulation governing plant introductions or pest risk analysis applied to insects or pathogens that might be introduced. I suggested that we need a separate analysis of whether fungi, oomycetes, nematodes, and other pathogens show the same association with plant diversity in the receiving environment.
Studies of plant-insect relationships continue to be published. I welcome this!
Bonnamour et al. (2023) builds on the earlier studies. They also found that the presence of non-native plant species was a better predictor of insect invasions than such more widely discussed socioeconomic variables as trade volumes generally or even trade in plant products. However, detection of the associated insect invasions occurs years after detection of the plant invasions. Indeed, numbers of established non-native insect species corresponded more closely to plant introduction volumes in 1900 than current or recent import volumes.
Bonnamour et al. note that while the insect taxa that respond most directly to the non-native plant diversity are those that rely on those plants as hosts, pollinators, and plant visitors, over time those non-native herbaceous insects support introduced predators and parasites also.
Because of the “invasion debt” associated with that lag, Bonnamour et al. estimate that newly detected insect invasions will increase by 35% worldwide as a result of only recent plant introductions. They differentiate this “invasion debt” from “future invasions”, meaning the actual introduction of additional species resulting from future trade activities.
The model developed by Bonnamour et al. points to the highest numbers of newly introduced insect species occurring in areas with less capacity to deal with bioinvasions. Thus, the Afrotropics are anticipated to receive 869 new insect species, or a 10-fold increase over the number currently known to be established in the region. The Neotropics are projected to be invaded by 809 insect species, also a 10-fold increase. The Indomalayan region will probably detect 776 new insect species, a startling 20-fold increase. In reality, the “invasion debt” might not be quite this severe, since – as Bonnamour et al. note several times – the low numbers of introduced insects currently reported for these tropical regions probably partially reflect limited sampling. They note that already a high proportion of insect species intercepted by biosecurity services on imports arriving from Africa and South America are not yet recorded as established in the exporting regions.
Although both the European Palearctic and Australasia have already received many non-native insect species, their “invasion debt” is relatively high: 417 species for Europe, 317 species for Australasia.
The Neotropics are expected to be the greatest source of insect invasions in the future (904 exported species), followed by the European Palearctic (732 species).
Bonnamour et al. did not include non-native plant species used in agriculture, forestry, or ornamental horticulture. As noted above, these widespread deliberate plantings also affect insect fauna and higher trophic layers.
The greatest number of recorded insect introductions so far are in the Nearctic, Oceania (primarily Hawaii), Europe, and Australasia. While this imbalance is probably caused in part by the significantly limited sampling of non-native insect species in the Asian Palearctic and tropics, it is also true that these regions have received the majority of plant introductions through 1900. This factor has changed in the century since then; many non-native plant species have been recorded in the Afrotropics, Oceania, and Asia.
Eucalyptus plantation in Kwa-Zulu-Natal, South Africa; Kwa-Zulu-Natal Dept. of Transportation
Bonnamour et al. offer several potential explanations for the lag in detecting introduced insects compared to detecting introduced plants. First, it might be necessary for non-native host plants to reach a threshold of abundance before the associated insects are able to establish and spread. Second, reaching that threshold might require repeated introductions of the insect’s host plant species. Third, since only some of the imported plants are transporting insects, repeated imports of host plants might be necessary for the insect to achieve sufficient numbers to establish. Fourth, while their analysis included all non-native insect species, only some insect feeding guilds – herbivores and pollinators – are probably directly facilitated by introduced host plants. Fifth, plant species’ presence tends to be more quickly recorded than insects’ presence. Indeed, MacLaughlin et al. reported a median delay of 80 years between establishment and discovery of plant-feeding Hemiptera. This suggests that the actual time lag between plant and insect establishments might be shorter than the period discussed in Bonnamour et al.
Many insects from the European Palearctic have been introduced to the Nearctic; fewer insects have been introduced in the opposite direction. There is no consensus on the explanation. Thirty years ago Mattson et al. argued that there might be fewer niches for non-native insects in Europe due to the lower host plant diversity in this region caused by the Pleistocene/Holocene glaciations. On the other hand, more plant species from the European Palearctic to the Nearctic than the opposite.
Bonnamour et al. call for further research on:
1) time lags at the scale of individual insect species with their host plants.
2) effects of non-native plants used in agriculture, forestry, or ornamental horticulture.
3) whether time lags between plant and insect invasions vary among taxonomic groups, feeding guilds, or among regions.
4) effect of non-native plant abundance, rather than just species richness, on non-native insect establishment.
Recommendations
Writers about interactions of non-native plant species and insect introductions make a common plea: limit the introduction and spread of non-native plants in order to prevent future invasions of both plants and insects. Bonnamour et al. suggest including the risk of insect introductions in plant invasion risk screening tools. Earlier, the Tallamy and Lalk teams called for ending widespread planting of non-native plants.
USDA Secretary Tom Vilsack
Will policy-makers accept this advice?
I believe that these same interaction of plant host and “pest” introductions presumably applies to pathogens, too. I reiterate my frequent complaint that regulators have not responded to two or more decades of criticism of the failures of the international phytosanitary system re: insect and pathogen introductions via the international nursery trade. Examples include Brasier 2008; Liebhold el. al. 2012; Santini et al. 2013; Roy et al. 2014; Eschen et al. 2015; Jung et al. 2015; Meurisse et al. 2019; O’Hanlon et al. 2021.
As I have said earlier, I appreciate that some scientists are trying to reduce scientific uncertainty about the invasive potential of pathogens native to regions other than North America; I refer here to Jiri Hulcr (see Li et al.), Mech, and Schultz. Many more such studies are needed, addressing potential impacts on a wider variety of North American host trees and shrubs.
The late (& very much lamented!) Gary Lovett of the Cary Institute had advocated halting imports of plants that are congenerics of important North American tree species, in order to minimize the risk that pests that damage those genera will be introduced.
In January I suggested that at the global level we need:
National agricultural agencies, stakeholders, FAO & International Plant Protection Convention (IPPC) should consider amending the IPPC requirement that scientists identify a disease’s causal agents before regulating it. Experience shows that this policy virtually guarantees that pathogens will continue to enter, establish, & damage natural and agricultural environments.
National governments & FAO / IPPC should fund greatly expanded research to identify microbes resident in regions that are important sources of origin for traded plants, vulnerability of hosts in importing countries, and new technologies for detecting pathogens (e.g., molecular tools, volatile organic compounds [VOCs]).
Researchers & agencies should expand international “sentinel plants” networks; incorporate data from forestry plantations, urban plantings, etc. of non-native trees.
NPPOs should adopt regulations that apply the “systems approach” or HACCP programs outlined in ISPM#36. I had discussed these approaches in my Fading Forests III report – link at end of this blog.)
I suggested further that Americans need to
Evaluate the efficacy of current regulations – that is, implementing NAPPRA & Q-37 revision. This evaluation should be based on AQIM data, not port interception data. It should include arthropods, fungal pathogens, oomycetes, bacteria, viruses, nematodes. It should include threats to U.S. tropical islands (Hawai`i, Puerto Rico, Guam, etc.) which are centers of plant endemism.
Apply existing programs (e.g., NAPPRA, Clean Stock Network, post-entry quarantine) to strictly regulate trade in plant taxa most likely to transport pests that threaten our native plants; e.g., plants belonging to genera shared between North American trees & plants on other continents.
Recognize that plant nurseries are incubators for microbial growth, hybridization, and evolution; require nurseries to adopt sanitary operation procedures regardless of whether they sell in inter-state or intra-state commerce
SOURCES
Bonnamour, A., R.E. Blake, A.M. Liebhold, H.F. Nahrung, A. Roques, R.M. Turner, T. Yamanaka, and C. Bertelsmeier. 2023. Historical plant intros predict current insect invasions. PNAS 2023 Vol. 120 No. 24 e2221826120 https://doi.org/10.1073/pnas.2221826120
Burghardt, K. T., D. W. Tallamy, C. Philips, and K. J. Shropshire. 2010. Non-native plants reduce abundance, richness, and host specialization in lepidopteran communities. Ecosphere 1(5):art11. doi:10.1890/ES10-00032.
