Category: other realms

New Sirex established in South America … threat to pine plantations + threat to native conifer from North American aphid

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

Or

https://fadingforests.org

Invasive shot hole borers – global travellers

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

Or

https://fadingforests.org

Shothole borer & associated fungus – demonstrating threat in South Africa & possibly beyond

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/

or

www.fadingforests.org

“Ecological memory” determines a forest’s resilience — implications of bioinvasion to New Zealand’s unique flora

kauri dieback

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 Leptospermum bullata.

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 if Leptospermum scoparium 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

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/

or

www.fadingforests.org

More pests in Europe & Mideast – hazard to North American trees

giant sequoia; photo by Matthew Dillon via Flickr

The pest alert system “PestLens” has again alerted us to plant pests in Europe or Asia that feed on species closely related to tree species native to North American forests.  Two of the insects named in the alert apparently pose a hazard to icons of the forests of America’s Pacific coast forests, giant sequoia and redwood.

I hope APHIS is using this information to alert port and on-the-ground staff and perhaps initiating more in-depth risk assessments.

The posting on February 27, 2025 reported that cotton jassid, Jacobiasca lybica (Hemiptera: Cicadellidae), affects not just cotton and citrus but also Cupressus sempervirens (Mediterranean cypress) [Cupressaceae]. More than a dozen North American trees species are in this family, including

  • Sequoiadendron giganteum or giant sequoia. Giant sequoia is listed as an endangered species by the IUCN with fewer than 80,000 remaining in its native California.
  • Chamaecyparis thyoides and C. lawsoniana (Port-Orford cedar). Port-Orford cedar has been decimated in its native range by an introduced pathogen, Phytopthora lateralis. A major breeding effort has developed trees that are resistant to the pathogen; they are now available for people to plant.
  • Thuja occidentalis, also known as northern white-cedar, eastern white-cedar, or arborvitae,
  • Taxodium ascendens, also known as pond cypress
  • several Juniperus
  • Hesperocyparis macrocarpa also known as Cupressus macrocarpa, or the Monterey cypress. NatureServe ranks the cypress as GI – critically imperiled.

Cotton jassid been reported from several countries in Europe, Africa, and the Middle East.

China has reported the existence of a previously unknown bark beetle species, Phloeosinus metasequoiae (Coleoptera: Curculionidae). It was found infesting Metasequoia glyptostroboides (dawn redwood) trees in China. Affected trees exhibited reddened leaves and holes and tunnels in branches.

China has also discovered a several new hosts utilized by the fungus Pestalotiopsis lushanensis (Sordariomycetes: Amphisphaeriales). Formerly known to infect tea (Camellia sinensis) and several other plant species, P. lushanensis has now been found shoot causing blight and leaf drop on a conifer, deodar cedar (Cedrus deodara) and leaf spots on an angiosperm with congeners in North America — the rare Chinese species, Magnolia decidua. There are eight species of Magnolia native to North America.

Magnolia grandiflora; photo by DavetheMage via Wikimedia

APHIS’ ability to respond to alerts remains uncertain.

The agency’s probationary employees have been fired – just as at other agencies. APHIS staff were prohibited from participating in last week’s annual USDA Invasive Species Research Forum – the 33rd such meeting. The bird flu emergency is demanding all the attention and funds.

So – how can the rest of us fill in?

At the USDA Research Forum I again presented a poster urging greater attention to tree-killing pathogens. Scientists have made considerable progress in identifying factors that indicate whether a non-native insect might pose a significant threat (see blogs on conifer and deciduous species; more to come!). However, USDA had not funded a similar effort to improve understanding of pathogens. The most promising strategy so far are sentinel plantings. However, these systems have weaknesses; I will blog in the near future about another analysis.

I propose that APHIS start by working with independent scientists to determine the actual, current level of pathogens associated with various types of incoming goods. Contact me directly if you wish to read the text of my poster.

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/

or

www.fadingforests.org

Pest Alerts – is USDA (able to) pay attention?

redbay (Persea borbonia killed by laurel wilt

The pest alert system “PestLens” provides information about new reports of plant pests around the world. Notices are published weekly. These provide North American stakeholders advance notice of pests to be on the lookout for. While I have followed these postings for several years, I have been alarmed by recent notices report documenting the presence of insects or pathogens that feed on species in the same genera as tree species native to North American forests. The alerts cover pests of all types of crops, not just trees.

I note that several of these not-yet-introduced pests attack the genus Persea, which contains several native tree and shrub species that are already severely affected by laurel wilt disease.

The report for 19 December, 2024, provided information about two pathogens.

flowering dogwood Cornus florida; photo by F.T. Campbell
  1. The bacterium Pectobacterium aroidearum (Gammaproteobacteria: Enterobacteriales) was detected in China. The bacterium infests several crops and Persea americana (avocado). Although the detection in China is new, the bacterium is apparently already widespread, since it has been earlier been reported from parts of Africa, the Middle East, Asia, Brazil, and Jamaica.
  2. The dagger nematodes Xiphinema simile and X. zagrosense (Longidoridae) were reported in Syria.  X. simile is associated with economically important plants, including Cornus spp. (dogwood; North American species already decimated by the introduced pathogen dogwood anthracnose), Malus spp.(apple), Prunus spp. (stone fruit), Quercus spp. (oaks – already under attack by many non-native organisms), and Vitis vinifera (grape). X. zagrosense is also associated with Poaceae. X. simile has earlier been reported from parts of Europe, Kenya, Iran, and Russia. X. zagrosense has also been reported from Iran.

