drivers of extinction – Page 12 – Center for Invasive Species Prevention

Climate Change & Habitat Disruptions: Connected by Carbon Dioxide

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 CO₂has 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).

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 C₃or a C₄ photosynthetic mechanism. C₄plants 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 CO₂relative 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.

Habitat Disruptions: Changing of Plants’ C/N Ratio and Nutrient Content

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).

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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Belote, R., J. Weltzin, and R. Norby, 2004, Response of an Understory Plant Community to Elevated [CO₂] Depends on Differential Responses of Dominant Invasive Species and Is Mediated by Soil Water Availability, New Phytologist 161, 827-835.

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.

Cardoso, P., et al. 2020, Scientists’ warning to humanity on insect extinctions, Biological Conservation 242, 108426. https://doi.org/10.1016/j.biocon.2020.108426

Dukes, J., N. Chiariello, S. Loarie, and C. Field, 2011, Strong Response of an Invasive Plant Species (Centaurea solstitialis L.) to Global Environmental Changes, Ecological Applications 21, 1887-1894.

Hall, M., P. Stiling, B. Hungate, B. Drake, and M. Hunter, 2005a, Effects of elevated CO₂ and herbivore damage on litter quality in a scrub oak ecosystem, Journal of Chemical Ecology, 31, 2343-2356.

Hall, M., P. Stiling, D. Moon, B. Drake, and M. Hunter, 2005, Effects of elevated CO₂ of foliar quality and herbivore damage in a scrub oak ecosystem. Journal of Chemical Ecology 31, 267-286.

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Hattenschwiler, S. and C. Korner, 2003, Does Elevated CO₂ Facilitate Naturalization of the Non-indegenous Prunus laurocerasus in Swiss Temperate Forests?, Functional Ecology 17, 778-785.

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Knepp, R., J. Hamilton, A. Zangeri, M. Berenbaum, and E. Delucia, 2007, Foliage of oaks grown under elevated CO₂ reduces performance of Antherae Polyphemus (Lepidoptera: Saturnidae), Environmental Entomology 36, 609-617.

Kolbert, E., 2014, The Sixth Extinction, Henry Holt & Co., NY

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Ziska, L.H., J.S. Pettis, J. Edwards, J.E. Hancock, M.B. Tomecek, A. Clark, J.S. Dukes, I. Loladze, and H.W. Polley, 2016, Rising atmospheric CO₂ is reducing the protein concentration of a floral pollen source essential for North American bees, Proc. R. Soc. B, 283, 20160414, http://dx.doi.org/10.1098/rspb.2016.0414

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Global Weirding: Rising CO2 Impacts Plants & People

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”.

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 (37-38^oC), 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	28-35^oC	18-22^oC	35^oC
Soybean	25-37^oC	22-24^oC	39^oC
Wheat	20-30^oC	15^oC	34^oC
Rice	28-35^oC	23-26^oC	36^oC
Sorghum	26-34^oC	25^oC	35^oC
Cotton	34^oC	25-26^oC	35^oC
Peanut	31-35^oC	20-26^oC	39^oC

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 20^th century is now ~8 billion at the beginning of the 21^st. 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.

Now.

EAB in Eastern Europe – Worse (+ war!)

A special issue of the journal Forests (Vol. 13 2022) seeks to improve understanding of the root causes of exacerbated threats from insect pests. The issue contains 15 papers; most focus on geographic areas other than North America. The journal is open access!

Choi and Park (full citations below) link increased pest risk to climate change and increased international trade. They provide brief summaries of all 15 papers. My focus here is on two articles that provide updates on the status of the emerald ash borer (EAB Agrilus planipennis) in Russia and Ukraine. The article by Davydenko et al. also examines interactions between EAB and the invasive pathogen Hymenoscyphus fraxineus, which causes ash dieback disease. In other blogs I will look at insects linked to North America (both species from North America that threaten forests on other continents, and species in Russia that pose a threat to North America) and at the overall Russian experience.

I blogged about EAB invasion of Russia in April 2021 so this is an update.

Musolin et al. (2022) (full citations below) remind us that the EAB invasions of North America and Russia were detected almost simultaneously: in Michigan and Ontario in 2002 and in European Russia (Moscow) in 2003. They conclude that both invasions probably originated from a common source (most probably China). They date the introduction to the late 1980s or early 1990s; pathways might have been wooden crafts, wood packaging, or ash seedlings. Nate Siegert used dendrological studies to estimate a similar introduction date for the North American invasion.

European ash (*Fraxinus excelsior*) specimen in Belgium; photo by Jean-Pol Granmont

EAB has spread far in the intervening 30 + years. By early 2022, outbreaks were recorded in five Canadian provinces, 35 US states, 18 provinces and several cities in European Russia, and two provinces in Ukraine (Musolin et al. 2022) Davydenko et al. report that EAB had also established in eastern Belarus, but provide no details.

As demonstrated in the earlier blog and confirmed by Musolin et al. (2022) and Davydenko et al., the EAB has spread much faster to the southwest than directly West and to the Northwest. Davydenko et al. attribute the slower spread in the St. Petersburg area to the colder and wetter climate of this region – which is ~1200 km north of Ukraine. While the EAB reproduces in two cohorts in Eastern Ukraine, to the north the beetle requires more than one year to complete its life cycle, at least two years in the St. Petersburg area. In 2021, Musolin et al. 2021 speculated that pressure by the native parasitoid Spathius polonicus Niezabitowski might also be slowing EAB’s spread in the North. In 2022, Musolin does not address this possibility. (I note that APHIS has approved two Spathius species as biocontrol agents in the U.S.).

Musolin et al. (2022) and Davydenko et al. agree that the EAB poses real threat to ash in central and western Europe. In both the south (Davydenko et al.) and in the northwestern area around St. Petersburg ash grows in continuous stretches, linking Russia or Ukraine to Romania, Hungary, Slovakia, and Poland. These ash consist of both natural woodlands, and extensive plantings of both one of the European ash species, F. excelsior and the highly-susceptible North America green ash (F. pennsyvanica). Furthermore, the EAB is an excellent hitchhiker on vehicles & railway cars. Davydenko et al. also consider the beetle to be a strong flyer. Musolin et al. (2022) cite a separate analysis in stating that EAB can probably invade most European countries. Only some regions of Norway, Sweden, Finland, Ireland, and Great Britain are probably protected by their low temperatures.

Both articles were written too early to consider how the current war in the relevant area of Ukraine might affect spread of the EAB, although we know Ukrainians are cutting firewood. The war has certainly interrupted monitoring and other efforts.

The sources agree on EAB’s severe impacts. Musolin et al. (2022) notes that the beetle has killed millions of trees in the forests and urban plantings in North America, European Russia, and Eastern Ukraine. Davydenko et al. note that the Fraxinus genus is one of the most widely distributed tree genera in North America. They then assert that the EAB could virtually eliminate it. I know that North American scientists agree that the beetle threatens many species in the genus; but do they agree that the genus would be “virtually eliminated”? Davydenko et al. think the EAB could pose similar threat to Euro ash F. excelsior.

Musolin et al. 2022 estimate that potential economic losses in Europe could reach US$1.81 billion. By this indicator, the species ranks fourth among the most “costly” invasive pests. Russia spent an estimated US$258.9 million between 2011 and 2016.

areas of Ukraine where studies conducted

Species’ varying vulnerability

Musolin et al. (2022) cite experience in the Moscow Botanical Garden as showing that only two Asian species — Chinese ash, F. chinensis, and Manchurian ash, F. mandshurica — are were resistant to the EAB. The beetle killed both North American ash (i.e., F. pennsylvanica and F. americana) and European ash (i.e., F. excelsior, F. angustifolia, and F. ornus).

Experience in the field in Ukraine (Davydenko et al.) suggests that F. excelsior is less vulnerable to EAB than F. pennsyvanica. The overwhelming majority of EAB infestations were found on the American species. Furthermore, although similar densities of EAB larvae were found in colonized branches of both species, the proportion of larvae that were viable was significantly higher on F. pennsyvnica (91.4%) than on F. excelsior (76.1%). However, the reverse was found in the Moscow and St. Petersburg regions. Davydenko et al. don’t address directly whether they think this discrepancy is attributable to climatic factors or to differences in vulnerability between trees growing in native forests vs. human plantings. They did note that all observed cases of infestation of the native F. excelsior in Ukraine occurred in artificial plantings rather than in natural woodlands.

Interactions with Ash Decline

ash dieback disease – unfortunately pictured in the UK.
cc-by-sa/2.0 – © Adrian Diack – geograph.org.uk/p/6497286

Davydenko et al. studied parts of Eastern Ukraine where EAB was entering areas already infected by the invasive ascomycete fungus Hymenoscyphus fraxineus (cause of ash dieback, ADB). [Two of these regions — Luhansk and Kharkiv – have been the very center of the current war.] Other studies have shown that ~1 to 5% of F. excelsior trees exhibit some resistance to ADB. These trees are thus a potential foundation for future propagation and restoration of ash in Europe – if enough of them survive attack by EAB.

They found that F. excelsior is more resistant to EAB than F. pennsylvanica, but more susceptible to ADB.

The Luhansk and Kharkiv regions have both EAB and ADB; the Sumy region has only the pathogen. EAB probably invaded the Luhansk region by 2016 (although it was detected only in 2019). The proportion of ash trees (both native and introduced species) infested rose from ~ 10–30% in 2019 to 60 – 90% by 2020–2021. The EAB arrived later in the Kharkiv region, to the Northwest, but the proportion of infested trees was similar by 2021. Combining the two regions, 75% of F. pennsylvanica trees were EAB-infested, whereas only 31% of F. excelsior trees were.

Frequencies of infections by ADB were the reverse. Pooled data from all three study regions showed 21% of F. pennsylvanica trees were infected vs. 42% of F. excelsior. In the plots invaded by both EAB and ADB (in Luhansk and Kherson regions), 4%of F. pennsylvanica were affected by both invasive species vs. 14% of F. excelsior trees. Davydenko et al. conclude that ADB facilitates EAB attack on F. excelsior trees

The impact of EAB is seen in the fact that overall mortality rates were higher in F. pennsylvanica despite the fact that in the Sumy region mortality rates were higher in F. excelsior because of the disease (EAB was absent from this region). On the other hand, EAB infests and kills F. pennsylvanica trees regardless of their prior health condition (i.e., regardless of presence/absence of ADB).

Still, fewer than half the F. excelsior trees in sites affected by both EAB & ADB (in Luhansk and Kherson regions) have died. Davydenko et al. think the survivors constitute a source of material for eventual propagation. These trees need to be carefully mapped – a task certainly not facilitated by the war!

Davydenko et al. conclude that

1. Invasion of EAB in Ukraine occurred 2–3 years before detection in 2019 [I think this is actually quite prompt for detection of EAB invasions]; the invasion is currently expanding both in terms of newly infested trees and invaded geographic area.