Lalk, S. J. Hartshorn, and D.R. Coyle. 2021. IAS Woody Plants and Their Effects on Arthropods in the US: Challenges and Opportunities. Annals of the Entomological Society of America, 114(2), 2021, 192–205 doi: 10.1093/aesa/saaa054
Li, Y., C. Bateman, J. Skelton, B. Wang, A. Black, Y-T. Huang, A. Gonzalez, M.A. Jusino, Z.J. Nolen, S. Freeman, Z. Mendel, C-Y. Chen, H-F. Li, M. Kolařík, M. Knížek, J-H. Park, W. Sittichaya, T-H.
Pham, S. Itoo, M. Torii, L. Gao, A.J. Johnson, M. Lur, J. Sun, Z. Zhang, D.C. Adams, J. Hulcr. 2022. Pre-invasion assessment of exotic bark beetle-vectored fungi to detect tree-killing pathogens. https://apsjournals.apsnet.org/doi/full/10.1094/PHYTO-01-21-0041-R
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. www.frontiersinecology.org
Liebhold, A.M., T. Yamanaka, A. Roques, S. August, S.L. Chown, E.G. Brockerhoff & P. Pyšek. 2018. Plant diversity drives global patterns of insect invasions. Sci Rep8, 12095 (2018). https://doi.org/10.1038/s41598-018-30605-4
MacLachlan, M.J., A. M. Liebhold, T. Yamanaka, M. R. Springborn. 2022. Hidden patterns of insect establishment risk revealed from two centuries of alien species discoveries. Sci. Adv. 7, eabj1012 (2021).
Mattson, W. J., P. Niemela, I. Millers, and Y. Ingauazo. 1994. Immigrant phytophagous insects on woody plants in the United States and Canada: an annotated list. USDA For. Ser. Gen. Tech. Rep. NC-169, 27 pp.
Mech, A.M., K.A. Thomas, T.D. Marisco, 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, and P.C. Tobin. 2019. Evolutionary history predicts high-impact invasions by herbivorous insects. Ecol Evol. 2019 Nov; 9(21): 12216-12230.,
Richard, M., D.W. Tallamy and A.B. Mitchell. 2019. Intro plants reduce species interactions. Biol Invasions https://doi.org/10.1007/s10530-018-1876-z
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.
Tallamy, D.W., D.L. Narango and A.B. Mitchell. 2020. Ecological Entomology (2020), DOI: 10.1111/een.12973 Do NIS plants contribute to insect declines? Conservation Biology DOI: 10.1111/j.1523-1739.2009.01202.x
Uden, D.R, A.M. Mech, N.P. Havill, A.N. Schulz, M.P. Ayres, D.A. Herms, A.M. Hoover, K.J. K. Gandhi, R.A. Hufbauer, A.M. Liebhold, T.D. M., K.F. Raffa, K.A. Thomas, P.C. Tobin, C.R. Allen. 2023. Phylogenetic risk assessment is robust for forecasting the impact of European insects on North American conifers. Ecological Applications. 2023; 33:e2761.
Posted by Faith Campbell
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
black locust – one of the most widespread invasive tree species on Earth; photo via Flickr
In recent years there has been an encouraging effort to examine bioinvasions writ large see earlier blogs re: costs of invasive species – here and here. One of these products is the Routledge Handbook of Biosecurity and Invasive Species (full citation at end of this blog). I have seen only the chapter on bioinvasion in forest ecosystems written by Sitzia et al. While they describe this situation around the globe, their examples are mostly from Europe.
Similar to other overviews, this article re-states the widely-accepted attribution of rising numbers of species introductions to globalization, especially trade. In so doing, Sitzia et al. assert that the solution is not to curtail trade and movement of people, but to improve scientific knowledge with the goal of strengthening biosecurity and control programs. As readers of this blog know, I have long advocated more aggressive application of stronger restrictions on the most high-risk pathways. Still, I applaud efforts to apply science to risk assessment.
Sitzia et al. attempt to provide a global perspective. They remind readers that all major forest ecosystems of Earth are undergoing significant change as a result of conversion to different land-uses; invasion by a wide range of non-native introduced species—including plants, insects, and mammals; and climate change. These change agents act individually and synergistically. Sitzia et al. give greater emphasis than other writers to managing the tree component of forests. They explain this focus by asserting that forest management could be either the major disturbance favoring spread of non-native species or, conversely, the only way to prevent further invasions. They explore these relationships with the goal of improving conservation of forest habitats.
Japanese stiltgrass invasion; photo by mightyjoepye via Flickr
Sitzia et al. focus first on plant invasions. They contend that – contrary to some expectations – plants can invade even dense forests despite competition for resources. They cite a recent assessment by Rejmánek & Richardson that identified 434 tree species that are invasive around Earth. Many of these species are from Asia, South America, Europe, and Australia. These non-native trees can drive not only changes in composition but also in conservation trajectories in natural forests. However, the example they cite, Japanese stilt grass (Microstegium vimineum) in the United States, is not a tree! Sitzia et al. note that in other cases it is difficult to separate the impacts of management decisions, native competitive species, and non-native species.
Sitzia et al. note that plant invasions might have a wide array of ecological impacts on forests. They attempt to distinguish between
“drivers” of environmental change – including those with such powerful effects that they call them “transformers”;
“passengers” whose invasions are facilitated by other changes in ecosystem properties; and
“backseat drivers” that benefit from changes to ecosystem processes or properties and cause additional changes to native plant communities.
An example of the last is black locust (Robinia pseudoacacia). This North American tree has naturalized on all continents. It is a good example of the management complexities raised by conflicting views of an invasive species’ value, since it is used for timber, firewood, and honey production.
Sitzia et al. then consider invasions by plant pathogens. They say that these invasions are one of the main causes of decline or extirpations in tree populations. I applaud their explicit recognition that even when a host is not driven to extinction, the strong and sudden reduction in tree numbers produces significant changes in the impacted ecosystems.
American chestnut – not extinct but ecological role gone; photo by F.T. Campbell
Sitzia et al. contend that social and economic factors determine the likelihood of a species’ transportation and introduction. Specifically, global trade in plants for planting is widely recognized as being responsible for the majority of introductions. Introductions via this pathway are difficult to regulate because of the economic importance (and political clout) of the ornamental plants industry, large volumes of plants traded, rapid changes in varieties available, and multiple origins of trade. As noted above, the authors seek to resolve these challenges by improving the scientific knowledge guiding biosecurity and control programs. In the case of plant pathogens, they suggest adopting innovative molecular techniques to improve interception efficiency, esp. in the case of latent fungi in asymptomatic plants.