The report for 9 January, 2025, conveyed information about 1 pathogen and 1 insect.

  1. It noted the presence in Thailand of the fungus Pseudoplagiostoma perseae (Sordariomycetes: Diaporthales) on Persea americana.
  2. The South American palm borer, Paysandisia archon (Lepidoptera: Castniidae), is infesting several palms at multiple locations in Switzerland. It attacks several economically important palm species and the native genus Washingtonia spp. (fan palm).
native California fan palm, Washingtonia filifera; photo by F.T. Campbell

The report for 13 February, 2025, gave information about 1 pathogen and 1 insect.

  1. An anthracnose fungus Colletotrichum aenigma (Sordariomycetes: Glomerellales) infecting Carya illinoinensis (pecan) and Ilex cornuta (Chinese holly) in China. Colletotrichum aenigma infects other economically important plants. These include the following genera with native species in North America: Vaccinium (blueberry), Malus (apple), Persea americana (avocado), and Vitis vinifera (grape). Colletotrichum aenigma is also widespread; it has been reported from parts of Europe, the Middle East, Asia, New Zealand, and South America.
  2. South African citrus thrips, Scirtothrips aurantii (Thysanoptera: Thripidae) in a greenhouse in the Netherlands. The thrips infests several woody plants, including Ilex crenata (Japanese holly), Rosa spp., Malus (apple), Persea americana (avocado), Prunus spp. (stone fruit), and Vitis vinifera (grape). S. aurantii  it is under eradication in Portugal and Spain. It has also been reported from parts of Africa, Yemen, and Australia.
Scirtothrips aurantii; photo by Pablo Alvarado Aldea, Spain

A few weeks ago I wanted to conclude this blog by stating my hope that APHIS is using this information to alert port and on-the-ground staff and perhaps initiating more in-depth risk assessments. Now – as we learn about mindless firings of USDA staff, I fear I must limit my hopes to a future for APHIS’ programs and skilled staff in more general terms.

Do we face shut-down of pest prevention/response efforts across the board?

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 here or here.

Coming to an Ecosystem Near You??

Xylotrechus chinensis; EPPO photograph

Europe has been invaded by two insect species that North Americans should be watching out for. First, a Cerambycid, the wasp-mimicking tiger longicorn beetle, Xylotrechus chinensis. And second,the Buprestid cypress jewel beetle, Lamprodila festiva. We should also ensure that none of the other 500+ beetles introduced to Europe poses a threat to our trees. These are summarized in a 2024 paper by Bunescu et al.

Tiger Longicorn Beetle

This beetle is native to eastern Asia. It feeds on and kills mulberry trees (Moraceae: Morus spp.). It might also attack apple and pear trees and grapevines – Asian sources report these as hosts. The status of grapevines has been questioned by a Spanish experiment, in which artificial inoculations failed. I have seen no further information about the vulnerability of apple (Malus spp.) and pear (Pyrus spp.) (Saarto i Monteyu, Costa Ribeu, and Savin 2021)

In Europe, the pest threatens mulberry trees which are commonly planted for shade and ornamentation, especially in southern France, Spain and Greece (Saarto i Monteyu, Costa Ribeu, and Savin 2021). For example, there are more than 20,000 mulberry trees in Athens (EFSA 2021). The trees’ abundance contributes to spread of any associated pests, the level of damage caused by falling branches, and the expense of tree removal. Economic damages are those typically associated with wood-borer invasions of urban areas. That is, the cost of tree removals, loss of shade and amenity values, and increased risk of injury from falling branches.

We Americans should be concerned, too. Wild red mulberry (Morus rubra) occupies much of the eastern United States, from southern New England west to southeastern Minnesota, then south along the eastern edge of the Great Plains to central Texas, and east to southern Florida. It is also found in Bermuda. It grows primarily in flood plains and low moist hillsides. . Presumably it would also be attacked by Xylotrechus chinensis, although I don’t know whether anyone has tested this. As a native tree, red mulberry plays a role in natural ecosystems, including wildlife food supplies. Thus, America would see even more significant losses if Xylotrechus chinensis were to establish.

Morus rubra in Fairfax County, Virginia; photo by Fmartin via Wikipedia

Red mulberry is already declining in parts of its central range, possibly due to a bacterial disease. The effects and extent of this disease have not been investigated thoroughly.

Apples and pears are important crops across North America; the farm-gate value is estimated at $3.2 billon.

Introductions of the beetle to Spain, France, and Greece might have resulted from inadequately-treated wood packaging or other wood products. Detections of the species in wood imports were reported in Germany in 2007 and 2017 (Saarto i Monteyu, Costa Ribeu, and Savin 2021). The U.S. has also intercepted X. chinensis at least once, at the port of Philadelphia, in 2011 (EFSA 2021).

These detections have raised questions to which no-one yet has answers. First, can X. chinensis develop in cut logs? The European Food Safety Agency concluded that it can (EFSA 2021). Second, one detection involved a shipment of wooden items made from birch (Betula spp.) and willow (Salix spp). It is not yet clear whether these taxa are also hosts (EFSA 2021). (The wood species were not specified in the case of the other interceptions.) I have blogged often about how “leaky” the wood packaging pathway has been; to see these blogs, scroll below the “archives” section of the webpage, then click on the category “wood packaging”.

European scientists believe X. chinensis might also be transported in shipments of plants for planting. However, the beetle prefers to oviposit on large trees. This pathway is less viable for the United States since USDA APHIS allows imports of mulberries (Morus) and pears (Pyrus) only from Canada. Apple trees (Malus spp.), however, may be imported from France – which hosts an introduced population of X. chinensis – and other European countries.  

