Prostanthera cuneata – member of a genus endemic to Australia. Photo by Leonora (Ellie) Enking via Flickr
In 2023 a global meeting of plant conservation experts convened by The Royal Botanic Gardens, Kew (U.K.) released the 5th edition of a report on the State of the World’s Plants and Fungi.
Associate Professor of Plant Ecology and Conservation Science Rachael Gallagher from Western Sydney University had led the global evaluation of conservation assessments for unique flora species. She is also the lead author of an article (2023; full citation at the end of this blog) evaluating how well countries around the world met their treaty obligation to assess the conservation status of endemic plant species native to their territories. The analysis identified 221,399 endemic plant species in a total of 173 countries. The treasure is not distributed evenly. Five countries harbor a third of the endemic plant species: in descending order, Brazil, Australia, China, Mexico, and South Africa. (The United States, including its islands, ranks 8th.)
On average, countries completed assessments of just 34% of their endemic species. New Zealand and here and South Africa shone: they assessed 87% of their unique species. China assessed 71%. One of the world’s poorest countries, Madagascar, evaluated 42% of its ~10,000 endemic plant species. Reminder: tiny Madagascar ranks 6th in the number of endemic plants. Australia – one of the richest countries– carried out the process for 39% — slightly more than the global average. Other countries that are stewards of numerous endemic plants were below the average: Brazil reviewed 29%, Mexico assessed only 24%.
Rachael Gallagher and her colleagues in the Australian Biodiversity Council were quite critical of Australia’s low level of performance. They called on their countrymen to do much more to prevent the decline and extinction of the country’s unique plant species. Australia, as party to the Convention on the Conservation of Biological Diversity, has a treaty obligation to prevent extinction of species which occur nowhere else. Remember, Australia’s flora and fauna rank extremely high on a scale of phylogenetic distinctness as an heir of the isolated continent of Gondwanaland.
Gallagher and colleagues concede that many endemic plant taxa in Australia have huge ranges — averaging 235,829 km2. But these vast expanses do not prevent sudden population crashes caused by calamities. They mention the megafires of 2019–2020 and – over the longer term – climate change. I think of the invasion by the rust fungus Austropuccinia psidii.
When we think about Australia, we wonder at the kangaroos and koalas. I assume Australians consider their unusual fauna to be iconic symbols of their country. Why are they not equally committed to their flora – 88% of their plant species are endemic. Do they suffer from the same “plant blindness” I have encountered in the United States? South Africa undertook an assessment of her endemic flora that concluded that a quarter of these species are threatened. Sixty percent of the country’s 20,000 plant species are endemic.
a protea in South Africa’s fynbos; photo by Michael Wingfield
[I have found no parallel analysis of America’s endemic plant species. Our nation’s rank of 8th in number of endemic species is explained by the highly unique floras of the islands, especially the Hawaiian archipelago. More than 95% of native species on the Islands are endemic. This includes 67% of the large trees still present in the forests (Potter et al. 2023).]
Sources
Gallagher, R.V., S. P. Allen, R. Govaerts, M.C. Rivers, A.P. Allen, D.A. Keith, C. Merow, B. Maitner, N. Butt, T.D. Auld, B.J. Enquist, W.L. Eiserhardt, I.J. Wright, J.C.O. Mifsud, S. Espinosa-Ruiz, H. Possingham, V.M. Adams. 2023. Global shortfalls in threat assessments for endemic flora by country. Plants, People, Planet. DOI: 10.1002/ppp3.1036
Potter, K.M., C. Giardina, R.F. Hughes, S. Cordell, O. Kuegler, A. Koch, E. Yuen. 2023. How invaded are Hawaiian forests? Non-native understory tree dominance signals potential canopy replacement. Landsc Ecol 2023 https://doi.org/10.1007/s10980-023-01662-6
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
an aye-aye – one of the highly endangered lemurs dependent on moist tropical forests of Madagascar; photo by Andrew Ciscel via Wikimedia
A forthcoming study examines two important issues: interactions of pathogens’ spread and changing climate, and invasive species threats to tropical islands’ forests.
Underwood et al. (in press) analyzed how an introduced vascular wilt pathogen — Leptographium calophylli – is likely to affect a tree endemic to Madagascar’s already threatened mid-level elevation humid & subhumid forests, Calophyllum paniculatum (sorry; I can find no photographs of the tree species).
Climate change is expected to cause substantial shifts in temperature and precipitation patterns on the island. These temperature and moisture regimes in turn govern pathogen sporulation, infection efficiency, and survival. They also affect the host’s levels of stress and defenses. The direction of change is not certain, however. In some cases, warming and other changes to the climate might facilitate a pathogen’s spread, allowing it to track shifts in the host’s range and expand into previously unoccupied refugia. In other cases, these changes might erect environmental thresholds that limit the pathogen’s survival and spread, thereby creating spatial refugia for the host.
diademed lemur, courtesy of Animalia
Environmental change increases the area of suitable landscape, that is, it weakens climatic barriers to establishment. Continued anthropogenic movement of some vector (biological or not) generates multiple introductory events over time. As a result, the likelihood of a successful establishment also increases, even if the probability per individual introduction is unchanged. Underwood et al. say that invasion outcomes thus become increasingly dependent on propagule pressure.
On many other tropical islands the threat from climate change is exacerbated by deforestation. On Madagascar, clearing driven by slash-and-burn agriculture and fuelwood harvesting has already reduced natural forest cover to less than 10% of its original extent. [For more on this topic, see e.g., Mittermeier et al. (2011).] Underwood et al. cite a determination by the ForestAtRisk model that humid forest in Madagascar could be almost entirely lost by 2100.
Loss of Madagascar’s forest has global implications. The island is one of 36 global biodiversity hotspots for both flora and fauna (e.g., lemurs). Its flora exceeds 12,000 plant species, of which 83% are endemic. In this case, the host tree species — Calophyllum paniculatum — is already considered vulnerable by the International Union for the Conservation of Nature (IUCN). Thus it is of global importance to understand the relative importance of several threats so that conservations can adopt the most effective countermeasures.
While they do not say so explicitly, it appears that Underwood et al. worry that too few of the conservationists active on Madagascar are paying attention to the possible impact of introduced pathogens. They note that pathogen-driven mortality of dominant or functionally unique trees can rapidly alter community structure and ecosystem function, potentially triggering local extinctions and cascading ecological consequences. For example, if an infection removes mature trees, their loss reduces fruit and nectar availability and so depresses populations of dependent wildlife. The trees’ death also diminishes above-ground carbon stocks and litter inputs. In combination, these impacts can shift community composition toward disturbance-tolerant states and heighten susceptibility at forest margins. These changes difficult to reverse once thresholds crossed.
red-bellied lemur in Ranomafana National Park – site of the first detection of Leptographium calphylli; via Flickr
This threat is not hypothetical. Since 2016 mature C. paniculatum at one site – a National Park – have been dying from a vascular wilt disease caused by a species in the Leptographium genus, probably Leptographium (formerly Verticillium) calophylli. While the species hasnot yet officially been recorded in Madagascar, it is established on neighboring Indian Ocean islands and across much of mainland Africa. Various species in the fungal genus are known to cause disease in other woody hosts. Underwood et al. suggest it was probably transported to Madagascar on infected wood, although they present no data.
Inside forests, Leptographium spp. are vectored by bark beetles in the Cryphalus genus. At least 25 Cryphalus species occur on the African Continent; some are vectoring disease on Seychelles and Mauritius.
The analysis by Underwood et al. indicates that future climatic conditions are likely to worsen the Leptographium calophylli infection over coming decades. The causal agent is likely to retain two-thirds of its current probable distribution and expand into previously uninhabited regions. The suitable habitat is expected to stretch across the entire north-south humid belt – the entire distribution of the host tree. Underwood et al. (in press) say it is even possible that the pathogen might remain in the forest, subsisting on other hosts, after C. paniculatum becomes functionally extinct across its range.
Meanwhile, that host – Calophyllum paniculatum – is projected to experience severe range shifts, with an overall net contraction across all climate change scenarios. It is forecast up to 67% of its current area by 2100. This range contraction will be compounded by fragmentation and dispersal limitation resulting from from deforestation. The refugia will be few and geographically isolated by late in the 21st century.
red-veined swallowtail; photographed in Ranomafana National Park by Frank Vassen, via Wikimedia
Are conservationists considering the implications of Leptographium calophylli’s probable persistence? Underwood et al. imply they are not; they say the impact of this and related pathogens on Madagascar & nearby islands is “still an unknown to the conservation community”. They urge their colleagues to conduct a set of research actions to identify, monitor, & limit the fungus’ spread – – and thereby improve the effectiveness of conservation efforts.
Host range & other targets: determine whether L. calophylli infects other taxa in Madagascar – especially the endemic species and genera. They suggest systematic field sampling of multiple species across sites within the core probable range of L. calophylli. A trained pathologists should be consulted to officially identify the pathogen.
Determine the spread phase of the pathogen. They suggest random sampling of species & sites within & outside of the fungus’ probable distribution, mapping the possible start point & dispersal patterns, including both anthropogenic & natural spread routes.
Assess applicability of IPBES tools & suggestions for invasive species management to the case of a fatal pathogen in the context of tropical islands’ characteristics. How might Madagascar implement prevention, early detection & rapid response systems?
I applaud Underwood et al. for trying to alert the conservation community active on tropical islands to the simultaneous impacts of multiple global & regional change drivers on vulnerable species. Probably other host-pathogen systems are experiencing the same diverging trajectories that might intensify their biodiversity loss, particularly when compounded by deforestation.
SOURCES
Mittermeier, R.A., E.E. Louis Jr., M. Richardson, C. Schwitzer, O. Langrand, A.B. Rylands. 2010. Lemurs of Madagascar. Conservation International, Arlington, USA. ISBN 9781934151235
Underwood, E.L., K.A Brown, A. Ronnfeldt, M. Mulligan, N. Walford, R. Allgayer. In press. Climate change facilitates fungal pathogen expansion while driving endemic host range contractions in a tropical biodiversity hotspot. Research Square.
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
sorting coffee beans; photo by Niels Van Iperen via Wikimedia
Many have recognized that preventing introduction of invasive species is the most efficient approach to minimizing their ecological and economic impacts. Prevention requires many capacities, including control over a country’s borders, strong border biosecurity agencies and policies, and foreknowledge of probable pathways of introduction and high-impact species that might arrive.
Horizon scanning is one tool for gathering information about non-native species likely to enter, how they might arrive, and their probable impact. Horizon scanning involves a systematic search for potential invaders, assessment of their potential to harm BD, economic activities and human health, and opportunities for impact mitigation. It thus supports choice of prevention policies, targetting of efforts, and implementation of early identification and eradication procedures (Kenis et al. 2022; Martinou et al. 2026)
I have reviewed two case studies of the application of horizon scans.