The likelihood that a pathogen transported to a new region will establish is determined by biogeographic and ecological factors. Like other recent studies, Sitzia et al. attempt to identify important factors. They name a large and confusing combination of pathogen- and host-specific traits and ecosystem conditions. These include the fungus’ virulence, host specificity, and modes of action, reproduction, and dispersal, as well as the host’s abundance, demography, and phytosociology. A key attribute is the non-native fungus’ ability to exploit micro-organism-insect interactions in the introduced range. (A separate study by Raffa et al. listed Dutch elm disease as an example of this phenomenon.) I find it interesting that they also say that pathogens that attack both ornamental and forest trees spread faster. They do not discuss why this might be so. I suggest a possible explanation: the ornamental hosts are probably shipped over wide areas by the plant trade.
surviving elms in an urban environment; photo by F.T. Campbell
Sitzia et al. devote considerable attention to bioinvasions that involve symbiotic relationships between bark and ambrosia beetles and their associated fungi. These beetles are highly invasive and present high ecological risk in forest ecosystems. Since ambrosia beetle larvae feed on symbiotic fungi carried on and farmed by the adults inside the host trees, they are often polyphagous. Bark beetles feed on the tree host’s tissues directly, so they tend to develop in a more restricted number of hosts. Both can be transported in almost all kinds of wood products, where they are protected from environmental extremes and detection by inspectors. Sitzia et al. specify the usual suspects: wood packaging and plants for planting, as ideal pathways. These invasions threaten indigenous species by shifting the distribution and abundance of certain plants, altering habitats, and changing food supplies. The resulting damage to native forests induces severe alterations of the landscape and causes economic losses in tree plantations and managed forests. The latter losses are primarily in the high costs of eradication efforts – and their frequent failure.
Eucalyptus plantation in Kwa-Zulu-Natal, South Africa; photo by Kwa-Zulu-Natal Department of Transportation
Perhaps their greatest contribution is their warning about probable damage caused by invasive forest pests in tropical forests. (See an earlier blog about invasive pests in Africa.) Sitzia et al. believe that bark and ambrosia beetles introduced to tropical forests threaten to cause damage of the same magnitude as climate change and clear cutting, but there is little information about such introductions. Tropical forests are exposed to invading beetles in several ways:
1) A long history of plant movement has occurred between tropical regions. Sitzia et al. contend that the same traits sought for commercial production contribute to risk of invasion.
2) Logging and conversion of tropical forests into plantation forestry and agriculture entails movement of potentially invasive plants to new areas. Canopies, understory plant communities, and soils are all disturbed. Seeds, insects, and pathogens can be introduced via contaminated equipment.
3) Less developed nations are often at a disadvantage in managing potential invasion. Resources may be fewer, competing priorities more compelling, or potential threats less obvious.
Sitzia et al. call for development of invasive species management strategies that are relevant to and realistic for less developed countries. These strategies must account for interactions between non-native species and other aspects of global environmental change. Professional foresters have a role here. One clear need is to set out practices for dealing with conflicts between actors driven by contrasting forestry and conservation interests. These approaches should incorporate the goals of shielding protected areas, habitat types and species from bioinvasion risk. Sitzia et al. also discuss how to address the fact that many widely used forestry trees are invasive. (See my earlier blog about pines planted in New Zealand.)
planted forest in Sardinia, Italy; photo by Torvlag via Flickr
In Europe, bark beetle invasions have damaged an estimated ~124 M m2 between 1958 and 2001. Sitzia et al. report that the introduction rate of non-native scolytins has increased sharply. As in the US, many are from Asia. They expect this trend to increase in the future, following rising global trade and climate change. Southern – Mediterranean – Europe is especially vulnerable. The region has great habitat diversity; a large number of potential host trees; and the climate is dry and warm with mild winters. The region has a legacy of widespread planting of non-native trees which are now important components of the region’s economy, history and culture. These include a significant number of tree species that are controversial because they are – or appear to be – invasive. Thus, new problems related to invasive plants are likely to emerge.
Noting that different species and invasion stages require different action, Sitzia et al. point to forest planning as an important tool. Again the discussion centers on Europe. Individual states set forest policies. Two complications are the facts that nearly half of European forests are privately owned; and stakeholders differ in their understanding of the concept of “sustainability”. Does it mean ‘sustainable yield’ of timber? Or providing multiple goods and services? Or sustaining evolution of forest ecosystems with restrictions on the use of non-native species? Resolving these issues requires engagement of all the stakeholders.
Sitzia et al. say there has recently been progress. The Council of Europe issued a voluntary Code of Conduct on Invasive Alien Trees in 2017 that provides guidelines on key pathways. A workshop in 2019 elaborated global guidelines for the sustainable use of non-native tree species, based on the Bern Convention Code of Conduct on Invasive Alien Trees. The workshop issued eight recommendations:
Use native trees, or non-invasive non-native trees;
Comply with international, national, and regional regulations concerning non-native trees;
Be aware of the risk of bioinvasion and consider global change trends;
Design and adopt tailored practices for plantation site selection and silvicultural management;
Promote and implement early detection and rapid response programs;
Design and adopt practices for invasive non-native tree control, habitat restoration, and for dealing with highly modified ecosystems;
Engage with stakeholders on the risks posed by invasive NIS trees, the impacts caused, and the options for management; and
Develop and support global networks, collaborative research, and information sharing on native and non-native trees.
SOURCE
Sitzia, T., T. Campagnaro, G. Brundu, M. Faccoli, A. Santini and B.L. Webber. 2021 Forest Ecosystems. in Barker, K. and R.A. Francis. Routledge Handbook of Biosecurity and Invasive Species. ISBN 9780367763213
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
spread of non-native conifers in mountains of New Zealand; photos by Richard Bowman; New Zealand government website
As those of us who want to “do something” to counter bioinvasions struggle to mobilize both the resources and the political will necessary, I rejoice that more studies are examining what factors affect “social license” [= public approva] for such programs. One such study was recently published in New Zealand — Mason et al. (full citation at the end of the blog). New Zealand enjoys a greater appreciation of the uniqueness of its biology and awareness of invasive species’ impacts than the United States. However, their findings might provide useful guidance in the US and elsewhere.
Mason et al. sought to understand motivations of, and constraints on, those local groups responsible for controlling the spread of non-native conifers into New Zealand’s remnant native ecosystems. Non-forest ecosystems across much of the country are at risk of rapidly transforming into exotic conifer forests. For these reasons, authorities are pressing for timely removal of existing seed sources, that is, mature non-native conifer trees of several species. The blog I posted earlier apparently describes effects of conifer invasions in lowland ecosystems, whereas the Programme described here is focused on high-elevation systems.
The eradication effort in the study is the National Wilding Conifer Control Programme, establishedin 2016. A large increase in funding provided during the COVID-19 lockdown made it practical to try to eradicate seed sources from large swathes of vulnerable land. The Programme coordinates control efforts across the country, working across property and land-tenure boundaries. Landowners are expected to cover 20% of the cost of removing conifers from their land. Since removing all seed sources of high-risk conifer species from the landscape is key to achieving long-term goals, success is unlikely if significant seed sources are allowed to persist.
Mason et al. combined workshops, questionnaires, and site visits to gather data on particular aspects of this Programme. They found that social resistance, rather than lack of scientific knowledge, was often the main barrier to success in managing widespread invasive species. The authors do not address whether the fact that only 30 people provided information for their study might undermine the reliability of their findings.
map of conifer wilding sites; adapted from Wilding conifers – New Zealand history and research background, a presentation by Nick Ledgard at the “Managing wilding conifers in New Zealand – present and future” workshop (2003)
The authors suggest that the main benefit of scientific information might be to increase stakeholders’ support for management interventions — rather than to guide manager’ decisions about which strategies to pursue. To support social license, invasive species research programs might need to focus not only on cost-effective control technologies and strategies, but – perhaps especially — the benefits (both tangible and intangible) of invasive species control for society.
Mason et al. found that people were motivated to combat conifer invasions by impacts with direct influence on humans or human activities (e.g., reduced water yield, damage to infrastructure from wildfires, reduced tourist activities due to landscape transformation) and also by impacts affect ecosystems (e.g., impacts on biodiversity, aquatic ecosystems and landscapes).
People objected to control or eradication programs primarily because of social concerns. These included the unwillingness of landowners to participate and regulatory frameworks that had perverse incentives.
Mason et al. called for greater efforts by scientists to persuade stakeholders[p1] to allow removal of “wilding” conifers from private land and development of more appropriate regulations. They found that forecasting models were particularly effective in persuading people to support these efforts. It seems to me that outreach teams might need “translators” to convert scientists’ findings to information that would be more useful by stakeholders.