Detection of any invasion by X. chinensis will pose the usual difficulties associated with woodborers. In some European cities, hundreds or even a thousand trees were infested before the outbreak was detected (EFSA 2021).

I am concerned that the Europeans appear to have been slow to respond to the threat from Xylotrechus chinensis. After several outbreaks were discovered in Greece, France, and Spain in 2017 and 2018, the European and Mediterranean Plant Protection Organization (EPPO) added X. chinensis to its Alert List. This action requires member states (which are not limited to European Union members) to report new outbreaks and inform about efforts to either stop or eradicate them (Saarto i Monteyu, Costa Ribeu, and Savin 2021).

Shortly afterwards the European Union Commission requested the European Food Safety Agency (EFSA) to conduct a risk assessment. This analysis was completed in 2021. (It contains lots of photos of the insect and its damage.) The analysis concluded that Xylotrechus chinensis could probably infest most areas in the Union and cause significant damage. The species meets the criteria for designation as a quarantine pest in the Union. However, as of December 2024, this action had not been taken. As a result, control measures for this species are not mandatory.

Introductions continue; an outbreak in Lombardy, Italy, was found in June 2023 (Sarto i Monteys, Savin, Torras i Tutusaus & Bedós i Balsach 2024).  European regulations – following IPPC standards – also are linked to named pests and known outbreak locations. Such restrictions almost guarantee that the pest will continue to spread from not-yet-detected outbreaks. (Decades ago, after the emerald ash borer invasion, Michigan’s State Plant Regulatory Official, Ken Rasher, noted that, to be effective, “slow the spread” efforts must apply to areas beyond the known limits of the pest’s range.) The EFSA risk assessment did suggest delimitation of buffer zones around known European outbreaks. I don’t know whether such zones have been set up.

The risk assessment also recommended [true?] improving detection of this insect by developing male pheromones as lures. These have not been acted on. Guidance on best timing for treatment [trunk injections of systemic insecticides] also appears to have been taken up by Greece but not by Spain (Sarto i Monteys, Savin, Torras i Tutusaus & Bedós i Balsach 2024).

These authors include more information about the Xylotrechus chinensis life cycle and trajectory of the invasion,. They note that climate change appears to be altering the insect’s phenology. Especially, the adult flight period is beginning earlier in the spring.

Lamprodila festiva; Udo Schmidt via flickr

Cypress jewel beetle

This second pest of concern is a buprestid that attacks trees in the Cupressaceae. Infested trees generally die within a few years.

In its native Mediterranean range, the beetle feeds on native Juniperus, Cupressus and Tetraclinis. In invaded urban landscapes of Europe it attacks primarily introduced Cupressaceae , particularly Thuja, Chamaecyparis, Platycladus, Callitris, and some hybrids (Cupressocyparis). It has also been recorded as damaging Sequoia sempervirens (Brunescu, et al., 2024). (Genera in bold are native to North America.)

Thuja occidentalis; photo by H. Zell via Wikimedia

White cedar, Thuja occidentalis is the focus of Brunescu, et al.’s article. It is native to eastern Canada and much of the north-central and northeastern United States. The European and Mediterranean Plant Protection Organization (EPPO) has identified eight species in the Lamprodila genus as important pests, (Brunescu et al. 2024) so the danger might be more widespread. The invasion of Europe is probably the result of adult flight or other short-range transport. The article does not suggest pathways that the species might exploit to cross oceans.

SOURCES

Bunescu, H., T. Florian, D. Dragan, A. Mara, I-B. Hulujan, X-D. Rau. 2024  The Cypress Jewel Beetle Lamprodila Festiva Linné, 1767 (Coleoptera: Buprestidae), an Invasive Major Pest of Thuja Occidentalis Linné in Romania Hop and Medicinal Plants, 2024 XXXII, No. 1-2, 2024.

Saarto i Monteyu V., A. Costa Ribeu. I. Savin. 2021a. The invasive longhorn beetle Xylotrechus chinensis, pest of mulberries, in Euro: Study on its local spread & efficacy of abamectin control Plos One January 29, 2021. https://doi.org/10.1371/journal.pone.0245527

Sarto i Monteys, V., I. Savin, G. Torras i Tutusaus & M. Bedós i Balsach. 2024b. New evidence on the spread in Catalonia of the invasive longhorn beetle, Xylotrechus chinensis, & the efficacy of abamectin control. Scientific Reports | (2024) 14:26754 | https://doi.org/10.1038/s41598-024-78265-x  www.nature.com/scientificreports/

Posted by Faith Campbell

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

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

or

www.fadingforests.org

Impacts of introduced rust on unique flora — New Zealand’s expectations

predicted community vulnerability from A. psidii mediated mortality of Kunzea ericoides & Leptospermum scoparium; from McCarthy et al.

Scientists in New Zealand have recently completed a study of the probable impact of myrtle rust – caused by Austropuccinia psidii – on plants in the plant family Myrtaceae. McCarthy et al. say their results should guide management actions to protect not only the unique flora of those islands but also on Australia and Hawai`i – other places where key dominant tree species are susceptible to myrtle rust. The disease attacks young tissue; susceptible Myrtaceae become unable to recruit new individuals or to recover from disturbance. Severe cases can result in tree death & localized extinctions

[I note that myrtle rust is not the only threat to the native trees of these biologically unique island systems. New Zealand’s largest tree, kauri (Agathis australis), is threatened by kauri dieback (caused by Phytophthora agathidicida). On Hawai`i, while the most widespread tree, ‘ōhi‘a (Metrosideros polymorpha) is somewhat vulnerable to the strain of rust introduced to the Islands, the greater threat is from a different group of fungi, Ceratocystis lukuohia and C. huliohia, collectively known as rapid ‘ōhi‘a death. On Australia, hundreds of endemic species on the western side of the continent are being killed by Phytophthora dieback, caused by Phytophthora cinnamomi. [I note the proliferation of tree-kiling pathogens; I will blog more about this in the near future.]

