Historical plant introductions predict current insect invasions.

Thousands of insect species have been introduced outside of their native ranges, and some of them strongly impact ecosystems and human societies. Because a large fraction of insects feed on or are associated with plants, nonnative plants provide habitat and resources for invading insects, thereby facilitating their establishment. Furthermore, plant imports represent one of the main pathways for accidental nonnative insect introductions. Here, we tested the hypothesis that plant invasions precede and promote insect invasions. We found that geographical variation in current nonnative insect flows was best explained by nonnative plant flows dating back to 1900 rather than by more recent plant flows. Interestingly, nonnative plant flows were a better predictor of insect invasions than potentially confounding socioeconomic variables. Based on the observed time lag between plant and insect invasions, we estimated that the global insect invasion debt consists of 3,442 region-level introductions, representing a potential increase of 35% of insect invasions. This debt was most important in the Afrotropics, the Neotropics, and Indomalaya, where we expect a 10 to 20-fold increase in discoveries of new nonnative insect species. Overall, our results highlight the strong link between plant and insect invasions and show that limiting the spread of nonnative plants might be key to preventing future invasions of both plants and insects.

Early Monitoring of Forest Wood-Boring Pests with Remote Sensing.

Wood-boring pests (WBPs) pose an enormous threat to global forest ecosystems because their early stage infestations show no visible symptoms and can result in rapid and widespread infestations at later stages, leading to large-scale tree death. Therefore, early-stage WBP detection is crucial for prompt management response. Early detection of WBPs requires advanced and effective methods like remote sensing. This review summarizes the applications of various remote sensing sensors, platforms, and detection methods for monitoring WBP infestations. The current capabilities, gaps in capabilities, and future potential for the accurate and rapid detection of WBPs are highlighted.

Global rise in emerging alien species results from increased accessibility of new source pools.

Our ability to predict the identity of future invasive alien species is largely based upon knowledge of prior invasion history. Emerging alien species-those never encountered as aliens before-therefore pose a significant challenge to biosecurity interventions worldwide. Understanding their temporal trends, origins, and the drivers of their spread is pivotal to improving prevention and risk assessment tools. Here, we use a database of 45,984 first records of 16,019 established alien species to investigate the temporal dynamics of occurrences of emerging alien species worldwide. Even after many centuries of invasions the rate of emergence of new alien species is still high: One-quarter of first records during 2000-2005 were of species that had not been previously recorded anywhere as alien, though with large variation across taxa. Model results show that the high proportion of emerging alien species cannot be solely explained by increases in well-known drivers such as the amount of imported commodities from historically important source regions. Instead, these dynamics reflect the incorporation of new regions into the pool of potential alien species, likely as a consequence of expanding trade networks and environmental change. This process compensates for the depletion of the historically important source species pool through successive invasions. We estimate that 1-16% of all species on Earth, depending on the taxonomic group, qualify as potential alien species. These results suggest that there remains a high proportion of emerging alien species we have yet to encounter, with future impacts that are difficult to predict.

Return of the moth: rethinking the effect of climate on insect outbreaks.

The sudden interruption of recurring larch budmoth (LBM; Zeiraphera diniana or griseana Gn.) outbreaks across the European Alps after 1982 was surprising, because populations had regularly oscillated every 8-9 years for the past 1200 years or more. Although ecophysiological evidence was limited and underlying processes remained uncertain, climate change has been indicated as a possible driver of this disruption. An unexpected, recent return of LBM population peaks in 2017 and 2018 provides insight into this insect's climate sensitivity. Here, we combine meteorological and dendrochronological data to explore the influence of temperature variation and atmospheric circulation on cyclic LBM outbreaks since the early 1950s. Anomalous cold European winters, associated with a persistent negative phase of the North Atlantic Oscillation, coincide with four consecutive epidemics between 1953 and 1982, and any of three warming-induced mechanisms could explain the system's failure thereafter: (1) high egg mortality, (2) asynchrony between egg hatch and foliage growth, and (3) upward shifts of outbreak epicentres. In demonstrating that LBM populations continued to oscillate every 8-9 years at sub-outbreak levels, this study emphasizes the relevance of winter temperatures on trophic interactions between insects and their host trees, as well as the importance of separating natural from anthropogenic climate forcing on population behaviour.