The authors concede that the “wilding conifer” situation has unique attributes. First, invading conifers present a stark, easily seen difference between native and invaded ecosystems. Second, some – but not all—stakeholders appreciate the uniqueness of New Zealand’s biomes. Third, the impacts of conifer invasion are sufficiently well known that they can be described succinctly and accurately.
Do these unique attributes undercut the relevance of this study to North America? It is still true that ongoing support from local stakeholders (including landowners and community groups) influences the effectiveness or profitability of managing invasive species. .It is also true that groups’ varying values affect willingness to support the activities.
Mason et al. think through the issue of stakeholders’ conflicting perspectives on the value of particular invasive species and the values threatened by that invader. These can include ethical or safety concerns around management methods, particularly regarding toxins and genetic modification. Economoic costs are also a factor – especially if the landowner must pay all or some of them.
I find it interesting that the government simultaneously funded a 5-year research program to study various issues regarding the spread, ecosystem impacts, and control of wilding conifers. The result is the Mason et al. study discussed here. I wish the U.S. funded independent analyses of its invasive species programs!
Pinus contorta – the most rapidly growing Pinus introduced to New Zealand; photo by Walter Siegmund / Wikimedia
More Details, Policy Suggestions
Workshop attendees unanimously identified landscape impacts as a reason for controlling wilding conifers. This primarily concerned the loss of New Zealand’s visual heritage or cultural identity rather than loss of native species’ habitats. When the landowner was raised in Europe, these cultural or heritage values sometimes had the opposite effect, since they see conifer forests as important components of “natural” landscapes.
Currently, landowner funding and permission is required for conifer removal. Some individual landowners want to establish new forestry plantings. Some resist removal of existing forestry plantations (which provide income) and shelter belts (which provide shelter for livestock in high country landscapes). Some landowners were unwilling to pay their 20% of removal costs. Or they objected to certain conifer control methods—particularly helicopter spraying of herbicides. New Zealand’s regulatory process also requires years of negotiating to remove standing trees – further delaying any action. In theory, landowners who resist removal could be prosecuted under the Biosecurity Act. However, this approach has never been tried for removing wilding conifers.
Mason et al. suggested several changes in policy to overcome some of these barriers.
First, forestry consultants can “game” the wilding conifer “risk calculator” to obtain government approval to establish conifer plantations in high-risk environments. The authors suggest that authorities create a “liability calculator.” Under this system, landowners wishing to retain conifers on their land for whatever reason would be liable for any subsequent containment costs. However, developing such a tool requires more finely-scaled models of conifer spread.
Second, given the high costs of combatting invading conifers if seed sources are allowed to persist, they suggested that it might be more cost-effective for the control program to pay for plantation removal under New Zealand’s Emissions Trading Scheme.
Given the overwhelmingly social and regulatory nature of barriers to success, the primary role for scientific information is providing assessments of outcomes in the absence of wilding conifer control. Preferred messages were return-on-investment estimates and forecasts of ecosystem impacts, particularly relating to biodiversity loss, water yield reduction, and wildfire hazard. Forecasts were key to demonstrating that management interventions reduced future control costs and avoided environmental impacts which large sections of the community value (i.e. biodiversity loss, reduction in water yield and agricultural productivity, increased wildfire risks). Practitioners felt that forecasting models might also channel research toward areas of high uncertainty. Mason et al. recognize the difficulties presented by inherent complexity of ecological systems. However, they think “good practice” guidelines on forecasting are emerging.
The authors find that information content and presentation need to be tailored to the various audiences – most of whom lack experience in interpreting data from environmental forecasting models. They suggest that outreach materials focus on clear illustration of the tangible and intangible benefits of wilding conifer management rather than detailed explorations of scenarios. Participants suggested ways to improve the web tool to make it more accessible to a non-expert audience.
Mason et al. mention aspects that require balancing, but don’t suggest criteria for making these choices. They say it is important to include all relevant stakeholders in invasive species management governance bodies. The absence of stakeholders with positive attitudes to wilding conifer invasions led to unanticipated external social resistance to the Programme. They recognize that including stakeholders with conflicting interests might obstruct the decision-making process. Also, in areas where there has been success in containing conifers’ spread, people can’t see invading trees, so they don’t recognize the problem. They also note that existing data do not adequately recognize risks of spread from deliberately planted seed sources such as shelter-belts, plantations and amenity plantings. The authors do not discuss how to integrate these data into analyses and public outreach.
Finally, Mason et al. recognize that many other factors strongly influence stakeholders’ willingness to support invasive species control programs, especially the level of trust and strength of relationships between bioinvasion program staff and stakeholders.
Also, they suggest topics for future research: assessing how well forecasting models are integrated with communications with stakeholders; how qualitative and quantitative research methods in different fields might support one another; and empirical tests to measure the relative effects on social license of a) involving stakeholders in developing models, b) using forecasts to assess the consequences of different management decisions and, c) the usefulness of different methods for incorporating scientific information in stakeholder engagement.
SOURCE
Mason, N.W.H., Kirk, N.A., Price, R.J. et al. Science for social license to arrest an ecosystem-transforming invasion. Biol Invasions25, 873–888 (2023). https://doi.org/10.1007/s10530-022-02953-w
see also https://www.doc.govt.nz/nature/pests-and-threats/weeds/common-weeds/wilding-conifers/
Posted by Faith Campbell
What do YOU think about the role “social license” plays in US invasive species programs? 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
spotted lanternfly; photo by Stephen Ausmus, USDA; establishment facilitated by extent of invasion by its preferred host, Ailanthus
Seven coauthors (full citation at end of blog) compared various factors associated with numbers of invasive insect species in 44 land areas.These ranged from small oceanic islands to entire continents in different world regions, Liebhold et al. determined that the numbers of established non-native insect species are primarily driven by diversity of plants, including both native and non-indigenous. Other factors, e.g., land area, latitude, climate, and insularity, strongly affect plant diversity. Through this mechanism these factors affect insect diversity as a secondary impact.
Seven coauthors (full citation at end of blog) compared various factors associated with numbers of invasive insect species in 44 land areas.These ranged from small oceanic islands to entire continents in different world regions, Liebhold et al. determined that the numbers of established non-native insect species are primarily driven by diversity of plants, including both native and non-indigenous. Other factors, e.g., land area, latitude, climate, and insularity, strongly affect plant diversity. Through this mechanism these factors affect insect diversity as a secondary impact.
At large spatial scales [greater than 10 km2], regions supporting more diverse plant communities offer greater opportunities for herbivore colonization. Thus, plant diversity promotes invasion through the “facilitation effect”. Since most insects – including most of those introduced to naïve ecosystems – are herbivores, a greater number of possible foods is a clear advantage. Those insects that prey on herbivores benefit by plant diversity indirectly.
Non-native coral tree, Erythrina, in Hawai`i; photo by Forrest and Kim Starr; did wide planting of exotic Erythrina facilitate invasion by Erythrina gall wasp?
At smaller spatial scales, plant diversity might impair the ability of insects to locate hosts because of the “dilution effect”. I have been asking for decades why so few of the Eurasian insects established in eastern North America have not also established along the Pacific coast from Oregon into British Columbia. The region has a plant-friendly climate and almost every plant species from temperate climates is grown there in cultivation. Perhaps the non-native plants – while numerous enough to become invaders themselves – are still sufficiently scarce or dispersed to impair introduced insects’ locating an familiar host?
According to the Smithsonian Institution, Hawai`i has approximately 2,499 taxa of flowering plants and 222 taxa of ferns and related groups. The native flora of the United States includes about 17,000 species of vascular plants; at least 3,800 non-native species of vascular plants are recorded as established outside cultivation. I don’t know how many non-native plant species are in cultivation.
horticultural viburnum invading riparian forest in Fairfax County, VA. photo by F.T. Campbell; did the widespread presence of many non-native viburnum species facilitate establishment of the viburnum leaf beetle?
