# Worldwide tests of generic attractants, a promising tool for early detection of non-native cerambycid species

Alain Roques, Lili Ren, Davide Rassati, Juan Shi, Evgueni Akulov, Neil

Audsley, Marie-Anne Auger-Rozenberg, Dimitrios Avtzis, Andrea Battisti,
Richard Bellanger, et al.

► To cite this version:

Alain Roques, Lili Ren, Davide Rassati, Juan Shi, Evgueni Akulov, et al. .. Worldwide tests of generic attractants, a promising tool for early detection of non-native cerambycid species. NeoBiota, 2023, 84, pp.169-209. 10.3897/neobiota.84.91096 . hal-04171539

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Worldwide tests of generic attractants,
a promising tool for early detection
of non-native cerambycid species

Alain Roques¹,², Lili Ren²,³, Davide Rassati⁴, Juan Shi²,³, Evgueni Akulov⁵,
Neil Audsley⁶, Marie-Anne Auger-Rozenberg¹,², Dimitrios Avtzis⁷, Andrea Battisti⁴,
Richard Bellanger⁸, Alexis Bernard¹, Iris Bernadinelli⁹, Manuela Branco¹⁰,
Giacomo Cavaletto⁴, Christian Cocquempot¹¹, Mario Contarini¹²,
Béatrice Courtial¹, Claudine Courtin¹, Olivier Denux¹, Miloň Dvořák¹³,
Jian-ting Fan¹⁴, Nina Feddern¹⁵, Joseph Francese¹⁶, Emily K. L. Franzen¹⁷,¹⁸,
André Garcia¹⁰, Georgi Georgiev¹⁹, Margarita Georgieva¹⁹, Federica Giarruzzo¹²,
Martin Gossner¹⁵, Louis Gross¹, Daniele Guarneri²⁰, Gernot Hoch²¹,
Doris Hölling¹⁵, Mats Jonsell²², Natalia Kirichenko²³,²⁴, Antoon Loomans²⁵,
You-qing Luo²,³, Deborah McCullough²⁶, Craig Maddox²⁷, Emmanuelle Magnoux¹,
Matteo Marchioro⁴, Petr Martinek¹³, Hugo Mas²⁸, Bruno Mériguet²⁹,
Yong-zhi Pan³⁰, Régis Phélut¹, Patrick Pineau¹, Ann M. Ray¹⁷, Olivier Roques¹,
Marie-Cécile Ruiz³¹, Victor Sarto i Monteys³², Stefano Speranza¹², Jiang-hua Sun²,³³,
Jon D. Sweeney³⁴, Julien Touroult³⁵, Lionel Valladares³⁶, Loïs Veillat¹, Yuan Yuan²,³,
Myron P. Zalucki³⁷, Yunfan Zou³⁸, Alenka Žunič-Kosi³⁹,
Lawrence M. Hanks⁴⁰, Jocelyn G. Millar³⁸

1 INRAE URZF, 45075, Orléans, France 2 IFOPE, Sino-French Joint Laboratory for Invasive Forest Pests in Eurasia, INRAE URZF and Beijing Forestry University, Orléans, France 3 Beijing Key Laboratory for Forest Pest Control, College of Forestry, Beijing Forestry University, Beijing, China 4 Department of Agronomy, Food, Natural Resources, Animals and Environment (DAFNAE), University of Padua, Padova, Italy 5 Russian Plant Quarantine Center, Krasnoyarsk Branch, Krasnoyarsk 660075, Russia 6 Fera Science Ltd., Sand Hutton, York, YO41 1LZ, UK, United Kingdom 7 Forest Research Institute, Hellenic Agricultural Organization Demeter, Thessaloniki 57006, Greece 8 INRAE UEVT, 06160 Antibes Juan les Pins, France 9 Plant Health and Research Service – ERSA, Via Sabbatini 5, 33050 Pozzuolo Del Friuli (UD), Italy 10 Forest Research Center (CEF), School of Agriculture (ISA), University of Lisbon, Lisbon, Portugal 11 55 rue du Questel, 29640 Plougonven, France 12 Department of Agriculture and Forest Sciences, University of Tuscia, Viterbo, Via S. Camillo de Lellis, 01100, Italy 13 Department of Forest Protection and Wildlife Management, Faculty of Forestry and Wood Technology, Mendel University in Brno, Czech Republic 14 School of Forestry and Biotechnology, Zhejiang Agriculture and Forestry University, Lin'an, China 15 Forest Health and Biotic Interactions, Swiss Federal Research Institute WSL, Zürcherstrasse 111, CH-8903 Birmensdorf, Switzerland 16 USDA APHIS PPQ S&T, Forest Pest Methods Laboratory, Buzzards Bay, MA 02542, USA 17 Department of Biology, Xavier University, Cincinnati, OH 45207, USA 18 USDA APHIS PPQ S&T, Bethel, OH 45106, USA 19 Forest Research Institute, Bulgarian Academy of Sciences, Sofia, Bulgaria 20 Ente Parco Nazionale del
Circeo, 04016 Sabaudia, Italy 21 BFW – Austrian Research Centre for Forests, Seckendorff-Gudent-Weg 8, 1131 Vienna, Austria 22 Department of Ecology, Swedish University of Agricultural Sciences, Box 7044, Uppsala 750 07, Sweden 23 Sukachev Institute of Forest, Siberian Branch of Russian Academy of Sciences, Federal Research Center «Krasnoyarsk Science Center SB RAS», Krasnoyarsk 660036, Russia 24 Siberian Federal University, Krasnoyarsk 660041, Russia 25 Netherlands Food and Consumer Product Safety Authority, Division Agriculture and Nature (NPPO) National Reference Centre, Geertjesweg 15, Wageningen, Netherlands 26 Department of Entomology and Dept. of Forestry, Michigan State University, East Lansing MI, 48824, USA 27 NSW Department of Primary Industries, Wollongbar & Centre for Tropical Horticulture, Alstonville NSW 2477, Australia 28 Laboratori de Sanitat Forestal – CIEF VAERSA- Generalitat Valenciana, 46930 Quart de Poblet (València), Spain 29 Office pour les Insectes et leur Environnement (OPIE), 78041 Guyancourt, France 30 Southwest Forestry College, Kunming, China 31 Office pour l'Environnement de la Corse (OEC), Corte, France 32 Institut de Ciència i Tecnologia Ambientals (ICTA), Entomology, Plants and Health, Universitat Autònoma de Barcelona, 08193 Bellaterra, Spain 33 College of Life Science, Institute of Life Science and Green Development, Hebei University, Baoding 071002, China 34 Natural Resources Canada, Canadian Forest Service, Atlantic Forestry Center, Fredericton, NB, E3C2G6, Canada 35 PatriNat (OFB, CNRS, MNHN), CP41, 36 rue Geoffroy Saint-Hilaire, Paris, France 36 INP Purpan, Toulouse University, Toulouse, France 37 School of Biological Sciences, The University of Queensland, Brisbane, 4072, Australia 38 Departments of Entomology and Chemistry, University of California, Riverside, CA 92521, USA 39 Department of Organisms and Ecosystems Research, National Institute of Biology, 1000 Ljubljana, Slovenia 40 Department of Entomology, University of Illinois at Urbana-Champaign, Urbana, IL 61801, USA

Corresponding author: Alain Roques (alain.roques@inrae.fr)

Academic editor: Marc Kenis | Received 31 July 2022 | Accepted 18 November 2022 | Published 18 May 2023

**Citation:** Roques A, Ren L, Rassati D, Shi J, Akulov E, Audsley N, Auger-Rozenberg M-A, Avtzis D, Battisti A, Bellanger R, Bernard A, Bernadinelli I, Branco M, Cavaletto G, Cocquempot C, Contarini M, Courtial B, Courtin C, Denux O, Dvořák M, Fan J-t, Feddern N, Francese J, Franzen EKL, Garcia A, Georgiev G, Georgieva M, Giarruzzo F, Gossner M, Gross L, Guarneri D, Hoch G, Hölling D, Jonsell M, Kirichenko N, Loomans A, Luo Y-q, McCullough D, Maddox C, Magnoux E, Marchioro M, Martinek P, Mas H, Mériguet B, Pan Y-z, Phélut R, Pineau P, Ray AM, Roques O, Ruiz M-C, Sarto i Monteys V, Speranza S, Sun J-h, Sweeney JD, Touroult J, Valladares L, Veillat L, Yuan Y, Zalucki MP, Zou Y, Žunič-Kosi A, Hanks LM, Millar JG (2023) Worldwide tests of generic attractants, a promising tool for early detection of non-native cerambycid species. In: Jactel H, Orazio C, Robinet C, Douma JC, Santini A, Battisti A, Branco M, Seehausen L, Kenis M (Eds) Conceptual and technical innovations to better manage invasions of alien pests and pathogens in forests. NeoBiota 84: 169–209. https://doi.org/10.3897/neobiota.84.91096

