Phytopthora ramorum-infected potted plants; photo by Washington State University
At this year’s USDA Invasive Species Forum I will be seeking to promote a discussion of what American and other stakeholders can do to suppress spread of forest pathogens. I have raised this issue many times before. To see my blogs about the P4P pathway, scroll down below the archives to the “categories”. See especially here and here.
I note that:
Non-native invasive pathogens and pests are decimating forests worldwide, threatening biodiversity & limiting efforts to rely on forests to alleviate impacts of climate change.
Many of the most damaging non-native organisms are pathogens that are especially difficult to detect at borders or to contain or eradicate once introduced.
A principal pathway by which pathogens are introduced is the international trade in living plants, or “plants for planting” (P4P).
Forest pathologists have long advocated a more pro-active approach – but national and international plant health officials have not taken up the challenge. [think Clive Brasier, Bitty Roy, Thomas Jung, Michael Winfield …]
Austropuccinia psidii on Melalecua in Australia; John Tann via Flickr
At the global level I suggest that we need:
National agricultural agencies, stakeholders, FAO & International Plant Protection Convention (IPPC) to consider amending IPPC requirement that scientists identify a disease’s causal agents before regulating it. I think experience shows that this policy virtually guarantees that pathogens will continue to enter, establish, & damage natural and agricultural environments.
National governments & FAO / IPPC to fund greatly expanded research to identify microbes resident in regions that are important sources of origin for traded plants, vulnerability of hosts in importing countries, and new technologies for detecting pathogens (e.g., molecular tools, volatile organic compounds [VOCs]).
Researchers & agencies to expand international “sentinel plants” networks; incorporate data from forestry plantations, urban plantings, etc. of non-native trees.
Application of ISPM#36 to promote use of HACCP programs for plants in trade. (See also my discussion in Fading Forests III – link at end of this blog.)
‘ohi‘a trees killed by rapid ‘ohi‘a death; photo by Richard Sniezko, USFS
We Americans need to
Evaluate efficacy of current regulations – incorporating NAPPRA & Q-37 revision. Rely on AQIM data. Include arthropods, fungal pathogens, oomycetes, bacteria, viruses, nematodes. Include threats to U.S. tropical islands (Hawai`i, Puerto Rico, Guam, etc.) which are centers of plant endemism.
Apply existing programs (e.g., NAPPRA, Clean Stock Network, post-entry quarantine) to strictly regulate trade in plant taxa most likely to transport pests that threaten our native plants; e.g., plants belonging to genera shared between North American trees & plants on other continents.
Recognize that plant nurseries are incubators for microbial growth, hybridization, and evolution; require nurseries to adopt sanitary operation procedures regardless of whether they sell in inter-state or intra-state commerce
I will explain my sense of urgency by noting the many recent introductions of pathogens – most probably via P4P or cut vegetation:
13 outbreaks of Phytophthora-caused disease in forests and natural ecosystems of Europe, Australia and the Americas. Three of four known strains of P. ramorum are established in U.S. forests.
Myrtle rust (Austropuccinia psidii) has been introduced to 27 countries, including the U.S., Australia, and South Africa.
Two new species of Ceratocystis (C. lukohia & C. huliohia)—causal agents of rapid ‘ohi‘a death (ROD) – spreading on the Hawaiian Islands. The former species appears to have originated in the Caribbean; the latter in Asia.
Since 2012, beech leaf disease has spread from northeastern Ohio to Maine.
Boxwood blight (caused by 2 ascomycete fungi, Calonectria pseudonaviculata & C. henricotiae) introduced to at least 24 countries in 3 geographic areas: Europe / western Asia; New Zealand, North America.
ash dieback fungus (Hymenoscyphus fraxineus) has spread across Europe after introduction from Asia.
What do you think? Can we find more effective methods to curtail introductions?
beech leaf disease
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
Platanus orientalis in Turkey; photo by Zeynek Zebeci
A current issue of the journal Forests (2022 Vol. 13) is a special issue focused on forest pests. This topic was chosen because of increased pest incursions. Choi and Park (full citations at the end of the blog) link this to climate change and increased international trade, as well as difficulties of predicting which pests will cause damage where.
The journal issue contains 15 papers. Several patterns appear throughout. First is the important role of international trade in living plants – “plants for planting” – in introductions. This is hardly news! A second pattern is that at least two North American species were introduced to Europe during the 1940s, probably in wood packaging used to transport military supplies during World War II.
This compilation provides the opportunity to review which organisms of North American origin have become damaging invaders in Eurasia — and sometimes other continents. For example, the journal carries four articles discussing pine wilt disease (PWD). It is caused by the North American nematode Bursaphelenchus xylophilus, and is vectored by wood-boring insects in the genus Monochamus. Beetles introduced from North America and those native to the invaded area are both involved. This disease is considered a severe threat to forest health globally. No apparent association with WWII exists for PWD.
Two fungal pathogens from North America cause serious damage in urban and natural forests of Europe and central Asia. Neither is discussed in the special issue:
Ceratocystis platani has devastated urban trees in the Platanus genus, especially the “London plane” hybrid, and the native European tree, Platanus orientalis. This fungus was accidentally introduced to southern Europe during WWII – as were the two insects described by Musolin et al. It was first reported in northern Italy and Mediterranean France in the early 1970s, but disease symptoms had been observed years earlier. C. platani is established across the northern rim of the Mediterranean and to the east in Armenia and Iran. The worst damage has been in Greece, especially in natural forest stands in riparian areas. Spread of the pathogen there is facilitated by root grafts and by tree wounds caused by floating wooden debris during floods (Tsopelas et al. 2017.)
Platanus orientalis along Voidomatis River in Greece; photo by Onno Zweers, via Wikimedia
Heterobasidion irregulare infects conifers. It has spread and killed large numbers of Italian stone pine (Pinus pinea). The disease was inadvertently introduced to central Italy in the 1940s. H. irregulare has greater sporulation potential and decays wood more quickly than the native congener H. annosum. H. irregulare appears to be replacing the European species; scientists fear it will exacerbate tree infection and mortality rates (Garbelotto, Leone, and Martiniuc. date?)
A third North American pathogen, sooty bark disease (Cryptostroma corticale) has been introduced to Europe. This disease, found on sugar maple in eastern North America, was detected in Great Britain in 1945; it is now throughout Europe (Tanney 2022). EPPO reports that it is widespread in western Europe and in some Balkan countries. The website provides no information on its impact in Europe.
Pests in Russia
A paper authored by Musolin, et al. discusses 14 species of invasive or emerging tree pests found in Russian forest and urban ecosystems. Of these, two are native to North America. Another eight pose a threat to North America if they are introduced here.
As Musolin et al. point out, Russia covers a huge territory across Europe and Asia – stretching 10,500 km, or 6,500 miles. These encompass a great variety of ecological zones. Russia is also actively involved in international trade. It is not surprising, then, numerous non-native organisms have been introduced.