Plant Pests in Ghana
One horizon scanning exercise aimed to identify and rank potential invasive non-native plant pest species that could be harmful to agriculture, forestry, and the environment in Ghana. The ultimate objective was to enable prioritization of actions aimed at preventing their introduction. As the participants in this exercise note (Kenis et al. 2022), the resource-poor farmers of Sub-Saharan Africa are particularly vulnerable to invasive pests that attack their crops, both those grown for subsistence e.g., maize and sorghum, and those grown for the international market, e.g., cacao and tomatoes. The continent’s vulnerability is increased by porous borders, weak cross border biosecurity, and inadequate capacity to limit or stop invasions. This exposes Africa both to repeated invasions and to continued spread across the continent once they have arrived.
Marc Kenis and 21 others assessed 110 arthropod and 64 pathogenic species using a simplified pest risk assessment. This set had been winnowed from an initial list of 1486 arthropods, nematodes and pathogens. Unfortunately, assessors were unable to agree on confidence levels for the assessments.
Sixteen of the assessed species – 14 arthropods and two pathogens – were thought at the time to not be on the African continent. Another 19 arthropod and 46 pathogenic species had been reported established in the neighboring countries of Burkina Faso, Côte d’Ivoire, and Togo. Seventy-seven species [62 of them pathogens] were recognized as established elsewhere in Africa.
Ninety-five percent of the arthropods were considered likely to arrive as contaminants on commodities, i.e. on their host plants; 23% were also likely to arrive as stowaways; some good fliers already present in neighboring countries could also enter unaided.
The 64 pathogen species included 14 bacteria, 16 fungi, 14 nematode, seven water moulds (Kingdom: Chromista), and 13 viruses. Sixty-two of these species have been detected on the African continent; 46 are reported in neighboring countries. Thirty-one (48.4%) of the pathogenic organisms were considered likely to arrive both as contaminants on commodities and/or as stowaways; Twenty-six (40.6%) probably arrive only as contaminants; five could arrive exclusively as stowaways. Kenis et al. (2022) specify which of the fungi, nematodes, viruses, bacteria, and water moulds fall into which category.
The most important input in the threat scoring process was likelihood of entry. The unsurprising result was that species known to be in neighboring countries or spreading rapidly in Africa received the highest overall scores. The likelihood of establishment was less important because the assessors had already excluded species they thought would encounter an unsuitable climate or absence of host plants. The impact score played an important role in the overall score; it was based primarily through their potential economic impact. There is little information about or attention to the potential threat of non-native plant pest species to non-commercial plants. Kenis et al. (2022) cite well-known examples to remind us that invasive plant pest species have had “huge impacts” on native tree species and biodiversity in North America and Europe. On the African continent, most non-native pests attack mostly concern exotic trees. They note one exception, Euwallacea fornicatus, DMF a wood-boring beetle from Asia killing many native trees in South Africa.
Bemisia tabaci; one of the arthropod pests in a country bordering Ghana; photo courtesy of INCTELUNI
Kenis et al. (2022) state that some of the several alien arthropods and pathogens identified in neighboring countries might already be present in Ghana although not yet recorded or identified to the species level. They say it is essential to clarify these species’ status by enhanced surveillance and applying morphological and molecular methods. Some of these possibly introduced species received high scores in the assessment. They threaten cocoa, a key crop in Ghana, and vegetable crops.
I am disappointed that Kenis et al. (2022)’s main actions suggested for both arthropod and pathogenic species that scored highly are to ramp up surveys and to conduct full pest risk analyses. It is true, as thy point out, that such assessments are required by international regulations before a country may implement phytosanitary measures. [See discussion of the requirements of the International Plant Protection Convention here.]
To some extent, the horizon scan echoed the obvious: most of species ranked high are already on the African continent, including 19 arthropod and 46 pathogenic species known to be established in neighboring countries. Plus, the recommended actions are minimal. Since Kenis et al. (2022) is essentially the scan itself, it provides no information on whether Ghana has implemented the recommendations. Still, given what I assume is lagging preparation across most of Africa, the horizon scan might be useful in encouraging countries to set priorities and take some action.
Cyprus
The second case study of applying horizon scanning is more encouraging. Scientists on Cyprus tried to assess the efficacy of their own horizon scanning exercise. I applaud their decision to do so. The horizon scan itself might have been undertaken on their own initiative? Or it might have been taken on in response to European Union regulations, which oblige Member States to enact measures to prevent or manage introduction and spread of invasive species designated as of Union Concern. The Union also encourages development of national invasive species lists and provides a legal basis for emergency measures in response to a detection.
Scientists carried out two horizon scan workshops in 2017 and 2019. The two workshops evaluated 225 and 352 species, respectively, to predict which are most likely to arrive and the level of provable impact to Cyprus’ biodiversity, human health, and economy. In 2023, four to six years after the workshops, scientists evaluated the listed species to reveal the accuracy of the predictions and actions taken so far (Martinou et al. 2026).
During the period 2017 – 2023 there were 183 Martinou et al. (2026) found publications naming 183 non-native species not previously officially detected in Cyprus. (As I will discuss later, a significant number of these species had been present on the island in 2017 but knowledge of their presence did not reach the assessors.) Of the 183 newly reported species, 31 had been included on some list of invasive species (e.g., EPPO or European Union list of species “of Concern”) or predicted by the horizon scanning exercises to rank amongst the top 100 riskiest species.
Cyprus’ horizon scans highlighted the risk posed by 10 of these 26 species. Martinou et al. (2026) focused on seven of them as having been ranked as high risk to the nation’s BD, human-health or economy. They added an eighth species, a venomous marine fish.
A further 10 species that were detected in the country had received lower impact scores, so they had not been included on the high priority lists of the horizon scans.
One of the species allotted a lower impact score, Spodoptera frugiperda, is under eradication, although it is widely distributed on the island. This action might be in response to the species’ inclusion on the EPPO A2 list.
As I noted above, scientists learned that 17 of the species had been present in Cyprus before the scanning exercises were undertaken but since their presence was then unknown to the participants, they were assessed as if still had not been introduced. This points to the country’s non-native species checklists not being fully up to date at the time.
Nine plant species common in the plant trade were most certainly present on Cyprus before the horizon scans (2017), but there were no published reports of their escape from cultivation. Nevertheless, they might have already been present in the wild. It is also possible that at least some escaped since the scans. Always tricky; always depends on who looking where.
Actions upon detection of specific taxa Detection of the common myna (Acridotheres tristis) – a species widely recognized as invasive – occurred in January 2022, close to a port. Eradication measures were implemented by the wildlife agency. Martinou et al. (2026) believe the introduction was facilitated by shipping. They think there is an extremely high risk of repeated introductions of mynas.
Aedes aegypti; photo by James Gathany via Flickr
Two mosquitoes were detected in 2022. A pilot project to eradicate The yellow fever mosquito, Aedes aegypti, was begun in 2023. There is no information about its success. The Asian tiger mosquito, Aedes albopictus, has been documented by citizen scientists as spreading rapidly in the suburbs of Limassol and Nicosia. To date the proposed interventions have been unsuccessful, possibly due to focusing on public land while the mosquitoes can also breed on private properties. Detection of the little fire ant Wasmannia auropunctata (in 2022) was not surprising since it had already invaded other regions of the Mediterranean. Martinou et al. (2026) believe the introduction was probably facilitated by the plant trade. The scientists note that ant management and eradication efforts are both challenging and costly, but do not report whether any has been initiated. Detection of several marine invasive species was reported, some by citizens, e.g., divers or fishermen. Among the 17 species determined to have been present on the island since before 2017 were some fairly conspicuous vertebrates: brown rat (Rattus norvegicus), raccoon Procyon lotor, two tortoise species, house crow (Corvus splendens) ruddy duck (Oxyura jamaicensis). Also two more ant species, Solenopsis geminata and Trichomyrmex destructor. There were also several non-native plant species, including the notorious seaweed Caulerpa taxifolia.
Value of the Horizon Scan I am surprised that Martinou et al. (2026) do not explore why so many detections were published in 2022 since they assert that horizon scanning helped raise awareness amongst the authorities, scientists and the public. They do note that this awareness led, in some cases, to a rapid response by the competent authorities. Martinou et al. (2026) assert further that the exercise facilitated communication between invasive species experts, policy makers and society, encouraged active engagement and raised awareness regarding the importance of early warning, rapid response, and management of IAS. They therefore propose that the horizon scanning process for the island of Cyprus be repeated regularly – every five to 10 years – since new introductions continue. These efforts should include development pathway management plans and contingency planning that would be shared with local authorities and stakeholders.
Martinou et al. (2026) note two detections that have not, apparently, resulted in establishment. A dead specimen of brown marmorated stink bug (Halyomorpha halys) was reported in luggage in May 2022, the result of ‘Bug Alert Cyprus’ awareness campaign. The Colorado potato beetle (Leptinotarsa decemlineata) was detected in 2010 by Department of Agriculture inspectors in a consignment of potatoes. The agency ordered immediate destruction. Imports of potatoes are subject to special phytosanitary requirements for protected zones. It is not clear that this measure was implemented by Cyprus or is a European Union decree.
brown marmorated stinkbug; courtesy of Oregon Department of Agriculture
Martinou et al. (2026) are worried that no introductions have been reported at border crossings across the ‘Green Line’ [the United Nations-controlled buffer zone between Greek and Turkish portions of the island]. They call for enhanced cross-community collaboration and improved information and data sharing for border control staff and customs officers about invasive species. They suggest that border order inspections and pathway monitoring could be supported by local experts offering identification services for a variety of taxa. They suggest that the horticultural industry is a major pathway for the introduction of plants and insects such as ants.
Martinou et al. (2026) also advocate efforts to improve communication among the various institutions and authorities that discover bioinvasions and are responsible for taking action. While researchers + experts from government departments involved in the horizon scans are informed, the findings of the horizon scanning needs to be provided to e.g., customs officers, fishers, ship crews, pet shop owners, and school teachers. Much of this information might be exchanged through informal networks and through a growing body of web-based databases and other resources.
Early detection and rapid response depends increasingly on efforts by citizen scientists to report observations of IAS of concern. Martinou et al. (2026) note that six of the invasive species identified in the horizon scanning exercise were reported by citizen scientists. They express the hope that artificial intelligence and deep learning models could help identify species from photographs collected by citizen scientists on platforms such as iNaturalist. Such platforms also facilitate rapid dissemination of information to decision-makers who can take appropriate action. Martinou et al. (2026) also hope eDNA can help detect cryptic bionvaders, including freshwater or marine taxa.
As I blogged earlier, Mark Hoddle had endorsed several components of prevention programs: * Early research to identify natural enemy species that might “self-introduce” along with the invading host. * Collaborating with non-U.S. scientists to identify and mitigate invasion bridgeheads. * Sentinel plantings. These plantings can also support research on natural enemies of key pests. [A year ago, Eliana Torres Bedoya of Ohio State alerted participants in the annual USDA research forum on invasive species that fungi, including potential pathogens, were isolated from asymptomatic plants; Detection of the full range of fungal pathogens requires that samples must be collected throughout the growing season; microbes present differ. Need to expand surveillance beyond symptomatic plants – at both sentinel gardens and plant health border inspection stations. *Integrating online platforms, networks, professional meetings, and incursion monitoring programs into “horizon scans” for potential invasive species. He mentions specifically PestLens, (https://pestlens.info/); online community science platforms, e.g., iNaturalist; international symposia; and official pest surveillance, e.g., U.S. Forest Service’s bark beetles survey and surveys done by the California Department of Food and Agriculture and border protection stations.