Deciphering the worldwide invasion of the Asian long-horned beetle: A recurrent invasion process from the native area together with a bridgehead effect.

Retracing introduction routes is crucial for understanding the evolutionary processes involved in an invasion, as well as for highlighting the invasion history of a species at the global scale. The Asian long-horned beetle (ALB) Anoplophora glabripennis is a xylophagous pest native to Asia and invasive in North America and Europe. It is responsible for severe losses of urban trees, in both its native and invaded ranges. Based on historical and genetic data, several hypotheses have been formulated concerning its invasion history, including the possibility of multiple introductions from the native zone and secondary dispersal within the invaded areas, but none have been formally tested. In this study, we characterized the genetic structure of ALB in both its native and invaded ranges using microsatellites. In order to test different invasion scenarios, we used an approximate Bayesian "random forest" algorithm together with traditional population genetics approaches. The strong population differentiation observed in the native area was not geographically structured, suggesting complex migration events that were probably human-mediated. Both native and invasive populations had low genetic diversity, but this characteristic did not prevent the success of the ALB invasions. Our results highlight the complexity of invasion pathways for insect pests. Specifically, our findings indicate that invasive species might be repeatedly introduced from their native range, and they emphasize the importance of multiple, human-mediated introductions in successful invasions. Finally, our results demonstrate that invasive species can spread across continents following a bridgehead path, in which an invasive population may have acted as a source for another invasion.

Developing a list of invasive alien species likely to threaten biodiversity and ecosystems in the European Union.

The European Union (EU) has recently published its first list of invasive alien species (IAS) of EU concern to which current legislation must apply. The list comprises species known to pose great threats to biodiversity and needs to be maintained and updated. Horizon scanning is seen as critical to identify the most threatening potential IAS that do not yet occur in Europe to be subsequently risk assessed for future listing. Accordingly, we present a systematic consensus horizon scanning procedure to derive a ranked list of potential IAS likely to arrive, establish, spread and have an impact on biodiversity in the region over the next decade. The approach is unique in the continental scale examined, the breadth of taxonomic groups and environments considered, and the methods and data sources used. International experts were brought together to address five broad thematic groups of potential IAS. For each thematic group the experts first independently assembled lists of potential IAS not yet established in the EU but potentially threatening biodiversity if introduced. Experts were asked to score the species within their thematic group for their separate likelihoods of i) arrival, ii) establishment, iii) spread, and iv) magnitude of the potential negative impact on biodiversity within the EU. Experts then convened for a 2-day workshop applying consensus methods to compile a ranked list of potential IAS. From an initial working list of 329 species, a list of 66 species not yet established in the EU that were considered to be very high (8 species), high (40 species) or medium (18 species) risk species was derived. Here, we present these species highlighting the potential negative impacts and the most likely biogeographic regions to be affected by these potential IAS.

Are traded forest tree seeds a potential source of nonnative pests?

The international seed trade is considered relatively safe from a phytosanitary point of view and is therefore less regulated than trade in other plants for planting. However, the pests carried by traded seeds are not well known. We assessed insects and fungi in 58 traded seed lots of 11 gymnosperm and angiosperm tree species from North America, Europe, and Asia. Insects were detected by X-raying and molecular methods. The fungal community was characterized using high-throughput sequencing (HTS) and by growing fungi on non-selective agar. About 30% of the seed lots contained insect larvae. Gymnosperms contained mostly hymenopteran (Megastigmus spp.) and dipteran (Cecidomyiidae) larvae, while angiosperms contained lepidopteran (Cydia latiferreana) and coleopteran (Curculio spp.) larvae. HTS indicated the presence of fungi in all seed lots and fungi grew on non-selective agar from 96% of the seed lots. Fungal abundance and diversity were much higher than insect diversity and abundance, especially in angiosperm seeds. Almost 50% of all fungal exact sequence variants (ESVs) found in angiosperms were potential pathogens, in comparison with around 30% of potentially pathogenic ESVs found in gymnosperms. The results of this study indicate that seeds may pose a greater risk of pest introduction than previously believed or accounted for. A rapid risk assessment suggests that only a small number of species identified in this study is of phytosanitary concern. However, more research is needed to enable better risk assessment, especially to increase knowledge about the potential for transmission of fungi to seedlings and the host range and impact of identified species.