2. Fraxinus excelsior (at least when growing in the interior of forest stands) is more resistant to EAB than F. pennsylvanica (when growing in field shelterbelts).

3. Fraxinus excelsior is more susceptible to ADB than F. pennsylvanica.

4. Infection by ADB is likely to predispose F. excelsior to infestation by EAB.

5. Ash trees infected by ADB are predisposed for the colonization by ash bark beetles Hylesinus spp. [I did not discuss these data.]

6. Inventory and mapping of surviving F. excelsior, affected by both ADB and EAB, is necessary to acquire genetic resources for the work on strategic, long-term restoration of devastated areas, thereby tackling a possible invasion of EAB to the EU.

I was surprised that Musolin et al. (2022) think EAB’s host shift from local Asian ash species to introduced North America ash planted in the Russian Far East and China triggered EAB outbreaks in Eastern China that contributed to the beetle’s introduction to North America and European Russia. American scientists apparently agree — Haack et al. (2022) refer to both this episode and a similar to one posited for Asian longhorned beetle (Anoplophora glabripennis) — that widespread planting of Populus plantations led to rapid expansion of ALB in northern China, and the pest-weakened wood was then used in wood packaging.

SOURCES

Choi, W.I.; Park, Y.-S. Management of Forest Pests and Diseases. Forests 2022, 13, 1765. https://doi.org/10.3390/f13111765

Davydenko, K.; Skrylnyk, Y.; Borysenko, O.; Menkis, A.; Vysotska, N.; Meshkova, V.; Olson, Å.; Elfstrand, M.; Vasaitis, R. Invasion of emerald ash borer Agrilus planipennis and ash dieback pathogen Hymenoscyphus fraxineus in Ukraine-A concerted action. Forests 2022, 13, 789.

Haack RA, Hardin JA, Caton BP and Petrice TR (2022) Wood borer detection rates on wood packaging materials entering the United States during different phases of ISPM#15 implementation and regulatory changes. Front. For. Glob. Change 5:1069117. doi: 10.3389/ffgc.2022.1069117

Musolin, D.L.; Selikhovkin, A.V.; Peregudova, E.Y.; Popovichev, B.G.; Mandelshtam, M.Y.; Baranchikov, Y.N.; Vasaitis, R. North-Westward Expansion of the Invasive Range of EAB, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) towards the EU: From Moscow to Saint Petersburg. Forests 2021, 12, 502. https://doi.org/10.3390/f12040502

Musolin, D.L.; Kirichenko, N.I.; Karpun, N.N.; Aksenenko, E.V.; Golub, V.B.; Kerchev, I.A.; Mandelshtam, M.Y.; Vasaitis, R.; Volkovitsh, M.G.; Zhuravleva, E.N.; et al. Invasive insect pests of forests and urban trees in Russia: Origin, pathways, damage, and management. Forests 2022, 13, 521.

Siegert, N.W. 2006. 17th USDA Interagency Research Forum on Gypsy Moth and Other Invasive Species. Annapolis, MD. January 10-13, 2006.

Posted by Faith Campbell

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

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

www.fadingforests.org

US invasive species — updated USGS database now on-line

ōhiʻa rust on Hawai`i; photo by J.B. Friday

The U.S. Geological Survey (USGS) has published an updated register of introduced species in the United States. The master list contains 14,700 records, of which 12,571 are unique scientific names. The database is divided into three sub-lists: Alaska, with 545 records; Hawai`i, with 5,628 records; and conterminous (lower 48) United States, with 8,527 records.

The project tracks all introduced (non-native) species that become established, because they might eventually become invasive. The list includes all taxa that are non-native everywhere in the locality (Alaska, Hawai`i, or 48 conterminous states) and established (reproducing) anywhere in that locality.

Each record has information on taxonomy, a vernacular name, establishment means (e.g., unintentionally, or assisted colonization), degree of establishment (established, invasive, or widespread invasive), hybrid status, pathway of introduction (if known), habitat (if known), whether a biocontrol species, dates of introduction (if known; currently 47% of the records), associated taxa (where applicable), native and introduced distributions (when known), and citations for the authoritative source(s) from which this information is drawn.

The 2022 version is more complete re: plant pathogens than earlier iterations; I thank the hard-working compilers for their efforts!

Hawai`i

wiliwili tree (Erythrina *sandwicensis*); photo by Forest and Kim Starr

Among the non-native species listed as being in Hawai`i are 3,603 Arthropods, including the following about which I have blogged:

three Euwallaceae,
Erythrina gall wasp (Quadrastichus erythrinae),
Klambothrips myopori,
eight species of mosquito in the Hawaiian islands, including the Culex and Aedes species that vector the diseases that have caused extinction of numerous endemic bird species on the Islands.

The list also includes 25 fungi, among them the two species of Ceratocystis that cause rapid ʻōhiʻa death; DMF & blog 270 and the ʻōhiʻa or myrtle rust, Austropuccinia psidii.

Also listed are 95 mollusk species and 20 earthworm species. I wonder who is studying the worms’ impacts? I doubt any is native to the Islands.

The Hawaiian list contains 1,557 non-native plant species. Families with largest representation are Poaceae (grass) – 223 species; Fabaceae (beans) – 156 species; and Asteraceae – 116 species. About a third of the plant species – 529 species – are designated as “widespread invaders”. This number is fifteen times higher than the numbers in lists maintained by either the Hawaiian Ecosystems At Risk project (106 species) [HEAR unfortunately had to shut down a decade ago due to lack of funds]; or Hawaiian Invasive Species Council (80 species). Furthermore, some of the species listed by HEAR and HISC are not yet widespread; the lists are intended to facilitate rapid responses to new detections. We always knew Hawai`i was being overrun by invasive species!

Among the 529 most “widespread invaders” are the following from the most introduced families:

Poaceae – Agrostis stolonifera, 6 Cenchrus spp, 2 Cortaderia spp, 3 Eragrostis,8 Paspalum, 4 Setaria spp, 2 Urochloa (Poacae)
Fabaceae – 3 Acacia, 2 Prosopis

Other families have fewer introduced species overall, but notable numbers of the most widespread invaders:

Euphorbiaceae – 8 spp. of Euphorbia
Cyperaceae – 6 spp. of Cyperus
Myrtaceae – Melaleuca quinquenervia, 2 Psidium, Rhodomyrtus tomentosa rose myrtle, 3 Syzygium [rose myrtle has been hard-hit by the introduced myrtle rust fungus]
Zingiberaceae – 3spp. Hedychium (ginger)
Anacardiaceae — Schinus molle (Peruvian peppertree); USGS considers congeneric S. terebinthifolia to be somewhat less widespread.

Plus many plant taxa familiar to those of us on the continent: English ivy, privet, castor bean, butterfly bush, Ipomoea vines … and in more limited regions, Japanese climbing fern Lygodium japonicum.

*Rhus sandwicensis*; photo by Forest and Kim Starr

I learned something alarming from the species profiles posted on the HISC website: the Hawaiʻi Division of Forestry and Wildlife and Hawaiʻi Department of Agriculture are considering introduction of a species of thrips, Pseudophilothrips ichini, as a biocontrol agent targetting S. terebinthifolia. I learned in early 2019, when preparing comments on Florida’s proposed release of this thrips, that Pseudophilothrips ichini can reproduce in low numbers on several non-target plant species, including two native Hawaiian plants that play important roles in revegetating disturbed areas. These are Hawaiian sumac Rhus sandwicensis and Dodonea viscosa. The latter in particular is being propagated and outplanted in large numbers to restore upland and dryland native ecosystems. While the environmental assessment prepared by the USDA Animal and Plant Service says the thrips causes minimal damage to D. viscosa, I am concerned because of the plant species’ ecological importance. Of course, the two Schinus species are very damaging invasive species in Hawai`i … but I think introducing this thrips is too risky. [To obtain a copy of CISP’s comments, put a request in comments section. Be sure to include your email address in your comment; the section algorithm does not include email addresses (how inconvenient!).]

Continental (lower 48) states

Among the 8,500 species listed in the USGS Register for the 48 continental states are 4,369 animals, among them 3,800 arthropods; 3,999 plants; and just 89 fungi. Among the arthropods, there are 1,045 beetles and 308 lepidopterans. The beetles listed include 12 Agrilus (the genus which includes emerald ash borer and goldspotted oak borer.) It does not include the elm zig-zag sawfly USGS staff have not found any publications documenting its U.S. occurrences. Among the microbes are six Phytophthora (P. cinnamomi, P. lateralis, P. pseudocryptogea, P. quercina, P. ramorum, P. tentaculata). Profiles of several of these species are posted at www.dontmovefirewood.org; click on “invasive species”, then scroll using either Latin or common name.

elm zig-zag sawfly; photo by Gyorgy Czoka via Bugwood

Citation:

Simpson, Annie, Pam Fuller, Kevin Faccenda, Neal Evenhuis, Janis Matsunaga, and Matt Bowser, 2022, United States Register of Introduced and Invasive Species (US-RIIS) (ver. 2.0, November 2022): U.S. Geological Survey data release, https://doi.org/10.5066/P9KFFTOD

United States Register of Introduced and Invasive Species; US-RIIS ver. 2.0, 2022

If you would like to contribute to future versions of the US-RIIS, please email the project leaders at us-riis@usgs.gov.

Posted by Faith Campbell

www.fadingforests.org

New Publication on Threats to World’s Forests – including Invasives

In a new paper, “Forest Resources of the World: Present Status and Future Prospects,” Singh et al. affirm the importance of forests for terrestrial biodiversity, provision of multiple ecosystem services, and supporting the economic well-being of approximately 1.6 billion people directly. This equals about a quarter of Earth’s population. The authors conclude that achieving global Sustainable Development Goals (SDGs), including poverty reduction, food security, and mitigating and adapting to climate change — all depend on sustaining forests.

According to the 2020 Global Forest Resource Assessment, Earth’s forested area comprises ~4.06 billion hectares, or 31% of the total land surface.More than half (54%) of all global forest area is found in five countries: the Russian Federation, Brazil, Canada, the United States, and China. Tropical forests constitute 45% of this total; boreal forests, 27%; temperate forests, 16%; and subtropical forests, 11%. An estimated 93% (3.75 billion ha) regenerate through natural processes; 7% (290 million ha) is planted forest.

The extent of global forest area has been declining for decades but the rate of loss slowed significantly between 1990 and 2020. This reflects decreased deforestation in some countries and an increase in forest area in others. The latter is due to both afforestation and also natural forest growth. However, conversion of tropical forests to agriculture continues apace. From 2010 to 2020, the net loss of forest area was highest in Africa (3.9 million ha) and South America (2.6 million ha). Increases in net forest area occurred in Asia, Oceania and Europe. The status of the top 10 countries or territories in global forest resources as of 2020 is given in Table 1.2 of the chapter. [News sources document that rapid deforestation continues in Brazil, at least.]