I note that this article appeared more than four years ago. However, its important findings do not appear to have been integrated into either policy formulation governing plant introductions or pest risk analysis applied to insects or pathogens that might be introduced. (Indeed, we probably need a separate analysis of whether fungi, oomycetes, nematodes, and other pathogens show the same association with plant diversity in the receiving environment.)
How do we – government agencies, academics, conservation organizations, plant industry representatives — use this information to help curtail introductions of plant pests? Can it be integrated into APHIS’ NAPPRA process?
SOURCE
Liebhold, A.M., T. Yamanaka, A. Roques, S. August, S.L. Chown, E.G. Brockerhoff & P. Pyšek. 2018. Plant diversity drives global patterns of insect invasion. www.nature.com/scientificreports/
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
Wildfire: one of the widely recognized results of climate change (The Pioneer Fire located in the Boise National Forest near Idaho City, ID began on Jul. 18, 2016 and the cause is under investigation. The Pioneer Fire has consumed 96,469 acres. U.S. Forest Service photo. Original public domain image from Flickr)
A guest blog by Michael Aucott. Mike is a retired research scientist of the NJ Department of Environmental Protection. He has also taught chemistry at the College of New Jersey. He is currently a member of the NJDEP Science Advisory Board Standing Committee on Climate and Atmospheric Sciences, and on the board of directors of the PA/NJ Chapter of the American Chestnut Foundation. If you wish to contact Mike, use the contact button on this website. You MUST include your email address; it is not recorded automatically.
Two major perturbations affect Earth and its living systems, climate change and habitat disruptions. Emerging data show that these are more closely related than previously realized; they are connected by carbon dioxide, CO2.
Climate change basics: the physics
Climate change concerns have focused on the alteration of weather and climate due to the increase in atmospheric concentrations of greenhouse gases, primarily carbon dioxide, CO2. The impact of CO2 on climate has been understood for at least 120 years. In 1896 the Nobel-Prize-winning Swedish chemist Svante Arhennius published calculations demonstrating that human emission of CO2, when combined with the positive feedback effects of water vapor, would warm the Earth (Arhennius, 1896). His equation, ΔF = α ln(C/C0), relates the change in climate “forcing” (the degree to which temperature change is forced) to the ratio of the concentration of CO2 currently in the atmosphere (C) to a previous concentration (C0). This equation is still in use today. Arhennius estimated that a doubling of CO2 would warm the Earth by about 4 degrees C. This estimate is not far off from current estimates based on much more elaborate calculations.
This warming impact is caused by the physics of CO2, water vapor, and other “greenhouse” gases. Infrared radiation causes the CO2, water, and other greenhouse gas molecules to vibrate, leading to the absorption of the energy carried by that radiation. Much of the solar energy coming from the sun is not in the infrared frequency range, so it passes through the atmosphere without being absorbed. However, when this energy is absorbed by the surfaces of the Earth and its biota, and is re-radiated as infrared radiation, it is then absorbed by greenhouse gases, warming the planet.
The amount of water vapor in the atmosphere is directly related to the atmosphere’s temperature: warm air holds more water vapor. Human activity hasn’t directly changed the concentration of water vapor in the atmosphere significantly. But by burning fossil fuels, humans have dramatically increased the atmospheric concentration of CO2 and in so doing, also indirectly increased the concentration of water vapor. Just as Arhennius predicted over 120 years ago, this increase in CO2 is warming the Earth.
Ramifications of this warming include increased heat episodes, the intensification of the hydrological cycle (greater frequency of both heavy precipitation events and of droughts), sea level rise due to the melting of land-based glaciers and the thermal expansion of ocean water, and, almost certainly, intensification of storms and an increase in extreme weather. These climate-warming- based perturbations have the potential to influence the functioning of Earth’s biota in many deleterious ways, and clearly can be associated with the many facets of habitat disruption.
Climate change amplifications: the chemistry
But there’s another aspect of CO2 that may be more important insofar as habitat disruption is concerned and that has been largely ignored: chemistry. CO2 is a trace gas as far as we humans and other animals are concerned, unnoticed by our bodies in normal life. But to plants it is a vital food. It is taken up by plants as an essential input to photosynthesis. In this chemical reaction, using the energy of sunlight, plants combine CO2 and water vapor to make oxygen and carbohydrates, represented with a generic formula of CH2O, according to the equation CO2 + H2O → CH2O + O2. Without this reaction, life as we know it would not exist.
The atmospheric concentration of CO2 has varied over time; some 50 million years ago it was considerably higher than today. However, for at least the last three million years, the concentration of CO2 has been in the range of 280 ppm. Over these millions of years biota have adapted to this concentration. But within the last 300 years, one ten thousandth of this period – a blink of an eye in the geological or evolutionary time scale – the concentration of CO2 has shot up to 420 ppm, a 50% increase, due to humanity’s burning of fossil fuels and forests.
Imagine what might happen to a person who had been on a tight dietary budget for most of his or her life but suddenly got access to 50% more carbohydrates, but no more protein or minerals? We could expect major changes in the metabolism of that person. This dramatic change is what has, in effect, happened to the whole of life on Earth. Our planet’s primary biota, plants, now suddenly have the opportunity to gorge on CO2. But their access to other important substances such as nitrogen has not changed. Evidence is accumulating that CO2 at its elevated level of 420 ppm is not, as has been proclaimed by some, a healthy influence but is instead throwing Earth’s ecosystem into disarray.
Much of the recent experimental evidence on the impacts of enriched atmospheric CO2 has been assembled by Lewis Ziska and presented in his new book, Greenhouse Planet: How Rising CO2 Changes Plants and Life as We Know It (Ziska, 2022; see full citation at the end of the blog). The findings documented in this book reveal a variety of impacts of elevated CO2. These impacts include stimulation of growth of invasive plants, decreases in the nutrient content of major crops, and changes in plants’ production of insecticidal, allergenic, and other compounds. The changing chemistry of plants may be contributing to a major die-off of insects and insect-eating animals including birds. Below are some details.
Habitat Disruptions: Stimulation of Invasive Plants
The generally accepted explanation for why some plants are invasive is that they have been introduced to new regions where their historic predators and parasites aren’t present. Without these drags on their growth, they have flourished. That some alien plants are not browsed by white-tailed deer contributes significantly to their invasiveness in Eastern North America. Other factors are clearly involved as well, including changes in the temperature regime and the availability of water and other resources such as nitrogen.
But elevated CO2 is also a factor. In recent years, techniques for measuring responses of organisms in situ under elevated CO2 conditions have been developed, making possible investigations of the impacts of CO2 concentrations that could exist in the future under otherwise relatively realistic conditions. What the actual atmospheric CO2 concentration will be in 2050 or 2100 is difficult to predict; it depends on what humanity does to control emissions. Various scenarios suggest that levels could exceed 500 ppm by 2050 and might exceed 1000 ppm by 2100 (Tollefson, 2020).
cheatgrass; photo by Jaepil Cho
One study found that the invasive weed Canada thistle, Cirsium arvense, responds more strongly to elevated CO2 than soybean, a crop that it often plagues. In a high CO2 environment, this weed’s root system grows strongly enough to make it significantly more resistant to herbicides (Ziska, et al., 2004) (Ziska, 2010). The highly invasive and dangerously flammable cheatgrass (Bromus tectorum), nicknamed “grassoline” by the U.S. Forest Service because of its propensity to intensify wildfires, also responds strongly to elevated CO2 (Ziska, et al., 2005). Also found to be boosted by enriched CO2 is yellow star-thistle (Centaurea solstitialis), considered one of California’s worst weeds. In one study (Dukes, et al., 2011) this plant grew 600% larger in elevated CO2 relative to ambient, while native plants responded much less strongly or not at all. Japanese honeysuckle, Lonicera japonica, which plagues many areas in the U.S., was found to increase in biomass by 135% at a CO2 concentration of 675 ppm while a similar native plant, coral honeysuckle (Lonicera sempervirens) increased by only 40% (Sasek & Strain, 1991). In a field study also involving Japanese honeysuckle (Belote, et al., 2004), researchers found that its above ground net production (ANPP) approximately tripled under enriched CO2 while other plants in the trial showed showed lesser increases or actual decreases.