## Abstract

A large proportion of the insects which have invaded new regions and countries are emerging species, being found for the first time outside their native range. Being able to detect such species upon arrival at ports of entry before they establish in non-native countries is an urgent challenge. The deployment of traps baited with broad-spectrum semiochemical lures at ports-of-entry and other high-risk sites could be one such early detection tool. Rapid progress in the identification of semiochemicals for cerambycid beetles during the last 15 years has revealed that aggregation-sex pheromones and sex pheromones are often conserved at global levels for genera, tribes or subfamilies of the Cerambycidae. This possibly allows the development of generic attractants which attract multiple species simultaneously, especially when such
pheromones are combined into blends. Here, we present the results of a worldwide field trial programme conducted during 2018–2021, using traps baited with a standardised 8-pheromone blend, usually complemented with plant volatiles. A total of 1308 traps were deployed at 302 sites covering simultaneously or sequentially 13 European countries, 10 Chinese provinces and some regions of the USA, Canada, Australia, Russia (Siberia) and the Caribbean (Martinique). We intended to test the following hypotheses: 1) if a species is regularly trapped in significant numbers by the blend on a continent, it increases the probability that it can be detected when it arrives in other countries/continents and 2) if the blend exerts an effective, generic attraction to multiple species, it is likely that previously unknown and unexpected species can be captured due to the high degree of conservation of pheromone structures within related taxa. A total of 78,321 longhorned beetles were trapped, representing 376 species from eight subfamilies, with 84 species captured in numbers greater than 50 individuals. Captures comprised 60 tribes, with 10 tribes including more than nine species trapped on different continents. Some invasive species were captured in both the native and invaded continents. This demonstrates the potential of multipheromone lures as effective tools for the detection of ‘unexpected’ cerambycid invaders, accidentally translocated outside their native ranges. Adding new pheromones with analogous well-conserved motifs is discussed, as well as the limitations of using such blends, especially for some cerambycid taxa which may be more attracted by the trap colour or other characteristics rather than to the chemical blend.

### Keywords

Cerambycidae, early detection, Holarctic, invasion, multi-pheromone blend, pheromone trapping

## Introduction

During the last several decades, the unprecedented development of worldwide trade has resulted in increasing translocation and establishment of non-native insects outside their native ranges, with little evidence of saturation (Seebens et al. 2017, 2021). Insect herbivores, accidentally introduced as plant contaminants, appear to be mainly responsible for this sharp increase, at least in Europe (Roques 2010; Pergl et al. 2017). Amongst these non-native herbivores, species associated with woody plants largely dominate, accounting for 76.5% of all herbivore species newly recorded in Europe from 2000 to 2014, while species of importance to agricultural plants and products are a minority (Roques et al. 2016). The increased extent of trade in ornamental plants has been suggested as a major driver of this increase (Liebhold et al. 2012; Eschen et al. 2014; Essl et al. 2015; Roques et al. 2020). Additionally, wood packaging material (e.g. pallets, crating, dunnage) transported with international cargo shipments represents another significant pathway for introduction of non-native phloem- and wood-boring insects (Aukema et al. 2010; Haack et al. 2014; Lovett et al. 2016). An average of 6.1 non-native insect species attacking woody plants became newly established in Europe per year from 2000–2019, compared to 2.4 cases per year from 1950–1970 (Roques et al. 2020). Similar trends were observed in North America (Aukema et al. 2010), New Zealand (Brockerhoff and Liebhold 2017) and at a slower rate in China (Roques et al. 2020).
Another key attribute of this recently-arrived, non-native entomofauna is the increasing presence of “emerging” species, which have not been reported previously as invaders and are not considered to be pests in their native ranges. Arrival of these species probably results from evolving changes in trade routes and imported goods, which leads to accessibility to new pools of species (Seebens et al. 2018). For example, the emerald ash borer, *Agrilus planipennis* Fairmaire, was not considered a significant pest until it invaded North America, where it has caused massive damage (Dang et al. 2022). The same is true for a number of other xylophagous cerambycid beetle species which have recently invaded Europe, such as the Asian mulberry longhorned beetle, *Xylotrechus chinensis* (Chevrolat) (Sarto i Monteys and Torras i Tutusaus 2018), the round-headed apple-tree borer, *Saperda candida* Fabricius (Nolte and Krieger 2008) and the Asian redneck longhorned beetle *Aromia bungii* (Faldermann) (Russo et al. 2020). At first, such species were typically not subject to regulatory measures or strict phytosanitary inspections at borders because their invasive potential had not been recognised. For example, only seven of the 117 non-native insect species that infest woody plants that established in Europe during the period 1995–2012 had been intercepted in such inspections (Eschen et al. 2015). In Australia, 61 of the 135 non-native species established in forests during the period 2003–2016 had never been intercepted, despite relatively intensive border controls (Nahrung and Carnegie 2021). Therefore, the development of new strategies to detect such unanticipated and unregulated species as early as possible is essential to implement rapid and effective eradication or containment measures (Nahrung et al. 2023).

Deployment of traps baited with broad-spectrum semiochemical lures at ports-of-entry (Brockerhoff et al. 2006; Rassati et al. 2014, 2015a; Hoch et al. 2020) or other high-risk sites (e.g. urban wood-waste landfills and industrial sites, Rassati et al. 2015b; Rabaglia et al. 2019) could be one such early detection tool. Given the difficulty of predicting which species may arrive and in what numbers (i.e. propagule pressure), such lures should be efficient even at low population densities and should ideally attract multiple species from different taxa (family, subfamily, tribe). Combining pheromones of several species into blends could be expected to result in such a generic attraction when antagonistic effects amongst blend components are relatively minor, for example, reduced attraction of relatively few species, such that the net effect of blending multiple components is an increase in the number of target taxa detected. The addition of plant volatiles, acting as kairomones, may further enhance the attraction. For instance, a pine specialist, *Monochamus galloprovincialis* (Olivier), was significantly more attracted when its pheromone, monochamol, was combined with volatiles from its pine hosts (Alvarez et al. 2016). Similarly, ethanol had a synergist effect on the capture of species related to broadleaved trees in Eurasia (*Phymatodes testaceus* [L.]; Sweeney et al. 2014; Fan et al. 2019) and in southern USA (Miller et al. 2017). However, the addition of plant volatiles did not affect, either positively or negatively, the captures of several other cerambycid species (Fan et al. 2019). Overall, relationships between host volatiles and cerambycids are probably more dependent on the exploited host and less on insect taxonomy. Potential for using blended lures for detection would be further
enhanced if each component of the blend was attractive to multiple related species, i.e. a pheromone or kairomone shared by species within a genus or tribe as occurs in the longhorned beetle family Cerambycidae.

This large family of Coleoptera includes between 34,000 and 38,000 described species (Rossa and Goczał 2021; Tavakilian and Chevillotte 2022). Although recent molecular studies using a multigene approach revealed that the phylogeny at the upper taxonomic levels is not completely resolved and still under debate (Lee and Lee 2020; Nie et al. 2020), Tavakilian and Chevillotte (2022) recognised 13 subfamilies. The subfamily Lamiinae is by far the most diverse with more than 21,000 species, 3,002 genera and 86 tribes, followed by Cerambycinae (> 12,000 species, 1,848 genera, and 119 tribes), Lepturinae (> 1,830 species, 232 genera, 11 tribes), Prioninae (> 1,250 species, 311 genera, 26 tribes) and Spondylidinae (> 150 species, 32 genera, seven tribes); other subfamilies are smaller and much less diverse. Cerambycid larvae of many species develop as endophytic borers concealed beneath the bark of woody plants or, much less frequently, within herbaceous plants. This cryptic lifestyle, coupled with the usual long duration of the hidden larval stages, facilitates the transport of these insects around the world in logs and wooden packing materials (Eyre and Haack 2017), but also via trade in living plants if the plants have a sufficiently large diameter. For example, larvae of the citrus longhorned beetle, *Anoplophora chinensis* (Forster), were detected in Japanese maples, *Acer palmatum* Thunb., shipped to Europe (Eschen et al. 2015). Thus, a steadily increasing number of cerambycid species have become globally important as invasive forest and orchard pests (Venette and Hutchison 2021).