Several trends are concerning to those of us who value primary or undisturbed forests. First, the area of naturally regenerating forest has decreased, while the area of planted forest has expanded – but only by 123 million ha. In the last decade, the rate of increase in the area of planted forests has also slowed.

Second, total carbon stock in forests declined from 668 gigatons to 662 gt in 1990–2020. This is only 6%, but it is trending in the wrong direction. As we know, forest conservation counters climate change in two ways: conserved forests are a carbon sink, while degraded or destroyed forests are a significant source of atmospheric CO₂. In fact, forests are the 2^nd largest storehouses of carbon, after oceans. Global forests sequester about one-third of total CO₂ emission from the combustion of fossil fuels. Almost all forest carbon is found in living biomass (44%) and soil organic matter (45%).

Costa Rican rainforest; photo by eflon via Flickr

Third, primary forests are already severely reduced and continue to shrink. Primary forests are those composed of native species, and supporting relatively undisturbed ecological processes. They are irreplaceable for sustaining biological diversity. These forests are already severely reduced – they cover only ~ 1 billion ha. Since 1990, the extent of primary forest has decreased by 81 million ha. More than half are in Brazil, Canada, and Russia.

Singh et al. report that only about 10% of the world’s forests are set aside for biodiversity conservation. Again, trends are in the wrong direction. The rate of increase in the area of forest designated largely for biodiversity conservation has slowed. On the other hand, forest areas designated for other non-extractive purposes have increased: soil and water conservation, recreation, tourism, education, research, and the protection of cultural and spiritual sites.

Singh et al. are cheered by the fact that more than 2 billion hectares are under management with well-defined management plans. The extent of forests under management plans has increased by 233 million ha since 2000.

Singh et al. say that continuously increasing anthropogenic pressure is the main cause of deforestation and forest degradation in unmanaged forests. Citing projections that the world’s population will reach almost 10 billion by 2050, they say this growth will make reconciling the need for forest conservation with the basic requirements of humans for food, shelter, and fuel more difficult than ever.

I appreciate this honesty. Too many experts interviewed on the day that the global population was estimated at 8 billion made optimistic statements about the consequences. They mentioned Earth’s carrying capacity only in reference to First World people demanding excessive resources. There was minimal discussion of humanity’s carbon footprint and no reference to ever-increasing threats to biological diversity. Nor to the fact that people in developing countries want to raise their standards of living – which entails higher demand for resources, including energy. For an example, see The Washington Post editorial, here.

On the other hand, Ruby Mellen in the Post on 15 November mentioned that, according to the World Wildlife Fund, 75% of Earth’s ice-free land has been significantly altered by people, and two-thirds of mammal, fish, reptile, and amphibian species have become endangered in the last ~50 years. Unfortunately, the on-line version of the paper doesn’t have this specific article!

fires in Siberian forest in 2016; European Space Agency

Threats to Forests: Fire

Singh et al. rank fire as the most disastrous threat, affecting biodiversity and carbon sequestration potential. According to the U.N. Food and Agriculture Organization, about 29% of the total geographical area in the world was affected by forest fires during 2001–2018; more than two-thirds of these fires occurred in Africa. U.S. media, however, focused on fires in the Amazon, temperate areas (U.S., Europe), and, sometimes, boreal forests or Australia. Singh et al. say that areas that are frequently affected by fire are prone to other types of disturbances like drought and outbreaks of insect pests.

tanoaks killed by *Phytophthora ramorum* in Oregon; photo by Oregon Department of Forestry

Threats to Forests: Diseases and Pests

I am glad that Singh et al. recognize the damage to forest productivity caused by disease and pest infestations. In doing so, they cite familiar sources – Clive Brasier, Peter Vitousek, Juliann Aukema, Gary Lovett, Sandy Liebhold, Kerry Britton, Bitty Roy, Hanno Seebens – regarding surges in pest attacks; the growing diversity of damaging pests; resulting changes in forest species composition and structure that impede ecosystem functions and productivity. Singh et al. follow these sources in calling for improved hygiene in nurseries, adoption of scientific silvicultural practices reducing physical damage to the vegetation, selection of genotypes that are resistant, and reinforcing national and international policies on quarantine and biosecurity measures to minimize pest impacts in the future. They also mention adoption of remote sensing technologies to detect the trees under stress and use of sentinel plantings. They list the 10 most important international agreements dealing with invasive species issues as the International Plant Protection Convention, Ramsar Convention, Convention on International Trade in Endangered Species of Wild Fauna and Flora, Convention on Migratory Species, Convention on Biological Diversity and its Cartagena Protocol on Biosafety, IUCN Invasive Species Specialist Group, World Trade Organization Agreement on Sanitary and Phytosanitary Measures, Global Invasive Species Program, and International Civil Aviation Organization, and Cartagena.

slash and burn agriculture in Bolivia; photo Neil Palmer

Threats to forests: Development Projects

Singh et al. consider development projects to be the third threat to forest conservation. Their roads, powerlines, and other linear developments cause habitat loss and fragment landscapes. In their view, environmental impact assessments and other similar requirements are not yet sufficient to safeguard sustainable use of forest resources.

Policy Responses

Singh et al. call for more inclusive forest management structures to respond to the threat climate change poses to forests, industries, and forest-dependent communities. They all for partnerships that bring together researchers from several disciplines with forest managers and local stakeholders. Geoffrey M. Williams and others (including me) advocate for similar conservation approaches. (See pre-print here.)

In this context, Singh et al. mention several reports, plans, and agreements aimed at global forest conservation. Participants in global fora have recognized the importance of forests in contributing to food security and sustainable development. Among agreements mentioned are the UN’s Strategic Plan for Forests 2030 and recommendations of the International Institute for Sustainable Development (IISD) published in 1994. The former tries to generate greater coherence, collaboration, and synergy across UN programs aimed at encouraging volunteer forest conservation by countries, international, regional, and local organizations, partners, and stakeholders. Unfortunately, they do not discuss the extent to which the 30-year old IISD recommendations have – or have not – been implemented.

They also describe Forest Landscape Restoration as an effective strategy to restore the functionality of forests.Again, the focus is on a collaborative approach aimed at integrating efforts by all forestry-related stakeholders, e.g., scientific and academic organizations, local communities, indigenous peoples, and private sectors, including forest-based enterprises and NGOs.

Also praised is rising attention to trees outside forest. This includes fostering use of trees in agroforestry systems ranging from home gardens to farm forestry systems, shelterbelts, and woodlots. This approach helps to sustain the livelihoods of rural communities and maintain a stable and secure food supply. Meanwhile, it reduces dependence on natural forests

Singh et al. say community forest management and decentralized governance have gained acceptance. They describe examples from Gambia and Rwanda. They concede that such decentralization has its own risks and challenges. For example, e the most marginalized sections of the community must be ensured adequate capacity for robust conflict resolution.

Singh et al. advocate that all nations seek to increase their forest cover; affluent countries that are hampered by physical and climatic conditions should aid poorer nations in increasing and upgrading their forest cover. They suggest “recognition” and encouragement of countries that maintain forest cover above 30% of territory.

See also about loss of floral diversity and blog about IUCN’s global forest assessment.

SOURCE

Singh, M., N.N. Shahina, S. Das, A. Arshad, S. Siril, D. Barman, U. Mog, P. Panwar, G. Shukla, and S. Chakravarty. 2022. Forest Resources of the World: Present Status and Future Prospects. In Panwar, P., G. Shukla, J.A. Bhat, S. Chakravarty. 2022. Editors. Land Degradation Neutrality: Achieving SDG 15 by Forest Management; ISBN 978-981-19-5477-1 ISBN 978-981-19-5478-8 (eBook)

https://doi.org/10.1007/978-981-19-5478-8

Posted by Faith Campbell

www.fadingforests.org

Australia Builds Capacity to Address Forest Pests

Australian Eucalypts; photo by John Turnbull via Flickr

I congratulate Australian scientists for bringing about substantial improvements of their country’s biosecurity program for forest pests. While it is too early to know how effective the changes will be in preventing new introductions, they are promising. What can we Americans learn from the Australian efforts? [I have previously praised South Africa’s efforts – there is much to learn there, too.]

Australia has a reputation of being very active in managing the invasive species threat. However, until recently biosecurity programs targetting forest pests were minimal and ad hoc. Scientists spent 30 years trying to close those gaps (Carnegie et al. 2022). Their efforts included publishing several reports or publications (listed at the end of the blog) and an international webinar on myrtle rust. Scientists are hopeful that the new early detection program (described below) will greatly enhance forest protection. However, thorough pest risk assessments are still not routinely conducted for forest pests. (Nahrung and Carnegie 2022).

The native flora of Australia is unique. That uniqueness has provided protection because fewer of the non-native insects and pathogens familiar to us in the Northern Hemisphere have found suitable hosts (Nahrung and Carnegie 2020). Also – I would argue – the uniqueness of this flora imposes a special responsibility to protect it from threats that do arise.

Only 17% of Australia’s landmass is covered by forests. Australia is large, however; consequently, these forests cover 134 million hectares (Nahrung and Carnegie 2020). This is the 7th largest forest estate in the world (Carnegie et al. 2022).

Australia’s forests are dominated by eucalypts (Eucalyptus, Corymbia and Angophora). These cover 101 million ha; or 75% of the forest). Acacia (11 million ha; 8%); and Melaleuca (6 million ha) are also significant. The forest also includes one million ha of plantations dominated by Pinus species native to North America (Carnegie et al. 2022). A wide range of native and exotic genera have been planted as amenity trees in urban and peri-urban areas, including pines, sycamores, poplars, oaks, and elms (Carnegie et al. 2022). These urban trees are highly valued for their ecosystem services as well as social, cultural, and property values (Nahrung and Carnegie 2020). Of course, these exotic trees can support establishment and spread of the forest pest species familiar to us in the Northern Hemisphere. On the positive side, they can also be used as sentinel plantings for early detection of non-native species (Carnegie et al. 2022 and Nahrung and Carnegie 2020).

Despite Australia’s geographic isolation, its unique native flora, and what is widely considered to be one of the world’s most robust biosecurity system, at least 260 non-native arthropods and pathogens of forests have established in Australia since 1885 (Nahrung and Carnegie 2020). [(This number is about half the number of non-native forest insects and pathogens that have established in the United States over a period just 25 years longer (Aukema et al. 2010).] As I noted, forest scientists have cited these introductions as a reason to strengthen Australia’s biosecurity system specifically as it applies to forest pests.

What steps have been taken to address this onslaught? For which pests? With what impacts? What gaps have been identified?

Which Pests?