That blog also cites Weber et al.’s support for sentinel plant nurseries because accidental plant and herbivore invasions often occur at the same points of entry.
At the 2026 meeting of the annual USDA Research Forum on Invasive Species, Ashley Schulz (Mississippi State) reported findings of study analyzing establishment of insects imported deliberately as biocontrol agents as clues to bioinvasion. She found that generalist phytophagous insects might be more likely to find a suitable host and survive after introduction. The “goldilocks” standard applies: the host must be sufficiently closely related to the insect’s native host to be recognizable but sufficiently distant so that it lacks defenses. Considering impact, phytophagous insects that feed on structures not easily restored – e.g., main stem or root, cause more damage than those that feed on easily replaced leaves. Entomopagous insect, on the other hand, must be able to find hosts that can hide or defend themselves. This means that highly specialized insects might be more likely to establish.
Kenis et al. 2022. Horizon scanning for prioritizing invasive alien species with potential to threaten agriculture and biodiversity in Ghana. Neobiota 71: 129-148 (2022) doi: 10.3897
Martinou, A.F., J. Demetirou, I. Angelidou, N. Kassinis, A. Melifronidou, J.M. Peyton, H.E. Roy, A.N.G. Kirschel. 2026. Multiple introductiions of invasive alien species on a Mediterranean Island predicted by horizon scanning. Biological Invasions (2026) 28:41 https://doi.org/10.1007/s10530-025-03729-8
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 Or https://fadingforests.org/
pines in a plantation in Argentina killed by Sirex noctilio; photo by J. Villacide
A decade ago, Payn et al. (2015) compiled studies from around the globe to evaluate threats to widespread tree plantations. At that time, they said climate change posed the greatest threat to plantation forestry globally, in the forms of storm and flood damage and simultaneous warming and drying trends with extreme temperatures.
Still, the authors warned that forest health would be an increasingly important constraint to plantation productivity. They were optimistic, however, that modern breeding and other technologies could offset losses.
What is the current situation? The countries that depend on these plantations for fiber production are not demanding that leaders of the international phytosanitary structure build a more effective system to protect their investments. Instead, individual scientists struggle to better understand threats. Mostly, they propose expanded research.
Economic Importance of these Species
Eucalypts
“Eucalypts” comprises three genera in the family Myrtaceae: Angophora, Corymbia and Eucalyptus. These include more than 700 tree species native primarily to Australia. A few species are native to Indonesia, New Guinea and the Philippines (Paine et al. 2011; Crous et al. 2019). Some of these species have been extensively planted outside their native ranges for more than 100 years. These plantations have expanded rapidly in recent decades, especially in Southeast Asia and the Southern Hemisphere (Crous et al. 2019). Eucalypts are now the most widely planted hardwood timber in the world (Paine et al. 2011).
Eucalypt plantation in Brazil; photo by Jonathan Wilken via Wikimedia
Eucalypts’ popularity has been driven chiefly by their rapid growth; short rotation times including through coppicing; and adaptability to a very wide variety of sites and climatic conditions (Paine et al. 2011; Crous et al. 2019). Also, these trees are an important source of the short-fiber pulp required for production of high-quality paper used in modern office copiers and printers (Paine et al. 2011). Plantations are increasing even in Australia, where harvesting of native forests is increasingly being restricted (Paine et al. 2011).
Pines
Pines – a genus restricted naturally to the Northern Hemisphere – is second in global popularity. South America hosts 4.6 million hectares of pine plantations (Lantschner and Villacide 2025). South America is more dependent on forestry plantations for wood production than any other region. In 2012, 88% of its industrial roundwood was produced by non-native plantations. This far exceeded the global proportion of approximately 19%.
These intensively managed plantations have enabled Brazil and Chile to become “planted forest powerhouses.” Uruguay and, more slowly, Argentina are following the same path (Payn et al. 2015).
Documentation of the Damage
Euclaypts
The highly diverse eucalypts host an even greater diversity of fungi. As of 30 years ago, scientists were aware of more than 500 species of just one type, the leaf-infecting fungi. Additional fungi are associated with seeds, capsules, twigs, branches, and stems. Little is known about the vast majority of these fungi. Even species considered causal agents of important diseases have not yet been confirmed using Koch’s Postulates. Areas of origin for most is also unknown (Crous et al. 2019).
Crous et al. (2019) compiled information on 110 genera of fungi found on eucalypt foliage. Some genera include well-recognized primary pathogens. They name Austropuccinia and Calonectria, Coniella, Elsinoe, Pseudocercospora, Quambalaria and Teratosphaeria. Other genera are thought to include species that are opportunists that develop on stressed or dying tissues. Many other leaf fungi are putative pathogens, but unstudied. Additional fungi cause vascular wilts (e.g. Ceratocystidaceae), stem canker diseases (Cryphonectriaceae, Botryosphaeriaceae) and root diseases (e.g. Armillaria, Ganoderma) of eucalypts.
Crous et al. (2019) state that the rust Austropuccinia psidii is one of the most damaging of the foliage fungal pathogens. They consider it to be a greater threat to eucalypt plantations outside the trees’ native ranges. (The Myrtaceous species in Australia most damaged by A. psidii are in other genera.)
Two families of leaf fungi – Mycosphaerellaceae and Teratosphaeriaceae – include species that cause serious diseases. Pérez, et al. report a study in plantation in Uruguay that detected six new species. They also discovered new hosts for some known species. (Such initial detections of new fungal species in out-of-native-range plantations is a usual occurrence.)
Over the 100-year history of planting eucalyptus outside Australasia, dozens of leaf pathogens have been transported to novel regions. Crous et al. 2019 report the wide geographic breadth of many of these introductions. For example, Mycosphaerellaheimii is crippling plantation forestry in five global regions – South America (Brazil and Venezuela); Asia (Indonesia and Thailand); Africa (Madagascar), Europe (Portugal); and in its presumably native Australia. A second species, M. marksii, has a similarly wide introduced range: Portugal, China and Indonesia, South Africa, Ethiopia, and Uruguay. Pérez et al. calls Mycosphaerella leaf diseases one of the most important impediments to Eucalyptus plantation forestry in Uruguay.
Although Crous et al. do not provide dates of detection, it appears that many of these leaf pathogens were introduced outside Australasia before the mid-990s, when the World Trade Organization (WTO) and International Plant Protection Convention (IPPC) came into force. Together, these agreements govern what actions phytosanitary officials may take to curtail international movement of plant pests. (To see my critique of the WTO/IPPC system, visit here.) The possible exception might be Kirramyces gauchensis, a well-known pathogen of Eucalyptus grandis in South America (Argentina and Uruguay), Hawai`i, and Africa (Uganda and Ethiopia) (Pérez, et al. 2009). Crous et al. (2019) expect another genus, Quambalaria species, to become a threat to eucalypt plantation forestry globally in the future.
Phoracantha semipunctata; photo by Umo Schmidt via Flickr
Arthropod pests have also been spread to many Eucalyptus-growing regions in North and South America, Europe and Africa since the 1980s. Some species have colonized virtually all eucalypt-growing regions, e.g.,Phoracantha semipunctata. Some have – so far – appeared on only one continent.
In an effort to determine how many of these introductions have occurred after adoption of the WTO/ IPPC system, I Googled the species named by Paine et al. (2011). I used the year 2000 as the cutoff date, to allow for detection lag. Among the insect species that fit this criterion are a lerp psyllid, a leaf beetle, and two gall wasps detected in North America; a true bug, two galling insects, and a leaf beetle in South Africa; and three psyllids in Europe.
Asia stands out as having very few introduced Australian insects plaguing eucalyptus plantations. Only one insect of Australian origin is causing significant damage in this region, Leptocybe invasa. It was detected after 2000, so it might have been introduced under the WTO/IPPC regime. Many widespread species, e.g., Phoracantha semipunctata, are notably absent. Instead, large numbers of endemic insects use these trees. This contrasts with the situation in the Southern Hemisphere, where few of the numerous native insects have shifted onto eucalypts.
New Zealand has detected only two new species of Australian origin since 1999 — two psyllids. This is despite the two nations’ proximity, the large volume of trade that passes between them, and the likelihood that at least some small sap-suckers might be introduced via aerial dispersal. New Zealand is famous for its strict phytosanitary (and sanitary) policies and programs.
Eucalyptus plantation in Kwa-Zulu, South Africa
Plantations’ vulnerability has been increased by expanding reliance on clonal, artificially-induced hybridization. Developers’ goals – and initial results – are enhanced adaptation to specific environments, desired fiber characteristics, and hybrid vigor. However, these vast areas planted in genetically identical trees are sitting ducks. An insect or pathogen that overcomes the host’s defenses can spread rapidly across the entire planting.
These hybrids also can act as “bridges,” facilitating spread of fungi to formerly resistant host species. Crous et al. (2019) fear that this process will undermine resistance in Eucalyptus pellita to the pathogen Teratosphaeria destructans. Plantations in Southeast Asia and South Africa now comprise hybrids between this resistant species and the highly susceptible Eucalyptus brassiana.
Pines
As with the eucalypts, the intensively managed pine plantations are comprised of fast-growing exotic species, all at the same developmental stage, and with minimal genetic diversity, planted to maximize wood production. These practices again lead to biological homogenization and reduced resilience to pests (Villacide and Fuetealba, 2025)
In the Southern Hemisphere, Sirex noctilio has become the most significant economic pest of Pinus species. These attacks can cause up to 80% mortality. Several other Sirex species have also been introduced, all apparently in the 1980s or earlier (Wilcken et al., 2025) – before adoption of the current international phytosanitary regime. However, in 2023, a new species, Sirexobesus, was discovered causing tree mortality in pine plantations in southeastern Brazil. This species is indigenous to the United States and Mexico.
Stazione et al. (2026) discuss two other non-native pine pests that established recently in South America.
Analysis of mitochondrial DNA of Orthotomicus erosus points to a western Eurasian lineage. The low genetic diversity of the introduced population in Argentina and Uruguay suggests a single or limited introduction event followed by regional spread.
The source region of Cyrtogenius luteus is more difficult to determine but is probably somewhere in China. The higher haplotype diversity might reflect multiple introductions. Again, shared haplotypes between Argentina and Uruguay countries indicates a contiguous regional spread, possibly driven by extensive pine plantations & intra-regional trade (Stazione et al. 2026)
Policy Aspects
Some scientists express concern about the failure of international phytosanitary measures. But are their countries speaking up in regulatory bodies, especially the International Plant Protection Convention?
Studies by Crous et al. (2019) and Pérez et al. (2009) clearly show that pathogens from Australia continue to be transported to regions where eucalypt plantations are grown. This happens despite most of the movement of genetic material being in the form of seeds – which should be less likely to transport pathogens than trade in plants. Pérez et al. (2009) explicitly raise concerns about the effectiveness of current quarantine procedures. Crous et al. (2019) state that quarantinescontinue to fail in many parts of the world.