As of 2011, 192 species of phytophagous non-native insects from 48 families and eight orders were documented in the European part of Russia. This number does not include the vast areas in Asian Russia. Additional introductions have probably occurred in the most recent decade. Some of these introduced species have cause significant economic losses. Still, Russia appears to rarely mount a serious control effort.
Of course, the opposite is also true: pests native to some part of Russia can be transported to new regions of Russia or beyond its borders. We North Americans have focused on various species of tussock moths (Lymantria spp., etc.). There are many others. Musolin et al. describe eight in detail. All the information in this blog are from that article unless otherwise indicated.
Two North American Species’ Damage in Eurasia
Both these introductions were detected around the year 2000. Was there some event – other than simply expanding trade – that might explain these introductions?
Leptoglossus occidentalis; photo by nutmeg66 via Flickr
Western Coniferous Seed Bug, Leptoglossus occidentalis
This insect from western North America has invaded Eurasia, North Africa, and Central America. The first detection in Europe was in 1999 in Italy. It spread quickly and is present now from Morocco to Japan, as well as in South Africa and South America. The seed bug is spreading northward in European Russia, including into the forest-steppe zone. Its ability to spread to the East is uncertain.
L. occidentalis attacks a wide range of Pinaceae and Cupressaceae. In the Mediterranean region it has had serious impacts on the pine nut supply (Ana Farinha, IUFRO, Prague, September 2021). In southern parts of Russia it has caused “significant damage”. L. occidentalis also vectors a pathogenic fungus Sphaeropsis sapinea (=Diplodia pinea), which causes diplodia tip blight. The cumulative damage of insect and pathogen to pines can be significant.
The introduction pathway to Russia is unknown. It might have flown from established populations in Europe, or it might have been transported on plants for planting or Christmas decorations.
Oak Lace Bug, Corythucha arcuata
This insect is widespread in the United States and southern Canada. It was first detected in Europe – again, Italy – in 2000. Twenty years later it has spread to almost 20 countries.
Russia was invaded relatively recently; the first outbreak was detected in 2015 in the subtropical zone along the Black Sea coast and Caucasus. Musolin et al. expect the lace bug to spread to natural forests of Central Asia and other countries of the Caucasus. Its spread will be assisted by air currents and movement of plants for planting. The insect is causing considerable aesthetic damage, but other impacts have not been estimated.
Hosts include many species of oak (Quercus spp.), European and American chestnuts (Castanea spp.) plus trees from other botanical families: willows and maples (Salicaceae), redbay (Fagaceae), and alder (Betulaceae).
Pests in Russia that Could Damage North America if Introduced Here
Malus sierversii; photo by Lukacz Szczurowski via Wikimedia
Threat to Apples — Apple Buprestid, Agrilus mali
This Asian beetle has caused extensive mortality of wild apple (Malus sieversii) forests in Xinjiang, China. Wild apple trees are important components of deciduous forests in the Central Asian mountains. The species is also an ancestor of the domestic apple tree. Consequently, the borer is considered a potential threat to cultivated apple trees – presumably everywhere. A. mali might also attack other fruit trees in the Rose family, i.e., Prunus (plums, cherries, peaches, apricots, almonds) and Pyrus (pears).
Unlike most of the other species described here, A. mali is a quarantine pest in Russia and across Europe and the Mediterranean regions – the region where phytosanitary policies are coordinated by the European and Mediterranean Plant Protection Organization (EPPO). Russia bans imports of apple seedlings from infested areas.
China is reported to be experimenting with a possible biocontrol agent, Sclerodermus pupariae (a parasitoid of emerald ash borer).
Threat to Pines and Firs, Already Under Invasive Species Threats
Small Spruce Bark Beetle, Ips amitinus
This European beetle has been considered a secondary pest of dying conifers. Over the last 100 years, it has moved farther North. The first Russian record was 100 years ago, in the region where Russia, Belarus, and Ukraine meet. (Did military action during World War I play a role? This is not discussed by the authors.) By 2022, the beetle occupies 31 million ha. It is probably spread through transport of logs by rail.
In Western Siberia, the spruce beetle has attacked a new host, Siberian pine (Pinus sibirica).
The danger to North America arises from this beetle’s preference for five-needle pines (genus Pinus section Quinquefoliae). North America’s five-needle pines are already under severe pressure from the introduced pathogen white pine blister rust (Cornartium ribicola) and the native mountain pine beetle (Dendroctonus ponderosae).
Four-Eyed Fir Bark Beetle, Polygraphus proximus
This East Asian beetle feeds on firs (Abies spp.). Less commonly, it feeds on other genera in the Pinaceae: spruce (Picea ), pines (Pinus), larch (Larix), hemlock (Tsuga).
This beetle has been spreading west; the first substantiated record in European Russia was 2006 in Moscow. The beetle was probably present in western Siberia in the 1960s, although it was not detected until 2008. Again, the probable pathway of spread is movement of lumber by railroad.
P. proximus vectors an obligate symbiotic fungus, which can rapidly weaken the host. Musolin et al. comment on the beetle’s impacts – which they rarely do in this article. (Does this signify more damaging impacts, or availability of past studies?) They note significant changes in the forests’ ecosystem structure and microclimate, vegetation cover, and local insect fauna.
The danger to North America arises from this beetle’s preference for firs from the sections Balsamea and Grandis. Many North American firs are in these sections, including Fraser fir (Abies fraseri), balsam fir (A. balsamea), subalpine fir (A. lasiocarpa), grand fir (A. grandis), white fir (A. concolor), and others. Several of these firs already are challenged by the introduced balsam woolly adelgid. Firs in central and western Europe are less vulnerable since they are in the section Abies, which the beetle prefers less.
Threats to Poplars
Spotted Poplar Borer,Agrilus fleischeri
This boring beetle is native to northern Asia. It has caused significant mortality in native and exotic Populus plantations in China. Although there have been no reports of this beetle moving beyond its native range, many other Agrilus species have. Canada has twice intercepted adult spotted poplar borers on wood packaging. Musolin et al. fear that the adoption of non-native hosts might trigger an outbreak that would facilitate spread.
Poplar Leafminer, Phyllonorycter populifoliella
balsam poplar; photo by Matt Lavin via Flickr
This micromoth is widely distributed across the Palearctic. It was recently detected on introduced poplars growing in India.
The danger to North America arises from the beetle’s preference for black and balsam poplars. Several species in these taxonomic groups are common in North America, including Populus balsamifera, P. trichocarpa, P. deltoides, and Populus × Canadensis.
Threat to Oaks — Leaf Blotch Miner Moth, Acrocercops brongniardella
This micromoth is widely distributed in Europe and expanding to the north. The pest mines the leaves of several oak species (Quercus spp.), especially English oak, Q. robur; and sometimes European chestnut (Castanea sativa). Leaf blotch miner is considered one of the most important folivore insect pests of oaks in Russia. Damage has been greater in Omsk Oblast (Siberia), where both English oak and the micromoth are introduced species, than in St. Petersburg, which is on the northern limit of their natural range. Musolin et al. fear that the warming climate will lead to the pest causing greater damage in the northern portions of its range.