Nahrung and Carnegie (2020) compiled the first comprehensive database of tree and forest pests established in Australia. The 260 species of non-native forest insect pests and pathogens comprise 143 arthropods, 117 pathogens. Nineteen of them (17 insects and 2 fungal species) had been detected before 1900. These species have accumulated at an overall rate of 1.9 species per year; the rate of accumulation after 1955 is slightly higher than during the earlier period, but it has not grown at the exponential rate of import volumes.

While over the entire period insects and pathogens were detected at an almost equal rate (insects at 1.1/year; pathogens at 0.9/year), this disguises an interesting disparity: half of the arthropods were detected before 1940; half of the pathogens after 1960 (Nahrung and Carnegie (2020). By 2022, Nahrung and Carnegie (2022) said that, on average, one new forest insect is introduced each year. Some of these recently detected organisms have probably been established for years. More robust surveillance has just detected them recently. I have blogged often about an apparent explosion of pathogens being transported globally in recent decades.

In a more recent article (Nahrung and Carnegie, 2022), gave 135 as the number of non-native forest insect pests. The authors don’t explain why this differs from the 143 arthropods listed before.

damage to pine plantations caused by *Sirex noctilio*; photo courtesy of Helen Nahrung

Eighty-seven percent of the established alien arthropods are associated with non-native hosts (e.g., Pinus, Platanus, Populus, Quercus, Ulmus) (Carnegie et al. 2022). Some of these have escaped eradication attempts and caused financial impact to commercial plantations (e.g., sirex wood wasp, Sirex noctilio) and amenity forests (e.g., elm leaf beetle, Xanthogaleruca luteola) (Carnegie and Nahrung 2019).

About 40% of the alien arthropods were largely cosmopolitan at the time of their introduction in Australia (Carnegie et al. 2022). Only six insects and six fungal species are not recorded as invasive elsewhere (Nahrung and Carnegie 2020). Of the species not yet established, 91% of interceptions from 2003 to- 2016 were known to be invasive elsewhere. There is strong evidence of the bridgehead effect: 95% of interceptions of three species were from their invaded range (Nahrung and Carnegie 2022). These included most of the insects detected in shipments from North America, Europe and New Zealand. These ubiquitous “superinvaders” have been circulating in trade for decades and continue to be intercepted at Australia’s borders. This situation suggests that higher interception rates of these species reflect their invasion success rather than predict it (Nahrung and Carnegie 2021).

I find it alarming that most species detected in shipments from Africa, South America, and New Zealand were of species not even recorded as established in those regions (Nahrung and Carnegie 2021; Nahrung and Carnegie 2022).

*Arhopalus ferus*, a Eurasian pine insect often detected in wood from New Zealand; photo by Jon Sullivan – in New Zealand; via Flickr

Half of the alien forest pests established in Australia are highly polyphagous. This includes 73% of Asian-origin pests but only 15% of those from Europe (Nahrung and Carnegie 2021). Nahrung and Carnegie (2022) confirm that polyphagous species are more likely to be detected during border inspections.

PATHWAYS

As in North America and Europe, introductions of Hemiptera are overwhelmingly (98%) associated with fresh plant material (e.g. nursery stock, fruit, foliage). Coleoptera introductions are predominantly (64%) associated with wood (e.g. packaging, timber, furniture, and artefacts). Both pathways are subject to strict regulations by Australia (Nahrung and Carnegie 2021).

Eradication of High-Priority Pests

Eight-five percent of all new detections were not considered high-priority risks. Of the four that were, two had not previously been recognized as threats (Carnegie and Nahrung 2019). One high-priority pest – expected to pose a severe threat to at least some of Australia’s endemic plant species – is myrtle rust, Austropuccinia psidii. Despite this designation, when the rust appeared in Australia in 2010, the response was confused and ended in an early decision that eradication was impossible. Myrtle rust has now spread along the continent’s east coast, with localized distribution in Victoria, Tasmania, the Northern Territory, and – in 2022, Western Australia. `

*Melaleuca quinquenervia* forest; photo by Doug Beckers via Wikimedia

There have been significant impacts to native plant communities. Several reviews of the emergency response criticized the haste with which the initial decision was made to end eradication (Carnegie and Nahrung 2019). (A review of these impacts is here; unfortunately, it is behind a paywall.)

A second newly introduced species has been recognized as a significant threat, but only after its introduction to offshore islands. This is Erythina gall wasp Quadrastichus erythrinae (Carnegie and Nahrung 2019). DMF Although Australia is home to at least one native species in the Erythrina genus, E. vespertilio,, the gall wasp is not included on the environmental pest watch list.

Four of the recently detected species were considered to be high impact. Therefore eradication was attempted. Unfortunately, these attempts failed in three cases. The single success involved a pinewood nematode, Bursaphelenchus hunanesis. See Nahrung and Carnegie (2021) for a discussion of the reasons. This means three species recognized as high-impact pests have established in Australia over 15 years (Nahrung and Carnegie (2021). In fact, Australia’s record of successful forest pest eradications is only half the global average (Carnegie and Nahrung (2019).

Carnegie and Nahrung (2019) conclude that improving early detection strategies is key to increasing the likelihood of eradication. They discuss the strengths and weaknesses of various strategies. Non-officials (citizen scientists) reported 59% of the 260 forest pests detected (Carnegie and Nahrung 2019). Few alien pests have been detected by official surveillance (Carnegie et al 2022). However, managing citizen scientists’ reports involves a significant workload. Futhermore, surveillance by industry, while appreciated, is likely to detect only established species (Carnegie and Nahrung 2019).

Interception Frequency Is Not an Indicator of Likelihood of Establishment

Nahrung & Carnegie (2021) document that taxonomic groups already established in Australia are rarely detected at the border. Furthermore, only two species were intercepted before they were discovered to be established in Australia.

Indeed, 76% of species established in Australia were either never or rarely intercepted at the border. While more Hemiptera species are established in Australia, significantly more species of Coleoptera are intercepted at the border. Among beetles, the most-intercepted family is Bostrichid borers (powderpost beetles). Over the period 2003 – 2016, Bostrichid beetles made up 82% of interceptions in wood packaging and 44% in wood products (Nahrung and Carnegie 2022). This beetle family is not considered a quarantine concern by either Australian or American phytosanitary officials. I believe USDA APHIS does not even bother recording detections of powderpost beetles. Nahrung and Carnegie (2021) think the high proportion of Bostrichids might be partially explained by intense inspection of baggage, mail, and personal effects. While Australia actively instructs travelers not to bring in fruits and vegetables because of the pest risk, there are fewer warnings about risks associated with wood products.

Nahrung & Carnegie (2021) concluded that interception frequencies did not provide a good overall indicator of likelihood of risk of contemporaneous establishment.

Do Programs Focus on the Right Species?

Although Hemiptera comprise about a third of recent detections and establishments, and four of eight established species are causing medium-to-high impact, no Hemiptera are currently listed as high priority forestry pests by Australian phytosanitary agencies (Nahrung & Carnegie (2021). On the other hand, Lepidoptera make up about a third of the high-priority species, yet only two have established in Australia over 130 years. Similarly, Cerambycidae are the most frequently intercepted forest pests and several are listed as high risk. But only three forest-related species have established (Nahrung and Carnegie 2020). (Note discussion of Bostrichidae above.).

Unlike the transcontinental exchanges under way in the Northern Hemisphere, none of the established beetles is from Asia; all are native to Europe. This is especially striking since interceptions from Asia-Pacific areas account for more than half of all interceptions Nahrung and Carnegie (2021).

Interestingly, 32 Australian Lepidopteran and eight Cerambycid species are considered pests in New Zealand. However, no forest pests native to New Zealand have established in Australia despite high levels of trade, geographic proximity, and the high number of shared exotic tree forest species (Nahrung and Carnegie 2020).

STRUCTURE OF PROGRAM

The structure of Australia’s plant biosecurity system is described in detail in Carnegie et al. (2022). These authors call the program “comprehensive” but to me it looks highly fragmented. The federal Department of Agriculture and Water Resources (DAWR,[recently renamed the Department of Agriculture, Fisheries, and Forestry, or DAFF) is responsible for pre-border (e.g., off-shore compliance) and border (e.g., import inspection) activities. The seven state governments, along with DAFF, are responsible for surveillance within the country, management of pest incursions, and regulation of pests. Once an alien pest has become established, its management becomes the responsibility of the land manager. In Australia, then, biosecurity is considered to be a responsibility shared between governments, industry and individuals.

Even this fragmented approach was developed more recently than one might expect given Australia’s reputation for having a stringent biosecurity system. Perhaps this reflects the earlier worldwide neglect of the Plant Kingdom? Carnegie and Nahrung (2019) describe recent improvements. Until the year 2000, Australia’s response to the detection of exotic plant pests was primarily case-by-case. In that year Plant Health Australia (PHA) was incorporated. Its purpose was to facilitate preparedness and response arrangements between governments and industry for plant pests. In 2005, the Emergency Plant Pest Response Deed (EPPRD) was created. It is a legally-binding agreement between the federal, state, and territorial governments and plant industry bodies. As of 2022, 38 were engaged. It sets up a process to implement management and funding of agreed responses to the detection of exotic plant pests – including cost-sharing and owner reimbursement. A national response plan (PLANTPLAN) provides management guidelines and outlines procedures, roles and responsibilities for all parties. A national committee (Consultative Committee on Emergency Plant Pests (CCEPP) works with surveys to determine invaded areas (delimitation surveys) and other data to determine whether eradicating the pest is technically feasible and has higher economic benefits than costs..

*Austropuccinia psidii* on *Melaleuca quinquenervia*; photo by John Tann via Flickr

Even after creation of EPPRD in 2005, studies revealed significant gaps in Australia’s post-border forest biosecurity systems regarding forest pests (Carnegie et al. 2022; Carnegie and Nahrung 2019). These studies – and the disappointing response to the arrival of myrtle rust – led to development of the National Forest Biosecurity Surveillance Strategy (NFBSS) – published in 2018; accompanied by an Implementation Plan. A National Forest Biosecurity Coordinator was appointed.

The forest sector is funding a significant proportion of the proposed activities for the next five years; extension is probable. Drs. Carnegie and Nahrung are pleased that the national surveillance program has been established. It includes specific surveillance at high-risk sites and training of stakeholders who can be additional eyes on the ground. The Australian Forest Products Association has appointed a biosecurity manager (pers. comm.)

This mechanism is expected to ensure that current and future needs of the plant biosecurity system can be mutually agreed on, issues identified, and solutions found. Plant Health Australia’s independence and impartiality allow the company to put the interests of the plant biosecurity system first. It also supports a longer-term perspective (Carnegie et al. (2022). Leading natural resource management organizations are also engaged (Carnegie, pers. comm.).

Presumably the forest surveillance strategy (NFBSS) structure is intended to address the following problems (Carnegie and Nahrung 2019):

Alien forest pests are monitored offshore and at the border, but post-border surveillance is less structured and poorly resourced. Australia still lacks a surveillance strategy for environmental pests.
Several plant industries have developed their own biosecurity programs, co-funded by the government. These include the National Forest Biosecurity Surveillance Strategy (NFBSS).