Burgess and Wingfield (2017) list pathogens that have spread widely since the beginning of the 21st Century: Austropuccinia psidii, Calonectria (= Cylindrocladium) eudonaviculata (=Cylindrocladium buxicola), Ceratocystis lukuohia and C. huliohia introduced to Hawai`i. I add that insect-vectored diseases such as Euwallacea species carryingFusarium fungi have also experienced a burst of introductions around the globe since 2000.
Crous et al. (2019) attribute this failure partially to the enormous difficulty of applying effective quarantine to the huge volumes of planting material traded globally. Another factor is undoubtedly the poor understanding of microbial species, their pathogenicity, hosts, pathways of spread, even taxonomies. Some genera cannot be grown in culture.
Furthermore, pathogens’ impacts vary, possibly due to environmental conditions of the location or differing virulence on different hosts. Finally, with so many fungi and so little knowledge, it is difficult to separate true disease agents from multiple secondary infections.
Crous et al. (2019) express the hope that increased recognition of the importance of pathogens, along with improved detection and identification tools, will clarify patterns of spread. But is that enough? Are there no policy changes needed?
Crous et al. (2019) also warn us about additional pathways for spreading pathogens. Some potential pathogens of eucalypts have been moved on plants of other, related genera. Furthermore, Botryosphaeriaceae have been detected in the skins of mangoes (Mangifera indica) and avocados (Persea americana). Both of these fruits move globally in large volumes.
mangoes; photo by Obsidian Soul via Wikimedia
Regarding insects, Paine et al. (2011) focus on a concern that species native to the plantation countries and generalist herbivores from other parts of world will invade Australia and threaten eualypts in their native ranges. See other blog They also call for research to understand international pathways, develop detection methods, improve understanding of patterns of host suitability, susceptibility, and selection.
Villacide and Fuetealba (2025) note that while the introductory pathway for that new species, Sirex obesus, has not been determined, they suspect it might have been wood packaging materials. Villacide and another colleague (Lantschner and Villacide 2025) suggest an initial step would be for Argentina and other countries in the region to negotiate with Brazil to adopt more protective protocols governing trade in wood products, including wood packaging.
I have repeatedly advocated strengthening regulation of wood packaging. Such measures could improve protection of Earth’s forests from pests that use a well-documented high-risk introductory pathway. To see my arguments and underlying data, scoll down below the “archives” to “Categories” and click on “wood packaging”.
SOURCES
Burgess, T.I. and M.J. Wingfield. 2017. Pathogens on the Move: A 100-Year Global Experiment with Planted Eucalypts. Bioscience. Volume 67, Issue 1, January 2017. https://doi.org/10.1093/biosci/biw146
Crous, P.W., M.J. Wingfield, R. Cheewangkoon, A.J. Carnegie, T.I. Burgess, B.A. Summerell, J. Edwards, P.W.J. Taylor, and J.Z. Groenewald. 2019. Folia pathogens o eucalypts. Studies in Mycology 94:125-298 (2019).
Lantschner, V. and J. Villacide. 2025. Invasion Potential of the Recently Established Woodwasp Sirex obesus. Neotropical Entomology. (2025) 54:117 https://doi.org/10.1007/s13744-025-01347-6
Paine, T.D., M.J. Steinbauer, and S.A. Lawson. 2011. Native and Exotic Pests of Eucalyptus: A Worldwide Perspective. Annu. Rev. Entomol. 2011. 56:181-201
Payn, T., J-M. Carnus, P. Freer-Smith, M. Kimberley, W. Kollert, S. Liu, C. Orazio, L. Rodriguez, L. Neves Silva, M.J. Wingfield. 2015. Changes in planted forests and future global implications. Forest Ecology and Management 352 (2015)
Pérez,, C.A., M.J. Wingfield, N.A. Altier, and R.A. Blanchette. 2009. Mycosphaerellaceae and Teratosphaeriaceae associated with Eucalyptus leaf diseases and stem cankers in Uruguay For. Path. 39 (2009) 349–360 doi: 10.1111/j.1439-0329.2009.00598.x www3.interscience.wiley.com
Stazione, L., Soliani, C., Cognato, A. et al. Reconstructing the invasion history of the bark beetles Orthotomicus erosus & Cyrtogenius luteus (Coleoptera, Curculionidae, Scolytinae) in South America. Biol Invasions28, 49 (2026). https://doi.org/10.1007/s10530-026-03779-6
Villacide, J. and A. Fuetealba. 2025. Pests in plantations: Challenging traditional productive paradigms in the Southern Cone of America. Forest Ecology and Management 597 (2025) 123127
Wilcken, C.F., T.A. da Mota, C.H. de Oliveir, V.R. de Carvalho, L.A. Benso, J.A. Gabia, S.R.S. Wilcken, E.L. Furtado, N.M. Schiff, M.B. de Camargo, M.F. Ribeiro. 2025. Sirex obesus (Hymenoptera: Siricidae) as invasive pest in pine plantations in Brazil. Scientific Reports. 2025. 15:22522 https://doi.org/10.1038/541598-025-06418-7
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
Blepharocalyx salicifolius – a tree in the Myrtaceae native to South America on which found symptoms similar to those caused by Mycosphaerellaceae or Teratosphaeriaceae; photo by Pablo di Flores via Wikimedia
Pests that have followed their hosts to plantations outside the trees’ native ranges might threaten native plants in their new, introduced ranges. That is, the countries where the plantations are located.
Eucalypts
Eucalypts are now the most widely planted hardwood timber taxon in the world (Paine et al 2011). The 700 – 800 species in the three genera considered “eucalypts” (Angophora, Corymbia, and Eucalyptus) host a highly diverse fungal community — more than 500 species have been identified of just one type, leaf-infecting fungi (Crous et al. 2019).
As I described in a related blog, link dozens of leaf pathogens have been transported to countries hosting eucalypt plantations. Among them, two families – Mycosphaerellaceae and Teratosphaeriaceae – are prominent in both numbers of introductions and potential to cause serious diseases.
Nunez Chapa
Pérez et al. (2009) reported that a relatively large number of Mycosphaerellaceae and Teratosphaeriaceae are found on Eucalyptusin Uruguay. The authors cite one troubling case of host shifting: Mycosphaerella lateralis is causing leaf disease on a Musa cultivar (banana!) which is not in the Myrtaceae.
A follow-up study by the same authors (Pérez et al. 2013) surveyed several native forests, paying special attention to those located close to Eucalyptus plantations. They found five species belonging to the Mycosphaerellaceae and Teratosphaeriaceae clades on native Myrtaceous trees; three of these had previously been reported on Eucalyptus in Uruguay. Those occurring on both Eucalyptus and native Myrtaceae included Pallidocercospora heimii, Pseudocercospora norchiensis, and Teratosphaeria aurantia. A fourth species, Mycosphaerella yunnanensis, not previously recorded in Uruguay, was found on the leaves of two native Myrtaceous hosts. Pérez et al. (2013) believe circumstances indicate that all these fungi have been introduced. They warn that these apparent jumps to new hosts have the potential to result in serious disease problems and they should be carefully monitored. This finding is more than a decade old; I have not found a more recent report.
On the global level, Pérez et al. (2013) report, at least 23 species of Mycosphaerellaceae and Teratosphaeriaceae have been found on non-Eucalyptus species in the Myrtaceae. These hosts are in several plant orders, including Myrtales, Proteales, Fabaes and Apiales. The authors express “considerable concern” about the apparent ease of movement in these fungi between hosts. I have been unable to learn more details about these introductions.
Arthropod pests have also been spread to many Eucalyptus-growing regions in North and South America, Europe, and Africa since the 1980s – but not to Asia or New Zealand (Paine et al. 2011). blog
Myrrhinium atropurpureum – another South American plant in the Myrtaceae on which symptoms found; photo by Prof. Atilio L, Botanical Garden of Uruguay
Pines
Pines – a genus restricted naturally to the Northern Hemisphere – is second in popularity for intensively managed plantations. South America has 4.6 million hectares of pine plantations (Lantschner and Villacide 2025). Most are in Brazil, Chile, Uruguay, and Argentina (Payn et al. 2015).
Cinara cupressi; photo by LBM via Wikimedia
As I reported in an earlier blog, some of the insect pests that followed pines to South America have entered native forests. The most alarming of which I am aware is the aphid Cinara cupressi. It attacks the native conifer Austrocedrus chilensis, which forms pure and mixed stands with southern hemisphere beech (Nothofagus spp.) across approximately 160,000 hectares (Villacide and Fuetealba 2025). Cordilleran cypress is also under attack by the oomycete Phytophthora austrocedri, an oomycete of unknown origin.
Some scientists express concern about phytosanitary measures … but are their countries speaking up in meetings of the International Plant Protection Convention?
Studies by Crous et al. and Pérez et al. clearly show that pathogens from Australia continue to be transported to regions where eucalypt plantations are grown – despite the fact that most of the movement of tree genetic material is in the form of seeds – which should be less likely to transport pathogens than trade in plants. Pérez et al. (2009) explicitly raise concerns about the effectiveness of current quarantine procedures. Crous et al. (2019) state that the quarantinescontinue to fail in many parts of the world.
See my critique of the international phytosanitary system under the IPPC by visiting the Fading Forest II report (see link below) and reading other blogs under the categories “invasive species policy” and “plants as vectors of pests”.
SOURCES
Crous, P.W., M.J. Wingfield, R. Cheewangkoon, A.J. Carnegie, T.I. Burgess, B.A. Summerell, J. Edwards, P.W.J. Taylor, and J.Z. Groenewald. 2019. Foliar pathogens of eucalypts. Studies in Mycology 94:125-298 (2019)
Lantschner, V. and J. Villacide. 2025. Invasion Potential of the Recently Established Woodwasp Sirex obesus. Neotropical Entomology. (2025) 54:117 https://doi.org/10.1007/s13744-025-01347-6
Paine, T.D., M.J. Steinbauer, and S.A. Lawson. 2011. Native & Exotic Pests of Eucalyptus: A Worldwide Perspective. Annu. Rev. Entomol. 2011. 56:181-201
Payn, T., J-M. Carnus, P. Freer-Smith, M. Kimberley, W. Kollert, S. Liu, C. Orazio, L. Rodriguez, L. Neves Silva, M.J. Wingfield. 2015. Changes in planted forests & future global implications. Forest Ecology and Management 352 (2015)
Pérez, C.A., M.J. Wingfield, N. Altier, and R.A. Blanchette. 2013. Species of Mycosphaerellaceae and Teratosphaeriaceae on native Myrtaceae in Uruguay: evidence of fungal host jumps. Fungal Biology Volume 117, Issue 2, February 2013.