Threat to Basswood — Lime Leaf Miner, Phyllonorycter issikii
This Asian moth has been moving west since the mid-1980s. It now occupies most of European Russia with some outbreaks in Siberia. In Europe, it is a conspicuous pest of Tilia species.
In these invaded regions, the leaf miner has shifted to novel hosts, including American basswood (T. americana). Basswood is a common plant in the eastern deciduous forest of North America.
Threat to Horse Chestnuts & Urban Trees — Horse-Chestnut Leaf Miner, Cameraria ohridella
This tiny moth was unknown to science before the first recorded outbreak in the late 1980s. Over the next three decades it spread to most of Europe, where horse chestnut (Aesculus hippocastanum)has been widely planted for three centuries. It has caused significant damage.
The first Russian detection was in Kaliningrad, on the shores of the Baltic Sea, in 2003. The leaf miner now occupies 69% of administrative units of European Russia. It is considered one of the Top 100 most dangerous invasive species in Russia.
In North America, the moth might attack native horse chestnuts, Ae. octandra (=flava) and Ae. glabra. Urban plantings are at particular risk because the leaf miner might attack both European horse chestnuts and two non-native maples that have been planted widely, sycamore maple (Acer pseudoplatanus) and Norway maple (A. platanoides). Data cited by Musolin et al. are contradictory regarding larval development on the maples. Once introduced, the leaf miner is difficult to contain because it spreads through natural flight of adults, wind-blown leaves, hitchhiking on vehicles, and movement of infected plants.
Shared Pests
Russia has been invaded by two species that have been introduced in many countries (beyond pine wilt nematode). These two entered the country on plants for planting being imported to landscape venues for the XXII Winter Olympic Games – held in Sochi in 2014.
First to arrive was the Box Tree Moth, Cydalima perspectalis. This East Asian species was first detected outside its native range in Germany in 2006. By 2011 it was widespread in European and Mediterranean countries. In 2021, the boxwood moth was found in North America (first Canada, then the United States). [I discuss the boxwood moth briefly here.]
boxtree moth; photographer unknown
In Russia, box tree moth larvae were first recorded in 2012 on the planting stock of its principal host, Buxus sempervirens. The moth quickly spread around the Black Sea region and to the North Caucasus. It spread farther, too: it reached the Kaliningrad Oblast (southeast coast of the Baltic Sea) in 2020. The main pathway of C. perspectalis invasion was the introduction of infested box-wood planting material.
Further spread of C. perspectalis is likely from Russia into the natural forests across the Caucasus (Transcaucasia) and to countries located further south. This is most distressing because the region has extensive natural forests of Buxus sempervirens. In 2015–2017, C. perspectalis almost completely destroyed the natural boxwood populationsin these regions of Russia and further eastwards in Abkhazia. Boxwood stands in Georgia and northern Iran are already suffering intensive defoliation as the result of infection by two non-native pathogens, Calonectria pseudonaviculata [synonym Cylindrocladium buxicola] and Calonectria henricotiae. Damage to these forests could lead to reductions in soil stability and subsequent declines in water quality and flood protection, changes in forest structure and composition, and declines in Buxus-associated biodiversity (at least 63 species of lichens, fungi, chromista and invertebrates might be obligate). (In December 2022, Iryna Matsiakh presented a compelling overview of threats to these forests in a webinar sponsored by the Horticulture Research Initiative; apparently no recording is available.)
The second global invader to appear was the Brown Marmorated Stink Bug, Halyomorpha halys.
This insect from southeast and east Asia invaded the United States in 1996. The first detection in Europe was in Liechtenstein in 2004. In both cases, it spread quickly across these continents.
Russia’s first detection of stinkbug was in 2014 in parks in Sochi and elsewhere along the Black Sea coast. The spread in Russia appears to have been limited to the Black Sea – Caucasus area.
The brown marmorated stinkbug is highly polyphagous, feeding on more than 300 species of plants. In southern Russia, 107 species have been documented as hosts. At times, stinkbug feeding has caused severe losses in yields of fruit and vegetable crops.
Patterns
Musolin et al. stress the importance of the pest shifting to new hosts–usually from the same or a closely related genus. They cite several examples of these shifts occurring in the pest’s native range, including Agrilus planipennis (from local Asian ash species to introduced North American ash species); Phyllonorycter populifoliella and Agrilus fleischeri (from local poplars to widely cultivated introduced North American poplars and hybrids); Agrilus mali (from cultivated to wild apples).
As I noted above, the introduction and spread pathways are the usual ones: plants for planting (three species) and shipments of logs. There is one indication of wood packaging – Spotted Poplar Borer, Agrilus fleischeri at the Canadian border.
Garbelotto, M., G. Lione, and A.V. Martiniuc. date? The alien invasive forest pathogen Heterobasidion irregulare is replacing the native Heterobasidion annosum. Biological Invasions https://doi.org/10.1007/s10530-022-02775-w
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.
Tanney, J. Forest Health Challenges Exacerbated by a Changing Climate: Swiss Needle Cast and Sooty Bark Disease in B.C. 65th ANNUAL FOREST PEST MANAGEMENT FORUM (Canada). December 7, 2022.
Tsopelas, P., A. Santini, M.J. Wingfield, and Z.W. de Beer. Canker Stain: A Lethal Disease Destroying Iconic Plane Trees. Plant Disease 2017. 101-645-658 American Phytopathological Society
Spruce budworm (Choristoneura fumiferana); photo by Jerald E. Dewey, USFS; via Bugwood; populations of several forest birds, including Cape May, Tennessee and Bay-Breasted warblers, become more numerous during budworm outbreaks
Now that Drs. Ziska and Aucott have educated us about the strong impact atmospheric CO2 can have on both plants and phytopagous insects, I have asked the experts whether these interactions have been incorporated in the models scientists are using to forecast pest activity in American forests as the climate changes.
The answer is no.
bay-breasted warbler; photograph by Dave Inman at Presque Isle State Park, PA; via Flickr
Dr. Bethany A. Bradley, Co-Director, Northeast Climate Adaptation Science Center at the University of Massachusetts, says empirical models of species range shifts typically only use climate and sometimes other environmental factors (like soils or topography) as predictors of potential geography. Inclusion of demographic processes like how plant growth is affected by more or less water, CO2, competition with other plants etc. would require a lot of data. It is currently impossible since there are tens of thousands of plant species interacting in the forests of eastern North America – and perhaps these factors have been analysed for only a hundred of them.
Mike Aucott points to the same difficulty: inclusion of CO2 in models of the future populations of specific plants would be difficult since the impacts vary from species to species and are compounded by other factors such as soil nitrogen levels, moisture levels, temperature, presence of competing plants, etc.