Processionary Moths and Associated Urtication Risk: Global Change-Driven Effects.

Processionary moths carry urticating setae, which cause health problems in humans and other warm-blooded animals. The pine processionary moth Thaumetopoea pityocampa has responded to global change (climate warming and increased global trade) by extending its distribution range. The subfamily Thaumetopoeinae consists of approximately 100 species. An important question is whether other processionary moth species will similarly respond to these specific dimensions of global change and thus introduce health hazards into new areas. We describe, for the first time, how setae are distributed on different life stages (adult, larva) of major groups within the subfamily. Using the available data, we conclude that there is little evidence that processionary moths as a group will behave like T. pityocampa and expand their distributional range. The health problems caused by setae strongly relate to population density, which may, or may not, be connected to global change.

IgE sensitization to Thaumetopoea pityocampa : diagnostic utility of a setae extract, clinical picture and associated risk factors.

BACKGROUND: Setae from Thaumetopoea pityocampa larvae (the pine processionary moth or PPM) can induce hypersensitivity reactions, but their clinical role in IgE-mediated responses is still subject to discussion. The aim of this study was to evaluate a setae extract for in vivo and in vitro diagnosis in nonhospitalized patients with reactions to PPM. METHODS: Forty-eight adult patients presenting with PPM cutaneous reactions were studied by skin prick test (SPT) and specific IgE using setae and whole larval (WL) extracts. Biological standardized extracts were used for skin tests. RESULTS: A total of 47.9% patients had a positive SPT for PPM (70% to both extracts, 17% only to the WL extract and 13% only to the setae extract). IgE immunoblotting detected several reactive bands in 91% of the SPT-positive cases. In multivariate analysis, male sex, immediate latency (<1 h) and duration of skin symptoms (<24 h) were independent predictors of a positive SPT. CONCLUSIONS: IgE sensitization to PPM was found in 48% of the study patients, which was associated with immediate reactions and evanescent cutaneous lesions. Most of these patients reacted to both WL and setae extracts, but some reacted to only one of them. According to our data, skin and in vitro tests to PPM should be performed using both extracts.

Socioeconomic legacy yields an invasion debt.

Globalization and economic growth are widely recognized as important drivers of biological invasions. Consequently, there is an increasing need for governments to address the role of international trade in their strategies to prevent species introductions. However, many of the most problematic alien species are not recent arrivals but were introduced several decades ago. Hence, current patterns of alien-species richness may better reflect historical rather than contemporary human activities, a phenomenon which might be called "invasion debt." Here, we show that across 10 taxonomic groups (vascular plants, bryophytes, fungi, birds, mammals, reptiles, amphibians, fish, terrestrial insects, and aquatic invertebrates) in 28 European countries, current numbers of alien species established in the wild are indeed more closely related to indicators of socioeconomic activity from the year 1900 than to those from 2000, although the majority of species introductions occurred during the second half of the 20th century. The strength of the historical signal varies among taxonomic groups, with those possessing good capabilities for dispersal (birds, insects) more strongly associated with recent socioeconomic drivers. Nevertheless, our results suggest a considerable historical legacy for the majority of the taxa analyzed. The consequences of the current high levels of socioeconomic activity on the extent of biological invasions will thus probably not be completely realized until several decades into the future.

Improving the efficiency of lepidopteran pest detection and surveillance: constraints and opportunities for multiple-species trapping.

Surveillance using attractants for invasive species can allow early detection of new incursions and provide decision support to response programs. Simultaneous trapping for multiple species, by baiting traps with several lures, is expected to increase the number of species that can be targeted in surveillance programs and improve the cost-effectiveness without affecting surveillance coverage. We tested this hypothesis by choosing four potential forest and urban lepidopteran pest species that are present in Europe but not yet in New Zealand and many other countries. We deployed traps in central and southern Europe with single lures or all possible species combinations (up to four lures per trap). There was only limited interference, apparently due to trap saturation, but no evidence for interspecific repellency among lures for gypsy moth, Lymantria dispar, fall webworm, Hyphantria cunea, pine processionary moth, Thaumetopoea pityocampa, and pine shoot moth, Rhyacionia buoliana. To assess what factors may be important in species compatibility/suitability for multiple-species trapping, we combined our results with those of previous studies conducted by the United States Department of Agriculture. For 75 combinations of pheromones, tested singly or in combination, 19 % showed no effect on trap catch for any of the species tested. In the other cases, either one or both species showed a reduction in trap catch. However, few lure combinations caused complete or nearly complete suppression. For most combinations, catches were still sufficiently high for detection purposes. Species from the same superfamily exhibited more interference than more distantly related species. Together, these results suggest that there are opportunities to improve the range of exotic pests under surveillance, at little additional cost, by multiple-species trapping for which compatibility has been demonstrated.