Villacide, J. and A. Fuetealba. 2025. Pests in plantations: Challenging traditional productive paradigms in the Southern Cone of America. Forest Ecology and Management 597 (2025) 123127
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
Chilecomadia valdiviana – one of the South American moths that attack Eucalyptus; photo by Natural History Museum of London via Wikimedia
Fifteen years ago, Paine, Steinbauer, and Lawson (2011) worried that insects in South America, Africa, Asia, and Europe that adapt to attacking Eucayptus trees planted there might be introduced to Australasia and threaten the genus in its native range. Their analysis applies to species in all three genera considered to be “eucalypts” — Angophora, Corymbia and Eucalyptus.
Some insects native to those continents have made this host shift already. Paine, Steinbauer, and Lawson reported that such host switching was especially prevalent among lepidopterans. They name several from Brazil, the Chilean cossid moth, Chilecomadia valdiviana, and southern African Coryphodema tristis. In their view, Brazilian eucalypt plantations’ proximity to native vegetation facilitates host-switching. Still, at that time they thought that there were no established pathways for introduction of the South American moths to Australia.
Host-switching is exceptionally common in Asia. Paine, Steinbauer, and Lawson (2011) thought the risk was greatest from insects on native eucalypts in near-neighbors Papua New Guinea, Timor, and The Philippines. An earlier risk assessment evaluating 10 insect species from the region concluded that most are polyphagous and probably switched to eucalypts. Two woodborers – Agrilus opulentus and A. sexsignatus –seem to have coevolved with Eucalyptus deglupta in New Guinea and The Philippines.
According to the same authors, most of the insects that have switched hosts are either polyphagous or normally feed on other myrtaceous species native to these regions. Thus, the Brazilian moth Thyrinteina arnobia feeds on Psidium guajava and several other Myrtaceae. Sarsina violascens is also a pest of Psidium species, as well as species in the Asteraceae, and Oleaceae. And the foliar rust Austropuccinia psidii was first described from Psidium guajava in Brazil and boasts a wide host range in the Myrtaceae in South America. It has been introduced to many regions with plants in the Myrtaceae, notably Hawai`i, Australia, South Africa, New Caledonia, and New Zealand. At least 15 Myrtaceae species in Australia are threatened with extinction.
Still, few non-native insects were damaging eucalypts in Australia’s native forests or plantations as of 2011. Those few are highly polyphagous. Several, if not most, were introduced in the first half of the 20th Century.
Why so few? Paine, Steinbauer, and Lawson (2011) suggest three possibilities: (a) Australia’s diverse endemic insects already occupy most niches, so they exclude new, foreign competitors; (b) most introduced insects were not previously exposed to Myrtaceae in their native range; and (c) Australia has strong quarantine procedures aiming to limit introductions of non-native herbivores.
The fact that none of the introduced insects has adapted to feed significantly on mature eucalypts’ above-ground tissues seems to me to point to protection provided by the adult trees’ phytochemicals and leaf structure. Paine, Steinbauer, and Lawson (2011) discuss some aspects of leaf structure and wax coatings.
As to Australia’s quarantine procedures, as I reported before, the country has been much less proactive regarding plant pests and diseases that threaten tree species rather than agricultural crops. Significant new programs were established only after 2000, when Plant Health Australia (PHA) was incorporated. The PHA is supposed to facilitate preparedness and response arrangements between governments and industry for plant pests (once an alien pest has become established, management becomes responsibility of the land manager). In 2005, federal, state, and territorial governments and plant industry bodies signed a legally-binding agreement — the Emergency Plant Pest Response Deed (EPPRD). 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.
Still, studies documented significant gaps in post-border forest biosecurity systems and the country’s response to the anticipated introduction of the foliar rust Austropuccinia psidii was disappointing. This prompted yet another initiative: development of the National Forest Biosecurity Surveillance Strategy (NFBSS) in 2018. The strategy was; accompanied by an Implementation Plan and appointment of a National Forest Biosecurity Coordinator. The forest sector fund a significant proportion of the proposed activities for the first five years. Still, Drs. Carnegie and Nahrung thought that in-country forest pest surveillance was still too fragmented.
Paine, Steinbauer, and Lawson (2011) consider the Asian spongy mothsLymantria dispar and Orgyia thyellina to pose serious threats. Five eucalypt species were assessed to be at risk of attack as are two preferred host oaks in Europe, Quercus pubescens and Q. robur. They note high volumes of imports from East Asia of containers, vehicles, and machinery, which are known to transport spongy moth egg-masses. It is not known whether the numerous natural enemies of Australia’s diverse lymantriid fauna [which includes four in the genus Lymantria] might provide some protection. These experts also worried that the highly polyphagous Asian longhorned beetle (Anoplophora glabripennis) might arrive in Australia. Eucalypts are not recognized as hosts.
Australia has adopted an enhanced surveillance program for ships arriving from Asian and European Lymantria ranges during female flight periods. Described here. Nahrung and Carnegie (2021) though that the high priority assigned to Lepidoptera exceeded the actual risk; only two non-native species had established in Australia over 130 years.
Paine, Steinbauer, and Lawson (2011) suggest several research topics aimed at reducing the risk to eucalypts in Australia. These include interactions between these insects and mechanisms by which insects adapt to new hosts; host chemistry and resistance mechanisms), chemical ecology (including host selection), population and community dynamics, including possible biocontrol agents, and pathway and risk analysis.
On the other hand, 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 noted that surveillance and management programs must expect and be prepared to respond to introductions of unanticipated species. They had found that 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.
SOURCES
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
Nahrung, H.F. and A.J. Carnegie. 2021. Border interceptions of forest insects established in Australia: intercepted invaders travel early and often. NeoBiota 64: 69–86. https://doi.org/10.3897/neobiota.64.60424
Paine, T.D., M.J. Steinbauer, and S.A. Lawson. 2011. Native & Exotic Pests of Eucalyptus: A Worldwide Perspective. Annu. Rev. Entomol. 2011. 56:181-201
Native & Exotic Pests of Eucalyptus: A Worldwide Perspective
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
pine plantation near Buenos Aires; photo by Biologicadero via Wikimedia
I have learned about the introduction of a North American woodwasp, Sirex obesus, in Brazil. Forestry interests in South America are worried that this woodwasp will cause significant damage to the pine plantations occupying 4.6 million hectares on the continent.
In July 2023, experts at the Estação Experimental de Ciências Florestais at ESALQ/USP in Itatinga, São Paulo, Brazil, investigated dead and symptomatic trees of several Pinus species and subspecies. They expected the causal agent to be Sirex noctilio – a woodwasp native to Europe and North Africa that has caused considerable damage to South American pine plantations since the 1980s (Wilcken et al.).
However, the pine species attacked were not typical hosts for S. noctilio (in Brazil, loblolly pine Pinus taeda). Instead, the infected trees were Caribbean pines, i.e., Pinus caribaea hondurensis, P. caribaea bahamensis, P. caribaea caribaea, P. maximinoi, P. tecunumani. The responsible woodwasp was identified as Sirex obesus. This species is native to the southwestern United States and northern and central Mexico (Wilcken et al.). This species is closely related to S. californicus (Wilcken et al.).
A second outbreak was found in November ~ 130 km away (still in São Paulo state). Scientists have not determined whether the two São Paulo outbreaks are related. Dr. Villacide reports (pers. comm.) that the two populations genetics have been compared, but he does not have the results.
A third population has been detected in a second, neighboring, state, Minas Gerais (Wilcken to Lantschner and Villacide).
Dr. Villacide (pers. comm.) reports that Brazilian scientists are trying to delimit the extent of the outbreaks. Public and private scientists in other countries with pine plantations have begun developing responses.
This is the first record of S. obesus outside of North America (Wilckens et al.).
Little is known yet about this woodwasp’s probable impact. It is clear that it can oviposit in a wide range of pines. In its native range, S. obesus has been reported on three host species: Pinus ponderosa, P. teocote (twisted-leaf pine), and P. leiophylla (no common name; native to Chihuahua – mostly in Mexico, and border areas of New Mexico and Arizona]. In Brazil, as noted, it has been recorded on other species as well as the hybrids P. caribaea x P. elliottii and P. caribaea x P. tecunumanii (Wilcken et al.).
So for purposes of their risk assessment, Lantschner and Villacide assumed that S. obesus can affect any of the species commonly planted in the region: P. taeda, P. elliottii, P. ponderosa, P. contorta, P. caribaea, P. oocarpa, P. patula, P. radiata, and P. tecunumanii (Lantschner and Villacide).
The risk assessment predicts suitable climatic conditions for invasion by S. obesus in 48% of the areas where South American pine plantation occur, particularly in montane and high-altitude regions along the Andean corridor and central-eastern Brazil. Incorporating other factors – host distribution, proximity to invaded areas, and volume of wood imports from Brazil – identified the most vulnerable areas as in southern Brazil, northeast Argentina, the Argentine Patagonia, and central Chile (Lantschner and Villacide).
pine plantation in Argentina; photo by Tomas Asurmendi via pexels
Preliminary sampling (Wilcken et al.) indicates the impacts could be severe. Mortality varies by species: in the worst cases average mortality approached 43% on P. caribaea hondurensis but only 11% on loblolly pine (P. taeda). They expect mortality rates to increase. Another 30% of P.c. hondurensis trees are dripping resin, a sign of woodwasp oviposition. If these eggs hatch, those larvae will probably kill the affected trees. Such a result would increase total mortality of P.c. hondurensis from 43% to ~ 73%. For P. taeda, the current mortality rate of 11% could rise to 49% as an additional 38% of trees succumb. Following this logic, these areas could experience complete tree mortality within a few years. Given the extent of pine plantations, and possible mortality rates, even a partial spread of S. obesus could lead to significant econ losses.
As second factor is the number of generations per year; the higher the number, the faster woodwasp populations can increase. Wilckens et al. report that adult emergence in Pinus logs maintained in cages indicates that S. obesus could have two or three generations per year.
S. obesus seems to prefer a different climate than S. noctilio. As noted, S. obesus seems to prefer montane and high-altitude climates. S. noctilio is concentrated in lowland temperate and humid regions (Lantschner and Villacide). The newly introduced species might substantially broaden the geographic area where pine plantations might be at risk – although further research is needed to clarify this point.
S. obesus also appears to be spreading at a rapid rate — ~46 km / year. At this rate, Lantschner and Villacide say it could spread throughout all major pine plantation areas in Brazil in less than years.
Sirex woodwasps kill trees by injecting a symbiotic wood decay fungus and a phytotoxic mucus into the tree when ovipositing. The toxin weakens the tree, allowing the fungus to spread, typically killing the tree in as little as three–four months. In North America S. obesus is associated with Amylostereum chailletti. While this species has not yet been confirmed in Brazil, (Wilckens et al.). Brazilian scientists are exploring whether S. obesus might adopt the fungus already present, Amylostereum areolatum, which is associated with S. noctilio.
Two insect species known to feed on woodwasps have emerged from logs infested with S. obesus: Ibalia leucospoides (Hymenoptera: Ibaliidae) and a species of Schlettererius (Hymenoptera: Stephanidae). While these two predators have not proved to be effective controls of woodwasps by themselves, they might become part of a control program. The parasitic nematode, Deladenus siricidicola (Nematoda: Neotylenchidae) used successfully in several South Hemisphere countries to control S. noctilio has not been found in Brazil (Wilckens et al.).