Recent advances in the chemical ecology of cerambycids and, particularly, the identification of volatile pheromones that act as long-range attractants, have provided new tools and opportunities for monitoring invasive woodborers. In total, pheromones or likely pheromones have been identified for more than 400 cerambycid species worldwide (Millar and Hanks 2017). Furthermore, field experiments have shown that these pheromones can be deployed in blends, with a potential generic attraction for both native and non-native species (Hanks et al. 2012; Hanks and Millar 2016; Hanks et al. 2018; Fan et al. 2019; Flaherty et al. 2019; Rassati et al. 2019). Currently, the aggregated data suggest that species in the subfamilies Cerambycinae, Lamiinae and Spondylidinae use male-produced aggregation-sex pheromones to attract both sexes, whereas species in the subfamilies Prioninae and Lepturinae use female-produced pheromones that attract only males (Hanks and Millar 2016). This research has revealed striking patterns in pheromone chemistry. Pheromone components are frequently highly conserved amongst species within genera, tribes and even at the subfamily level (Hanks and Millar 2013, 2016). For example, in the subfamily Lamiinae, hydroxyethers are used as aggregation-sex pheromones by many species native to different continents. Thus, 2-(undecyloxy)ethanol, or monochamol, is a pheromone component shared by European, North American and Asian species in the genus *Monochamus*, all of which vector the pine wood nematode (*Bursaphelenchus xylophilus* [Steiner & Buhrer]) (Pajares et al. 2010; Hanks and Millar 2016; Boone et al. 2018; Lee et al. 2018). In addition, field trials in southern China showed that four lamiine species in genera other
than *Monochamus* were attracted to monochamol (Wickham et al. 2014). A number of other compounds are widely shared amongst species within a given subfamily in different world regions. For example, terpenoids such as fuscumol ([E]-6,10-dimethyl-5,9-undecadien-2-ol) and its acetate, are aggregation sex-pheromone components for many species in the subfamily Spondylidinae and Laminae (Mitchell et al. 2011; Hanks and Millar 2016). In contrast, many species in the subfamily Cerambycinae from different continents utilise short-chain (6–10 carbon) hydroxyketones, such as 3-hydroxyalkan-2-ones and 2-hydroxyalkan-3-ones and the corresponding *syn*- and *anti*-2,3-alkanediols as aggregation-sex pheromones (Hanks and Millar 2016). Prionic acid ([3R,5S]-3,5-dimethyldodecanoic acid) similarly appears to be shared as a sex pheromone by several genera of the subfamily Prioninae on different continents (Barbour et al. 2011; Wickham et al. 2016a). This sharing of pheromone components by species in different world regions suggests that traps baited with these compounds have a good chance of detecting non-native, phylogenetically-related invaders that are introduced to another continent. Moreover, combining several of these pheromone components in a single blend has the potential to detect a broader range of species.

During the last 10 years, the generic effectiveness of such multi-component blends has been tested on different continents, but using different pheromone combinations, either alone or in combination with kairomones, such as ethanol and α-pinene (e.g. Miller et al. 2017; Fan et al. 2019). In Illinois, USA, Hanks et al. (2012) first tested a six-component blend, which included racemic 3-hydroxyhexan-2-one, *syn*- and *anti*-2,3-hexanediols, fuscumol, fuscumol acetate, monochamol and racemic 2-methylbutan-1-ol. Ten cerambycid species were caught in significant numbers in these trials, including four species in the subfamily Cerambycinae and six in the subfamily Lamiinae. Hanks et al. (2018) then tested this 6-component blend at a larger scale in several regions of the USA, adding both prionic acid and plant volatiles to the traps. The pheromone blend attracted about twice as many species as any of the individual components and the species attracted by the blend included three subfamilies, whereas individual components attracted species within only one subfamily. The inclusion of prionic acid also resulted in the additional captures of *Prionus* spp. which were not trapped by the previous six-pheromone blend. In a natural reserve in Yunnan (China), Wickham et al. (2021) trapped 71 species with another generic lure comprised of six components, three of which were the same as those used in the USA (*anti*-2,3-hexanediol, racemic 3-hydroxyhexan-2-one and monochamol). In France, using an 8-pheromone blend consisting of the same compounds as Hanks et al. (2018) to which was added geranylacetone targeting Spondylininae (Halloran et al. 2018), Fan et al. (2019) trapped 118 species, of which 114 were native species that represented 48% of the French cerambycid fauna. Trapping more than 50% of the species in 25 of the 41 cerambycid tribes present in the country indicates a considerable generic attraction of this 8-pheromone blend, significantly higher than an earlier trial which tested a blend of four pheromones. By contrast, unbaited control traps deployed in the same French sites caught very few species. Other trials of potentially generic blends, including fewer or different
compounds, were carried out in Russia (Sweeney et al. 2014), Australia (Hayes et al. 2016), Brazil (Silva et al. 2017), Poland, Italy and Canada (Flaherty et al. 2019; Rassati et al. 2019, 2021). Results from Australia differed from those reported in other continents because the tested blend attracted no more species than 3-hydroxyhexan-2-one alone (Hayes et al. 2016).

When using multi-pheromone blends, antagonistic effects might occur with either pheromone components or host plant volatiles (e.g. Hanks et al. 2018; Rassati et al. 2021). The North American species *Neoclytus acuminatus acuminatus* (F.), for example, was strongly attracted by *syn*-2,3-hexanediol, but the addition of racemic 3-hydroxyhexan-2-one to the latter pheromone interrupted attraction (Rassati et al. 2021). Addition of host plant volatiles, such as ethanol, significantly enhanced attraction of some cerambycid species (Sweeney et al. 2014; Miller et al. 2017; Hanks et al. 2018), but, with the exception of *P. testaceus*, had little effect on catch of cerambycid species in other studies (Fan et al. 2019). However, as long as inhibition did not completely prevent attraction, one trap with a multi-pheromone lure may still be somewhat more cost-effective than deploying multiple traps baited with individual lures. This can be assessed by a cost-benefit analysis, i.e. estimating the labour and materials costs of deploying and servicing a network of traps baited with single components, versus the costs of deploying and servicing a single trap baited with a blend of the same components.

Results of these different experiments on various continents stimulated us to propose a worldwide trapping programme using a standardised 'generic' 8-pheromone blend in all countries/trapping sites. The blend included the following compounds known to be widely shared amongst cerambycids of related taxa: fuscumol, fuscumol acetate, monochamol, geranylacetone, *anti*-2,3-hexanediol, 3-hydroxyhexan-2-one (C6-ketol), 2-methylbutan-1-ol and prionic acid. The programme relied on the following hypotheses: 1) if a species is attracted in significant numbers by the blend in a region, it increases the probability that it can be detected when it arrives at ports-of-entry in other regions and 2) if the blend exerts an effective, generic attraction to multiple species, it is likely that previously unknown and unexpected species can be captured due to the high degree of conservation of pheromone structures within related taxa, as described above. Our overarching objective was to build a global database of cerambycid species trapped by the 8-pheromone blend. To this end, field trials were conducted during 2018–2021 using operational protocols that were standardised as much as possible at all sites worldwide to cover simultaneously or sequentially 13 European countries, 10 Chinese provinces and some regions of the USA, Canada, Australia, Russia (Siberia) and the Caribbean. Over the course of the study, we also tested the possibility of adding new compounds to enlarge the pool of species trapped. Therefore, in 2020, two additional pheromones, the sex-aggregation pheromones trichoferone (a hydroxyketone pheromone of the velvet longhorned beetle, *Trichoferus campestris* (Faldermann) (Ray et al. 2019) and (*E*)-2-*cis*-6,7-epoxynonenal, the pheromone of the invasive species *A. bungii* (Xu et al. 2017), were added to the original 8-pheromone blend and tested in France and China. In addition, ethanol and α-pinene were included in most trials as synergists for some cerambycids.
# Materials and methods