Some pilot projects targetting high risk sites were initiated in the early 2000s. By 2019, only one surveillance program remained — trapping for Asian spongy (gypsy) moth.

The states of Victoria and New South Wales have set up sentinel site programs. Victoria’s uses local council tree databases. It is apparently focused on urban trees and is primarily pest-specific – e.g., Dutch elm disease. The New South Wales program monitors more than 1,500 sentinel trees and traps insects near ports. This program is funded by a single forest grower through 2022.

Dr. Carnegie states: “With the start of the national forest biosecurity surveillance program in December 2022, the issues and gaps identified by Carnegie et al. 2022 are starting to be addressed. The program will conduct biosecurity surveillance specifically for forest pests and pathogens and be integrated with national and state biosecurity activities. While biosecurity in Australia is still agri-centric, a concerted and sustained effort from technical experts from the forest industry is changing this. And finally, the new Biosecurity Levy should ensure sustained funding for biosecurity surveillance.”

There is a separate National Environmental Biosecurity Response Agreement (NEBRA), adopted in 2012. It is intended to provide guidelines for responding, cost-sharing arrangements, etc. when the alien pest threatens predominantly the environment or public amenity assets (Carnegie et al. (2022). However, when the polyphagous shot hole borer was detected, the system didn’t work as might have been expected. While PSHB had previously been identified as an environmental priority pest, specifically to Acacia, the decision whether to engage was made under auspices of the the Emergency Plant Pest Response Deed (EPPRD) rather than the environmental agreement (NEBRA). As a result, stakeholders focused on environmental, amenity and indigenous concerns had no formal representation in decision-making processes; instead, industries that had assessed the species as a low priority (e.g., avocado and plantation forestry) did (Nahrung, pers.comm.).

Additional Issues Needing Attention

Some needs are not addressed by the National Forest Pest Strategic Plan (Carnegie et al. 2022) (Nahrung, pers. comm.):

1) The long-term strategic investment from the commercial forestry sector and government needed to maintain surveillance and diagnostic expertise;

2) Studies to assess social acceptance of response and eradication activities such as tree removal;

3) Studies to improve pest risk prioritization and assessment methods; and

4) Resolving the biosecurity responsibilities for pests of timber that has been cut and used in construction.

In 2019, Carnegie and Nahrung (2019) called for developing more effective methods of detection, especially of Hemiptera and pathogens. They also promoted national standardization of data collection. Finally, they advocated inclusion of technical experts from state governments, research organizations and industry in developing and implementing responses to pest incursions. They note that surveillance and management programs must be prepared to expect and respond to the unexpected since 85% of the pests detected over the last 20 years—and 75% of subsequently mid-to high-impact species established—were not on high-priority pest list. See Nahrung and Carnegie 2022 for a thorough discussion of the usefulness and weaknesses of predictive pest listing.

SOURCES

Aukema, J.E., D.G. McCullough, B. Von Holle, A.M. Liebhold, K. Britton, & S.J. Frankel. 2010. Historical Accumulation of Nonindigenous Forest Pests in the Continental United States. Bioscience. December 2010 / Vol. 60 No. 11

Carnegie A.J. and H.F. Nahrung. 2019. Post-Border Forest Biosecurity in AU: Response to Recent Exotic Detections, Current Surveillance and Ongoing Needs. Forests 2019, 10, 336; doi:10.3390/f10040336 www.mdpi.com/journal/forests

Carnegie A.J., F. Tovar, S. Collins, S.A. Lawson, and H.F. Nahrung. 2022. A Coordinated, Risk-Based, National Forest Biosecurity Surveillance Program for AU Forests. Front. For. Glob. Change 4:756885. doi: 10.3389/ffgc.2021.756885

Nahrung H.F. and A.J. Carnegie. 2020. NIS Forest Insects and Pathogens in Australia: Establishmebt, Spread, and Impact. Frontiers in Forests and Global Change 3:37. doi: 10.3389/ffgc.2020.00037 March 2020 | Volume 3 | Article 37

Nahrung, H.F. and A.J. Carnegie. 2021. Border interceps of forest insects estab in AU: intercepted invaders travel early and often. NeoBiota 64: 69–86. https://doi.org/10.3897/neobiota.64.60424

Nahrung, H.F. & A.J. Carnegie. 2022. Predicting Forest Pest Threats in Australia: Are Risk Lists Worth the Paper they’re Written on? Global Biosecurity, 2022; 4(1).

Posted by Faith Campbell

www.fadingforests.org

Plants for Planting – Major Pathway, Too Little Attention

*Phytopthora cinnamomi* on manzanita in California; photo courtesy of Ted Swiecki/Phytosphere

While I blog often about wood packaging the fact is that imports of live plant [= “plants for planting” in USDA’s terms] have historically posed a higher risk of introducing tree-killing pests. In 2012, Liebhold et al. found that nearly 70% of 455 damaging pests introduced to the continental U.S. as of 2006 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. Imported plants not only carry a greater variety of pests than wood packaging; they also carry many more.

Introductions on imported plants for planting is not a rare event. An analysis of data in the Agriculture Quarantine Inspection Monitoring (AQIM) during 2009 found that 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. This alarming statistic receives less attention than warranted because APHIS objected to the accuracy of other aspects of the study.

APHIS has adopted changes to its phytosanitary system for plants for planting in the decade since 2009. The question is, have these changes reduced the known risks associate with live plant imports – especially given skyrocketing imports? Are more measures necessary? Current data and analyses cannot provide a scientifically valid answer.

ohia rust on endangered Hawaiian native plant *Eugenia koolauensis*

First, most studies focus on insects – they even exclude pathogens. Among pathogens introduced in recent decades, probably by the plant trade, are several Phytophthoras, rapid ‘ōhi‘a death, beech leaf disease, boxwood blight. (I am assuming that the Fusarium dieback disease vectored by Euwallacea beetles was introduced via wood packaging.) There have been repeated detections of the Ralstonia solanacearum Race 3 biovar 2, a bacterium that attacks a range of herbaceous plants, despite APHIS requiring specific integrated pest management programs in producing nurseries located in Central America. Examples of recently introduced leaf feeders include the European beech leaf-mining weevil and elm zigzag sawfly.

I concede that it is difficult to study introduced pathogens. It is nearly impossible to compile a complete list of introduced fungi and related organisms since only the most damaging are typically detected and their native ranges are frequently undeterminable. However, European forest pathologists are much more active on these questions. Why? What can we do to focus Americans on the threats these organism pose?

Second, most studies analyzing the pest risk associated with plant imports use port inspection data. However, port inspection data 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). Thus, most of the analyses carried out by Liebhold et al. and MachLachlan et al. (2022) are based on the pests found by APHIS inspectors: actionable pests were detected on only 2.6% of the incoming plants that they inspected.

Here I discuss two recent discussions of the risk associated with imported plant for planting. One is an analysis of establishments of one order of insects in the United States over 200 years (MacLachlan et al. 2022; full citation at the end of the blog). Again, the focus is on insects! The other is a discussion of the pathway during the recent annual meeting of the Continental Dialogue on Non-Native Forest Insects and Diseases. link to posting of presentations This discussion raised some of the key questions, although no answers were provided.

U.S. imports of plants have increased by more than 400% since the 1960s; 35% in just the last 15 years (in 2007 the U.S. imported approximately 3.7 billion plants [Liebhold et al. 2012]; in 2021 it was about 5 billion [MacLachlan et al. 2022]. Yet establishments of new non-native insects associated with this pathway have not risen commensurately. MacLachlan et al. (2022) attempt to answer why this is so. However, pests are often not detected for several years or a decade after their introduction. Furthermore, I doubt that an analysis based on inspection data, not the more reliable AQIM data, can provide an accurate assessment.

To clarify the pest risk associated with plant imports, studies of some insect types, excluding pathogens, is not sufficient. Again, APHIS should update the Liebhold et al. 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 usually excluded from analyses of imported pests, including Liebhold et al. 2012. 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!

*Cycas micronesica* endemic to Guam; threatened by cycad scale & cycad blue butterfly; photo courtesy A. Gawel

MacLachlan et al. (2022) focused their analysis on the insect order Hemiptera, including the so-called true bugs, including cicadas, aphids, planthoppers, and leafhoppers. This is the insect order most frequently transported with imported plants. In addition, establishments of Hemiptera can be attributed to plant imports rather than to wood or other vectors. Of the 3,500 species of non-native insects established in North America (including the contiguous U.S. states, Alaska, and Canada), about 27% are Hemiptera. Many are serious pests, e.g., hemlock woolly adelgid and balsam woolly adelgid). Complicating the analysis, however, is the fact that some Hemiptera are inconspicuous so they are difficult to detect. In fact, MacLaughlan et al. 2022 estimate the median delay between introduction and detection to be 80 years! They believe that many introduced species remain undiscovered, ranging from 21% for Eurasian regions to 38% for the Neotropics and 52% for Australasia.

eastern hemlocks killed by hemlock woolly adelgied; Linville Gorge, NC; photo by Steven Norman, USFS

MacLachlan et al. (2022) compare the relationship between plant imports and discoveries of Hemiptera from 1800 to the present in an attempt to answer the puzzle of why new Hemiptera establishments have remained relatively steady despite quadrupled plant imports. Perhaps the pool of novel insect species in the source region has been depleted. Or other factors might have changed, such as

the commodities imported (plant species or types; or geographic source)
phytosanitary measures applied by the U.S.

MacLachlan et al. (2022) tracked plant imports since 1854 from seven ecological regions: Afrotropic, Asian Palearctic, Australasia, European Palearctic, Indomalaya, Nearctic, Neotropic. In the early decades, both imported plants and introduced Hemiptera detected in the U.S., came predominantly from European and Asian Palearctic regions. Now, however, almost no new Hemiptera species are being introduced on plants imported from the European and Asian Palearctic regions. Since the 1950s, estimated establishments from the Indomalaya region have remained relatively stable. Establishments from the Neotropic and Afrotropic regions rose following World War II and have remained relatively high. After also declining in the first half of the 20th century, establishments of new species from Australasia have recently increased.

Generally, the regions associated with declining establishments of new species (Eurasia) are experiencing relatively gradual increases in their exports to the U.S. Those regions which contribute relatively steady or increasing establishments (Neotropics, Indomalaya, Australasia, and Afrotropic) have each undergone rapid increases in exports to the U.S.

Establishment Risk Among Regions

Source regions vary in the type of plants they export (e.g., rootless cuttings v. whole plants) and in the volume of exports. They also differ in the composition of their indigenous and introduced insect populations. Imports from areas with an abundance of species capable of establishing and adapted to environmental conditions in North America pose greater establishment risk, although it is challenging to determine the risk associated with individual species.