Regarding insects, Dr. Aucott thinks it is clear that some orders, such as Lepidoptera, don’t fare as well when feeding on plants grown under elevated CO2. He is not aware of efforts to model impacts of high CO2 on specific insects or even orders or feeding guilds.
juniper geometer (inchworm); Dr. Tallamy says inchworms are hairless & good tasting – so sought by birds
Dr. Ziska concurs about the difficulties. Dr. Ziska asks why there is so little funding to study these issues, especially given their probable impact on human food supplies and health – as described in his blog and an opinion piece published in Scientific American two years ago.
I hope that scientists, decision-makers, readers of this blog … maybe even the media! – take into consideration these complexities, even if they cannot be defined.
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 – [but do not address climate or CO2 aspects] review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
Wildfire: one of the widely recognized results of climate change (The Pioneer Fire located in the Boise National Forest near Idaho City, ID began on Jul. 18, 2016 and the cause is under investigation. The Pioneer Fire has consumed 96,469 acres. U.S. Forest Service photo. Original public domain image from Flickr)
A guest blog by Michael Aucott. Mike is a retired research scientist of the NJ Department of Environmental Protection. He has also taught chemistry at the College of New Jersey. He is currently a member of the NJDEP Science Advisory Board Standing Committee on Climate and Atmospheric Sciences, and on the board of directors of the PA/NJ Chapter of the American Chestnut Foundation. If you wish to contact Mike, use the contact button on this website. You MUST include your email address; it is not recorded automatically.
Two major perturbations affect Earth and its living systems, climate change and habitat disruptions. Emerging data show that these are more closely related than previously realized; they are connected by carbon dioxide, CO2.
Climate change basics: the physics
Climate change concerns have focused on the alteration of weather and climate due to the increase in atmospheric concentrations of greenhouse gases, primarily carbon dioxide, CO2. The impact of CO2 on climate has been understood for at least 120 years. In 1896 the Nobel-Prize-winning Swedish chemist Svante Arhennius published calculations demonstrating that human emission of CO2, when combined with the positive feedback effects of water vapor, would warm the Earth (Arhennius, 1896). His equation, ΔF = α ln(C/C0), relates the change in climate “forcing” (the degree to which temperature change is forced) to the ratio of the concentration of CO2 currently in the atmosphere (C) to a previous concentration (C0). This equation is still in use today. Arhennius estimated that a doubling of CO2 would warm the Earth by about 4 degrees C. This estimate is not far off from current estimates based on much more elaborate calculations.
This warming impact is caused by the physics of CO2, water vapor, and other “greenhouse” gases. Infrared radiation causes the CO2, water, and other greenhouse gas molecules to vibrate, leading to the absorption of the energy carried by that radiation. Much of the solar energy coming from the sun is not in the infrared frequency range, so it passes through the atmosphere without being absorbed. However, when this energy is absorbed by the surfaces of the Earth and its biota, and is re-radiated as infrared radiation, it is then absorbed by greenhouse gases, warming the planet.
The amount of water vapor in the atmosphere is directly related to the atmosphere’s temperature: warm air holds more water vapor. Human activity hasn’t directly changed the concentration of water vapor in the atmosphere significantly. But by burning fossil fuels, humans have dramatically increased the atmospheric concentration of CO2 and in so doing, also indirectly increased the concentration of water vapor. Just as Arhennius predicted over 120 years ago, this increase in CO2 is warming the Earth.
Ramifications of this warming include increased heat episodes, the intensification of the hydrological cycle (greater frequency of both heavy precipitation events and of droughts), sea level rise due to the melting of land-based glaciers and the thermal expansion of ocean water, and, almost certainly, intensification of storms and an increase in extreme weather. These climate-warming- based perturbations have the potential to influence the functioning of Earth’s biota in many deleterious ways, and clearly can be associated with the many facets of habitat disruption.
Climate change amplifications: the chemistry
But there’s another aspect of CO2 that may be more important insofar as habitat disruption is concerned and that has been largely ignored: chemistry. CO2 is a trace gas as far as we humans and other animals are concerned, unnoticed by our bodies in normal life. But to plants it is a vital food. It is taken up by plants as an essential input to photosynthesis. In this chemical reaction, using the energy of sunlight, plants combine CO2 and water vapor to make oxygen and carbohydrates, represented with a generic formula of CH2O, according to the equation CO2 + H2O → CH2O + O2. Without this reaction, life as we know it would not exist.
The atmospheric concentration of CO2 has varied over time; some 50 million years ago it was considerably higher than today. However, for at least the last three million years, the concentration of CO2 has been in the range of 280 ppm. Over these millions of years biota have adapted to this concentration. But within the last 300 years, one ten thousandth of this period – a blink of an eye in the geological or evolutionary time scale – the concentration of CO2 has shot up to 420 ppm, a 50% increase, due to humanity’s burning of fossil fuels and forests.
Imagine what might happen to a person who had been on a tight dietary budget for most of his or her life but suddenly got access to 50% more carbohydrates, but no more protein or minerals? We could expect major changes in the metabolism of that person. This dramatic change is what has, in effect, happened to the whole of life on Earth. Our planet’s primary biota, plants, now suddenly have the opportunity to gorge on CO2. But their access to other important substances such as nitrogen has not changed. Evidence is accumulating that CO2 at its elevated level of 420 ppm is not, as has been proclaimed by some, a healthy influence but is instead throwing Earth’s ecosystem into disarray.
Much of the recent experimental evidence on the impacts of enriched atmospheric CO2 has been assembled by Lewis Ziska and presented in his new book, Greenhouse Planet: How Rising CO2 Changes Plants and Life as We Know It (Ziska, 2022; see full citation at the end of the blog). The findings documented in this book reveal a variety of impacts of elevated CO2. These impacts include stimulation of growth of invasive plants, decreases in the nutrient content of major crops, and changes in plants’ production of insecticidal, allergenic, and other compounds. The changing chemistry of plants may be contributing to a major die-off of insects and insect-eating animals including birds. Below are some details.
Habitat Disruptions: Stimulation of Invasive Plants
The generally accepted explanation for why some plants are invasive is that they have been introduced to new regions where their historic predators and parasites aren’t present. Without these drags on their growth, they have flourished. That some alien plants are not browsed by white-tailed deer contributes significantly to their invasiveness in Eastern North America. Other factors are clearly involved as well, including changes in the temperature regime and the availability of water and other resources such as nitrogen.