## Study sites

The successive or parallel development of three European research projects (HOMED, MULTITRAP, SAMFIX) and two French projects (CANOPEE, PORTRAP) during 2018–2021 allowed us to carry out field trials at 302 sites distributed as follows: 244 in Europe (164 in France, 22 in Italy, 13 in Spain and Switzerland, 6 in Portugal, 5 in Austria and England, 4 in Greece and Slovenia, 3 in the Netherlands, 2 in Bulgaria and the Czech Republic and 1 in Sweden), 38 in Asia (35 in China and three in Siberia, Russia), 11 in North America (10 in the USA and one in Canada), five in the Caribbean (Martinique) and four in Australia (see Table 1 and Suppl. material 1 for details per country, coordinates and the relevant research project). A total of 1308 traps were deployed in stands of broad-leaved and/or coniferous trees in natural or managed environments, but also within and nearby potential ports-of-entry (maritime and fluvial ports, airports, national markets). Experiments in these latter sites usually included two traps placed on trees planted within the port and two traps placed in woody areas located within a 1 km-radius from the port, except in 2019 when a larger experiment was carried out (see below).

In 2018, trials were limited to four European countries (Austria, England, France, the Netherlands), including 41 sites with 143 traps. The 2019 trials were much more extensive and involved 12 European countries (the four from 2018, supplemented by Bulgaria, the Czech Republic, Greece, Italy, Portugal, Spain, Sweden and Switzerland), five provinces of China (Beijing, Hebei, Liaoning, Yunnan and Zhejiang), two States of the USA (Michigan and Ohio) and one site in Canada (Nova Scotia), resulting in a total of 79 sites and 626 traps. These 2019 trials included a large trapping programme targeting semi-urban forests located close to ports-of-entry in Europe, USA and Canada where 16 (Czech Republic, Portugal, Sweden) or 32 traps (France, Italy, Nova Scotia, Ohio, Switzerland) were deployed at each target site. The 2020 trials were substantially impacted by the COVID-19 pandemic, but were carried out for at least a part of the spring-summer season in six European countries (France, Greece, Italy, Portugal, Spain and Switzerland), eight provinces of China (those of 2019, except Beijing, to which were added Hunan, Inner Mongolia, Jiangxi and Shandong) and extended to Australia (New South Wales) and the Caribbean (Martinique), resulting in a total of 78 sites and 256 traps. The 2021 trials were deployed in the same countries as in 2020, supplemented by an additional European country (Slovenia), Russia (Siberia) and an additional province of China (Gansu), resulting in a total of 104 sites and 283 traps.

## Trapping protocol and 8-pheromone blend

Trials at all sites used either multifunnel or cross-vane panel traps supplied by different companies depending on the country (Econex, Spain; ChemTica Internacional, S.A., Heredia, Costa Rica; Alpha Scents Inc., West Linn, Oregon, USA). Cross-vane traps used in Italy (Colli Euganei area) in 2019 were hand-made (see Cavaletto et al. 2021 for
details). Black traps were generally deployed, but other colours were also used in France, Italy, Nova Scotia, Ohio and Switzerland (see Table 1). To improve trapping efficiency, all traps were coated with Fluon (AGC Chemicals Europe Ltd., Thornton Cleveleys, UK) diluted in 1:6 in water (Graham et al. 2010). In forests, traps were usually hung from tree branches or between two trees in the lower canopy, at ~ 3–5 m high. Exceptions are some of the countries involved in the 2019 trapping programme targeting semi-urban forests (i.e. Czech Republic, France, Canada, Italy, Portugal, Sweden, Switzerland and the USA) where traps were placed both in the understory and in the upper canopy (> 20 m), the trial carried out in Italy (Colli Euganei area) in 2019 where traps were placed at 5–7 m above the ground and the trappings carried out in some forests of north-central France during 2019–2021 where traps were placed in the upper canopy (> 20 m). Each trap was separated from the next by 50 m at least and traps were preferentially placed at the forest edge. In ports-of-entry, the traps were attached to branches of available trees, at least 2 m above ground with a minimum distance between traps of 100 m. A similar design was used for the traps placed within the 1 km-radius from the ports-of-entry. Duration of trap deployments were variable amongst sites and years, but in the Northern Hemisphere, experiments were mostly conducted from mid-April at the earliest to mid-October at the latest, except in 2020 when the COVID-19 pandemic delayed the onset of trapping until mid-June or mid-July. In Australia, traps were deployed from December to March, depending on the year. Detailed trap heights and trapping duration by site are provided in the Suppl. material 1.

All lures were prepared by INRAE before being shipped to all study participants. These lures consisted of a blend designed by Fan et al. (2019), which contains eight cerambycid pheromones (fuscumol, fuscumol acetate, monochamol, 3-hydroxyhexan-2-one, *anti*-2,3-hexanediol and 2-methylbutan-1-ol, all at 50 mg/ml; geranylacetone-25 mg/ml; and prionic acid-0.5 mg/ml; Table 2) dissolved in isopropanol as a carrier to a total volume of 1 ml per lure. The blend composition was expected to attract a large number of cerambycid subfamilies and tribes according to Hanks et al. (2012), (Table 2). Dispensers consisted of a cotton dental pad (to serve as a reservoir and stabilise release rate) placed into a polyethylene sachet (Minigrip, 4 cm × 6 cm × 60 µ; Dutscher, Brumath, France) and dosed with 1 ml of the lure solution. The release rate was estimated by Fan et al. (2019) as 0.0263 ± 0.002 g/d under 20 °C conditions. A dispenser was hung in the centre of each trap.