Myrtle rust arrived in New Zealand in 2017, probably blown on the wind from Australia (where it was detected in 2010). In New Zealand, myrtle rust infects at least 12 of 18 native tree, shrub, and vine species in the Myrtaceae plant family. Several of these species are important in the structure and succession of native ecosystems. They also have enormous cultural significance.

McCarthy et al. note that species differ in their contribution to forest structure and function. They sought to determine where loss of vulnerable species might have the greatest impact on community functionality. They also explored whether compensatory infilling by co-occurring, non-vulnerable species in the Myrtaceae would reduce the community’s vulnerability. Even when co-occurring Myrtaceae are relatively immune to the pathogen, only some of them – the fast-growing species – are likely to fill the gaps. They might lack the functional attributes of the decimated species.

To identify areas at greatest risk, McCarthy et al. took advantage of a nationwide vegetation plot dataset that covers all the country’s native forests and shrublands. The plot data enabled McCarthy et al. to determine which plant species not vulnerable to the rust are present and so are likely to replace the rust host species as they are killed.

Leptospermum scoparium; photo by Alyenaa Buckles via Flickr

McCarthy et al. concluded that forests and shrublands containing Kunzea ericoides and Leptospermum scoparium are highly vulnerable to their loss. Ecosystems with these species are found predominantly in central and southeastern North Island, northeastern South Island, and Stewart Island. While compensatory infilling by other species in the Myrtaceae would moderate the impact of the loss of vulnerable species, if these co-occurring species were unable to respond for various reasons, such as also being infected by the rust pathogen, community vulnerability almost always increased. In these cases the infilling species would probably have different functional attributes. In many areas the species most likely to replace the rust-killed native species would be non-native shrubs. Consequently, early successional woody plant communities, where K. ericoides and L. scoparium dominate, are at most risk.

Because the risk of A. psidii infection is lower in cooler montane and southern coastal areas, parts of inland Fiordland, the northwestern South Island and the west coast of the North Island might be less vulnerable.

Austropuccinia psidii has been spreading in Myrtaceae-dominated forests of the Southern Hemisphere since the beginning of the 21st Century. It was detected in Hawai`i in 2005; in Australia in 2010; in New Caledonia in 2013, and finally in New Zealand in 2017. Within 12 months of its first detection in the northern part of the North Island it had spread to the northern regions of the South Island.

Specific types of Threat

Succession

The ecosystem process most at risk to loss of Myrtaceae species to A. psidii is succession. About 10% of once-forested areas of New Zealand are in successional shrublands, mostly dominated by Kunzea ericoides and Leptospermum scoparium. Both species are wind dispersed, grow quickly, are resistant to browsing by introduced deer, and are favored by disturbance, especially fire. Both are tolerant of exposure and have a wide edaphic range (including geothermal soils). Still, K. ericoides prefers drier, warmer sites while L. scoparium tolerates saturated soils, frost hollows and subalpine settings.

Kunzea ericoides; photo by Tony Foster via Flickr

Loss of these two species would result in a considerable change in stand-level functional composition across a wide variety of locations. Their extensive ranges mean that it would be difficult for other species – even if functionally equivalent – to expand sufficiently quickly. Second, non-native species are common in these communities. All of these invaders – Ulex europaeus, Cytisus scoparius and species of Acacia, Hakea and Erica – promote fire. Some are nitrogen fixers. While they can facilitate succession, the resulting native forest will differ from that formed via Leptospermeae succession. Furthermore, compensatory infilling by the invasive species might also reduce carbon sequestration. Successional forests dominated by K. ericoides are significant carbon sinks owing to the tree’s size (up to 25 m under favorable conditions), high wood density, and long lifespan (up to ~150 years). In contrast, shrublands dominated by at least one of the non-native species, U. europaeus, are significant carbon sources.

Northern and central regions of the North Island and the northeastern and interior parts of the South Island are most vulnerable to the loss of these species since these successional shrub communities are widespread and the area’s climate is highly suitable for A. psidii infection. The southern regions of the South Island, including Stewart Island, are somewhat protected by the cooler climate.

Fortunately, neither Kunzea ericoides nor Leptospermum scoparium has yet been infected in nature. Laboratory trials indicate that some families of K. ericoides are resistant. Vulnerability also varies among types of tissue – i.e., leaf, stem, seed capsule.  

Metrosideros umbellata; photo by Stan Shebs via Wikimedia

Forest biomass

Although from the overall community perspective loss of species in the Metrosidereae would have a lower impact than loss of those in the Leptospermeae, there would be significant changes associated with loss of Metrosideros umbellata. This species can grow quite large (dbh often > 2 m; heights up to 20 m). That size and its exceptionally dense wood means that M. umbellata stores high amounts of carbon. Also, its slow decomposition provides habitat for decomposers. Lessening the potential impact of loss of this species are two facts: its litter nutrient concentrations and decomposition rates do not differ from dominant co-occurring trees; and, most important, it grows primarily in the south, where weather conditions are less suitable for A. psidii infection. One note of caution: if A. psidii proves able to spread into these regions, not only M. umbellata but also susceptible co-occurring Myrtaceae species are likely to be damaged by the pathogen.