Disentangling the role of environmental and human pressures on biological invasions across Europe.

The accelerating rates of international trade, travel, and transport in the latter half of the twentieth century have led to the progressive mixing of biota from across the world and the number of species introduced to new regions continues to increase. The importance of biogeographic, climatic, economic, and demographic factors as drivers of this trend is increasingly being realized but as yet there is no consensus regarding their relative importance. Whereas little may be done to mitigate the effects of geography and climate on invasions, a wider range of options may exist to moderate the impacts of economic and demographic drivers. Here we use the most recent data available from Europe to partition between macroecological, economic, and demographic variables the variation in alien species richness of bryophytes, fungi, vascular plants, terrestrial insects, aquatic invertebrates, fish, amphibians, reptiles, birds, and mammals. Only national wealth and human population density were statistically significant predictors in the majority of models when analyzed jointly with climate, geography, and land cover. The economic and demographic variables reflect the intensity of human activities and integrate the effect of factors that directly determine the outcome of invasion such as propagule pressure, pathways of introduction, eutrophication, and the intensity of anthropogenic disturbance. The strong influence of economic and demographic variables on the levels of invasion by alien species demonstrates that future solutions to the problem of biological invasions at a national scale lie in mitigating the negative environmental consequences of human activities that generate wealth and by promoting more sustainable population growth.

Setae from the pine processionary moth (Thaumetopoea pityocampa) contain several relevant allergens.

BACKGROUND: Pine processionary larvae produce urticating hairs (setae) that serve for protection against predators. Setae induce cutaneous reactions in animals and humans. The presence of toxic or allergic mechanisms is a matter of debate. OBJECTIVES: To detect the presence of allergens in setae and to characterize them. MATERIALS AND METHODS: Setae extracts were characterized by gel staining and immunoblot, with sera from patients with immediate reactions and positive prick test reactions, as well as a rabbit antiserum raised against setae. Setae proteins were fractionated by high-performance liquid chromatography. The most relevant allergen was analysed by matrix-assisted laser desorption/ionization (MALDI) mass spectrometry (MS), and its sequence was deduced from an expressed sequence tag bank. Results. Setae contained at least seven different allergens. The most intense detection corresponded to a protein of MW ~ 14,000 that was similar to thaumetopoein, a previously described protein with mast cell-degranulating properties. MALDI-MS-based de novo sequencing provided a partial amino acid sequence different from that of the previously described allergen Tha p 1, and it was named Tha p 2. This allergen was detected in 61% of patients, and it is therefore a new major caterpillar allergen. CONCLUSIONS: Penetration of the setae from the pine processionary caterpillar delivers their allergenic content in addition to causing mechanical or toxic injury.

New species and new records of exotic Scolytinae (Coleoptera, Curculionidae) in Europe.

Background: Bark and ambrosia beetles (Coleoptera, Scolytinae) are amongst the most important wood-boring insects introduced to Europe. During field investigations conducted between 2019 and 2021 in different countries and regions of Europe, many exotic species have been recorded providing new and relevant data. New information: Dryoxylononoharaense (Murayama, 1933) is recorded in Europe for the first time. Xyleborinusattenuatus (Blandford, 1894) is a species new to Italy, while Xylosandrusgermanus (Blandford, 1894), Hypothenemuseruditus (Westwood, 1836) and Amasa sp. near A.truncata are new country records for Portugal. Cnestusmutilatus (Blandford, 1894), Phloeotribusliminaris (Harris, 1852) were collected in Italy and Amasa sp. near A.truncata was collected in France after the first discovery, confirming their establishment and their dispersal into new areas.

Human exposure to larvae of processionary moths in France: study of symptomatic cases registered by the French poison control centres between 2012 and 2019.