But elevated CO2 is also a factor. In recent years, techniques for measuring responses of organisms in situ under elevated CO2 conditions have been developed, making possible investigations of the impacts of CO2 concentrations that could exist in the future under otherwise relatively realistic conditions. What the actual atmospheric CO2 concentration will be in 2050 or 2100 is difficult to predict; it depends on what humanity does to control emissions. Various scenarios suggest that levels could exceed 500 ppm by 2050 and might exceed 1000 ppm by 2100 (Tollefson, 2020).
cheatgrass; photo by Jaepil Cho
One study found that the invasive weed Canada thistle, Cirsium arvense, responds more strongly to elevated CO2 than soybean, a crop that it often plagues. In a high CO2 environment, this weed’s root system grows strongly enough to make it significantly more resistant to herbicides (Ziska, et al., 2004) (Ziska, 2010). The highly invasive and dangerously flammable cheatgrass (Bromus tectorum), nicknamed “grassoline” by the U.S. Forest Service because of its propensity to intensify wildfires, also responds strongly to elevated CO2 (Ziska, et al., 2005). Also found to be boosted by enriched CO2 is yellow star-thistle (Centaurea solstitialis), considered one of California’s worst weeds. In one study (Dukes, et al., 2011) this plant grew 600% larger in elevated CO2 relative to ambient, while native plants responded much less strongly or not at all. Japanese honeysuckle, Lonicera japonica, which plagues many areas in the U.S., was found to increase in biomass by 135% at a CO2 concentration of 675 ppm while a similar native plant, coral honeysuckle (Lonicera sempervirens) increased by only 40% (Sasek & Strain, 1991). In a field study also involving Japanese honeysuckle (Belote, et al., 2004), researchers found that its above ground net production (ANPP) approximately tripled under enriched CO2 while other plants in the trial showed showed lesser increases or actual decreases.
Other plants have been found to be selectively encouraged by enriched CO2 including cherry laurel (Prunus laurocerasus), invasive in the Pacific Northwest U.S. and the U.K., (Hattenschwiler & Korner, 2002); dalmation toadflax (Linaria dalmatica), invasive in much of North America (Blumenthal, et al., 2013); honey mesquite (Prosopis glandulosa) , invasive in Australia and parts of Africa (Polley, et al., 1996); and kudzu (Pueraria lobata), which afflicts the Southeast U.S. (Sasek & Strain, 1988). Three plants invasive in China or Southeast Asia, American rope (Mikania micrantha), Creeping oxeye (Wedelia trilobata), and a morning glory species (Ipomoea cairica), were found to produce 70.3% greater biomass when grown at a CO2 concentration of 700 ppm while three corresponding indigenous plants Paederia scandens, Wedelia chinensis and Ipomoea pescaprae, produced only 30.5% more biomass (Song, et al., 2009).
yellow star thistle; photo by Eugene Zelenko
The list goes on of studies showing increased growth of some plants under enriched CO2 conditions. As more in situ investigations are undertaken, it seems likely it will become clearer that today’s enriched level of CO2 is helping some plants to become invasive.
Complexities and contradictory findings exist however. Not all plants are stimulated by enriched CO2. An important difference in the response to higher levels of CO2 is whether a plant has a C3 or a C4 photosynthetic mechanism. C4 plants contain a biochemical pump that concentrates CO2, making them more adapted to low CO2 conditions (Hager, et al., 2016). At current levels of CO2, such plants’ need for CO2 is easily met. C3 plants do not have this CO2 concentrating ability, and so higher levels boost their growth. In a broad meta-analysis of literature, the average response to elevated CO2 of 365 C3 plant species and 37 C4 plant species was noted; the response was significantly increased in C3 species but was unchanged in C4 species (Robinson, et al. 2012). One striking example of such a difference was observed in the field study noted above (Belote, et al., 2004), wherein researchers found that Japanese honeysuckle (a C3 plant) was significantly encouraged by elevated CO2 relative to other plants at the same locale. The same study found that another aggressive invader, Japanese stiltgrass (Microstegium vimineum), a C4 plant, was unaffected or even slightly inhibited relative to competing plants’ growth by elevated CO2.
One finding is widespread; most plants studied under enriched CO2 regimes show an increase in biomass and evince a higher ratio of carbon to nitrogen (C/N ratio) in their tissues and an overall decline in nitrogen concentrations than when grown under ambient conditions. Since nitrogen is a key component of protein, this change can be expected to lead to lowered protein content of critical food crops. Some impacts of this change are already well underway because of today’s elevated CO2 concentration. Changes since 1850 in the C/N ratio and in the estimated protein content of an important plant product, pollen, were discovered in a striking study by Lewis Ziska and colleagues (Ziska, et al., 2016). Using archived museum samples, these researchers determined the nitrogen content of pollen of Solidago canadensis (Canada goldenrod) going back to the 1850s. They estimated that the protein content of goldenrod pollen, a vital nutrient for North American bees, has declined in inverse proportion to the rise in atmospheric CO2, dropping from a concentration of approximately 18% in the mid-1800s to approximately 12% today. They pointed out that it is possible that bees are now unable to provide sufficient protein and other nutrients to larvae, and that one of the main reasons for bee declines is malnutrition caused by enriched atmospheric CO2. Other studies also indicate that elevated CO2 could cause lower nitrogen concentrations in plants and lead to less proteinaceous plant parts, including pollen, being available to plant-feeding insects (Hall, et al., 2005; Knepp, et al., 2007).
bumblebee on goldenrod; photo by Keila
The changing C/N ratio is almost certainly already affecting the human food supply. As documented in an extensive review of published findings (Soares, et al., 2019), elevated CO2 has a considerable impact on the accumulation of minerals and protein in plants, with many plant species showing declines in both quality and quantity of key nutrients. These changes have worrisome implications for human nutrition and may already be responsible for increasing incidences of dietary deficiency in some areas. Lewis Ziska discusses the likely impact of rising CO2 on the future human food supply in his recent post. A number of studies showing declines in protein and also other nutrients such as zinc in food crops important to humanity are also described in Ziska’s new book, Greenhouse Planet, noted above.
Habitat Disruptions: Other Changes in Plant Chemistry
Other changes in plants besides nutritional content may be driven by enriched CO2. Plants produce a variety of secondary metabolites. Most plants use the C3 mechanism; with 50% more available of a key input, some changes in these plants’ production of such chemicals can be expected. Some changes have been observed. Mohan et al. (2006) report that enriched CO2 in an intact forest system increased water use efficiency, growth, and population biomass of poison ivy (Toxicodendron radicans) and that high-CO2 plants also produced a more toxic form of the allergenic compound urushiol.
Quercus chapmanii; photo by Mary Keim at Seminole State Forest, Florida
Many of the phytochemicals plants produce function as defenses against insect predation, and changes in such production have been found to impact herbivore feeding. For example, Landosky and Karowe (2014) suggest that specialist herbivores may have to contend with more effective chemical defenses by plants under elevated CO2. Hall, et al. (2005), in a study involving several oak and one legume species in a scrub oak ecosystem in Florida (see photo above), found that 700 ppm CO2 levels led to decreased damage to plants by four of six insect groups investigated. They did not see increases in plants’ production of carbon-based secondary metabolites, including total phenolic compounds, condensed tannins, hydrolyzable tannins, cellulose, hemicellulose, and lignin however. They concluded that the primary driver of decreased insect predation under elevated CO2 was lower overall plant nitrogen levels. They stated that the decline of nitrogen levels in foliage under elevated CO2 indicated lower foliar quality and hypothesized that the reductions in insect feeding stemmed from the combined effects of nutrient limitation and increases in parasitism and predation on the nutrient-constrained insects. They further stated that although insects try to compensate for lower nutrient content of leaves by eating more, they did not see an increased portion of damaged leaves in their study. These researchers did not report measurements of terpenoid compounds however, which are reported to represent the largest class of secondary metabolites (Wikipedia, 2022). In another study (Hall, et al., 2005a) found that concentrations of condensed tannins were higher in oak leaf litter under elevated CO2, which suggests that enhanced production of insecticidal compounds or other changes to plant tissues could affect not only insects that consume living plant tissue, but also detritivores.