Establishment risk of shipments from a particular region also changes over time. The number of potential new species of invaders might shrink as more and more arrive in North America. (This situation has no effect on the continued introduction of insect species already established in North America. These reintroductions might arrive in new areas – so expanding the area at risk; or their increasing number contributes to propagule pressure at establishment sites.) Another factor might be phytosanitary policies. Strengthening of phytosanitary measures might suppress the number of organisms that travel with the plant shipment, enter North America, and establish. The opposite might happen if phytosanitary measures are relaxed or if the sourcing or type of imports diversifies in ways that connect additional species in source regions with trade pathways.

Considering all regional plant sources, MacLachlan et al. (2022) estimate that establishments per unit of additional imports – of Hemipterans – have shrunk because of a combination of increased imports, accumulated introductions associated with past imports, and the passage of time. These decreases are substantial – between 75.2% and 99.8% for the various regions from 1962 to 2012. For the Asian Palearctic and Neotropic regions, MacLachlan et al. (2022) determined that depletion of species pools is a contributing factor. Other factors are thought to explain the substantial decline in establishment likelihood for the other regions. However, note the caveats above re: lag times in detecting introductions.

However, despite that significant decrease in risk per unit of imports, the number of establishments has remained relatively constant over the past century. MacLachlan et al. (2022) attribute this pattern to the decreases in marginal risk from additional imports being offset by 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) suggest that APHIS should target biosecurity resources to the specific commodity-country pairs associated with a demonstrated higher relative risk of introducing additional insect species.

MacLachlan et al. (2022) are unable to evaluate the efficacy of APHIS’ most important policy change: creation of the “Not Authorized for Importation Pending Pest Risk Assessment” (NAPPRA) program because it was adopted in 2011 and they analyzed data only through 2012. A decade later this policy restricts imports of about 250 taxa (Regelbrugge to Continental Dialogue). It is certainly time to evaluate its efficacy through a new study of pest approach rates in the “plants for planting” trade.

I do not think that U.S. phytosanitary policy should be based on an analysis of just one of at least three types of pests that travel via the pathway. We need analysis of the risk from pathogens, nematodes, viruses … and other orders of arthropods.

The Continental Dialogue on Non-Native Forest Insets and Pathogens

The Continental Dialogue on Non-Native Forest Insects and Pathogens hosted a discussion of the risk of pest introduction via the plant trade during its recent annual meeting. Participants asked: How can the international phytosanitary system curtail introductions of unknown organisms when it is based on risk assessments that address only species that are fully known and – usually – have proven to be invasive elsewhere.

*Rhodomyrtos psidioides* in eastern Australia killed by myrtle rust; photo by Peter Entwistle

In recent decades, tens of species of Phytophthora have been introduced to countries around the world. Myrtle rust (Austropuccinia psidii) has been introduced to 27 countries from the U.S. to Australia and South Africa. The two causal agents of boxwood blight has been introduced to at least 24 countries in three geographic areas: Europe and western Asia; New Zealand; and North America. The ash decline fungus has been introduced across Europe. Most of these species were unknown to science at the time of their introduction. Other species were known – but not believed to pose a threat because, in their native regions, their co-evolved hosts are not harmed.

For more than a decade, scientists have noted that the international phytosanitary system has failed to prevent this rapid worldwide spread of significant pathogens 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.

During the Continental Dialogue discussion, Craig Regebrugge, Vice President of AmericanHort (the principal nursery trade association) noted the economic importance of greenhouse and nursery production and the importance of offering novel plants to their customers. Also, he noted that U.S. retail nurseries import primarily unrooted plant cuttings. In so doing, they have a strong incentive to ensure that they are pest-free in order to avoid delays arising during inspections. Those delays would probably kill these highly perishable products. Most U.S. imports of “finished” plants come from Canada. There have been pest problems; one of the most recent examples is a moth that attacks boxwoods (Buxus), which is the top-selling shrub crop in the U.S. Earlier there was confusion over whether plants shipped from British Columbia had been infected by the sudden oak death pathogen.

Regelbrugge noted that the industry’s voluntary integrated pest management program – Systems Approach to Nursery Certification (SANC) – currently has about two dozen participating nurseries. Hoped-for adoption by more of the hundreds of production nurseries in the country has been delayed by COVID-related travel restrictions, but he hopes to restore momentum. The industry is looking for opportunities to strengthen the program through marketing messages.

Regelbrugge and a second speaker, Rebecca Epanchin-Niell of the University of Maryland, warned that prohibitions on imports will stimulate smuggling. Both raised concerns about direct-to-consumer sales by e-commerce vendors and sought ideas on how to change the behavior of both exporters and consumers.

Later Sarah Green of British Forest Research asked the APHIS representative whether the agency’s import procedures are working to prevent introductions. She pointed to the issues raised by the scientific sources I cited above: pest risk analyses address only known organisms, so this process cannot protect importers from unknown organisms. She noted that the United Kingdom is struggling to contain a number of introductions of previously unknown pathogens. Gary Lovett of the Cary Institute noted that this weakness of pest risk assessments also hampers U.S. attempts to prevent introductions – especially of pathogens. He called on the Dialogue to focus on the resource at risk – native and urban forests – and change our phytosanitary programs on this basis. He has 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.

an American elm that has survived DED – at Longwood Gardens; photo by F.T. Campbell

Jiri Hulcr of the University of Florida tried to reassure Dialogue participants by stating that recent research has substantially reduced the threat from “unknown unkowns”. I applaud Dr. Hulcr’s efforts to reduce scientific uncertainty about the invasive potential of pathogens native to regions other than North America. His study might be the largest attempted by U.S.-based scientists. However, I note that his study assessed the threat posed by 55 insect-vectored fungi to two species of oak and two species of pines. The forests of the southeastern U.S. comprise many other tree genera! He also set a very high bar for defining a threat as serious: the damage to the host must be equivalent to that caused by Dutch elm disease or laurel wilt. Both have devastated their respective hosts. I believe U.S. phytosanitary policy must aim at protecting the full range of native species. Furthermore, levels of damage that affect the host’s role in the ecosystem – not just rapid mortality — should not be acceptable.

SOURCES

Epanchin-Niell, R., M. Springborn, an A. Lindsay. 2016. Resources No. 193 Fall 2016. http://www.rff.org/files/document/file/RFF_Resources_193_Web.pdf

Li, Y. C. Bateman, J. Skelton, B. Want, A. Black, Y-T. Huang, A. Gonzalez, M.A. Jusino, Z.J. Nolen, S. Freemen, Z. Mendel, C-Y. Chen, H-F. Li, M. Kolarik, M. Knizek, J-H. Park, W. Sittichaya, P.H. Thai, S-I. Ito, M. Torii, L. Gao, A.J. Johnson, M. Lu, J. Sun, Z. Zhang, D.C. Adams, J. Hulcr. 2021. Pre-invasion assessment of exotic bark beetle-vectored fungi to detect tree-killing pathogen. Phytopathology. https://doi.org/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

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).

Posted by Faith Campbell

www.fadingforests.org

West Coast Steps Up Efforts to Protect Ash

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

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

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

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

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

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

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

The newsletter is “Ash across the West”.

The first issue of the newsletter provides the following information:

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

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

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

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

Fresnillo (Fraxinus gooddingii) AZ

Gregg’s ash (Fraxinus greggii) AZ

OR ash (Fraxinus latifolia) WA, OR, CA

Chihuahuan ash (Fraxinus papillosa) AZ, NM, TX

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

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

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

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

Posted by Faith Campbell

www.fadingforests.org

Invasive shot hole borers: global threat; will international phytosanitary system prevent further spread?

ISHB-infested California sycamore; photo by Beatriz Nobua-Behrmann, University of California Cooperative Extension

Numerous ambrosia beetles have become introduced species. Their invasions are facilitated by their cryptic habits and ecologies, wide host ranges, and specialized breeding systems – all of which allow extremely low populations to start an infestation. The way they breed often results in low genetic diversity in their introduced ranges, but this has not hampered their success. [Bierman et al. 2022]

Also, ambrosia beetles carry fungi, which provide food needed by their larvae. While most of these fungi don’t harm living trees, some do. The United States has been invaded by three damaging ambrosia beetle-fungal complexes: laurel wilt in the Southeast, and Fusarium dieback disease, carried to southern California with polyphagous and Kuroshio shot hole borers.

These shot hole borers and their fungi represent an especially high risk to our forests because they can be transported in both living and dead wood. So not only massive U.S. imports of live plants but also the global movement of goods enclosed in solid wood packaging offer ready pathways for them to arrive and spread here. Neither pathway is regulated effectively enough to prevent either pest imports or interstate spread.

Invasive ambrosia beetles in California and Hawai’i

The invasive ambrosia beetles introduced to California are in the genus Euwallacea. This genus has undergone several taxonomic revisions. Now, the Euwallacea are divided into four species (Stouthammer 2017), of which three are in the U.S.:

Euwallacea fornicatus s.s. – common name polyphagous shot hole borer; first came to attention in southern California in 2012; formerly known as E. whitfordiodendrus.
E. perbrevis – common name tea shot hole borer; formerly known as E. fornicatus s.l.
E. kuroshio – unchanged nomenclature since detected in California in 2013;
E. fornicatior — apparently has not invaded outside of its native range in Asia.

Those now in the U.S. have been introduced to naïve habitats here and elsewhere, often with dire consequences. E. perbrevis, and possibly other species in the complex, are established on the Hawaiian islands.

For an extensive discussion of their introduction history go here

The Fungi: U.S. and Worldwide

Several fungal associates are vectored by the polyphagous shot hole borer (PSHB) and Kuroshio shot hole borer (KSHB). The most important are Fusarium euwallacea and Fusarium kuroshium, respectively. These fungi were only described after they appeared in California in the 2010s. They cause Fusarium dieback disease.

Because the two beetle species are difficult to distinguish and the associated diseases cause very similar impacts, Californians studying them and educating stakeholders now speak of the two beetle-fungus complexes as one unit, “invasive shot hole borers”.

Both PSHB and KSHB have numerous genetic strains, or haplotypes. For PSHB, the greatest haplotype diversity is in Asia – Thailand, Vietnam and China. Remember that these same regions are also a center of diversity for the huge genus Phytophthora, blog a genus widely recognized as containing many plant pathogens. https://www.dontmovefirewood.org/pest_pathogen/sudden-oak-death-syndrome-html/ One of the PSHB haplotypes, H33, has invaded many more regions than the others, including Israel, California, and South Africa. It has also been detected in several tropical plant greenhouses in Europe (where it has been eradicated). H33apparently is native to Vietnam – near Hanoi and Ho Chi Minh City – the country’s major ports (Rugman-Jones et al 2020 and pers. comm.). Does this haplotype’s spread to three continents reflect circumstances, such as the proximity of its native range to major ports and a “bridgehead effect” from its multiple introductions (the insects can be introduced to new regions on shipments from invaded regions established earlier)? Or does it point to an unknown genetic superiority (Bierman et al. 2022). This issue seems worth exploring.