CONTEXT: Contact with the setae of larvae (caterpillars) of pine or oak processionary moths could induce urticarial or allergic reactions in humans. These species are present in France and presently expanding towards highly populated areas due to climate change and/or human-mediated translocations. We aimed to describe the symptomatic cases of exposure to processionary larvae in France. METHODS: We conducted a retrospective study of symptomatic cases of exposure to processionary larvae registered by the French poison control centres between 1 January 2012 and 31 July 2019. We reviewed all medical records coded with the agent "larvae". RESULTS: Of the 1274 included cases, 59% and 27% corresponded to pine and oak processionary larvae, respectively; the 14% remaining cases concerned unspecified processionary larvae. While the annual number of cases due to pine processionary larvae fluctuated during the study period, cases associated with oak processionary larvae increased steadily. Most of the annual cases occurred between January and May for pine processionary larvae, and April and August for oak processionary larvae (with a peak in March or June, respectively). Among the 1022 cases for which information was available, the sex ratio was 1.2 and the median age was 11 years old. Skin symptoms were reported by 96,3% of the cases, such as pruritus or urticaria. The severity was mild in 96.3% of cases, moderate in 3.5%, and severe in 0.2% (two cases). Ocular or oral exposures led more frequently to severe symptoms than dermal ones (respectively 31% and 18% vs. 2% of cases, p < 10-3). CONCLUSION: Since processionary moth larvae exposure is a growing health concern, which can cause severe injuries particularly after ocular or oral exposures, the population, and the professionals should be informed of existing recommendations to avoid exposure and measures to be taken after being exposed.

Prevalence of cutaneous reactions to the pine processionary moth (Thaumetopoea pityocampa) in an adult population.

BACKGROUND: Thaumetopoea pityocampa [pine processionary moth (PPM)] is one of the most important lepidopteran agents causing urticant cutaneous reactions in humans in Mediterranean countries. This species is also expanding northwards, because of global warming. OBJECTIVES: To investigate the prevalence, distribution by habitat group and possible risk factors of PPM cutaneous reactions in adults. METHODS: A randomly designed survey was carried out on 1224 adults. RESULTS: A point prevalence, estimated after corrections, of 8.7% was obtained (12% rural areas, 9.6% for semi-urban areas, and 4.4% for urban areas). The data showed a significantly higher risk of self-reported symptoms according to sex [p < 0.005; males, adjusted odds ratio (aOR) 1.84], habitat (p < 0.0005; rural, aOR 1.8; semi-urban, aOR 1.2), frequency of visits to pinewood areas (p < 0.005; daily exposure, aOR 2.1), and occupational exposure (p < 0.0001; aOR 5.04, 90% were males). Airborne contamination was the most important cause of reactions (83.3% of 48 participants who visited the hospital and fulfilled the criteria for a convincing reaction presented with symptoms after walking on/passing by pine tree areas). CONCLUSIONS: These findings show that PPM cutaneous reactions are common in this southern European population, including peripheral urban areas, and that the main risk is related to exposure to this insect.

Invasion History of Sirex noctilio Based on COI Sequence: The First Six Years in China.

Sirex noctilio F. (Hymenoptera: Siricidae: Siricinae), a new invasive species in China, is a significant international forestry pest which, transported via logs and related wood packing materials, has led to environmental damage and substantial economic loss in many countries around the world. It was first detected in China in 2013, and since then infestations have been found in 18 additional sites. Using a 322 bp fragment of the mitochondrial barcode gene COI, we studied the genetic diversity and structure of S. noctilio populations in both native and invaded ranges, with a specific focus in China. Twelve haplotypes were found across the native and invaded distribution of the pest, of which three were dominant; among these there were only one or two mutational steps between each pair of haplotypes. No obvious genetic structure was found other than in Chinese populations. China has a unique and dominant haplotype not found elsewhere, and compared with the rest of the world, the genetic structure of Chinese populations suggested a multiple invasion scenario.

Bark beetles and pinhole borers recently or newly introduced to France (Coleoptera: Curculionidae, Scolytinae and Platypodinae).