Robinson et al. (2012) also investigated plants’ production of secondary metabolites in their literature review. Looking at all plant groups, they found that under elevated CO2 the production of nitrogen-based secondary metabolites (e.g., alkaloids, cyanogenic glycosides, and glucosinolates) decreased by 16% while the carbon-based secondary metabolites total phenolics, condensed tannins, and flavonoids increased by 19%, 22%, and 27% respectively. Another carbon-based metabolite, terpenoids, declined by 13%. They further divided plants into grasses, shrubs, herbs/forbs, and trees and found differing responses to elevated CO2. Trees, for example, showed increased production of total glycosides and total phenolics, little change in production of total flavonoids, and a decline in the production of total terpenes. Like Hall et al., (2005), Robinson et al. found a strong and significant decrease in nitrogen concentrations under elevated CO2 for C3 plants. A decrease did not show up for C4 plants.
In addition to chemical defenses, plants have physical characteristics such as surface waxes, trichomes, secretory canals, and tissue toughness-enhancing substances such as lignin. All of these features can reduce the edibility of plants for arthropod herbivores. Robinson et al. (2012) found consistent responses to these characteristics under elevated CO2; leaf toughness and specific leaf weight increased by 11% and 18%, respectively, while specific leaf area did not show a significant change. They concluded that there is an increase in general “toughness” of leaves under elevated CO2. As did Hall et al., (2005), Robinson et al. concluded that elevated CO2 will induce changes in plant chemistry, physiology, and morphology that are likely to impact the nutritional quality of host plants for insect herbivores.
Habitat Disruptions: Changes in Plant Chemistry and Insect Decline
Numerous studies have documented a recent and dramatic decline in insect populations and discussed the probable cascading impacts of such declines through the food chain, affecting spiders, lizards, birds, and other organisms (Samways, et al., 2020; Cardoso, et al., 2020; Sánchez-Bayoa & Wyckhuys, 2019; Tallamy & Shriver, 2021). It has been argued that the main drivers of insect species declines are habitat loss and conversion to intensive agriculture and urbanization; pollution, mainly by synthetic pesticides and fertilizers; biological factors, including pathogens and introduced species; and climate change.
But a puzzling aspect is that some insect declines have been observed in nature preserves (Vogel, 2017) that presumably are not greatly affected by most of the above drivers. One example is a study spanning 27 years that found a 76% decline in flying insect biomass at several of Germany’s protected areas subject to rather low levels of human disturbance (Hallmann et al., 2017). Another study in rainforests of Puerto Rico, apparently not subject to influences such as light pollution, habitat loss, pesticides, or agriculture, reported biomass losses between 98% and 78% for ground-foraging and canopy-dwelling arthropods over a 36-year period, (Lister and Garcia, 2018). This leaves climate change as the likely culprit. But although the varied impacts of climate change, including heat episodes, drought, and other episodes of extreme weather could impact insect populations in remote as well as populated areas, the trends observed appear to far exceed the magnitude of such climate-related disturbances over the last several decades.
tent caterpillars; Shiela Brown, Public Domain Pics
Another puzzling aspect is that not all insect orders or feeding guilds seem to be equally affected. Sanchez-Bayoa & Wychuys (2019) whose article reports on a review of 73 historical reports, state that Lepidoptera, Hymenoptera and dung beetles (Coleoptera) appear to be the taxa most affected in terrestrial environments. Robinson et al. (2012) found that phloem feeders such as Homoptera respond positively to elevated CO2 while foliage feeders/Lepidoptera respond negatively. Lepidoptera were among the most impacted; relative growth rate, fecundity, and abundance all declined under high CO2 conditions, while relative consumption rate, total consumption, and development time all increased.
Most Lepidoptera are herbivorous, feeding in their larval stage, caterpillars, on plants. Caterpillars are key components of the terrestrial ecology; in most forests of the world, caterpillars consume more living leaves than all other animals combined (Janzen, 1988). Insect herbivores such as caterpillars are near the hub of most terrestrial food webs, comprising essential food for insect predators and parasitoids, spiders, amphibians, lizards, rodents, bats, birds, and even higher predators such as foxes and bears (Burghardt et al., 2010). At least 310 North American bird species are known to feed extensively on caterpillars, and the majority of terrestrial birds rely on insects during reproduction and other nutrient-limited periods in the annual cycle (Narango, Tallamy & Marra, 2018). Caterpillars and moths are the largest component of nestling diets in hundreds of species of migrant bird species (Tallamy & Shriver, 2021); they are among the “little things that run the world” (Wilson, 1987).
Carolina chickadee; one of the birds Dr. Tallamy focuses on because it feeds its young on caterpillars; photo by Dan Pancamo; through Wikimedia
Habitat Disruptions and Climate Change: Connected by CO2
The apparently heightened rate of decline of insect herbivores such as caterpillars compared to some other insects, and the findings that many declines have been observed in areas relatively unimpacted by direct human influences such as light pollution, pesticides, and land-use change, point to the likelihood of a broad, perhaps ubiquitous, cause. Climate change is such a broad cause. Even broader and more ubiquitous is the main driver of climate change, CO2. Every plant in the world is constantly bathed in an enriched concentration of this gas. A conclusion seems likely: CO2 is not only causing global warming and climate change but is also affecting life on this planet in ways that, while still poorly understood, are already reducing the nutritive value of food crops, may be a significant cause of the spread of invasive plants, and may be the main driver of insect declines and the cascading impacts of such declines on insect-eating animals such as birds.
What to do?
To mitigate climate change and, as argued here, to mitigate habitat disruption, the steady rise in the atmosphere’s burden of CO2 must be halted, and then steps must be taken to lower the current concentration to a healthier level. These are not hopeless tasks. Although what has been a relentless rise in CO2 emissions at the global level continues, increases have slowed and even stopped in some parts of the world. Accelerating the development of low- and zero-carbon energy sources and encouraging energy conservation, as will be done through the U.S.’s Inflation Reduction Act, will further this progress.
More will be needed. Putting a significant and steadily increasing price on the carbon in fossil fuels is arguably the most important next step. Fossil fuels enjoy a free ride. The byproduct of their combustion, CO2, is dumped with little or no restrictions into the world’s atmosphere. A price on carbon would end this inequity. There are ways this could be done in a revenue-neutral (“fee and rebate”) manner that would avoid harm to economies and those with low- and moderate-incomes. A major step forward in pricing carbon by the European Union, a carbon border adjustment mechanism, is close to implementation. For more on this and other developments in cutting CO2 emissions, see the analyses and insights of the Carbon Tax Center and learn more about actions you can take to influence governments with Citizens’ Climate Lobby.