Highly specific habitats

McCarthy et al. note that their study might underestimate the impact of loss of species with unique traits that occupy specialized habitats. They focus on the climber Metrosideros excelsa. This is an important successional species that helps restore ecosystems following fire, landslides, or volcanic eruptions. The species’ tough and nutrient poor leaves promote later successional species by forming a humus layer and altering the microenvironment beneath the plant. Its litter has high concentrations of phenolics and decomposes more slowly than any co-occurring tree species.  [They say its role is analogous to that of M. polymorpha in primary successions on lava flows in Hawai`i.] M. excelsa dominates succession on many small offshore volcanic islands, rocky coastal headlands and cliffs.

Another example is Lophomyrtus bullata, a small tree that is patchily distributed primarily in forest margins and streamside vegetation. This is the native species most affected by A. psidii; the pathogen is likely to cause its localized extinction. McCarthy et al. call for assessment of ex situ conservation strategies for this species.

Each of these species is represented in only seven of the plots used in the analysis, so community vulnerability to their loss might be underestimated.

Another habitat specialist, Syzygium maire, is found mostly in lowland forests, usually on saturated soils. It currently occupies only a fraction of its natural range due to deforestation and land drainage. Evaluating the impact of loss of S. maire is complicated by its poor representation in the database (only six plots), and the fact that many of the co-occurring species are also Myrtaceae.

Lack of data similarly prevents detailed assessment of the impacts from possible loss of other species, including M. parkinsonii, M. perforata and L. obcordata. McCarthy et al. say only that their disappearance will “take the community even further from its original state”.

McCarthy et al. warn that the risk could increase if more virulent strains of A. psidii were introduced or evolved through sexual recombination of the current pandemic strain. Other scientists have discovered strong evidence that the many strains of A. psidii attack different host species (see Costa da Silva et al. 2014).

New Zealand bell bird (Anthornis melanura); photo from https://animalia.bio/new-zealand-bellbird

McCarthy et al. note that other factors are also important in determining the impact of loss of a plant species. Especially significant is the host plant species’ association with other species. They say these relationships are poorly understood. One example is that only four Myrtaceae species produce fleshy fruits. Loss or decline of these four species might severely affect populations of native birds, many of which are endemic. Many invertebrates – also highly endemic — are dependent on nectar from other plants in the family.

In their conclusion, McCarthy et al. note that A. psidii has been introduced relatively recently so there is still time to reduce the disease’s potential consequences. They suggest such management interventions as identifying and planting resistant genotypes and applying chemical controls to protect important individual specimens. They hope their work will guide prioritization of both species and spatial locations. They believe such efforts have substantial potential to reduce myrtle rust’s overall functional impact to New Zealand’s unique ecosystems.

SOURCES

Costa da Silva, A; P.M. Teixeira de Andrade, A. Couto Alfenas, R. Neves Graca, P. Cannon, R. Hauff, D. Cristiano Ferreira, and S. Mori. 2014. Virulence and Impact of Brazilian Strains of Puccinia psidii on Hawaiian Ohia (Metrosideros polymorpha). Pacific Science 68(1):47-56.  doi: https://dx.doi.org/10.2984/68.1.4

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 & Distributions, 2024; https://doi.org/10.1111/ddi.13928

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/

or

www.fadingforests.org

Non-Native Moths in England: Ever Upward

Platyperigea kadenii — one of the moth species that feeds on native plant species introduced recently to Great Britain. Photo by Tony Morris via Flickr

Will phytosanitary agencies and the international system respond to continuing introductions of non-native species?

A new study confirms that introductions of insects continue apace, links this pattern to the horticultural trade, and examines the role of climate change in facilitating introductions. This study focuses on moths introduced to the United Kingdom (Hordley et al.; full citation at the end of the blog). The study sought to detect any trends in numbers of species establishing and the relative importance of natural dispersal vs. those assisted – intentionally or inadvertently – by human activities.

The authors determined that moths continue to be introduced by both processes; there is no sign of “saturation”. This finding agrees with that of Seebens and 44 others (2017; citation below), which analyzed establishments of all types of non-native species globally. The British scientists found that rapidly increasing global trade is the probable driver of the recent acceleration of human-assisted introductions. They emphasize the horticultural trade’s role specifically. Climate change might play a role in facilitating establishment of species entering the UK via human activities.

Hordley et al. found that long-term changes in climate, not recent rapid anthropogenic warming, was important in facilitating introductions of even those moth species that arrived without human assistance. As they note, temperatures in Great Britain have been rising since the 17th Century. These changes in temperature have probably made the British climate more suitable for a large number of Lepidoptera. The data show that the rate of natural establishments began rising in the 1930s, 60 years before anthropogenic changes in temperatures became evident. Hordley et al. point out that an earlier study that posited a more significant role for climate change did not distinguish between insect species which have colonized naturally and those benefitting from human assistance.

The authors expect introductions to continue, spurred by ongoing environmental and economic changes. Fortunately, very few of the introduced moths had any direct or indirect negative impacts. (The box-tree moth (Cydalima perspectalis) is the exception. [Box-tree moth is also killing plants in North America.]

boxtree moth; photo by Tony Morris via Flickr

Still, they consider that introductions pose an ongoing potential risk to native biodiversity and related human interests. Therefore, they advocate enhanced biosecurity. Specifically, they urge improved monitoring of natural colonizations and regulation of the horticultural trade.