Scientists don’t know the pathway by which S. obesus entered Brazil. Wilckens believes it was via wood packaging; technicians from the Ministry of Agriculture have found some pallets associated with imports that lacked the ISPM#15 mark (Wilckens et al.).
Both Lantschner and Villacide and Wilcken et al. stress the vulnerability of South American pine plantations to introduction of damaging pests. The plantations are reportedly intensively managed, even-aged, regularly spaced monocultures. These conditions can facilitate invasive species establishment and spread by providing abundant host resources and reduced natural enemy pressure. Lantschner and Villacide cite Michael Wingfield that in plantation forestry, introduction of a single pest species can damage large areas of valuable timber.
mortality caused by Sirex noctilio in a pine plantation in Argentina; photo courtesy of Jose Villacide
The family Siricidae contains more than 120 species distributed across the forests of the Northern Hemisphere. In their native ranges they are typically minor or secondary pests (Wilckens et al.). Woodwasps have demonstrated that they can be transported in international commerce – S. noctilio alone has invaded pine stands (native or exotic) in nine countries in Oceania, Africa, and South and North America. Three other species in the family — Urocerus gigas, Urocerus flavicornis and Tremex fuscicornis – have been detected in South America (Wilckens et al.). If each represents a unique threat, countries with widespread pine plantations should enhance their phytosanitary programs. Exporting parties, e.g., the United States and European Union, should assist in efforts to prevent spread of these wood borers. One major step would be to strengthen regulations governing wood packaging material. [To see my criticisms of shortfalls of the ISPM#15 system, scroll down the list of blogs to “Categories” and click on “wood packaging”.]
Lantschner and Villacide cautionthat their assessment is based on a limited record of S. obesus occurrences in its native range. This range might be restricted by factors other than climate, including geographic barriers or biotic interactions (natural enemy pressure or interspecific competition). If so, the species’ potential invasive range might be larger than the climate-based models predict.
Recommendations for management strategies
I applaud Lantschner and Villacide for proposing immediate steps to improve management of the threat posed by introduction of S. obesus. These recommendations should prioritize enhanced phytosanitary inspections of wood products moving between high-risk regions and other South American countries. They suggest that Brazil adopt bilateral agreements with its major trading partners which would specify protocols for woodwaspdetection and quarantines. [Since many of these countries already have established populations of S. noctilio they probably do not have strong phytosanitary measures targeting wood borers at present.] Lantschner and Villacide advise creation of targeted surveillance programs in southern Brazil, northeastern Argentina, Argentine Patagonia, and central Chile. They should focus on sites near major transportation hubs and border crossings. Less intense surveillance should be instituted in regions they classified as medium risk. Again, the focus should be on major points of entry for imported goods and on plantations located near the Brazilian border. They note that preventing spread of S. obesus into new areas will require not only national efforts but also regionally coordinated monitoring, research, and forest health policies.
Lantschner and Villacide also identify priority areas for future research. These include clarifying S. obesus’shost range, the environmental conditions that enable the woodwasp to establish and persist beyond its native range, dispersal rates, and whether S. obesus exhibits pulse-like pop dynamics[long periods of low density interrupted by sudden outbreaks] seen in S. noctilio.
Dr. Villacide (pers. comm.) reports that Brazilian scientists are trying to delimit the extent of the outbreaks. Public and private scientists in other countries with pine plantations have begun developing responses. Dr. Villacide has posted a video from a recent online seminar sponsored by the Southern Cone Forest Health Group. Go to https://youtu.be/uVU6CpFNhlQ?si=lqXtwJTtz5rKXfL3 or https://sanidadforestalconosur.org/
A wider prespective
Dr. Villacide’s attention to Sirex obesus is part of his broader work on pest issues in South America’s commercial plantations. In another publication (Villacide and Fuetealba 2025; full citation at the end of this blog), he explores how to make these plantations sustainable in the face of rising threats from pests – both introduced and native to the region. Dr. Villacide and Alvaro Fuetealba report that every year 1.2 million hectares of plantations in the Southern Cone are affected by pests. Their vulnerability of will be worsened by the extreme weather events expected under climate change.
These plantations present vast areas of homogeneous stands: ~97% of the Southern Cone planted area consists of exotic tree species – mainly Pinus and Eucalyptus. Typical plantations are high density and managed intensively – including thinning, pruning, and fertilizing – to prompt rapid growth. As Villacide and Fuetealba point out, while these practices maximize wood production efficiency, they also lead to biological homogenization and reduced resilience to pests.
They report that pine plantations are under attack by wood and bark borers that have followed pines to the region, including Sirex noctilio, Orthotomicus erosus, and Cyrtogenius luteus; and now the newly detected Sirex obesus (above). At least two fungal pathogens — Fusarium circinatum and Dothistroma septosporum – have also been introduced. The principal threat to pine plantations from native pests comes from leaf-cutting ants (Atta and Acromyrmex).Eucalyptus plantations are plagued by several insects that have arrived from Australia, including Phoracantha semipunctata, Thaumastocoris peregrinus, and Leptocybe invasa. Pests native to the region that attack Eucalyptus are the Chilean carpenter worm (Chilecomadia valdiviana) and the leaf-cutting ants.
Cordilleran cypress; photo by LBM 1948 via Wikimedia
Threat to native conifer
More worrying to me is that introduced pests have entered native forests. Villacide and Fuetealba report that the aphid Cinara cupressi is attacking the native conifer Austrocedrus chilensis. Cordilleran cypress, also called Chilean or Patagonian cedar, is an endemic, monospecific tree in the Cupressaceae family. In southern Argentina and Chile the species forms pure and mixed stands with southern hemisphere beech (Nothofagus spp.) across ~ 160,000 ha. The profile Cinara cupressi on the Global Invasive Species Database is unclear about how many species are in the species complex and their places of origin.
Cordilleran cypress is also under attack by the oomycete Phytophthora austrocedri, an oomycete of unknown origin. This pathogen is of unknown origin. It is now thought to have been present in Argentina since at least the 1960s. P. austrocedri has also been ntroduced to Europe, western Asia, and North America.
Villacide and Fuetealba advocate several actions to might diversify tree species in the plantations to reduce their vulnerability to pests. They note that this recommendation builds on foundational ecological theory, including the resource concentration and natural enemy hypotheses. Diversity-promoting actions should reach beyond any plantation to the landscape level. Managers should consider connectivity of susceptible stands, the number of nutritionally optimal host trees in the landscape, and the availability and quality of hosts in adjacent stands.
Villacide and Fuetealba say mixed plantations can provide additional ecological and economic benefits, such as enhanced stand-level productivity; production of a wider range of commercial and subsistence products; and greater resistance and resilience to natural disturbances, e.g., extreme weather events.
They warn that designing and implementing mixed plantations must reflect ecological interactions and pest dynamics as well as management. There is need for regionally coordinated experimental plantations where scientist could test how variables such as tree species composition, density and spatial arrangement, and silvicultural practices influence pest dynamics, forest productivity, and ecosystem resilience under local conditions. They suggest incorporating sentinel plantings both early-warning systems and decision-support tools at plot and regional scales. Researchers should evaluate pest-specific responses, productivity trade-offs, long-term forest health outcomes under different scenarios.
Since the plantations extend across a multinational region with few natural barriers and uniform silvicultural practices, as well as high levels of trade, so do the pest problems. Therefore, the response must also be regional – e.g., regional experimental plantations and living laboratories. A collaborative approach linking researchers, forest managers, and policymakers is essential to translate experimental findings into practice and develop adaptive, ecol grounded silvicultural strategies. Long-term ecological trials must be embedded in operational contexts and aligned across countries.
SOURCES
Lantschner, V. and J. Villacide. 2025. Invasion Potential of the Recently Established Woodwasp Sirex obesus. Neotropical Entomology. (2025) 54:117 https://doi.org/10.1007/s13744-025-01347-6
Villacide, J. and A. Fuetealba. 2025. Pests in plantations: Challenging traditional productive paradigms in the Southern Cone of America. Forest Ecology and Management 597 (2025) 123127
Wilcken, C.F., T.A. da Mota, C.H. de Oliveir, V.R. de Carvalho, L.A. Benso, J.A. Gabia, S.R.S. Wilcken, E.L. Furtado, N.M. Schiff, M.B. de Camargo, M.F. Ribeiro. 2025. Sirex obesus (Hymenoptera: Siricidae) as invasive pest in pine plantations in Brazil. Scientific Reports. 2025. 15:22522 https://doi.org/10.1038/541598-025-06418-7
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
Erythrina caffra (native tree in South Africa)infested by PSHB; photo by J. Paap
A complex of closely related ambrosia beetles continues to be introduced to new places and cause increasing damage. The most widespread is the polyphagous shot hole borer (PSHB)Euwallacea fornicatus ss. Other members of the complex include the Kuroshio shot hole borer (KSHB) E. kuroshio and a third species, E. interjectus. Each beetle harbors its own plant-pathogenic fungus – all in the Fusarium genus.
Places invaded and impacts
The PSHB is established in the U.S. (southern and central California), Israel, South Africa, & Australia. The outbreak in South Africa covers the largest geographic area; the PSHB-Fusarium disease has been found in eight of the country’s nine provinces (every province except Limpopo) (Bierman et al. 2022). The South Africa outbreak is the most extensive geographically of all of them (Mudede et al. 2025).
The KSHB is established in southern California, from where it has spread to neighboring Mexico. E. interjectus is established in Santa Cruz County in California.
A fourth member of the species complex, E. perbrevis, has been established for decades on several Hawaiian islands and for at least 20 years in Florida. E. perbrevis has also been detected in nurseries in the Netherlands, but authorities reported it has been eradicated. E. perbrevis has long been known to be present in northern Queensland; this region might be part of its native range.
South America
In 2023 PSHB had been detected in Argentina – reported as E. fornicatus. A few weeks ago it was reported in neighboring Uruguay (PestLens for June 26, 2025). The beetle in South America is a different haplotype (genetic strain) than that introduced in South Africa, Israel, and California. It is more similar to specimens found in European greenhouses (Ceriani-Nakamurakare & others) As of 2022, scientists had identified 43 haplotypes (genetic variants) of E. fornicatus s.s. identified around the world; the greatest diversity is in several Asian countries (P. Rugman-Jones, pers. comm). The other species also comprise several haplotypes.
In South America the beetle has been observed attacking several new hosts. The most frequently attacked hosts are are Acer japonicum (Japanese maple) and a Ficus sp. Other hosts that support the full life cycle of the beetle and its associated fungus are Bauhinia forficata (cow’s foot), Ceiba speciosa (floss silk tree), Diospyros inconstans (jacuiba), Ficus aspera (mosaic fig), Fraxinus excelsior (European ash), Gardenia thunbergia (white gardenia), Geoffroea decorticans (chañar), Myrsine laetevirens, and Neltuma (Prosopis) caldenia (caldén) plants (PestLens June 26, 2025).