All primary compounds were obtained from ChemTica Internacional, except prionic acid, which was purchased from Alpha Scents Inc. Commercial high release rate ethanol (100 ml dose, 96% purity, release rate 2 g/day at 20 °C; Econex, Spain) and α-pinene lures (25 ml dose, 98% purity, release rate 0.3 g/day at 20 °C; Econex, Spain) were added to traps in most trials (1076 of the 1308 traps; Table 1). These compounds are known to synergise attraction of some cerambycid species to their pheromones (e.g. Allison et al. 2012; Ryall et al. 2015; Collignon et al. 2016; Miller et al. 2017), but are mildly repellent for others (Collignon et al. 2016). Trap catches were tabulated every 3–4 wk, at which time the pheromone lures were replaced. The ethanol UHR and α-pinene lures, when added, were replaced every 6 wk.
<table><caption>**Table 1.** Summary of the trapping design per country from 2018 to 2021. Research project: C: CANOPEE; H: HOMED; M: MULTITRAP; P: PORTRAP; S: SAMFIX. Blend: #8: 8-pheromone blend; #8+ET: 8-pheromone blend + ethanol UHR; #8+AP+ET: 8-pheromone blend + α-pinene + ethanol UHR; #10+AP+ET: 10- pheromone blend + α-pinene + ethanol UHR. Trap type: MF: multifunnel; CV: crossvane. Trap colour: B: black; G: Green; BG: Black base and green top; P: Purple; Y: Fluorescent yellow; others: brown, blue, red, grey (corresponding to data collected by Cavaletto et al. 2021). Collection type: W: wet; D: dry. Note that the sites could be similar in a country from one year to the next.</caption><thead><tr><th>Region</th><th>Year</th><th>Country/ Province</th><th>Project</th><th>No Sites</th><th>No Traps</th><th>#8</th><th>#8+ET</th><th>Blend</th><th>#8+AP+ET</th><th>#10+AP+ET</th><th>Trap type</th><th>MF</th><th>CV</th><th>B</th><th>BG</th><th>P</th><th>Y</th><th>Other</th><th>W</th><th>Collection Type</th></tr></thead><tbody><tr><td>Europe</td><td>2018</td><td>Austria</td><td>M</td><td>3</td><td>20</td><td>10</td><td>0</td><td>10</td><td>10</td><td>0</td><td>10</td><td>10</td><td>20</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>20</td><td></td></tr><tr><td>Europe</td><td>2018</td><td>England</td><td>M</td><td>3</td><td>8</td><td>4</td><td>0</td><td>4</td><td>4</td><td>0</td><td>6</td><td>2</td><td>8</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Europe</td><td>2018</td><td>France</td><td>M,P</td><td>32</td><td>97</td><td>18</td><td>0</td><td>79</td><td>79</td><td>0</td><td>41</td><td>56</td><td>78</td><td>7</td><td>7</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>97</td></tr><tr><td>Europe</td><td>2018</td><td>Netherlands</td><td>M</td><td>3</td><td>18</td><td>6</td><td>0</td><td>12</td><td>12</td><td>0</td><td>9</td><td>9</td><td>9</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>18</td><td>0</td><td></td></tr><tr><td>Europe</td><td>2019</td><td>Austria</td><td>H</td><td>2</td><td>4</td><td>0</td><td>0</td><td>4</td><td>4</td><td>0</td><td>4</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td><td></td></tr><tr><td>Europe</td><td>2019</td><td>Bulgaria</td><td>H</td><td>2</td><td>3</td><td>0</td><td>0</td><td>3</td><td>3</td><td>0</td><td>3</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td><td></td></tr><tr><td>Europe</td><td>2019</td><td>Czech Rep</td><td>H</td><td>2</td><td>32</td><td>0</td><td>0</td><td>32</td><td>32</td><td>0</td><td>32</td><td>32</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>32</td><td>0</td><td></td></tr><tr><td>Europe</td><td>2019</td><td>England</td><td>H</td><td>2</td><td>4</td><td>2</td><td>0</td><td>2</td><td>2</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td><td></td></tr><tr><td>Europe</td><td>2019</td><td>France</td><td>C,H,P,S</td><td>26</td><td>170</td><td>0</td><td>0</td><td>170</td><td>0</td><td>164</td><td>6</td><td>89</td><td>59</td><td>13</td><td>9</td><td>0</td><td>0</td><td>0</td><td>88</td><td>82</td><td></td></tr><tr><td>Europe</td><td>2019</td><td>Greece</td><td>H</td><td>2</td><td>3</td><td>0</td><td>0</td><td>3</td><td>3</td><td>0</td><td>3</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td><td></td></tr><tr><td>Europe</td><td>2019</td><td>Italy</td><td>H,S</td><td>19</td><td>192</td><td>0</td><td>128</td><td>64</td><td>0</td><td>64</td><td>128</td><td>48</td><td>48</td><td>0</td><td>16</td><td>16</td><td>64</td><td>192</td><td>0</td><td></td><td></td></tr><tr><td>Europe</td><td>2019</td><td>Portugal</td><td>H</td><td>2</td><td>32</td><td>0</td><td>0</td><td>32</td><td>32</td><td>0</td><td>32</td><td>32</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>32</td><td>0</td><td></td><td></td></tr><tr><td>Europe</td><td>2019</td><td>Spain</td><td>H</td><td>1</td><td>2</td><td>0</td><td>0</td><td>2</td><td>2</td><td>2</td><td>2</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td><td></td><td></td></tr><tr><td>Europe</td><td>2019</td><td>Sweden</td><td>H</td><td>1</td><td>16</td><td>0</td><td>0</td><td>16</td><td>0</td><td>16</td><td>0</td><td>16</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>16</td><td>0</td><td></td></tr><tr><td>Europe</td><td>2019</td><td>Switzerland</td><td>H</td><td>2</td><td>64</td><td>0</td><td>0</td><td>64</td><td>0</td><td>64</td><td>32</td><td>32</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>64</td><td>0</td><td></td></tr><tr><td>Europe</td><td>2020</td><td>France</td><td>C,H,P,S</td><td>48</td><td>166</td><td>2</td><td>0</td><td>64</td><td>100</td><td>160</td><td>6</td><td>77</td><td>35</td><td>17</td><td>17</td><td>0</td><td>0</td><td>0</td><td>18</td><td>148</td><td></td></tr><tr><td>Europe</td><td>2020</td><td>Greece</td><td>H</td><td>1</td><td>1</td><td>0</td><td>0</td><td>1</td><td>1</td><td>1</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td></td></tr><tr><td>Europe</td><td>2020</td><td>Italy</td><td>H</td><td>2</td><td>8</td><td>0</td><td>0</td><td>8</td><td>8</td><td>0</td><td>8</td><td>2</td><td>4</td><td>0</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td><td></td></tr><tr><td>Europe</td><td>2020</td><td>Portugal</td><td>H</td><td>2</td><td>8</td><td>0</td><td>0</td><td>8</td><td>0</td><td>8</td><td>0</td><td>4</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td><td></td></tr><tr><td>Europe</td><td>2020</td><td>Spain</td><td>H</td><td>6</td><td>22</td><td>0</td><td>0</td><td>22</td><td>0</td><td>22</td><td>0</td><td>22</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>22</td><td></td></tr><tr><td>Europe</td><td>2020</td><td>Switzerland</td><td>H</td><td>2</td><td>4</td><td>0</td><td>0</td><td>4</td><td>0</td><td>4</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td><td>0</td><td></td></tr><tr><td>Europe</td><td>2021</td><td>France</td><td>C,H,P,S</td><td>58</td><td>165</td><td>2</td><td>0</td><td>58</td><td>105</td><td>162</td><td>3</td><td>98</td><td>42</td><td>0</td><td>14</td><td>11</td><td>0</td><td>0</td><td>30</td><td>135</td><td></td></tr><tr><td>Europe</td><td>2021</td><td>Greece</td><td>H</td><td>1</td><td>1</td><td>0</td><td>0</td><td>1</td><td>1</td><td>1</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td></td></tr><tr><td>Europe</td><td>2021</td><td>Italy</td><td>H</td><td>1</td><td>4</td><td>0</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>0</td><td>2</td><td>0</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td><td></td></tr><tr><td>Europe</td><td>2021</td><td>Portugal</td><td>H</td><td>2</td><td>8</td><td>0</td><td>0</td><td>8</td><td>8</td><td>0</td><td>8</td><td>4</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td><td></td></tr><tr><td>Europe</td><td>2021</td><td>Slovenia</td><td>H</td><td>4</td><td>18</td><td>0</td><td>0</td><td>18</td><td>0</td><td>18</td><td>0</td><td>18</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>18</td><td></td></tr><tr><td>Europe</td><td>2021</td><td>Spain</td><td>H</td><td>6</td><td>16</td><td>0</td><td>0</td><td>16</td><td>0</td><td>16</td><td>0</td><td>12</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>16</td><td></td></tr><tr><td>Europe</td><td>2021</td><td>Switzerland</td><td>H</td><td>9</td><td>19</td><td>8</td><td>0</td><td>11</td><td>0</td><td>19</td><td>0</td><td>8</td><td>11</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>11</td><td>8</td><td></td></tr><tr><td>Europe</td><td>Total</td><td></td><td></td><td>244</td><td>1105</td><td>52</td><td>128</td><td>720</td><td>205</td><td>885</td><td>220</td><td>631</td><td>252</td><td>37</td><td>63</td><td>44</td><td>64</td><td>505</td><td>600</td><td></td><td></td></tr><tr><td>Asia</td><td>2019</td><td>China/Beijing</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td><td></td></tr></tbody></table>