I have blogged about the rising volume of imports from Vietnam, including to ports on the Gulf Coast –a region that has climatic similarities to Vietnam and known host species, so it seems quite vulnerable to invasion by either PSHB or KSHB.

A second species in the genus, KSHB, was detected in southern California in 2012; it has now spread to Mexico. So far, only one haplotype of this species has been detected in North America; this haplotype is widespread in Taiwan.

Finally, E. perbrevis (formerly known as E. fornicatus s.l.) has been detected in Florida, Hawai`i (island of Maui), and West Australia (to which it is probably native). This species has also been detected in nurseries in the Netherlands, where authorities report that it has been eradicated (Rugman-Jones et al. 2020).

Akacia koa – native tree in Hawai“i attacked by Euwallaceae; photo by David Eckhoff, via Flickr

Some species or haplotypes have been detected in only one introduced location: E. fornicatus H35 and E. kuroshio (H20) in California; H38 in South Africa; H43 on Oahu and the Big Island of Hawai`i; and an unnamed haplotype in West Australia (Rugman-Jones et al. 2020).

This is a brief guide to worldwide invasions by one or more Euwallacea-fungus complexes (Rugman-Jones et al. 2020):

Southern California — two haplotypes of E. fornicatus s.s. (H33 & H35) and E. kuroshio (one haplotype).
Hawai`i – a unique haplotype of E. fornicatus s.s. (H43) on Oahu, the Big Island, and possibly other islands; E. perbrevis on Maui and possibly other islands.
Israel — E. fornicatus s.s. haplotype H33 only.
South Africa — E. fornicatus s.s. haplotype H33 and a unique haplotype (H38).
Western Australia — a unique haplotype of E. fornicatus s.s. and E. perbrevis (which is probably native in northern Queensland).
Greenhouses in Europe – both E. fornicatus s.s. (haplotype not specified) and – in the Netherlands — E. perbrevis; both reported eradicated.

When a location has been invaded by two or more species or haplotypes, this is probably an indication of separate introductions. Multiple introductions thus are suspected in California (Stouthamer et al. 2017; Bierman et al. 2022); South Africa (Bierman et al. 2022); and Hawai`i (Bierman et al. 2022).

As is true of other pathogens, e.g., Phytophthoras, there appears to have been a spurt of introductions in recent decades, to, e.g., California, South Africa, and the second species in Hawai`i. Bierman et al 2022 note the constantly growing number of locations with introductions.

*Indigofera jucuna* – reproductive host of PSHB in South Africa; photo by Giardano de Barcelona

Impact and Spread

As is common in the case of forest pests, especially pathogens, detection occurred only years after the initial introduction. In South Africa this delay was five years – from 2012 to 2017 or 2018. In California, identification of the species as PSHB in 2012 was nine years after the organism was first detected in the state (2003).

Over the decade since 2012, PHSB, KSHB, and the pathogens they transmit have spread through large portions of southern California. KSHB has spread through “jumps” to distant locations in Orange, Los Angeles, and as far as Santa Barbara and Ventura counties. There have also been detections in even more distant San Luis Obispo and Santa Clara. These latter apparently have not become established.

A likely explanation for this pattern is the movement of firewood. (Rugman-Jones et al 2020 and pers. comm.) See the map here The two beetles and the plant pathogens they carry are expected to spread throughout much of California wherever their many host plants occur.

On Hawai`i, PSHB is attacking several endemic species including one of the largest forest trees, Acacia koa, as well as Pipturus albidus and Planchonella sandwicensis. Numerous non-native species growing on the Islandsare also attacked, including crops (Macadamia and Mangifera) and invasive species

In South Africa, PSHB has spread faster and farther. It has been present since at least 2012 (Stouthamer et al. 2017), although it was not identified until 2018. In about a decade it has spread to every province except Limpopo – PSHB’s largest geographical outbreak of this beetle [Bierman et al. 2022]

Hosts and Areas at Greatest Risk

Hundreds of plant species in at least 33 plant families support successful reproduction of both beetle and fungus. These include many species widespread in southern California, other parts of the U.S., and South Africa. Some California ecosystems are at particular risk because they are dominated by susceptible tree or shrub species. These vulnerable ecosystems are mixed evergreen forests, oak woodlands, foothill woodlands, and riparian habitats. In San Diego County alone, more than 58,000 acres of riparian woodlands are at risk (California Forest Pest Council).

Experience with the Kuroshio shot hole borer (KSHB) in the Tijuana River valley along the California-Mexico border demonstrates the importance of ecological factors in determining disease outcomes. Following introduction, the KSHB killed a high proportion of the willows near the main river channel. However, beginning in 2016, these trees have regrown to almost pre-infestation sizes. Lead researcher John Boland is not certain why these new, fast-growing trees have not been attacked by the KSHB which remains in the area. See links to the Boland studies below.

riparian forest in Tijuana River Valley after recovery from KSHB attack; photo by John Bolton

Urban forests are at particular risk. For example, in South Africa, conservative estimates were that 25% of urban trees would be lost (Bierman et al. 2022). In California, a model developed by Shannon Lynch found the cities at greatest jeopardy are San Diego, Los Angeles, the San Francisco Bay area, and Sacramento. In other areas in the state that lack data on city tree composition, Lynch applied climate models; this approach extended the list of threatened areas to the eastern half of southern California and other parts of the Central Valley. (Lynch presentation to ISHB webinar April 2022; 2^nd day.) In my view, this model should also be applied to cities in Arizona and Nevada with similar climates.

Management

Symptoms of PSHB attack and fungus infection differ among tree species. For illustrations of the symptoms on various species, visit here.

Most important, prevent the beetles’ spread through movement of dead or cut wood, e.g., green waste, firewood, and even large wood chips or mulch. Websites provide information on managing these sources.

Where the beetles have already established, California scientists recommend focusing management on heavily infested “amplifier trees”. On these trees, dead limbs should be pruned; dying trees and those with beetles infesting the main trunk should be removed. The wood must be disposed of properly.

Sources

Bierman, A., F. Roets, J.S. Terblanche. 2022. Population structure of the invasive ambrosia beetle, Euwallacea fornicatus, indicates multiple introductions into South Africa. Biol Invasions (2022) 24:2301–2312 https://doi.org/10.1007/s10530-022-02801-x

Boland, J.M. — all of Boland’s reports and articles on the KSHB are available at: The Ecology and Management of the Kuroshio Shot Hole Borer in the Tijuana River Valley — Tijuana Estuary : TRNERR]

California Forest Pest Council. 2015. 2015 California Forest Pest Conditions. http://bofdata.fire.ca.gov/hot_topics_resources/2015_california_forest_pest_conditions_report.pdf

Eskalen, A., Stouthamer, R., Lynch, S. C., Twizeyimana, M., Gonzalez, A., and Thibault, T. 2013. Host range of Fusarium dieback and its ambrosia beetle (Coleoptera: Scolytinae) vector in southern California. Plant Dis. 97:938-951.

Stouthamer, R., P. Rugman-Jones, P.Q. Thu, et al. 2017. Tracing the origin of a cryptic invader: phylogeography of the Euwallacea fornicatus (Coleoptera: Curculionidae: Scolytinae) species complex. Agric For Entomol 19:366-375. https://doi.org/10.1111/afe.12215

recordings of April 2022 webinar posted at https://youtu.be/RyqJYyLkshk day 1; and https://youtu.be/kWmtcbjTczw day 2

Posted by Faith Campbell

www.fadingforests.org

Hundreds of U.S. Tree Species Endangered, Most due to Non-Native Pests

Horton House on Jekyll Island, Georgia before laurel wilt killed the giant redbay trees; photo by F.T. Campbell

Close to four hundred tree species native to the United States are at risk of extinction. The threats come mainly from non-native insects and diseases – a threat we know gets far too little funding, policy attention, and research.

As Murphy Westwood, Vice President of Science and Conservation at the Morton Arboretum, which led the U.S. portion of a major new study, said to Gabriel Popkin, writing for Science: “We have the technology and resources to shift the needle,” she says. “We can make a difference. We have to try.”

Staggering Numbers

More than 100 tree species native to the “lower 48” states are endangered (Carrero et al. 2022; full citation at the end of this blog). These data come from a global effort to evaluate tree species’ conservation status around the world. I reported on the global project and its U.S. component in September 2021. This month Christina Carrero and colleagues (full citation at the end of this blog) published a summary of the overall picture for the 881 “tree” species (including palms and some cacti and yuccas) native to the contiguous U.S. (the “lower 48”).

This study did not address tree species in Hawai`i or the U.S. Pacific and Caribbean territories. However, we know that another 241 Hawaiian tree species are imperiled (Megan Barstow, cited here).

Assessing Threats: IUCN, NatureServe, and CAPTURE

Carrero and colleagues assessed trees’ status by applying methods developed by IUCN and NatureServe. (See the article for descriptions of these methods.) These two systems consider all types of threats. Meanwhile, three years ago Forest Service scientists assessed the specific impacts of non-native insects and pathogens on tree species in the “lower 48” states and Alaska in “Project CAPTURE” (Conservation Assessment and Prioritization of Forest Trees Under Risk of Extirpation). All three systems propose priorities for conservation efforts. For CAPTURE’s, go here.

Analyses carried out under all three systems (IUCN, NatureServe, and CAPTURE) concur that large numbers of tree species are imperiled. Both IUCN and CAPTURE agree that non-native insects and pathogens are a major cause of that endangerment. While the overall number of threatened species remained about the same for all three systems, NatureServe rated threats much lower for many of the tree species that IUCN and CAPTURE considered most imperiled.

This difference arises from the criteria used to rate a species as at risk. IUCN’s Criterion A is reduction in population size. Under this criterion, even extremely widespread and abundant species can qualify as threatened if the population declines by at least 30% over three generations in the past, present, and/or projected future. NatureServe’s assessment takes into account rapid population decline, but also considers other factors, for example, range size, number of occurrences, and total population size. As a result, widespread taxa are less likely to be placed in “at risk” categories in NatureServe’s system.

In my view, the IUCN criteria better reflect our experience with expanding threats from introduced pests. Chestnut blight, white pine blister rust, dogwood anthracnose, emerald ash borer, laurel wilt disease, beech leaf disease, and other examples all show how rapidly introduced pathogens and insects can spread throughout their hosts’ ranges. (All these pests are profiled here . ) They can change a species’ conservation status within decades whether that host is widespread or not.

Which Species Are at Risk: IUCN

Carrero and colleagues found that under both IUCN and NatureServe criteria, 11% to 16% of the 881 species native to the “lower 48” states are endangered. Another five species are possibly extinct in the wild. Four of the extinct species are hawthorns (Crataegus); the fifth is the Franklin tree (Franklinia alatamaha) from Georgia. A single specimen of a sixth species, an oak native to Texas (Quercus tardifolia),was recently re-discovered in Big Bend National Park.