Hordley et al. estimated the rate of establishment during the period 1900 – 2019 for (i) all moth species; (ii) immigrants (i.e., those introduced without any human assistance); (iii) immigrants which feed on native hosts; (iv) immigrants which feed on non-native hosts; (v) adventives (i.e., species introduced with human assistance); (vi) adventives which feed on native hosts; and (vii) adventives which feed on NIS hosts.

Their analysis used data on 116 moth species that have become established in Great Britain since 1900. Nearly two-thirds of these species – 63% – feed on plant species native to Great Britain; 34% on plant species that have been imported – intentionally or not. Data were lacking on the hosts of 3 species.

Considering the mode of introduction, the authors found that 67% arrived through natural colonization; 33% via human assistance. Sixty-nine percent of the 78 species that were introduced through natural processes (54 species) feed on plant species native to Great Britain; 31% (24 species) feed on non-native plants. Among the 38 species whose introduction was assisted by human activities, one-half (19 species) feed on native plant species; 42% (16 species) feed on introduced hosts.

Regarding trends, they found that when considering all moth species over the full period, 21.5% more species established in each decade than in the previous decade. This average somewhat obscured the startling acceleration of introductions over time: one species was reported as established in the first decade (1900–1909) compared to 18 species in the final decade (2010–2019).   

The rate of introduction for all immigrant (naturally introduced) species was 22% increase per decade. Considering immigrant species that feed on native plants, the rate of establishment was nearly the same – 23% increase per decade – when averaged over the 120-year period. However, a more detailed analysis demonstrated that these introductions proceeded at a steady rate until 1935, then accelerated by 11% per decade thereafter. In contrast, immigrants that feed on non-native plants have maintained a steady rate of increasing establishments – 13% per decade since 1900.

Adventive species (those introduced via human assistance) increased by 26% per decade. The data showed no signs of saturation. The rates of introduction were similar for adventives that feed on both native plants (22%) and non-native hosts (26%). Again, additional analysis demonstrated a break in rates for adventives that feed on native hosts. The rate was steady until the 1970s, then significantly increased during the years up to 2010. (The scientists dropped data from the final decade since lags in detection might artificially suppress that number.)

In summary, Hordley et al. found no significant differences in trends between

  • the number of species that established naturally (20%) vs. adventives (26%).
  • immigrant or adventive species that feed on native vs. non-native hosts.

The authors discuss the role of climate change facilitating bioinvasion by spurring natural dispersal, changing propagule pressure in source habitats, changing the suitability of receiving habitat, and changing in pathways for natural spread, e.g., altered wind and ocean currents. They recognize that the two modes of colonization – adventives and immigrants – can interact. They stress, however, that the two colonization modes require different interventions.

Although their findings don’t support the premise that a surge of natural colonizers has been prompted by anthropogenic warming, Hordley et al. assert that climate clearly links to increased moth immigration to Britain and increased probability of establishment. They note that even so assisted, colonists still must overcome both the natural barrier of the English Channel and find habitats that are so configured as to   facilitate breeding success. They report that source pools do not appear to be depleted — moth species richness of neighboring European countries greatly exceeds that in Great Britain.

I would have liked to learn what factors they think might explain the acceleration in both natural and human-assisted introductions of species that feed on plant species native to Great Britain. In 2023 I noted that scientists have found that numbers of established non-native insect species are driven primarily by diversity of plants – both native and non-indigenous.

Hordley et al. assert that Great Britain has advantages as a study location because as a large island separated from continental Europe by the sea – a natural barrier – colonization events are relatively easy to detect. However the English Channel is only 32 km across at its narrowest point. I wonder, whether this relatively narrow natural barrier might lead to a misleadingly large proportion of introduced species being natural immigrants. I do agree with the authors that moths are an appropriate focal taxon because they are sensitive to climate and can be introduced by international trade. Furthermore, Britain has a long tradition of citizen scientists recording moth sightings, so trends can be assessed over a long period.

Hordley et al. stress that they measured only the temporal rate of new species’ establishments, not colonization pressure or establishment success rate. They had no access to systematic data regarding species that arrived but failed to establish. Therefore, they could not deduce whether the observed increase in establishment rates are due to:

(1) more species arriving—due either to climate-driven changes in dispersal or to accessibility of source pools; or

(2) higher establishment success due to improved habitat and resource availability; or

(3) both.

Hordley et al. noted two limitations to their study. First, they concede that there is unavoidably some subjectivity in classifying each species as colonizing naturally or with human assistance. They tried to minimize this factor by consulting two experts independently and including in the analysis only those species on which there was consensus.

Second, increases in detection effort and effectiveness might explain the recent increases in establishment rates. They agree that more people have become “citizen scientists” since 1970. Also, sampling techniques and resources for species identification have improved considerably. They note, however, that Seebens et al. (2018) tested these factors in their global assessment and found little effect on trends.

Hordley et al. believe that they have addressed a third possible limitation – the lag between introduction and detection – by running their analyses both with and without data from final decade (2010-2019). The results were very similar qualitatively.

SOURCE

Hordley, L.A., E.B. Dennis, R. Fox, M.S. Parsons, T.M. Davis, N.A.D. Bourn. 2024. Increasing rate of moth species establishment over 120 years shows no deceleration. Insect Conserv. Divers. 2024;1–10. DOI: 10.1111/icad.12783

Seebens, H. et al. 2017. No saturation in the accumulation of alien species worldwide. Nature Communications. January 2017. DOI: 10.1038/ncomms14435

Seebens, H. et al. 2018. Global rise in emerging IAS results from increased accessibility of new source pools. Proceedings of the National Academy of Sciences. www.pnas.org/cgi/doi/10.1073/pnas.1719429115

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/

or

www.fadingforests.org

Phytophthora here, Phytopthora there … level of threat is unclear

Mt. Triglav – highest peak in the Slovenian (Julian) Alps; photo by Gunter Nuyts via Pexel

Scientists have discovered sizable diversity of pathogenic Phytophthora species in Europe, specifically in the Alps of northeastern Italy and western Slovenija.  They have also named a new species, and noted the need to change the definition of species previously named. See Bregant et al. – full citation at the end of this blog – open access!