<table><thead><tr><th>Region</th><th>Year</th><th>Country/Province</th><th>Project</th><th>No Sites</th><th>No Traps</th><th>#8</th><th>#8+ET</th><th>Blend</th><th>Trap type</th><th>MF</th><th>CV</th><th>B</th><th>G</th><th>BG</th><th>P</th><th>Y</th><th>Other</th><th>W</th><th>Collection Type</th></tr></thead><tbody><tr><td>Asia</td><td>2019</td><td>China/Hebei</td><td>H</td><td>2</td><td>6</td><td>0</td><td>0</td><td>6</td><td>6</td><td>0</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Asia</td><td>2019</td><td>China/InnerMongolia</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2019</td><td>China/Laoning</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2019</td><td>China/Yunnan</td><td>H</td><td>1</td><td>4</td><td>0</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Asia</td><td>2019</td><td>China/Zhejiang</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2020</td><td>China/Hebei</td><td>H</td><td>4</td><td>12</td><td>0</td><td>0</td><td>12</td><td>0</td><td>12</td><td>0</td><td>12</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>12</td></tr><tr><td>Asia</td><td>2020</td><td>China/Hunan</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2020</td><td>China/Inner Mongolia</td><td>H</td><td>2</td><td>6</td><td>0</td><td>0</td><td>6</td><td>0</td><td>6</td><td>0</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Asia</td><td>2020</td><td>China/Liangxi</td><td>H</td><td>2</td><td>6</td><td>0</td><td>0</td><td>6</td><td>0</td><td>6</td><td>0</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Asia</td><td>2020</td><td>China/Laoning</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2020</td><td>China/Shandong</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2020</td><td>China/Yunnan</td><td>H</td><td>1</td><td>4</td><td>0</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Asia</td><td>2020</td><td>China/Zhejiang</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2021</td><td>China/Beijing</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2021</td><td>China/Gansu</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2021</td><td>China/Hebei</td><td>H</td><td>2</td><td>6</td><td>0</td><td>0</td><td>6</td><td>0</td><td>6</td><td>0</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Asia</td><td>2021</td><td>China/Hunan</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2021</td><td>China/Inner Mongolia</td><td>H</td><td>2</td><td>6</td><td>0</td><td>0</td><td>3</td><td>3</td><td>6</td><td>0</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Asia</td><td>2021</td><td>China/Jiangxi</td><td>H</td><td>2</td><td>6</td><td>0</td><td>0</td><td>6</td><td>0</td><td>6</td><td>0</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Asia</td><td>2021</td><td>China/Lioaning</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2021</td><td>China/Shandong</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2021</td><td>China/Yunnan</td><td>H</td><td>3</td><td>5</td><td>0</td><td>0</td><td>0</td><td>3</td><td>5</td><td>0</td><td>5</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Asia</td><td>2021</td><td>China/Zhejiang</td><td>H</td><td>1</td><td>3</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>2021</td><td>Russia/Siberia</td><td>H</td><td>3</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Asia</td><td>Total</td><td></td><td></td><td></td><td>38</td><td>106</td><td>3</td><td>0</td><td>92</td><td>11</td><td>106</td><td>0</td><td>106</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>106</td></tr><tr><td>North America</td><td>2019</td><td>USA/ Michigan</td><td>H</td><td>9</td><td>18</td><td>18</td><td>0</td><td>0</td><td>0</td><td>0</td><td>18</td><td>0</td><td>18</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>18</td><td>0</td></tr><tr><td>North America</td><td>2019</td><td>USA/ Ohio</td><td>H</td><td>1</td><td>32</td><td>0</td><td>0</td><td>32</td><td>0</td><td>32</td><td>0</td><td>32</td><td>0</td><td>16</td><td>16</td><td>0</td><td>0</td><td>0</td><td>32</td><td>0</td></tr><tr><td>North America</td><td>2019</td><td>Canada/ Nova Scotia</td><td>H</td><td>1</td><td>32</td><td>0</td><td>0</td><td>32</td><td>0</td><td>32</td><td>0</td><td>32</td><td>0</td><td>16</td><td>16</td><td>0</td><td>0</td><td>0</td><td>32</td><td>0</td></tr><tr><td>North America</td><td>Total</td><td></td><td></td><td></td><td>11</td><td>82</td><td>18</td><td>0</td><td>64</td><td>0</td><td>82</td><td>0</td><td>50</td><td>32</td><td>0</td><td>0</td><td>0</td><td>0</td><td>82</td><td>0</td></tr><tr><td>The Caribbean</td><td>2020</td><td>France/ Martinique</td><td>H</td><td>2</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>The Caribbean</td><td>2021</td><td>France/ Martinique</td><td>H</td><td>3</td><td>4</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>The Caribbean</td><td>Total</td><td></td><td></td><td></td><td>5</td><td>7</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td><td>0</td><td>7</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td></tr><tr><td>Australia</td><td>2020</td><td>Australia</td><td>H</td><td>2</td><td>4</td><td>0</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Australia</td><td>2021</td><td>Australia</td><td>H</td><td>2</td><td>4</td><td>0</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Australia</td><td>Total</td><td></td><td></td><td></td><td>4</td><td>8</td><td>0</td><td>0</td><td>8</td><td>0</td><td>8</td><td>0</td><td>8</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Grand total</td><td></td><td></td><td></td><td>302</td><td>1308</td><td>80</td><td>128</td><td>884</td><td>216</td><td>1088</td><td>220</td><td>802</td><td>284</td><td>37</td><td>63</td><td>44</td><td>64</td><td>587</td><td>721</td></tr></tbody></table>
<table><caption>**Table 2.** Composition of the 8-pheromone and 10-pheromone blends and targeted sex and cerambycid tribes.</caption><thead><tr><th rowspan="2">Blend</th><th rowspan="2">Compound</th><th rowspan="2">Amount/lure (mg/ml)</th><th rowspan="2">Target Sex</th><th colspan="3">Target subfamily</th><th rowspan="2">Target tribe/ genus</th><th rowspan="2">References</th></tr><tr><th>Cerambycinae</th><th>Lamiinae</th><th>Aseminae</th><th>Prioninae</th></tr></thead><tbody><tr><td rowspan="2">8-pheromones</td><td rowspan="2">Racemic 3-hydroxyhexan-2-one (C6-ketol)</td><td rowspan="2">50</td><td rowspan="2">M/F</td><td>X</td><td></td><td></td><td>Callidiini</td><td>Millar et al. (2018)</td></tr><tr><td>X</td><td></td><td></td><td>Clytini</td><td>Hanks and Millar (2013), Wickham et al. (2014), Bobadoye et al. (2019)</td></tr><tr><td rowspan="2">8-pheromones</td><td rowspan="2">Racemic 2-methylbutan-1-ol</td><td rowspan="2">50</td><td rowspan="2">M/F</td><td>X</td><td></td><td></td><td>Hesperophanini</td><td>unpub data JGM</td></tr><tr><td>X</td><td></td><td></td><td>Hylotrupini</td><td>Reddy et al. (2005)</td></tr><tr><td>8-pheromones</td><td>Racemic fuscumol + fuscumol acetate</td><td>50+ 50</td><td>M/F</td><td>X</td><td></td><td></td><td>Callidiini</td><td>Hanks et al. (2018)</td></tr><tr><td rowspan="3">8-pheromones</td><td rowspan="3">2R*,3S*-2,3-hexanediol</td><td rowspan="3">50</td><td rowspan="3">M/F</td><td>X</td><td></td><td></td><td>Clytini</td><td>Hanks and Millar (2013), Wickham et al. (2014)</td></tr><tr><td>X</td><td></td><td></td><td>Obriini</td><td>Millar et al. (2018)</td></tr><tr><td>X</td><td></td><td></td><td>Acanthocinini</td><td>Millar et al. (2018)</td></tr><tr><td rowspan="2">8-pheromones</td><td rowspan="2">Monochamol</td><td rowspan="2">50</td><td rowspan="2">M/F</td><td>X</td><td></td><td></td><td>Acanthoderini</td><td>Hanks and Millar (2013)</td></tr><tr><td>X</td><td></td><td></td><td>Asemini</td><td>Millar et al. (2018)</td></tr><tr><td rowspan="2">8-pheromones</td><td rowspan="2">Geranylacetone</td><td rowspan="2">25</td><td rowspan="2">M/F</td><td>X</td><td></td><td></td><td>Monochamini</td><td>Hanks et al. (2018)</td></tr><tr><td>X</td><td></td><td></td><td>Lamiini</td><td>Wickham et al. (2014)</td></tr><tr><td rowspan="2">8-pheromones</td><td rowspan="2">Prionic acid (4 stereoisomers)</td><td rowspan="2">05</td><td rowspan="2">M</td><td>X</td><td></td><td></td><td>Acanthocinini</td><td>Meier et al. (2016, 2019)</td></tr><tr><td>X</td><td></td><td></td><td>Asemini</td><td>Halloran et al. (2018)</td></tr><tr><td>10-pheromones</td><td>Racemic trichoferone</td><td>25</td><td>M/F</td><td>X</td><td></td><td></td><td>Trichoferus</td><td>Ray et al. (2019)</td></tr><tr><td>10-pheromones</td><td>(E)-2-cis-6,7-epoxynonalen</td><td>50</td><td>M/F</td><td>X</td><td></td><td></td><td>Aromia</td><td>Xu et al. (2017)</td></tr></tbody></table>