Franklinia (with Bachman’s warbler); both are extinct in the wild; painting by John Jacob Audubon

The oak and hawthorn genera each has more than 80 species. Relying on the IUCN process, Carrero and colleagues found that a significant number of these are at risk: 17 oaks (20% of all species in the genus); 29 hawthorns (34.5% percent). A similar proportion of species in the fir (Abies), birch (Betula), and walnut (Juglans) genera are also threatened.

Other genera have an even higher proportion of their species under threat, per the IUCN process:

all species in five tree genera, including Persea (redbay, swampbay) and Torreya (yews);
two-thirds of chestnuts and chinkapins (Castanea), and cypress (Cupressus);
almost half (46.7%) of ash trees (Fraxinus).

Pines are less threatened as a group, with 15% of species under threat. However, some of these pines are keystone species in their ecosystems, for example the whitebark pine of high western mountains.

Carrero et al. conclude that the principal threats to these tree species are problematic and invasive species; climate change and severe weather; modifications of natural systems; and overharvest (especially logging). Non-native insects and pathogens threaten about 40 species already ranked by the IUCN criteria as being at risk and another 100 species that are not so ranked. Climate change is threatening about 90 species overall.

Considering the invasive species threat, Carrero and colleagues cite specifically ash trees and the bays (Persea spp.). In only 30 years, the emerald ash borer has put five of 14 ash species at risk. All these species are widespread, so they are unlikely to be threatened by other, more localized, causes. In about 20 years, laurel wilt disease threatens to cause extinction of all U.S. tree species in the Persea genus.

Carrero and colleagues note that conservation and restoration of a country’s trees and native forests are extremely important in achieving other conservation goals, including mitigating climate change, regulating water cycles, removing pollutants from the air, and supporting human well-being. They note also forests’ economic importance.

As I noted above, USFS scientists’ “Project CAPTURE” also identified species that deserve immediate conservation efforts.

Where Risk Assessments Diverge

All three systems for assessing risks agree about the severe threat to narrowly endemic Florida torreya and Carolina hemlock.

With three risk ranking systems, all can agree (as above), all can disagree, or pairs can agree in four different ways. Groups of trees fall into each pair, with various degrees of divergence. Generally, only two of the three systems agree on more widespread species:

black ash: IUCN and Project CAPTURE prioritize this species. NatureServe ranked it as “secure” (G5) as recently as 2016.
whitebark pine: considered endangered by IUCN, “vulnerable” (G3) by NatureServe. The US Fish and Wildlife Service has proposed listing the species as “threatened” under the Endangered Species Act. https://www.fws.gov/species-publication-action/endangered-and-threatened-wildlife-and-plants-threatened-species-18 However, Project CAPTURE does not include it among its highest priorities for conservation. Perhaps this is because there are significant resistance breeding and restoration projects already under way.
tanoak: considered secure by both IUCN and NatureServe, but prioritized by Project CAPTURE for protection.

dead tanoak in Curry County, Oregon; photo by Oregon Department of Forestry

Carrero notes the divergence between IUCN and NatureServe regarding ashes. Four species ranked “apparently secure” (G4) by NatureServe (Carolina, pumpkin, white, and green ash) are all considered vulnerable by IUCN. They are also prioritized by Project CAPTURE. I have described the impact of the emerald ash borer on black ash. Deborah McCullough, noted expert on ash status after invasion by the emerald ash borer, also objects to designating this species as “secure” (pers. comm.).

This same divergence appears for eastern hemlock.

Port-Orford cedar is currently ranked as at risk by IUCN and Project CAPTURE, but not NatureServe. Growing success of the restoration breeding project has prompted IUCN to change the species’ rank from “vulnerable” to “near threatened”. IUCN is expected to reclassify it as of “least concern” in about a decade if breeding efforts continue to be successful (Sniezko presentation to POC restoration webinar February 2022).

While these differing detailed assessments are puzzling, the main points are clear: several hundred of America’s tree species (including many in Hawai`i, which – after all – is our 50^th state!) are endangered and current conservation and restoration efforts are inadequate.

Furthermore, a tree species loses its function in the ecosystem long before it becomes extinct. It might still be quite numerous throughout its range – but if each individual has shrunken in size it cannot provide the same ecosystem services. Think of thickets of beech root sprouts – they cannot provide the bounteous nut crops and nesting cavities so important to wildlife. Extinction is the extreme. We should act to conserve species much earlier.

YOU CAN HELP!

Congress is considering the next Farm Bill – which is due to be adopted in 2023. Despite its title, this legislation has often provided authorization and funding for forest conservation (for example, the US Forest Service’ Landscape Scale Restoration Program).

There is already a bill in the House of Representatives aimed at improving the US Department of Agriculture’s prevention and early detection/rapid response programs for invasive pests. Also, it would greatly enhance efforts to restore decimated tree species via resistance breeding, biocontrol, and other strategies. This bill is H.R. 1389.

The bill was introduced by Rep. Peter Welch of Vermont, who has been a solid ally and led on this issue for several years. As of August 2022, the bill has seven cosponsors, most from the Northeast: Rep. Mike Thompson [CA], Rep. Chellie Pingree [ME], Reps. Ann M. Kuster and Chris Pappas [NH], Rep. Elise Stefanik [NY], Rep. Deborah K. Ross [NC], Rep. Brian Fitzpatrick [PA].

Please write your Representative and Senators. Urge them to seek incorporation of H.R. 1389 in the 2023 Farm Bill. Also, ask them to become co-sponsors for the House or Senate bills. (Members of the key House and Senate Committees are listed below, along with supporting organizations and other details.)

Details of the Proposed Legislation

The Invasive Species Prevention and Forest Restoration Act [H.R. 1389]

Expands USDA APHIS’ access to emergency funding to combat invasive species when existing federal funds are insufficient and broadens the range of actives that these funds can support.
Establishes a grant program to support research on resistance breeding, biocontrol, and other methods to counter tree-killing introduced insects and pathogens.
Establishes a second grant program to support application of promising research findings from the first grant program, that is, entities that will grow large numbers of pest-resistant propagules, plant them in forests – and care for them so they survive and thrive.
[A successful restoration program requires both early-stage research to identify strategies and other scientists and institutions who can apply that learning; see how the fit together here.]

Mandates a study to identify actions needed to overcome the lack of centralization and prioritization of non-native insect and pathogen research and response within the federal government, and develop national strategies for saving tree species.

Incorporating the provisions of H.R. 1389 into the 2023 Farm Bill would boost USDA’s efforts to counter bioinvasion. As Carrera and colleagues and the Morton Arboretum study on which their paper is based demonstrate, our tree species desperately need stronger policies and more generous funding. Federal and state measures to prevent more non-native pathogen and insect pest introductions – and the funding to support this work – have been insufficient for years. New tree-killing pests continue to enter the country and make that deficit larger –see beech leaf disease here. Those here, spread – see emerald ash borer to Oregon.

For example, funding for the USDA Forest Service Forest Health Protection program has been cut by about 50%; funding for USFS Research projects that target 10 high-profile non-native pests has been cut by about 70%.

H.R. 1389 is endorsed by several organizations in the Northeast: Audubon Vermont, the Maine Woodland Owners Association, Massachusetts Forest Alliance, The Nature Conservancy Vermont, the New Hampshire Timberland Owners Association, Vermont Woodlands Association, and the Pennsylvania Forestry Association.

Also, major forest-related national organizations support the bill: The American Chestnut Foundation (TACF), American Forest Foundation, The Association of Consulting Foresters (ACF), Center for Invasive Species Prevention, Ecological Society of America, Entomological Society of America, National Alliance of Forest Owners (NAFO), National Association of State Foresters (NASF), National Woodland Owners Association (NWOA), North American Invasive Species Management Association (NAISMA), Reduce Risk from Invasive Species Coalition, The Society of American Foresters (SAF).

HOUSE AND SENATE AGRICULTURE COMMITTEE MEMBERS – BY STATE

STATE	Member, House Committee	Member, Senate Committee	Key members * committee leadership # forestry subcommittee leadership @ cosponsor of H.R. 1389
Alabama	Barry Moore
Arizona	Tom O’Halleran
Arkansas	Rick Crawford	John Boozman*
California	Jim Costa Salud Carbajal Ro Khanna Lou Correa Josh Harder Jimmie Panetta Doug LaMalfa
Colorado		Michael Bennet #
Connecticut	Jahana Hayes
Florida	Al Lawson Kat Cammack
Georgia	David Scott * Sanford Bishop Austin Scott Rick Allen	Raphael Warnock Tommy Tuberville
Illinois	Bobby Rush Cheri Bustos Rodney Davis Mary Miller	Richard Durbin	Note that the report was led by scientists at the Morton Arboretum – in Illinois!
Indiana	Jim Baird	Mike Braun
Iowa	Cindy Axne Randy Feenstra	Joni Ernst Charles Grassley
Kansas	Sharice Davids Tracey Mann	Roger Marshall#
Kentucky		Mitch McConnell
Maine	Chellie Pingree @
Massachusetts	Jim McGovern
Michigan		Debbie Stabenow *
Minnesota	Angie Craig Michelle Fischbach	Amy Klobuchar Tina Smith
Mississippi	Trent Kelly	Cindy Hyde-Smith
Missouri	Vicky Hartzler
Nebraska	Don Bacon	Deb Fischer
New Hampshire	Ann McLane Kuster @
New Jersey		Cory Booker
New Mexico		Ben Ray Lujan
New York	Sean Patrick Maloney Chris Jacobs	Kristen Gillibrand
North Carolina	Alma Adams David Rouzer
North Dakota		John Hoeven
Ohio	Shontel Brown Marcy Kaptur Troy Balderson	Sherrod Brown
Pennsylvania	Glenn Thompson
South Dakota	Dusty Johnson	John Thune
Tennessee	Scott DesJarlais
Texas	Michael Cloud Mayra Flores
Vermont		Patrick Leahy
Virginia	Abigail Spanberger #
Washington	Kim Schreir

SOURCES

Christina Carrero, et al. Data sharing for conservation: A standardized checklist of US native tree species and threat assessments to prioritize and coordinate action. Plants People Planet. 2022;1–17. wileyonlinelibrary.com/journal/ppp3

Washington Post: Sarah Kaplan, “As many as one in six U.S. tree species is threatened with extinction” https://www.washingtonpost.com/climate-environment/2022/08/23/extinct-tree-species-sequoias/

Popkin, G. “Up to 135 tree species face extinction—and just eight enjoy federal protection”, Science August 25, 2022. https://www.science.org/content/article/135-u-s-tree-species-face-extinction-and-just-eight-enjoy-federal-protection

Posted by Faith Campbell

www.fadingforests.org

Posted by Faith Campbell