Other plants have been found to be selectively encouraged by enriched CO2 including cherry laurel (Prunus laurocerasus), invasive in the Pacific Northwest U.S. and the U.K., (Hattenschwiler & Korner, 2002); dalmation toadflax (Linaria dalmatica), invasive in much of North America (Blumenthal, et al., 2013); honey mesquite (Prosopis glandulosa) , invasive in Australia and parts of Africa (Polley, et al., 1996); and kudzu (Pueraria lobata), which afflicts the Southeast U.S. (Sasek & Strain, 1988). Three plants invasive in China or Southeast Asia, American rope (Mikania micrantha), Creeping oxeye (Wedelia trilobata), and a morning glory species (Ipomoea cairica), were found to produce 70.3% greater biomass when grown at a CO2 concentration of 700 ppm while three corresponding indigenous plants Paederia scandens, Wedelia chinensis and Ipomoea pescaprae, produced only 30.5% more biomass (Song, et al., 2009).
yellow star thistle; photo by Eugene Zelenko
The list goes on of studies showing increased growth of some plants under enriched CO2 conditions. As more in situ investigations are undertaken, it seems likely it will become clearer that today’s enriched level of CO2 is helping some plants to become invasive.
Complexities and contradictory findings exist however. Not all plants are stimulated by enriched CO2. An important difference in the response to higher levels of CO2 is whether a plant has a C3 or a C4 photosynthetic mechanism. C4 plants contain a biochemical pump that concentrates CO2, making them more adapted to low CO2 conditions (Hager, et al., 2016). At current levels of CO2, such plants’ need for CO2 is easily met. C3 plants do not have this CO2 concentrating ability, and so higher levels boost their growth. In a broad meta-analysis of literature, the average response to elevated CO2 of 365 C3 plant species and 37 C4 plant species was noted; the response was significantly increased in C3 species but was unchanged in C4 species (Robinson, et al. 2012). One striking example of such a difference was observed in the field study noted above (Belote, et al., 2004), wherein researchers found that Japanese honeysuckle (a C3 plant) was significantly encouraged by elevated CO2 relative to other plants at the same locale. The same study found that another aggressive invader, Japanese stiltgrass (Microstegium vimineum), a C4 plant, was unaffected or even slightly inhibited relative to competing plants’ growth by elevated CO2.
One finding is widespread; most plants studied under enriched CO2 regimes show an increase in biomass and evince a higher ratio of carbon to nitrogen (C/N ratio) in their tissues and an overall decline in nitrogen concentrations than when grown under ambient conditions. Since nitrogen is a key component of protein, this change can be expected to lead to lowered protein content of critical food crops. Some impacts of this change are already well underway because of today’s elevated CO2 concentration. Changes since 1850 in the C/N ratio and in the estimated protein content of an important plant product, pollen, were discovered in a striking study by Lewis Ziska and colleagues (Ziska, et al., 2016). Using archived museum samples, these researchers determined the nitrogen content of pollen of Solidago canadensis (Canada goldenrod) going back to the 1850s. They estimated that the protein content of goldenrod pollen, a vital nutrient for North American bees, has declined in inverse proportion to the rise in atmospheric CO2, dropping from a concentration of approximately 18% in the mid-1800s to approximately 12% today. They pointed out that it is possible that bees are now unable to provide sufficient protein and other nutrients to larvae, and that one of the main reasons for bee declines is malnutrition caused by enriched atmospheric CO2. Other studies also indicate that elevated CO2 could cause lower nitrogen concentrations in plants and lead to less proteinaceous plant parts, including pollen, being available to plant-feeding insects (Hall, et al., 2005; Knepp, et al., 2007).
bumblebee on goldenrod; photo by Keila
The changing C/N ratio is almost certainly already affecting the human food supply. As documented in an extensive review of published findings (Soares, et al., 2019), elevated CO2 has a considerable impact on the accumulation of minerals and protein in plants, with many plant species showing declines in both quality and quantity of key nutrients. These changes have worrisome implications for human nutrition and may already be responsible for increasing incidences of dietary deficiency in some areas. Lewis Ziska discusses the likely impact of rising CO2 on the future human food supply in his recent post. A number of studies showing declines in protein and also other nutrients such as zinc in food crops important to humanity are also described in Ziska’s new book, Greenhouse Planet, noted above.
Habitat Disruptions: Other Changes in Plant Chemistry
Other changes in plants besides nutritional content may be driven by enriched CO2. Plants produce a variety of secondary metabolites. Most plants use the C3 mechanism; with 50% more available of a key input, some changes in these plants’ production of such chemicals can be expected. Some changes have been observed. Mohan et al. (2006) report that enriched CO2 in an intact forest system increased water use efficiency, growth, and population biomass of poison ivy (Toxicodendron radicans) and that high-CO2 plants also produced a more toxic form of the allergenic compound urushiol.
Quercus chapmanii; photo by Mary Keim at Seminole State Forest, Florida
Many of the phytochemicals plants produce function as defenses against insect predation, and changes in such production have been found to impact herbivore feeding. For example, Landosky and Karowe (2014) suggest that specialist herbivores may have to contend with more effective chemical defenses by plants under elevated CO2. Hall, et al. (2005), in a study involving several oak and one legume species in a scrub oak ecosystem in Florida (see photo above), found that 700 ppm CO2 levels led to decreased damage to plants by four of six insect groups investigated. They did not see increases in plants’ production of carbon-based secondary metabolites, including total phenolic compounds, condensed tannins, hydrolyzable tannins, cellulose, hemicellulose, and lignin however. They concluded that the primary driver of decreased insect predation under elevated CO2 was lower overall plant nitrogen levels. They stated that the decline of nitrogen levels in foliage under elevated CO2 indicated lower foliar quality and hypothesized that the reductions in insect feeding stemmed from the combined effects of nutrient limitation and increases in parasitism and predation on the nutrient-constrained insects. They further stated that although insects try to compensate for lower nutrient content of leaves by eating more, they did not see an increased portion of damaged leaves in their study. These researchers did not report measurements of terpenoid compounds however, which are reported to represent the largest class of secondary metabolites (Wikipedia, 2022). In another study (Hall, et al., 2005a) found that concentrations of condensed tannins were higher in oak leaf litter under elevated CO2, which suggests that enhanced production of insecticidal compounds or other changes to plant tissues could affect not only insects that consume living plant tissue, but also detritivores.
Robinson et al. (2012) also investigated plants’ production of secondary metabolites in their literature review. Looking at all plant groups, they found that under elevated CO2 the production of nitrogen-based secondary metabolites (e.g., alkaloids, cyanogenic glycosides, and glucosinolates) decreased by 16% while the carbon-based secondary metabolites total phenolics, condensed tannins, and flavonoids increased by 19%, 22%, and 27% respectively. Another carbon-based metabolite, terpenoids, declined by 13%. They further divided plants into grasses, shrubs, herbs/forbs, and trees and found differing responses to elevated CO2. Trees, for example, showed increased production of total glycosides and total phenolics, little change in production of total flavonoids, and a decline in the production of total terpenes. Like Hall et al., (2005), Robinson et al. found a strong and significant decrease in nitrogen concentrations under elevated CO2 for C3 plants. A decrease did not show up for C4 plants.