In most cases, the trapped insects were killed using a section of mesh impregnated with α-cypermethrin insecticide (Storanet, BASF Pflanzenschutz Deutschland, Germany) placed into the trap basins, whose bottoms had been replaced with a wire mesh to allow drainage and to keep specimens dry. However, in the targeted 2019 experiment in forests near ports-of-entry and in the Colli Euganei area (Italy), “wet” trap basins were used, containing water-diluted propylene glycol (50%) to act as a surfactant and preservative. In the trials conducted in Ohio and Michigan, trap collection cups were filled with ~ 200–400 ml of undiluted propylene glycol.

Trapped cerambycids were identified to species by local specialists or sent to IN-RAE for identification. However, specimens trapped in Australia could not be sent due to restrictions by the customs agency and so most could only be identified to the genus level. Nomenclature used in this article follows the reference checklist of the world database Titan (Tavakilian and Chevillotte 2022).

## Preliminary tests of a 10-pheromone blend

In 2020 and 2021, two additional pheromones, trichoferone (the pheromone of *T. campestris*) and (E)-2-cis-6,7-epoxynonalen (the pheromone of *A. bungii*), were added to the 8-pheromone lures used in France and China, to test for a possible increase in
monitoring effectiveness with a 10-pheromone blend (Table 2). Both compounds were synthesised by YFZ and JGM at the University of California, Riverside, using previously-reported syntheses (Ray et al. 2019; Xu et al. 2017, respectively). Lure preparation and insect collection procedures were similar to those described above. Captures were compared with those of the 8-pheromone blend at six sites in France during 2020 by deploying five pairs of traps baited with each blend at each site. The traps were spaced ~100 m apart and rotated at each insect collection, which enabled the number of collection dates at each site to be used as replicates. The Student t-test for paired samples was then applied to compare the number of species trapped by each lure.

## Results

A total of 78,321 longhorned beetles were trapped, representing 376 species, including 373 Cerambycidae, two Vesperidae and one Disteniidae species (Table 3). The cerambycids belonged to eight subfamilies, including 156 species of Cerambycinae, 102 species of Lamiinae, 78 species of Lepturinae, 21 species of Spondylidinae, 12 species of Prioninae, two species of Necydalinae and one species of Parandrinae (Fig. 1). Captures comprised 60 tribes, with 10 tribes including more than nine species trapped on different continents; in decreasing order the tribe Clytini (64 spp.), followed by Lepturini (44 spp.), Rhagiini (32 spp.), Acanthocinini (31 spp.), Callidiini (20 spp.), Monochamini (18 spp.), Saperdini (10 spp.) and Aseminii, Pogonocherini and Prionini (nine spp.).

<figure><img src="image_2.png" /><figcaption>**Figure 1.** Number of species trapped per subfamily and their region of origin.</figcaption></figure>
each; Fig. 2). Generally, fewer species were trapped in the Caribbean and Australia, where only a limited number of traps had been deployed. Some of the captured species belonged to tribes other than those targeted, such as Callidiopini (*Curtomerus flavus* [F.] in Martinique and *Bethelium* sp. in Australia), Eburiini (*Eburia* spp. in Martinique) and Tillomorphini (*Gourbeyrella madininae* Chalumeau & Touroult in Martinique).

Most tribes included species from the same genera trapped on different continents (Fig. 2). For example, 19 species of the Clytini genus *Xylotrechus* were captured, including 10 in Asia, five in Europe and four in North America. In the same tribe, 12 species of *Chlorophorus* were captured, of which eight were caught in Europe and four in Asia. A total of 12 *Monochamus* species (Lamiinae, Monochamini) were trapped, including five species in Asia, four in North America and three in Europe. The Callidiini genus *Phymatodes* was represented by 11 species, including seven in Europe and four in North America. A number of these species had not been trapped before by any semiochemically-baited trap (e.g. *X. chinensis*, *Chlorophorus glabromaculatus* [Goeze] and *Phymatodes pusillus* [F.]).