Ceiba speciosa A.St.-Hil. via Flickr
South Africa
The beetle and disease have been present since 2012 or earlier although it was not detected until 2017 (Winzer et al.). (This delay in detection is typical; in California PSHB was present for probably nine years before it was detected.) Winzer et al. decry the communication failure that resulted in the delayed official detection in South Africa and propose a system to correct the breakdown.
The haplotype (genetic strain) is the same as that found in Vietnam and introduced in California and other sites (Mudede et al.).
The South Africa outbreak is the most extensive geographically of all the outbreaks globally; within five years of its official detection, the PSHB-Fusarium disease was confirmed to be present in eight of the country’s nine provinces (every province except Limpopo) (Bierman et al. 2022).
More than 100 tree species – native and exotic – have been confirmed as hosts. Sources differ on the specific number: Mudede et al. report 130 species; Townsend et al. 2025 report 162. Both figures include both hosts that support reproduction of the insect and those that do not.
Mudebe et al. cite other studies that project the Fusarium disease will cause a decline in tree populations over a 10-year period of between 3.5% and 15.5%. They estimate the cost of removing urban trees killed by the disease will be $USD18.45 billion.
The impact in South Africa might differ from other invaded areas. Mudebe et al. report that over the five years of the study none of the Platanus species or A. buergerianum was dying despite being heavily infested. They say this suggests that trees can survive for more than 5 years.
Townsend et al. present a more disturbing picture. Their study examined PSHB impacts in plots in native forests in two provinces — KNZ (where PSHB first detected) and Western Cape. Over five years, PSHB invaded seven forest types; the only forest type not invaded was mangroves. PSHB colonization was detected on 43 native tree species. Eighteen species were recorded as competent hosts (able to support PSHB reproduction), eight as kill-competent hosts (can be killed by PSHB).
Over the five years 11 individual trees belonging to seven species died as a result of PSHB infestations. Some died very rapidly (within 2–5 years of first infestation); some died after apparently minor levels of infestation.
Each year of the study trees had a 7.5% increased chance of PSHB infestations; the number of entrance holes rose by over 10%. This means – no surprise – that the longer PSHB is active in the enviro the more trees it will infest, the higher its impact will be on hosts, & the higher the # of dispersing individuals produced. This will substantially increase the chances & rates of additional areas becoming infested, especially in areas close to infestation borders. Townsend et al. state that PSHB populations might be increasing exponentially – as occurred in California and Israel.
Townsend et al. discuss factors that might explain differing levels of infestation. Currently, a higher proportion of trees in the study plots in KwaZulu-Natal were infested than in the plots in Western Cape. The most likely explanation is that PSHB established there first – before 2012 compared to possibly five years later in the Western Cape. Other factors might be that source populations in the Western Cape were often found in alien tree species in urban areas distant from the study plots, while in KwaZulu-Natal, the beetles were frequently found in indigenous trees within monitoring plots. Forests in KZN are also fragmented, unlike the nearly contiguous woodlands in the Western Cape, and closer to urban areas with high PSHB infestation levels.
Although the PSHB’s spread into natural forests seems to be slow, Townsend et al. warn that they expect an increase in the rate of infestations as progressively more competent host individuals are infested. They fear severe ecological effects from rapid mortality of some key tree species, especially those sensitive to comparatively few attacks. They mention the native Erythrina caffra (coral tree), which is an important component of coastal forest ecosystems, especially in KwaZulu-Natal.
Other native trees at particular risk of PSHB infestations are Diospyros glabra, Ficus, Sparmannia africana, Trichelia emetic, and Vepris lanceolata. Townsend et al. remind us that each native tree species has a specific role in normal ecosystem functioning and supports a unique suite of species. Even if attacked trees do not die, Fusarium infection might weaken them, thereby increasing their susceptibility to other pathogens and pests, decreasing their longevity, or reducing their ability to produce fruits and flowers which can have long-term direct & indirect effects on normal ecosystem functioning & resilience.
Remember, South Africa is a biodiversity hotspot, home to its own Floral Kingdom!
The South Africans are trying to find more efficient methods for tracking spread of PSHB. Mudede et al. 2025 tested whether Google street view (GSV) images can be used to monitor its spread in urban forests. The test took place in Johannesburg. The test demonstrated that GSV images can be useful for mapping and monitoring PSHB-FD infestation on Platanus trees – but not on trees with rougher bark, e.g., Acer. While there were no false positives for any host species, most of the maple trees were misclassified as non-infested (false negatives).
Vietnam
Even in its native range, PSHB is a threat – in this case, to plantations utilizing non-native or exotic tree species. Thu et al. describe a growing number of pests threatening reforestation efforts in Vietnam. Surveys over the period 2011 to 2020 revealed outbreaks by 14 new insect species and 2 pathogens. Only two of the trouble-causing species are themselves non-native to Vietnam. One of these is PSHB. Thu et al. report the species’ range has spread rapidly in the country.
Thu et al. inform us that Vietnam has a high diversity of forest trees – and that almost nothing is known about pests that attack these trees.
Neolamarkia cadamba – native tree in Vietnam that might be resistant to PSHB; via Flickr
I welcome their call for higher investment in selection and breeding of hosts resistant to the various pests. The limited effort so far has identified provences of Neolamarckia cadamba and Nauclea orientalis that display some resistance to PSHB. Thu et al. advocate breeding programs to address the main biotic threats. They also recommend several actions to improve biosecurity, including enhanced hygiene in tree nurseries; improved silvicultural practices to minimize damage to trees; diversification of tree species being grown; and strengthening biosecurity and quarantine programs. They note that early detection of pest outbreaks is critical, so the country should develop forest health monitoring protocols for extensive forest reserves – sentinel plantings and remote sensing to detect trees under stress.
On a personal level, I found it interesting that Mudede et al. report that Google street view imagery determined that the invasive tree Ailanthus altissima dominates the street tree population in Istanbul – despite not having been intentionally planted. I visited Istanbul in April – and saw evidence of invasive vines and possibly the North American tree Cercis canadensis.
SOURCES
Ceriani-Nakamurakare, E., A.J. Johnson, D.F. Gomez. 2023. Uncharted Territories: First report of Euwallacea fornicates (Eichhoff) in South America with new reproductive host records. Zootaxa, 5325 (2), 289-297. https://doi.org/10.11646/zootaxa.5325.10
Mudede, M.F., S.W. Newete, K. Abutaleb, M.J. Byrn. 2025 Monitoring a polyphagous shot hole borer infestation in an urban forest using Google street view in the City of Johannasburg, South Africa Biol Invasions (2025) 27:144 https://doi.org/10.1007/s10530-025-03595-4
Thu, P.Q., D.N. Quang, N.M. Chi, T.X. Hung, L.V. Binh, B. Dell. 2021. New and Emerging Insect Pest and Disease Threats to Forest Plantations in Vietnam. Forests 2021, 12, 1301. https://doi.org/10.3390/f12101301
Townsend, G., M. Hill, B.P. Hurley, and F. Roets. 2025 Escalating threat: increasing impact of the polyphagous shot hole borer beetle, Euwallacea fornicatus, in nearly all major South African forest types. Biol Invasions (2025) 27:88 https://doi.org/10.1007/s10530-025-03551-2
Winzer, L.F, K.T. Faulkner, T. Paap, and J.R.U. Wilson. Preprint. From detection to action—a proposed workflow to ensure first reports of alien spp from molecular analyses are acted upon DOI: https://doi.org/10.3897/arphapreprints.e162421
Pest-lens link: https://pestlens.info/
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
Erythrina caffra one of the native tree species in South Africa killed by PSHB. photo by Coana/Riti via Flickr
Introductions of bark and ambrosia beetles (Coleoptera: Curculionidae, Scolytinae) have significantly increased over the past century. Surveys conducted at borders and ports of entry around the world have shown the majority of beetles intercepted were scolytines. These insects are highly destructive on their own. Also, they can carry pathogenic fungal symbionts that can have devastating effects on the trees they attack.
One or more species in a complex in the Euwallacea genus have become established in countries around the world. One of these, the polyphagous shot hole borer (Euwallacea fornicatus; PSHB) and its associated fungus (renamed from Fusarium euwallaceae to Neocosmospora euwallaceae) is threatening havoc in South Africa about a decade after its establishment (Townsend, Hill, Hurley, and Roets. 2025).
Over this brief period PSHB/Fusarium disease has spread from two introduction sites – Pietermaritzburg, in KwaZulu-Natal Province, and Cape Town, in Western Cape Province – to all but one of the country’s nine provinces. It has become established in four of five forest types studied – Afrotemperate, coastal, sand, and swamp forests. It has not established in mangrove forests. (The Western Cape Province is home to its own “floral kingdom”. The kingdom’s charactersitic fynbos flora is a heathland habitat, not a forest one.)
Townsend and colleagues established a network of 78 monitoring plots in the Western Cape and KwaZulu-Natal provinces. The sites reflected a variety of natural and human impacts.
tree infested by PSHB/Fusarium disease in KwaZulu-Natal Botanical Garden, Pietermaritzburg. Photo from website of Greenpop.org
By monitoring these plots over five years (2019 – 2024), Townsend and colleagues have demonstrated that the beetle/fungus complex and resulting “Fusarium disease” is spreading and intensifying. The number of infected trees rose from 100 to 176 over the five years – a mean increase of 0.6% per year. The number of PSHB entry holes increased by over 10% annually. The number of plots containing infected trees roughly doubled from 23 in 2019 (29% of the 78 plots) to 48 (60%) in 2023.
By the end of the study, 29% of the 148 species sampled had been infected. This represented 43 species and 7 unidentified trees infected. Trees of eight native species died, , although one — Diospyros glabra (Ebenaceae) – resprouted after the main bole died.
In addition to the eight species known to suffer mortality, another 18 species were found to be able to support PSHB reproduction. Townsend and colleagues worry that, as the infestation spreads and intensifies, some of these species might also succumb. They mention specifically Erythrina caffra (coral tree), which is prevalent in coastal forest ecosystems across South Africa.
Most of the hosts are in the same families as those identified earlier by Lynch et al. (2021), e.g., Ebenaceae, Fagaceae, Fabaceae, Malvaceae, Podocarpaceae, Rutaceae, Sapindaceae and Stilbaceae.
Disease progress, speed of death, and visibility of symptoms varied not only between species, but sometimes among individuals of the same species. Some trees died rapidly. Townsend and colleagues say it is impossible to predict which individuals will succumb to infection.
There is, though, a clear frequency-dependent relationship between trees and beetles. Sites with higher relative abundance of host trees also had a higher proportion of infected trees, on average. The number of PSHB holes per species and per plot both increased to a larger extent at these same sites.
Individual trees’ traits influenced the severity of infestations (measured by the number of PSHB entry holes). Larger trees, those with a less healthy canopy, and those farther from a water source suffered more attacks. (This last finding differs from others’; Townsend et al. speculate that in the absence of flood-stressed trees, drought-stressed trees might be more attractive to ambrosia beetles.)
native tree in Tsitsikama National Park; photo by F.T. Campbell
Characteristics of the monitoring plots also affected disease progression. Higher proportions of trees became infected when they grew in plots that were closer to source populations, or that contained a higher proportion of host species as distinct from non-host species. The proportion of trees infected decreased in plots with higher tree densities or tree species richness.