Two of its findings are especially important for the US

First, the authors document the vulnerability of alpine areas to 18 Phythophthora species. Most of the plant hosts they studied have congenerics in mountainous areas of North America: Acer, Alnus, Betula, Fagus, Fragaria, Fraxinus, Ilex, Juniperus, Larix, Lonicera, Lycopodium, Pinus, Populus, Quercus, Rhododendron, Rubus, Salix, Sorbus, Taxus, and Vaccinium.

Second, the paper discusses how junipers are at particular risk. I remind you that P. austrocedrii has recently been detected in nurseries in Ohio and Oregon. This is another non-native Phythophthora that attacks junipers. I hope authorities are actively seeking to determine whether P. austrocedrii is present in nurseries or natural systems in other parts of the country.

The genus Phytophthora includes many serious plant pathogens, from the one that caused the disastrous potato blight of Ireland (Phytophthora infestans) to globally important forest-destroying invasive species, e.g., P. cinnamomi and “sudden oak death” P. ramorum.

Bregant et al. surveyed 33 small tree, shrub, and herbaceous plant species in 54 sites on the Italian island of Sardinia and the Alps of both northeastern Italy and western Slovenija. Altitudes varied from the valley bottom (700 m) to above tree line (2100 m). Sites included typical forests, riparian ecosystems, and heathlands.

The 360 isolates taken from 397 samples belonged to 17 known Phytophthora species. Some species are widespread and well-known, e.g., P. pseudosyringae. Three isolates belonged to a putative new species described by Bregant et al. – Phytophthora pseudogregata sp. nov. This total of 18 taxa was unexpectedly high. Many of the species are able to cause aerial infections via production of caducous sporangia. These can infect various organs of the plant host: fruits, leaves, shoots, twigs and branches; and cause necrosis and rots. They detected 56 new host–pathogen associations. All are listed, by type of host, in Tables 4 – 6 of the paper.

The surprising diversity and detection of taxa previously described in Australia (see below) illustrate   scientists’ still poor understanding of this genus. They also confirm fears that the global phytosanitary system is unable control intercontinental movement of Phytophthora.

The authors express concern because Alpine and subalpine regions are important hotspots for floral biodiversity. The great variation in altitude, aspect, moisture regimes, etc. – including extreme conditions – results in many different habitats on small spatial scales, with large numbers of both plant species and endemics in very confined spaces. The pathogens they discovered are spreading and compromising the biodiversity of these ecologically fragile habitats.

The authors say their study emphasizes the need to assess the full diversity of Phytophthora species and the factors driving the emergence and local spread of these invasive pathogens. They specify studying the Phytophthora communities on fallen leaves to evaluate host specificity, geographic distribution and survival strategies of the main Phytophthora species detected in this study. They report that scientists are currently mapping the distribution of the new species, P. pseudogregata, in the Alpine habitats and trying to establish its natural host range.

another view of the Julian Alps; photo via Rawpixl

Bregant et al. point out that increased scientific interest over the last 30 years has led to discovery of several previously unknown Phytophthora species and pathogen-host associations. They note that all but two of the taxa in one taxonomic grouping, Sub-clade 6b, have been described in the last 12 years. The majority of taxa have been described from forest ecosystems. This trend is depicted in Figure 8 of the article. This figure also displays which species were isolated from nurseries, agricultural systems, and forest ecosystems.

Results by Plant Type – Disease incidence was highest in shrub vegetation, alpine heathlands and along the mountain riparian systems. The most impacted ecosystems were heathlands dominated by common juniper & blueberry, and riparian systems dominated by alders. In these ecosystems, the Phytophthora-caused outbreaks had reached epidemic levels trend with a high mortality rate. On shrubs and heath formations, disease was initially observed in small areas and progressively spread in a concentric manner affecting more plant species.

Hosts and Diseases – Table 3 in the article lists the 33 host plant species, briefly describes the symptoms, and in some cases provides incidence and mortality rates. Those hosts described as suffering “sudden death” included Alnus viridis, Calluna vulgaris, Genista corsica, Juniperus communis, Lycopodium clavatum, Pinus mugo,Rhododendron ferrugineum, Salix alpine, Vaccinium myrtillus and Vaccinium vitis-idaea

Role of P. pseudosyringae – The most common and widespread species detected was P. pseudosyringae. It constituted more than half of the isolates (201 of the 360). Also, it infected the highest number of hosts (25 out of 33, including all three plant types). It was isolated at 36 of the 54 sites distributed throughout all geographic regions. Seventeen of the host–pathogen associations were new to science. (See Tables 4-6, in the paper.)

Vaccinium myrtillis – a vulnerable host; photo by Tatyana Prozovora via Wikimedia

P. pseudosyringae dominated disease agents in the shrub community, especially among high-altitude shrubs and heaths, e.g., blueberry, dwarf pine, juniper, rhododendron, and alpine willows. Bregant et al. note that these shrubs are extremely low-growing (an adaptation to high elevation conditions). This form might favor attack by Phytophthora sporangia and zoospores present in fallen leaves. Vaccinium myrtillus suffers particularly severe disease – as previously reported in Ireland. In their laboratory studies, Bregant et al. found P. pseudosyringae to be highly aggresse on common juniper (Juniperus communis), producing wood necrosis and shoot blight only four weeks after inoculation.