Not discussed here, but another stark example of habitat disruption is the increasing acidification of the world’s oceans caused by the dissolution of atmospheric CO2 in the waters. The ocean’s average pH has dropped from 8.2 to 8.1 within the last 200 years. Because pH is a logarithmic scale, this represents an increase in hydrogen ion concentration of over 25%, a change that is already threatening some marine creatures. More on this is available from many sources; e.g., Kolbert (2014).
References
Arhennius, Svante, 1896, On the Influence of Carbonic Acid in the Air upon the Temperature on the Ground, Philosophical Magazine and Journal of Science, 41, 237-276.
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.
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.
Hallmann, C.A., Sorg, M., Jongejans, E., Siepel, H., Hofland, N., Schwan, H., Stenmans, W., Müller, A., Sumser, H., Hörren, T., Goulson, D., de Kroon, H., 2017, More than 75 percent decline over 27 years in total flying insect biomass in protected areas. PLoS One 12, e0185809.
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.
Janzen, Daniel H., 1988, Ecological Characterization of a Costa Rican Dry Forest Caterpillar Fauna, Biotropica, 20, 120-135.
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
Landosky, J., and D. Karowe, 2014, Will chemical defenses become more effective against specialist herbivores under elevated CO2? Global Change Biology, 20, 3159–3176.
Lister, B., and A. Garcia, 2018, Climate-driven declines in arthropod abundance restructure a rainforest food web, PNAS 115, E10397–E10406
Mohan, J., L. Ziska, W. Schlesinger, R. Thomas, R. Sicher, K. George, and J. Clark, 2006, Biomass and toxicity responses of poison ivy (Toxicodendron radicans) to elevated atmospheric CO2. PNAS 103, 9086-9089.
Narango, D., D. Tallamy, and P. Marra, 2018, Nonnative plants reduce population growth of an insectivorous bird, PNAS 115: 11549–11554.
Polley, H., H. Johnson, H. Mayeux, C. Tischler, and D. Brown, 1996, Carbon Dioxide Enrichment Improves Growth, Water Relations, and Survival of Droughted Honey Mesquite (Prosopis glandulosa) Seedlings, Tree Physiology, 16, 817-823.
Robinson, E., G. Ryan, and J. Newman, 2012, A meta-analytical review of the effects of elevated CO2 on plant-arthropod interactions highlights the importance of interacting environmental and biological variables, New Phytologist 194, 321-336.
Sánchez-Bayoa, F. and K. Wyckhuys, 2019, Worldwide decline of the entomofauna: A review of its drivers, Biological Conservation 232, 8-27.
Sasek, T. and B. Strain, 1988, Effects of Carbon Dioxide Enrichment on the Growth and Morphology of Kudzu (Puerarialobata), Weed Science 36, 28 – 36, DOI: https://doi.org/10.1017/S0043174500074415
Sasek, T. and B. Strain, 1991, Effects of CO2 Enrichment on the Growth and Morphology of a Native and Introduced Honeysuckle Vine, American Journal of Botany 78, 69-75.
Soares, J., C. Santos, S Carvalho, M Pintado, and M. Vasconceios, 2019, Preserving the nutritional quality of crop plants under a changing climate: importance and strategies. Plant and Soil 443, 1-26. https://doi.org/10.1007/s11104-019-04229-0
Song, L., J. Wu, C. Li, F. Li, S. Peng, and B. Chen, 2009, Different responses of invasive and native species to elevated CO2 concentration, Acta Oecologica 35, 128-135.
Tallamy, D. and W.G. Shriver, 2021, Are declines in insects and insectivorous birds related? Ornithological Applications 123: 1–8, DOI: 10.1093/ornithapp/duaa059
Vogel, G., 2017, Where Have All the Insects Gone? Surveys in German nature preserves point to a dramatic decline in insect biomass. Key members of ecosystems may be slipping away, Science 356, 376-379.
Wilson, Edward O., 1987, The Little Things That Run the World (The Importance and Conservation of Invertebrates), Conservation Biology, 1, 344-346.
Ziska, L., 2010, Elevated Carbon Dioxide Alters Chemical Management of Canada Thistle in No-till Soybean, Field Crops Research 119, 299-303.
Ziska, L., Shaun Falukner, and John Lydon, 2004, Changes in biomass and root: Shoot Ratio of Field-grown Canada Thistle (Cirsium arvense), a Noxious, Invasive Wed, with Elevated CO2: Implications for Control with Glyphosate, Weed Science 52, 584-588.
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
Ziska, Lewis, 2022, Greenhouse Planet: How Rising CO2 Changes Plants and Life as We Know It, Columbia University Press, NY.
Guest blog by Lewis Ziska, Associate Professor, Environmental Health Sciences at the Columbia University
[Dr. Ziska has spent his career analyzing the impacts of CO2 and climate change on plants – and therefore on people. He served as Project Leader for global climate change at the International Rice Research Institute; then spent 24 years at the USDA’s Agricultural Research Service, where he worked primarily on documenting the impact of climate change and rising carbon dioxide levels on: Crop selection improves production; Climate and agronomic pests, including chemical management; Climate, plant biology and public health impacts on food security with a focus on nutrition and pesticide use.]
No question you’ve heard the term, “Climate Change” or “Global Warming”, or my personal favorite, “Global Weirding”. The consequences are talked and discussed in the media—as they should be—but often the media, like many Americans, is focus challenged. Or in more polite terms, they have the attention span of a hummingbird on crack. Which is to say, that simple physical consequences, like sea level rise (heat melts ice!), and stranded animals on ice (Poor polar bear!), or intense storms (newscaster whipped about in the rain, yelling to be understood) are repeated, over and over again. Understandable, makes for good TV.
But it also makes you feel separate from what is happening, these consequences of climate change are to the “other”. I don’t live near the ocean, I don’t interact with polar bears; sure we have storms, but I live in the Midwest, in one of those states that begins with a vowel. Shoot, I commute to work, try and make ends meet, I’m not some damn tree hugger. Why should I care?
To understand why, you need a bit more background, some science that isn’t always available on TV or social media when it comes to global weirding.
First, while you may not be a tree hugger, you do, in fact, interact with nature. Several times. Every day. We call those times, “breakfast”, “lunch” and “dinner”.
dinner; photo by davylin via Flickr
You depend on nature for food. And clothing. And paper. And medicine. And oxygen. And construction materials (wood), and many, many other things. So, if nature gets hinky, and the climate becomes uncertain, it might be worth your while to think about climate change, or global weirding, in a different light. What I want to do here then, is to illuminate two examples that I hope will help you see why climate could affect you, directly and significantly.
Let’s begin with plants. Those green living things that comprise the bulk of the natural world (literally, if you were to weigh the natural world, 97% would be plants, 3% animals). Then let’s look at them through two different lenses—how will climate weirding alter your food; shoot, how will it alter the air that I breathe?