As of 2023, “Fusarium disease” is more widespread and intense in KwaZulu-Natal than in the Western Cape. In KwaZulu-Natal 0.11% of monitored trees are infected compared to 0.06% in the Western Cape. The number of infected trees rose twice as fast over the five years in KwaZulu-Natal – ~6%, than in Western Cape – 3%. While all KwaZulu-Natal plots contained infected trees, three of 11 monitoring sites in the Western Cape did not. Townsend and colleagues believe that the most likely explanation is that PSHB arrived in KwaZulu-Natal earlier (as far back as 2012 as opposed to 2017 in Western Cape). Another possible factor is that source populations of infected trees are indigenous trees within the forest in KwaZulu-Natal whereas, in the Western Cape, they are often non-native trees planted in urban areas far from the study plots. Also, forests in KwaZulu-Natal are fragmented while, in Western Cape, the study forests are nearly contiguous. Townsend et al. conclude that the disease will spread and intensify in Western Cape as additional source populations become established in the forest.
locations of PHSB/Fusarium disease in Cape Town, South Africa – West of the study sites; map from City of Cape Town
As of 2023, the proportion of trees infected appears to be small — 7.6% of the 2,313 trees monitored. Only 11 trees in the monitored plots have died. However, the longer PSHB is active in the environment the more trees it will infest, the higher its impact will be on hosts, and the higher the number of dispersing individuals produced. This will substantially increase the chances and rates of additional areas becoming infected, especially in areas close to infestations – e.g., cities. They fear that in the future impacts will increase as progressively more competent host individuals are infected. Therefore, they emphasize the importance of mitigating PSHB increase in natural ecosystems, even in already infected areas.
Townsend and colleagues urge phytosanitary officials and resource managers to prioritize surveillance and management on the families containing several host species (above) and within plant communities in which they predominate. Managers must also be alert to new reproductive hosts for the beetle that appear as the infestation spreads and intensifies.
The situation could be worse than described; the Townsend et al. study did not examine how the invasion might affect eco-regions outside these two provinces. Because the PSHB has such a broad host range, hosts can die quickly, and South Africa provides ideal climatic conditions, this bioinvader could cause severe ecological effects on most indigenous forest types as well as agriculture and urban trees throughout Africa.
SOURCES
Lynch, S.C., A. Escalen, and G.S. Gilbert. 2021. Host evolutionary relationships explain tree mortality caused by a generalist pest-pathogen complex. Evol Appl 14:1083 – 1094. https://doi.org/10.1111/eva.13182
Townsend, G., M. Hill, B.P. Hurley, and F. Roets 2025. Escalating threat: increasing impact of the polyphagous shot hole borer beetle, Euwallacea fornicatus, in nearly all major South African forest types. Biol Invasions (2025) 27:88 https://doi.org/10.1007/s10530-025-03551-2
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 https://treeimprovement.tennessee.edu/
Scientists in New Zealand are saying explicitly that a forest’s unique mixture of species matters when considering the future. This mixture is the result of the forest’s evolutionary history. Losing members of the biological community reduces the forest’s ability to respond to current and future stresses – its resilience.
New Zealand’s forests are part of the broader legacy of the ancient supercontinent of Gondwanaland – the island nation’s plants have close relatives in South America, the Pacific Ocean islands, and Australia. Still, these forests are unique: 80% of New Zealand’s plant species are endemic. The forests are also species-rich. The warm temperate evergreen rain forests of the North Island are home to at least 66 woody plant species that can reach that reach heights above six meters (Simpkins et al. 2024).
These forests have been severely changed by human activity. In just ~ 750 years people have cut down approximately 80% of the original forest cover! (Simpkins et al. 2024) Of the eight million hectares of surviving native forest, a little over five million hectares is managed for the conservation of biodiversity, heritage, and recreation. Another 2 million hectares are plantations of non-native species.
sites in New Zealand where pine plantations are “wilding”
All these forests are challenged by introduced mammals – from European deer to Australian possums. Climate change is expected to cause further disturbance, both directly (through e.g., drought, extreme weather) and indirectly (e.g., by facilitating weed invasion and shifting fire regimes) (Simpkins et al. 2024).
Pathogen threats are also common threats to the native trees of the Pacific’s biologically unique island systems. For example, Ceratocystis lukuohia and C. huliohia (rapid ‘ōhi‘a death, or ROD). The latter is killing ‘ōhi‘a (Metrosideros polymorpha) on the Hawaiian Islands. More than 40% of native plant species in Western Australia are susceptible to Phytophthora cinnamomi. Here I focus on two pathogens, kauri dieback and myrtle rust, now ravaging New Zealand’s native flora. No landscape-level treatment is available for either pathogen.
When considering this suite of challenges, Simpkins et al. focus on these two pathogens’ probable impact on forest carbon sequestration. They worry in particular about erosion of the forests’ resilience due to loss of “ecological memory” – the life-history traits of the species (e.g., soil seed banks) and the structures left behind after individual disturbances.
one of the largest remaining kauri trees, “Tane Mahuta”, in Waipoua Kauri Forest; photo by F.T. Campbell
Kauri Dieback
The causal agent of Kauri dieback, Phytophthora agathidicida, is a soil-borne pathogen that spreads slowly in the absence of animal or human vectors. The disease affects a single species, Agathis australis (kauri, Araucariaceae). However, kauri is a long-lived, large tree that is a significant carbon sink. It probably modifies local soil conditions, nutrient and water cycles, and associated vegetation. Also, kauri has immense cultural significance.
Simpkins et al. note that kauri dieback threatens stand-level loss of A. australis – that is, local extinctions. In the absence of disturbance Kauri trees can grow to awe-inspiring size. In the 19th Century, before widespread logging, some were measured at 20 meters or more in circumference. Consequently, kauri dieback might cause a decline in aboveground live carbon storage of up to 55%. This loss would occur over a period of hundreds of years, not immediately.
Huge kauri are not likely to be replaced by other long-lived emergent conifers (based on an analysis of one species, Dacrydium cupressinum). Instead, kauri are probably going to be replaced by late-successional angiosperms. The authors discuss the ecological implications for levels of carbon storage and proportions of trees composed of Myrtaceae – exacerbating damage caused by myrtle rust (see below).
The expectation of Simpkins et al. that kauri will suffer at least local extinctions is based on an assumption that no kauri trees are resistant to the pathogen. Fortunately, this might not be true: different Agathis populations show various levels of tolerance to Agathis dieback. Identification and promotion of some levels of resistance could enable A. australis to retain a diminished presence in the landscape.
However, Lantham, et al. make clear that containing kauri dieback remains “challenging,” despite its discovery nearly 20 years ago (in 2006). Scientists and land managers have little information on the distribution of symptomatic trees, much less of the pathogen itself. This means they don’t know where infection foci are or how fast the disease is spreading.
As is often true, the pathogen is probably present in a stand for years, possibly a decade or more, before symptoms are noticed. This means that the current reliance on public reports of diseased trees, or targetting surveillance on easy-to-access sites (e.g., park entrances and along existing track networks), or at highly impacted areas readily identified through aerial methods, fails to detect early stages of infection. Indeed, it seems probable that P. agathidicida had been present in New Zealand’s ecosystems for decades before its formal identification.
The Waipoua forest is one of the largest areas of forest with old kauri stands in the country. A new analysis of aerial surveys done between 1950 and 2019, shows how the forest is changing. The number of dead trees increased more than four-fold and the number of unhealthy-looking trees increased 16-fold over these 70 years. Kauri dieback is now widespread in this forest, especially in areas near human activities like clearing for pasture or planting commercial pine plantations).
Lantham et al. have developed a model which they believe will help identify areas of higher risk so as to prioritize surveillance and inform responses. These could delimit the disease front and help implement quarantines or other measures aimed at limiting the spread of P. agathidicida to uninfected neighboring sites.
I hope New Zealand devotes sufficient resources to expand surveillance and management to levels commensurate with the threat to this ecologically and culturally important tree species.
Leptospermum scoparia; photo by Brian Gatwicke via Flickr
Myrtle Rust
Myrtle rust is a wind-borne disease that affecting numerous species in the Myrtaceae, including some of the dominant early successional species (e.g., Leptospermum spp.). Simpkins et al. expect that myrtle rust might hasten the decline of two such tree species (L. scoparium and Kunzea ericoides). However, these trees’ small size and rapid replacement by other species during succession minimizes the effect of their demise on carbon storage.
Because I am concerned about the irreplaceable loss to biodiversity, I note that Simpkins et al. also feared immediate threats to some trees in the host Myrtaceae family, specifically highly susceptible species such as Leptospermumbullata.
As I reported in a recent blog, a second group of scientists (McCarthy et al.) explored the threat from myrtle rust more broadly. Austropuccinia psidii has spread through Myrtaceae-dominated forests of the Pacific islands for about 20 years.
Trees in the vulnerable plant family, Myrtaceae, are second in importance (based on density and cover) in New Zealand’s forests. Successional shrub communities dominated by the two species named above, Kunzea ericoides and Leptospermum scoparium, are widespread in the northern and central regions of the North Island and in northeastern and interior parts of the South Island. These regions’ vulnerability is exacerbated by the area’s climate, which is highly suitable for A. psidii infection (Simpkins et al. 2024).
McCarthy et al. concluded that ifLeptospermumscoparium and Kunzea ericoides prove to be vulnerable to myrtle rust, their loss would cause considerable change in stand-level functional composition across these large areas. They probably would be replaced by non-native shrubs, which are already common on the islands. Any resulting forest will differ from that formed via Leptospermeae succession.
These authors also worry that the risk to native ecosystems would increase if more virulent strains of the myrtle rust pathogen were introduced or evolved. They note that A. psidii is known to have many strains and that these strains attack different host species.
SOURCES
Latham, M.C., A. Lustig, N.M. Williams, A. McDonald, T. Patuawa, J. Chetham, S. Johnson, A. Carrington, W. Wood, and D.P. Anderson. 2025. Design of risk-based surveillance to demonstrate absence of Phytophthora agathidicida in New Zealand kauri forests. Biol. Invasions (2025) 27, no. 26
McCarthy, J.K., S.J. Richardson, I. Jo, S.K. Wiser, T.A. Easdale, J.D. Shepherd, P.J. Bellingham. 2024. A Functional Assessment of Community Vulnerability to the Loss of Myrtaceae from Myrtle Rust. Diversity and Distributions, https://doi.org/10.1111/ddi.13928
Simpkins, C.E., P.J. Bellingham, K. Reihana, J.M.R. Brock, G.L.W. Perry. 2024. Evaluating the effects of two newly emerging plant pathogens on North Aotearoa-New Zealand forests using an individual-based model. Ecological Modelling, www.elsevier.com/locate/ecolmodel
Posted by Faith Campbell
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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 https://treeimprovement.tennessee.edu/