The importance of P. pseudosyringae in mountainous regions has been found in previous studies in Asia, Europe, and North and South America. However, the authors call for further study of certain aspects of the species. These regard infectivity and survival of the species’ sporangia in infected tissues fallen to the ground; and the ability of oospores to persist for years in environments subject to extreme low temperatures. The former could increase the risk of outbreaks and promote faster disease progression.

The authors suggest P. pseudosyringae’s survival stems from its production of very large and thick-walled chlamydospores. This reported feature is in contradiction with the original species description, which prompts Bregant et al. to call for a correction.

Other Species, Old and New – P. cactorum was the only Phytophthora species other than P. pseudosyringae detected on all three types of hosts (small trees, shrubs, and herbaceous plants). Phytophthora plurivora was the second-most isolated species. It was detected on 12 hosts in 24 sites.

The new putative species — Phytophthora pseudogregata sp. nov. – was detected on Alnus viridis, Juniperus communis, and Rhododendron ferrugineum. As noted above, scientists are now testing whether other plant species are also hosts. It was detected at two sites in Italy — Borso del Grappa and San Nicolò di Comelico; and one site in Slovenija.

Juniperus communis; photo by Joan Simon via Flickr

Diseases of Juniper – Koch’s postulates have been fulfilled, demonstrating that eight Phytophthora species – the new P. pseudogregata sp. nov. as well as P. acerina, P. bilorang, P. gonapodyides, P. plurivora, P. pseudocryptogea, P. pseudosyringae, P. rosacearum – are pathogenic on common juniper (Juniperus communis). The lesions caused by P. pseudosyringae were significantly larger than those caused by other species. Lesions caused by P. pseudosyringae, P. plurivora and acerina progressively girdled the twigs causing shoot blight, browned foliage & wilting symptoms.

Most Threatening Phytophthora clades – The most-frequently isolated Phytophthora species belong mainly to clades 1 and 3 – including P. pseudosyringae. Bregant et al. say these species have several advantages for surviving in mountainous ecosystems: they produce caducous sporangia useful for aerial infections and they tolerate relatively low temperatures. Twoother species in clade 3 were isolated only from the mountains of Sardinia. One, P. psychrophila, was isolated from bleeding cankers on an oak species, Quercus pubescens. Its geographic distribution and impact are still unknown. A second species, P. ilicis, is a well-known pathogen on various hollies in Europe and North America.

Four species belonging to subclade 1a were isolated in the Alps of northeastern Italy and Slovenija. P. cactorum is a widespread polyphagous pathogen found from tropical to temperate climates. It has been responsible for severe diseases on agricultural crops and forest trees. Its occurrence in cold areas has recently been reported in Europe and Australia. The recently described P. alpina has the highest ability to survive in extremely cold conditions. It was detected on four hosts – Alnus viridis, Lonicera alpigena, Vaccinium myrtillus, and V. vitis-idaea.

Some species, e.g., P. hedraiandra and P. idaei, were reported for the first time in natural ecosystems in Europe. They have previously been linked to root and foliar disease in agricultural and ornamental nurseries.

The second-most common species in the Bregant et al. study, P. plurivora, was isolated from 54 symptomatic samples from 12 plant species; eight of the hosts are new. It is common in forest ecosystems of Central Europe – which is now considered to be its region of origin. Little is known about the closely related P. acerina. To date, the latter has been detected widely in agricultural systems, nurseries, forests, and ornamental trees in northern Italy and Sardinia. It is much more rarely found elsewhere. Both P. acerina and P. plurivora are already known to be primary pathogens involved in decline of common and grey alder in Italy.

Five of the Phytophthora species in this study, including the new species P. pseudogregata, are in Clade 6. These include pathogens very common in European forests, e.g., P. bilorbang and P. gonapodyides. Others have more limited or still unknown distributions, e.g., P. amnicola and P. rosacearum. These five species’ ability to cause aerial infections on mountain vegetation might warrant re-evaluation of the reputation of species in this clade being saprophytes or only occasional weak opportunistic pathogens.

P. pseudogregata – in sub-clade 6a – was originally described in 2011 in wet native forests in Australia and on dying alpine heathland vegetation in Tasmania. It has recently been reported in the Czech Republic and Finland. The related P. gibbosa is known to occur only in Australia, where it is associated with dying native vegetation on seasonally wet sites.

Two species of clade 8 — P. kelmanii & P. syringae — have a very limited distribution. A third – P. pseudocryptogea — is widespread in Italian ecosystems from Mediterranean areas to the tree line in the Dolomites. One species from clade 7 (P. cambivora) isolated, mainly from stem bleeding cankers of small trees and shrubs. It has two mating types; bothoccurr in the Alps of northeastern Italy and neighboring Slovenija — on Alnus incana, Laburnum alpinum and Sorbus aucuparia.

SOURCE

Bregant, C., G. Rossetto, L. Meli, N. Sasso, L. Montecchio, A. Brglez, B. Piškur, N. Ogris, L. Maddau, B.T. Linaldeddu. 2024. Diversity of Phytophthora Species Involved in New Diseases of Mountain Vegetation in Europe with the Description of Phytophthora pseudogregata sp. nov. Forests 2023, 14, 1515. https://doi.org/10.3390/f14081515 https://www.mdpi.com/journal/forests

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/

or

www.fadingforests.org