In addition to chemical defenses, plants have physical characteristics such as surface waxes, trichomes, secretory canals, and tissue toughness-enhancing substances such as lignin. All of these features can reduce the edibility of plants for arthropod herbivores. Robinson et al. (2012) found consistent responses to these characteristics under elevated CO2; leaf toughness and specific leaf weight increased by 11% and 18%, respectively, while specific leaf area did not show a significant change. They concluded that there is an increase in general “toughness” of leaves under elevated CO2. As did Hall et al., (2005), Robinson et al. concluded that elevated CO2 will induce changes in plant chemistry, physiology, and morphology that are likely to impact the nutritional quality of host plants for insect herbivores.
Habitat Disruptions: Changes in Plant Chemistry and Insect Decline
Numerous studies have documented a recent and dramatic decline in insect populations and discussed the probable cascading impacts of such declines through the food chain, affecting spiders, lizards, birds, and other organisms (Samways, et al., 2020; Cardoso, et al., 2020; Sánchez-Bayoa & Wyckhuys, 2019; Tallamy & Shriver, 2021). It has been argued that the main drivers of insect species declines are habitat loss and conversion to intensive agriculture and urbanization; pollution, mainly by synthetic pesticides and fertilizers; biological factors, including pathogens and introduced species; and climate change.
But a puzzling aspect is that some insect declines have been observed in nature preserves (Vogel, 2017) that presumably are not greatly affected by most of the above drivers. One example is a study spanning 27 years that found a 76% decline in flying insect biomass at several of Germany’s protected areas subject to rather low levels of human disturbance (Hallmann et al., 2017). Another study in rainforests of Puerto Rico, apparently not subject to influences such as light pollution, habitat loss, pesticides, or agriculture, reported biomass losses between 98% and 78% for ground-foraging and canopy-dwelling arthropods over a 36-year period, (Lister and Garcia, 2018). This leaves climate change as the likely culprit. But although the varied impacts of climate change, including heat episodes, drought, and other episodes of extreme weather could impact insect populations in remote as well as populated areas, the trends observed appear to far exceed the magnitude of such climate-related disturbances over the last several decades.
tent caterpillars; Shiela Brown, Public Domain Pics
Another puzzling aspect is that not all insect orders or feeding guilds seem to be equally affected. Sanchez-Bayoa & Wychuys (2019) whose article reports on a review of 73 historical reports, state that Lepidoptera, Hymenoptera and dung beetles (Coleoptera) appear to be the taxa most affected in terrestrial environments. Robinson et al. (2012) found that phloem feeders such as Homoptera respond positively to elevated CO2 while foliage feeders/Lepidoptera respond negatively. Lepidoptera were among the most impacted; relative growth rate, fecundity, and abundance all declined under high CO2 conditions, while relative consumption rate, total consumption, and development time all increased.
Most Lepidoptera are herbivorous, feeding in their larval stage, caterpillars, on plants. Caterpillars are key components of the terrestrial ecology; in most forests of the world, caterpillars consume more living leaves than all other animals combined (Janzen, 1988). Insect herbivores such as caterpillars are near the hub of most terrestrial food webs, comprising essential food for insect predators and parasitoids, spiders, amphibians, lizards, rodents, bats, birds, and even higher predators such as foxes and bears (Burghardt et al., 2010). At least 310 North American bird species are known to feed extensively on caterpillars, and the majority of terrestrial birds rely on insects during reproduction and other nutrient-limited periods in the annual cycle (Narango, Tallamy & Marra, 2018). Caterpillars and moths are the largest component of nestling diets in hundreds of species of migrant bird species (Tallamy & Shriver, 2021); they are among the “little things that run the world” (Wilson, 1987).
Carolina chickadee; one of the birds Dr. Tallamy focuses on because it feeds its young on caterpillars; photo by Dan Pancamo; through Wikimedia
Habitat Disruptions and Climate Change: Connected by CO2
The apparently heightened rate of decline of insect herbivores such as caterpillars compared to some other insects, and the findings that many declines have been observed in areas relatively unimpacted by direct human influences such as light pollution, pesticides, and land-use change, point to the likelihood of a broad, perhaps ubiquitous, cause. Climate change is such a broad cause. Even broader and more ubiquitous is the main driver of climate change, CO2. Every plant in the world is constantly bathed in an enriched concentration of this gas. A conclusion seems likely: CO2 is not only causing global warming and climate change but is also affecting life on this planet in ways that, while still poorly understood, are already reducing the nutritive value of food crops, may be a significant cause of the spread of invasive plants, and may be the main driver of insect declines and the cascading impacts of such declines on insect-eating animals such as birds.
What to do?
To mitigate climate change and, as argued here, to mitigate habitat disruption, the steady rise in the atmosphere’s burden of CO2 must be halted, and then steps must be taken to lower the current concentration to a healthier level. These are not hopeless tasks. Although what has been a relentless rise in CO2 emissions at the global level continues, increases have slowed and even stopped in some parts of the world. Accelerating the development of low- and zero-carbon energy sources and encouraging energy conservation, as will be done through the U.S.’s Inflation Reduction Act, will further this progress.
More will be needed. Putting a significant and steadily increasing price on the carbon in fossil fuels is arguably the most important next step. Fossil fuels enjoy a free ride. The byproduct of their combustion, CO2, is dumped with little or no restrictions into the world’s atmosphere. A price on carbon would end this inequity. There are ways this could be done in a revenue-neutral (“fee and rebate”) manner that would avoid harm to economies and those with low- and moderate-incomes. A major step forward in pricing carbon by the European Union, a carbon border adjustment mechanism, is close to implementation. For more on this and other developments in cutting CO2 emissions, see the analyses and insights of the Carbon Tax Center and learn more about actions you can take to influence governments with Citizens’ Climate Lobby.
Not discussed here, but another stark example of habitat disruption is the increasing acidification of the world’s oceans caused by the dissolution of atmospheric CO2 in the waters. The ocean’s average pH has dropped from 8.2 to 8.1 within the last 200 years. Because pH is a logarithmic scale, this represents an increase in hydrogen ion concentration of over 25%, a change that is already threatening some marine creatures. More on this is available from many sources; e.g., Kolbert (2014).
References
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Blumenthal, D., V. Resco, J. Morgan, D. Williams, D. LeCain, E. Hardy, E. Pendall, and E. Bladyka, 2013, Invasive Forb Benefits from Water Savings by Native Plants and Carbon Fertilization Under Elevated CO2 and Warming, New Phytologist 200, 1156-1165.
Burghardt, Karin T., D. W. Tallamy, C. Philips, and K. Shropshire, 2010, Non-native plants reduce abundance, richness, and host specialization in lepidopteran communities, Ecosphere 1: 1-22.
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Kolbert, E., 2014, The Sixth Extinction, Henry Holt & Co., NY
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Ziska, Lewis, 2022, Greenhouse Planet: How Rising CO2 Changes Plants and Life as We Know It, Columbia University Press, NY.
Guest blog by Lewis Ziska, Associate Professor, Environmental Health Sciences at the Columbia University
[Dr. Ziska has spent his career analyzing the impacts of CO2 and climate change on plants – and therefore on people. He served as Project Leader for global climate change at the International Rice Research Institute; then spent 24 years at the USDA’s Agricultural Research Service, where he worked primarily on documenting the impact of climate change and rising carbon dioxide levels on: Crop selection improves production; Climate and agronomic pests, including chemical management; Climate, plant biology and public health impacts on food security with a focus on nutrition and pesticide use.]
No question you’ve heard the term, “Climate Change” or “Global Warming”, or my personal favorite, “Global Weirding”. The consequences are talked and discussed in the media—as they should be—but often the media, like many Americans, is focus challenged. Or in more polite terms, they have the attention span of a hummingbird on crack. Which is to say, that simple physical consequences, like sea level rise (heat melts ice!), and stranded animals on ice (Poor polar bear!), or intense storms (newscaster whipped about in the rain, yelling to be understood) are repeated, over and over again. Understandable, makes for good TV.