Let’s start with a basic food, rice. Obviously you don’t want to mess with its production, or its nutritional quality. But that is exactly what global weirding is doing.
Rice has flowers. Not big showy ones, but flowers none the less—ones that get fertilized with pollen, and seed is produced. The seed that feeds some two billion people– or about a quarter of the earth’s population.
Like all living things, plants are heat sensitive, and for rice, and many crop plants, the degree of sensitivity varies, depending on the part of the plant in question. Take a look at the table. The crops that are listed, including rice, are the core of what the world eats. Now notice the difference in temperature sensitivity. Vegetative parts of the plant, leaves and stems, are reasonably tolerant of higher temperatures, but flowers are not. Pollen, the plant equivalent of animal sperm, is highly temperature sensitive, and if the temperatures get into the high 90s (37-38oC), they become deformed, and the rice plant doesn’t produce seeds. Same for a number of plants, ones necessary to feed 8 billion people.
Crop
Opt. Temp. Vegetative
Opt. Temp. Flowering
Failure Temp.Flowering
Maize
28-35oC
18-22oC
35oC
Soybean
25-37oC
22-24oC
39oC
Wheat
20-30oC
15oC
34oC
Rice
28-35oC
23-26oC
36oC
Sorghum
26-34oC
25oC
35oC
Cotton
34oC
25-26oC
35oC
Peanut
31-35oC
20-26oC
39oC
Data are adapted from Hatfield et al., 2011.
Doubtful you’ve seen this climate threat to the global food supply on TV or a streaming service. I caught a glimpse once of temperature and agriculture on a CNN newscast, but with the “expert” calmly stating that we would just have to grow our corn in Canada, ha-ha. (Somehow, at least for rice, it’s hard to imagine India, one of the world’s largest rice producers, moving its rice production northward to the Himalaya’s, but I digress.)
Food is fundamental. If production, especially that of a global staple like rice, is impacted by rising temperatures there will be consequences. Rising prices, reduced availability, and wide-spread hunger.
But there is more to consider. Given the global dependence on rice, any change in its nutritional quality will also have effects, especially on poorer countries that rely heavily on rice as a major food source. And here we need to delve a little deeper into another aspect of climate weirding that doesn’t make it to the popular media—that rising carbon dioxide (CO2), the primary greenhouse gas, can also directly influence plant nutrition. The reasons are complicated, but in simple terms all living things consist of elements, carbon, nitrogen, phosphorous, sulfur, copper, etc., etc. A plant gets it’s carbon from the air (CO2), but everything else (nitrogen, potassium) from the soil.
And there is an imbalance. In the last 50 years, atmospheric CO2 has increased by about 30%, and is projected to increase another 50% by the end of the century. With more CO2, plants are becoming carbon rich, but nutrient poor. Nutrient poor, because while CO2 has increased in the air, nutrients in the soil have not kept pace. A perverse carb loading at the plant level.
As a consequence, rice, and many other plants, are shifting their chemistry. For example, there is a general decline in protein, in part because protein requires nitrogen. There are similar ubiquitous declines in iron and zinc, important micro-nutrients needed for human development.
Such nutritional degradation is of obvious global importance, and does, on occasion, show up on basic media when warming / weirding is mentioned, but you’d be hard pressed to find it.
Let’s move our light to another hidden bit of science. How plants can influence the air we breathe.
As humans, we like to trade things. And a large percentage of what we trade are living organisms, from fish to trees. But what began as local, regionalized trading has grown with the global population and the needs of that population—a population of 1.6 billion at the beginning of the 20th century is now ~8 billion at the beginning of the 21st. And we haven’t stopped trading. Biological trade is not inherently bad, but it represents a historically unprecedented global movement of DNA across continents, across countries, regions, towns, cities and ecosystems. And some of the DNA, when introduced, can do great harm to the environment, the economy and to human health. That harm has a name, “Invasive Species”.
Let us focus on one such plant species introduced to Eastern Europe, one that almost every American has personal experience (ACHOO!) come fall. The species is common ragweed. An invasive plant whose introduction and spread in Eastern Europe—introduced accidently through imported seeds or contaminated hay – has resulted in enormous environmental and economic losses in agriculture and public health in recent decades. In Hungary, the most important ambient biological air pollutant is: ragweed.
collecting ragweed pollen under different climates (Author’s photo)
The photo is from studies that I led looking at how ragweed pollen would respond to temperature and carbon dioxide. (If you’re curious, ragweed likes both.) Warmer temperatures, earlier Springs, later Autumns can extend its pollen season; not only extend, but increase the amount of pollen being generated. There is even some data suggesting that rising CO2 can alter pollen chemistry, making it more allergenic (REFS). Sadly, ragweed pollen doesn’t appear as temperature sensitive as that of rice, or other agricultural plants.
I wish I could say that ragweed was the exception among allergenic plants, but it’s the rule. Parthenium weed is a highly invasive species that has spread to more than 40 countries around the world. Like ragweed its pollen are highly allergenic, but it can also produce severe rashes, like poison ivy, and is known to be poisonous to livestock. It is highly aggressive, and arriving in a new location (where it has no natural enemies) can dominate landscapes, reducing biodiversity. And as with ragweed, high temperatures, longer growing seasons, heatwaves and droughts are expanding its range, and for that matter, make controlling its spread more difficult.
Such responses among invasive species will have direct impacts on air quality, especially among those (myself included) who suffer from seasonal allergies. Gasping for air is never fun.
Estimates are that pollen and seasonal asthma affects more than 24 million of us, including 6 million kids. And yet, when watching news reports of climate change, how many times have you seen a report on pollen and air quality? On increasing allergies or asthma? Once? Twice?
I could go on, (and if you need more examples, read “Greenhouse Planet”, my latest book). But my point is this: Not all of the consequences of rising carbon dioxide and climate change, warming, weirding, whatever, make for “good” TV. There is so much more to explore. So, do yourself a favor. Take a deeper dive, find out what is happening behind the scenes.
Because if we are going to rise to the challenge, we need to know what we are fighting against. Right now, the media is exemplary on showing some things, but silent on much else of importance. Watching news coverage of climate change is a bystander watching a cataclysm, and thinking, “Boy, glad I’m not experiencing THAT!”. Yet in the simplest and most basic of terms, you are, or will be, affected– from food choices to nutrition, even your allergies. And so much more.
It isn’t just about polar bears. It’s about you. Read, Understand, Act.
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.
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.
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
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:
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.
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:
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
We welcome comments that supplement or correct factual information, suggest new approaches, or promote thoughtful consideration. We post comments that disagree with us — but not those we judge to be not civil or inflammatory.
For a detailed discussion of the policies and practices that have allowed these pests to enter and spread – and that do not promote effective restoration strategies – review the Fading Forests report at http://treeimprovement.utk.edu/FadingForests.htm
In 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 2nd 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.
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)
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
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 waspQuadrastichus 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
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
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 representativewhether 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.
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
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