We present an annotated list of 11 Scolytinae and Platypodinae species newly or recently introduced to France. Four species are recorded for the first time as interceptions: Euplatypus hintzi (Schaufuss), Euplatypus parallelus (Fabricius), Xyleborus affinis Eichhoff and Xyleborus ferrugineus (Fabricius). Two are possibly naturalised: Xyleborus bispinatus Eichhoff and Cryphalus dilutus Eichhoff, while Cyclorhipidion distinguendum (Eggers) and Xyloterinus politus (Say) are confirmed as species newly established in Europe. Moreover, an unidentified species of Amasa Lea, collected previously in Spain, is recorded for the first time in France: Amasa sp. near truncata (Erichson). We point out that literature references to Amasa truncata as an invasive species in New Zealand and South America are incorrect, as the photographs of these non-native populations do not match the holotype of A. truncata. For each species we have updated its global distribution, detailed all French records, and summarized biology, ecology, host trees and potential risks as pests.

Diapause Regulation in Newly Invaded Environments: Termination Timing Allows Matching Novel Climatic Constraints in the Box Tree Moth, Cydalima perspectalis (Lepidoptera: Crambidae).

The association between indirect environmental cues that modulate insect diapause and the actual stressors is by no means granted when a species encounters new environments. The box tree moth, Cydalima perspectalis, is an Asian pest whose rapid invasion in Europe causes considerable economic and ecological impacts. Larvae enter a winter diapause induced by the photoperiod in both native and invaded ranges, but factors that trigger the return to an active phase are still unknown. Yet, identifying them is crucial to understand how diapause end synchronizes with the end of the winter stress encountered in Europe. To test whether activity resumption is regulated by thermal and/or photoperiodic thresholds, or additive effects between these factors often involved in diapause termination, diapausing caterpillars from an invaded area were exposed to crossed treatments at the laboratory. The evolution of diapause rate was monitored over time and compared to that of nearby field sites invaded. A strong positive effect of increasing temperature was found on the rate and dynamics of diapause termination, whereas no compelling effect of photoperiod appeared. Resuming development directly when main stressors fade, not in response to indirect photoperiodic cues that could be mismatched outside native areas, likely contributes to the good match observed between diapause and the new climates that this pest encountered in the invaded range.

Quaternary history and contemporary patterns in a currently expanding species.

BACKGROUND: Quaternary climatic oscillations had dramatic effects on species evolution. In northern latitudes, populations had to survive the coldest periods in refugial areas and recurrently colonized northern regions during interglacials. Such a history usually results in a loss of genetic diversity. Populations that did not experience glaciations, in contrast, probably maintained most of their ancestral genetic diversity. These characteristics dramatically affected the present-day distribution of genetic diversity and may influence the ability of species to cope with the current global changes. We conducted a range-wide study of mitochondrial genetic diversity in the pine processionary moth (Thaumetopoea pityocampa/T. wilkinsoni complex, Notodontidae), a forest pest occurring around the Mediterranean Basin and in southern Europe. This species is responding to the current climate change by rapid natural range expansion and can also be accidentally transported by humans. Our aim was to assess if Quaternary climatic oscillations had a different effect across the species' range and to determine if genetic footprints of contemporary processes can be identified in areas of recent introduction. RESULTS: We identified three main clades that were spatially structured. In most of Europe, the genetic diversity pattern was typical for species that experienced marked glaciation cycles. Except in refugia, European populations were characterized by the occurrence of one main haplotype and by a strong reduction in genetic diversity, which is expected in regions that were rapidly re-colonized when climatic conditions improved. In contrast, all other sub-clades around the Mediterranean Basin occurred in limited parts of the range and were strongly structured in space, as is expected in regions in which the impact of glaciations was limited. In such places, genetic diversity was retained in most populations, and almost all haplotypes were endemic. This pattern was extreme on remote Mediterranean islands (Crete, Cyprus, Corsica) where highly differentiated, endemic haplotypes were found. Recent introductions were typified by the existence of closely-related haplotypes in geographically distant populations, which is difficult to detect in most of Europe because of a lack of overall genetic structure. CONCLUSION: In regions that were not prone to marked glaciations, recent moth introductions/expansions could be detected due to the existence of a strong spatial genetic structure. In contrast, in regions that experienced the most intense Quaternary climatic oscillations, the natural populations are not genetically structured, and contemporary patterns of population expansion remain undetected.