But it also makes you feel separate from what is happening, these consequences of climate change are to the “other”. I don’t live near the ocean, I don’t interact with polar bears; sure we have storms, but I live in the Midwest, in one of those states that begins with a vowel. Shoot, I commute to work, try and make ends meet, I’m not some damn tree hugger. Why should I care?
To understand why, you need a bit more background, some science that isn’t always available on TV or social media when it comes to global weirding.
First, while you may not be a tree hugger, you do, in fact, interact with nature. Several times. Every day. We call those times, “breakfast”, “lunch” and “dinner”.
dinner; photo by davylin via Flickr
You depend on nature for food. And clothing. And paper. And medicine. And oxygen. And construction materials (wood), and many, many other things. So, if nature gets hinky, and the climate becomes uncertain, it might be worth your while to think about climate change, or global weirding, in a different light. What I want to do here then, is to illuminate two examples that I hope will help you see why climate could affect you, directly and significantly.
Let’s begin with plants. Those green living things that comprise the bulk of the natural world (literally, if you were to weigh the natural world, 97% would be plants, 3% animals). Then let’s look at them through two different lenses—how will climate weirding alter your food; shoot, how will it alter the air that I breathe?
Let’s start with a basic food, rice. Obviously you don’t want to mess with its production, or its nutritional quality. But that is exactly what global weirding is doing.
Rice has flowers. Not big showy ones, but flowers none the less—ones that get fertilized with pollen, and seed is produced. The seed that feeds some two billion people– or about a quarter of the earth’s population.
Like all living things, plants are heat sensitive, and for rice, and many crop plants, the degree of sensitivity varies, depending on the part of the plant in question. Take a look at the table. The crops that are listed, including rice, are the core of what the world eats. Now notice the difference in temperature sensitivity. Vegetative parts of the plant, leaves and stems, are reasonably tolerant of higher temperatures, but flowers are not. Pollen, the plant equivalent of animal sperm, is highly temperature sensitive, and if the temperatures get into the high 90s (37-38oC), they become deformed, and the rice plant doesn’t produce seeds. Same for a number of plants, ones necessary to feed 8 billion people.
Crop
Opt. Temp. Vegetative
Opt. Temp. Flowering
Failure Temp.Flowering
Maize
28-35oC
18-22oC
35oC
Soybean
25-37oC
22-24oC
39oC
Wheat
20-30oC
15oC
34oC
Rice
28-35oC
23-26oC
36oC
Sorghum
26-34oC
25oC
35oC
Cotton
34oC
25-26oC
35oC
Peanut
31-35oC
20-26oC
39oC
Data are adapted from Hatfield et al., 2011.
Doubtful you’ve seen this climate threat to the global food supply on TV or a streaming service. I caught a glimpse once of temperature and agriculture on a CNN newscast, but with the “expert” calmly stating that we would just have to grow our corn in Canada, ha-ha. (Somehow, at least for rice, it’s hard to imagine India, one of the world’s largest rice producers, moving its rice production northward to the Himalaya’s, but I digress.)
Food is fundamental. If production, especially that of a global staple like rice, is impacted by rising temperatures there will be consequences. Rising prices, reduced availability, and wide-spread hunger.
But there is more to consider. Given the global dependence on rice, any change in its nutritional quality will also have effects, especially on poorer countries that rely heavily on rice as a major food source. And here we need to delve a little deeper into another aspect of climate weirding that doesn’t make it to the popular media—that rising carbon dioxide (CO2), the primary greenhouse gas, can also directly influence plant nutrition. The reasons are complicated, but in simple terms all living things consist of elements, carbon, nitrogen, phosphorous, sulfur, copper, etc., etc. A plant gets it’s carbon from the air (CO2), but everything else (nitrogen, potassium) from the soil.
And there is an imbalance. In the last 50 years, atmospheric CO2 has increased by about 30%, and is projected to increase another 50% by the end of the century. With more CO2, plants are becoming carbon rich, but nutrient poor. Nutrient poor, because while CO2 has increased in the air, nutrients in the soil have not kept pace. A perverse carb loading at the plant level.
As a consequence, rice, and many other plants, are shifting their chemistry. For example, there is a general decline in protein, in part because protein requires nitrogen. There are similar ubiquitous declines in iron and zinc, important micro-nutrients needed for human development.
Such nutritional degradation is of obvious global importance, and does, on occasion, show up on basic media when warming / weirding is mentioned, but you’d be hard pressed to find it.
Let’s move our light to another hidden bit of science. How plants can influence the air we breathe.
As humans, we like to trade things. And a large percentage of what we trade are living organisms, from fish to trees. But what began as local, regionalized trading has grown with the global population and the needs of that population—a population of 1.6 billion at the beginning of the 20th century is now ~8 billion at the beginning of the 21st. And we haven’t stopped trading. Biological trade is not inherently bad, but it represents a historically unprecedented global movement of DNA across continents, across countries, regions, towns, cities and ecosystems. And some of the DNA, when introduced, can do great harm to the environment, the economy and to human health. That harm has a name, “Invasive Species”.
Let us focus on one such plant species introduced to Eastern Europe, one that almost every American has personal experience (ACHOO!) come fall. The species is common ragweed. An invasive plant whose introduction and spread in Eastern Europe—introduced accidently through imported seeds or contaminated hay – has resulted in enormous environmental and economic losses in agriculture and public health in recent decades. In Hungary, the most important ambient biological air pollutant is: ragweed.
collecting ragweed pollen under different climates (Author’s photo)
The photo is from studies that I led looking at how ragweed pollen would respond to temperature and carbon dioxide. (If you’re curious, ragweed likes both.) Warmer temperatures, earlier Springs, later Autumns can extend its pollen season; not only extend, but increase the amount of pollen being generated. There is even some data suggesting that rising CO2 can alter pollen chemistry, making it more allergenic (REFS). Sadly, ragweed pollen doesn’t appear as temperature sensitive as that of rice, or other agricultural plants.
I wish I could say that ragweed was the exception among allergenic plants, but it’s the rule. Parthenium weed is a highly invasive species that has spread to more than 40 countries around the world. Like ragweed its pollen are highly allergenic, but it can also produce severe rashes, like poison ivy, and is known to be poisonous to livestock. It is highly aggressive, and arriving in a new location (where it has no natural enemies) can dominate landscapes, reducing biodiversity. And as with ragweed, high temperatures, longer growing seasons, heatwaves and droughts are expanding its range, and for that matter, make controlling its spread more difficult.
Such responses among invasive species will have direct impacts on air quality, especially among those (myself included) who suffer from seasonal allergies. Gasping for air is never fun.
Estimates are that pollen and seasonal asthma affects more than 24 million of us, including 6 million kids. And yet, when watching news reports of climate change, how many times have you seen a report on pollen and air quality? On increasing allergies or asthma? Once? Twice?
I could go on, (and if you need more examples, read “Greenhouse Planet”, my latest book). But my point is this: Not all of the consequences of rising carbon dioxide and climate change, warming, weirding, whatever, make for “good” TV. There is so much more to explore. So, do yourself a favor. Take a deeper dive, find out what is happening behind the scenes.
Because if we are going to rise to the challenge, we need to know what we are fighting against. Right now, the media is exemplary on showing some things, but silent on much else of importance. Watching news coverage of climate change is a bystander watching a cataclysm, and thinking, “Boy, glad I’m not experiencing THAT!”. Yet in the simplest and most basic of terms, you are, or will be, affected– from food choices to nutrition, even your allergies. And so much more.
It isn’t just about polar bears. It’s about you. Read, Understand, Act.