<figure><img src="image_3.png" /><figcaption>**Figure 2.** Number of species trapped per tribe in decreasing order and their native region. Only the tribes where > 3 spp were captured are shown.</figcaption></figure>
<table><caption>**Table 3.** Names of trapped species, origin and specimen numbers captured per continent. Species in bold were trapped in non-native continents.</caption><thead><tr><th>Subfamily</th><th>Tribe</th><th>Species</th><th>Origin</th><th>Europe</th><th>Asia</th><th>North America</th><th>The Caribbean</th><th>Australia</th><th>Total</th></tr></thead><tbody><tr><td>Cerambycinae</td><td>Anaglyptini</td><td>Anaglyptus gibbosus (Fabricius, 1787)</td><td>Europe</td><td>105</td><td>0</td><td>0</td><td>0</td><td>0</td><td>105</td></tr><tr><td>Cerambycinae</td><td>Anaglyptini</td><td>Anaglyptus mysticus (Linnaeus, 1758)</td><td>Europe</td><td>116</td><td>0</td><td>0</td><td>0</td><td>0</td><td>116</td></tr><tr><td>Cerambycinae</td><td>Anaglyptini</td><td>Cyrtophorus verrucosus (Olivier, 1800)</td><td>North America</td><td>0</td><td>0</td><td>197</td><td>0</td><td>0</td><td>197</td></tr><tr><td>Cerambycinae</td><td>Anaglyptini</td><td>Microclytus compressicollis (Laporte de Castelnau & Gory, 1841)</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Bothiospilini</td><td>Chlorida festiva (Linnaeus, 1758)</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Callichromatini</td><td>Aromia bungii Faldermann, 1835</td><td>Asia</td><td>0</td><td>25</td><td>0</td><td>0</td><td>0</td><td>25</td></tr><tr><td>Cerambycinae</td><td>Callichromatini</td><td>Aromia moschata (Linnaeus, 1758)</td><td>Europe</td><td>30</td><td>0</td><td>0</td><td>0</td><td>0</td><td>30</td></tr><tr><td>Cerambycinae</td><td>Callichromatini</td><td>Aromia moschata orientalis Plavilstshikov, 1933</td><td>Asia</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Callidium aeneum (Deguer, 1775)</td><td>Holarctic</td><td>120</td><td>79</td><td>0</td><td>0</td><td>0</td><td>199</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Callidium violaceum (Linnaeus, 1758)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Lioderina linearis (Hampe, 1870)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes aereus (Newman, 1838)</td><td>North America</td><td>0</td><td>0</td><td>14</td><td>0</td><td>0</td><td>14</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes alni (Linnaeus, 1767)</td><td>Europe</td><td>2295</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2295</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes amoenus (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>3100</td><td>0</td><td>0</td><td>3100</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes dimidiatus (Kirby, 1837)</td><td>North America</td><td>0</td><td>0</td><td>55</td><td>0</td><td>0</td><td>55</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes fasciatus (Villers, 1789)</td><td>Europe</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes glabratus (Charpentier, 1825)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes lividus (Rossi, 1794)</td><td>Europe</td><td>7</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes pusillus (Fabricius, 1787)</td><td>Europe</td><td>37</td><td>0</td><td>0</td><td>0</td><td>0</td><td>37</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes rufipes (Fabricius, 1776)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes testaceus (Linnaeus, 1758)</td><td>Europe</td><td>15085</td><td>0</td><td>41</td><td>0</td><td>0</td><td>15126</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Phymatodes varius (Fabricius, 1776)</td><td>North America</td><td>0</td><td>0</td><td>29</td><td>0</td><td>0</td><td>29</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Physocnemum brevilineum (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>4</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Pyrrhidium sanguineum (Linnaeus, 1758)</td><td>Europe</td><td>4388</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4388</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Ropalopus clavipes (Fabricius, 1775)</td><td>Europe</td><td>69</td><td>0</td><td>0</td><td>0</td><td>0</td><td>69</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Ropalopus femoratus (Linnaeus, 1758)</td><td>Europe</td><td>35</td><td>0</td><td>0</td><td>0</td><td>0</td><td>35</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Ropalopus macropus (Germar, 1823)</td><td>Europe</td><td>21</td><td>0</td><td>0</td><td>0</td><td>0</td><td>21</td></tr><tr><td>Cerambycinae</td><td>Callidiini</td><td>Ropalopus varini (Bedel, 1870)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Callidiopiini</td><td>Bethelium sp. Bethelium sp. (Fabricius, 1775)</td><td>Australasia</td><td>0</td><td>0</td><td>0</td><td>5</td><td>0</td><td>5</td></tr><tr><td>Cerambycinae</td><td>Callidiopiini</td><td>Curtomerus flavus (Fabricius, 1775)</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td><td>7</td></tr><tr><td>Cerambycinae</td><td>Callidiopiini</td><td>Stenodryas clavigera Bates, 1873</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Cerambycini</td><td>Cerambyx cerdo Linnaeus, 1758</td><td>Europe</td><td>20</td><td>0</td><td>0</td><td>0</td><td>0</td><td>20</td></tr><tr><td>Cerambycinae</td><td>Cerambycini</td><td>Cerambyx miles Bonelli, 1812</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Cerambycini</td><td>Cerambyx scopoli Fueßlins, 1775</td><td>Europe</td><td>141</td><td>0</td><td>0</td><td>0</td><td>0</td><td>141</td></tr><tr><td>Cerambycinae</td><td>Cerambycini</td><td>Cerambyx welensii (Küster, 1845)</td><td>Europe</td><td>22</td><td>0</td><td>0</td><td>0</td><td>0</td><td>22</td></tr><tr><td>Cerambycinae</td><td>Cerambycini</td><td>Nadezhdiella cantori (Hope, 1842)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus figuratus (Scopoli, 1763)</td><td>Europe</td><td>42</td><td>0</td><td>0</td><td>0</td><td>0</td><td>42</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus glabromaculatus (Goeze, 1777)</td><td>Europe</td><td>1391</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1391</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus glaucus (Fabricius, 1781)</td><td>Europe</td><td>36</td><td>0</td><td>0</td><td>0</td><td>0</td><td>36</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus herbstii (Brahm, 1790)</td><td>Europe</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus miwai Gressitt, 1936</td><td>Asia</td><td>0</td><td>9</td><td>0</td><td>0</td><td>0</td><td>9</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus motschulskyi (Ganglbauer, 1887)</td><td>Asia</td><td>0</td><td>7</td><td>0</td><td>0</td><td>0</td><td>7</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus ruficornis (Olivier, 1790)</td><td>Europe</td><td>41</td><td>0</td><td>0</td><td>0</td><td>0</td><td>41</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus sartor Müller, 1766</td><td>Europe</td><td>482</td><td>0</td><td>0</td><td>0</td><td>0</td><td>482</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus signaticollis (Laporte de Castelnau & Gory, 1836)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus tredecimcimulatus (Chevrolat, 1863)</td><td>Asia</td><td>0</td><td>2</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus trifasciatus (Fabricius, 1781)</td><td>Europe</td><td>33</td><td>0</td><td>0</td><td>0</td><td>0</td><td>33</td></tr></tbody></table>
<table><thead><tr><th>Subfamily</th><th>Tribe</th><th>Species</th><th>Origin</th><th>Europe</th><th>Asia</th><th>North America</th><th>The Caribbean</th><th>Australia</th><th>Total</th></tr></thead><tbody><tr><td>Cerambycinae</td><td>Clytini</td><td>Chlorophorus varius (Müller, 1766)</td><td>Europe</td><td>36</td><td>0</td><td>0</td><td>0</td><td>0</td><td>36</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Clytoleptus albofasciatus (Laporte de Castelnau & Gory, 1841)</td><td>North America</td><td>0</td><td>0</td><td>6</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Clytus arietis (Linnaeus, 1758)</td><td>Europe</td><td>52</td><td>0</td><td>0</td><td>0</td><td>0</td><td>52</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Clytus lama Mulsant, 1850</td><td>Europe</td><td>123</td><td>0</td><td>0</td><td>0</td><td>0</td><td>123</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Clytus rhamni Germar, 1817</td><td>Europe</td><td>85</td><td>0</td><td>0</td><td>0</td><td>0</td><td>85</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Clytus ruricola (Olivier, 1800)</td><td>North America</td><td>0</td><td>0</td><td>25</td><td>0</td><td>0</td><td>25</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Clytus tropicus (Panzer, 1795)</td><td>Europe</td><td>73</td><td>0</td><td>0</td><td>0</td><td>0</td><td>73</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Cyrtoclytus capra (Germar, 1823)</td><td>Asia</td><td>0</td><td>24</td><td>0</td><td>0</td><td>0</td><td>24</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Cyrtoclytus caproides (Bates, 1873)</td><td>Asia</td><td>0</td><td>5</td><td>0</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Demonax diversefasciatus Pic, 1920</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Demonax nansenensis Pic 1903</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Demonax sp. 1</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Demonax sp. 2</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Glycobius speciosus (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Isotomus speciosus (Schneider, 1787)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Megacyllene caryae (Gahan, 1908)</td><td>North America</td><td>0</td><td>0</td><td>22</td><td>0</td><td>0</td><td>22</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Neoclytus acuminatus acuminatus (Fabricius, 1775)</td><td>North America</td><td>37</td><td>0</td><td>28</td><td>0</td><td>0</td><td>65</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Neoclytus caprea (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Neoclytus leucozonus (Laporte de Castelnau & Gory, 1841)</td><td>North America</td><td>0</td><td>0</td><td>15</td><td>0</td><td>0</td><td>15</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Neoclytus muconatus muconatus (Fabricius, 1775)</td><td>North America</td><td>0</td><td>0</td><td>323</td><td>0</td><td>0</td><td>323</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Neoclytus muricatulus (Kirby, 1837)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Neoclytus scutellaris (Olivier, 1790)</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Perissus paulonotatus (Pic, 1902)</td><td>Asia</td><td>0</td><td>21</td><td>0</td><td>0</td><td>0</td><td>21</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Plagionotus arcuatus (Linnaeus, 1758)</td><td>Europe</td><td>95</td><td>0</td><td>0</td><td>0</td><td>0</td><td>95</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Plagionotus christophi (Kraatz, 1879)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Plagionotus detritus (Linnaeus, 1758)</td><td>Europe</td><td>299</td><td>0</td><td>0</td><td>0</td><td>0</td><td>299</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Pseudosphegesthes cinerea (Laporte de Castelnau & Gory, 1841)</td><td>Europe</td><td>27</td><td>0</td><td>0</td><td>0</td><td>0</td><td>27</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Raphuma anongi Gressitt & Rondon, 1970</td><td>Asia</td><td>0</td><td>96</td><td>0</td><td>0</td><td>0</td><td>96</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Raphuma gracilipes (Faldermann, 1835)</td><td>Asia</td><td>0</td><td>24</td><td>0</td><td>0</td><td>0</td><td>24</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Raphuma laosica Gressitt & Rondon, 1970</td><td>Asia</td><td>0</td><td>22</td><td>0</td><td>0</td><td>0</td><td>22</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Raphuma sp.</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Rhabdoclytus acutivittis (Kraatz, 1879)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Sarosesthes fulminans (Fabricius, 1775)</td><td>North America</td><td>0</td><td>0</td><td>39</td><td>0</td><td>0</td><td>39</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus antilope (Schönherr, 1817)</td><td>Europe</td><td>1303</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1303</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus antilope var sekenai Podaný, 1970</td><td>Europe</td><td>16</td><td>0</td><td>0</td><td>0</td><td>0</td><td>16</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus arvicola (Olivier, 1800)</td><td>Europe</td><td>379</td><td>0</td><td>0</td><td>0</td><td>0</td><td>379</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus atronotatus Pic, 1917</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus buqueti (Laporte de Castelnau & Gory, 1841)</td><td>Asia</td><td>0</td><td>38</td><td>0</td><td>0</td><td>0</td><td>38</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus chinensis (Chevrolat, 1852)</td><td>Asia</td><td>41</td><td>3</td><td>0</td><td>0</td><td>0</td><td>44</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus clarinus Bates, 1884</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus colonus (Fabricius, 1775)</td><td>North America</td><td>0</td><td>0</td><td>484</td><td>0</td><td>0</td><td>484</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus gratus Viktora, 2020</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus integer (Haldeman, 1847)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus latefasciatus ochroceps Gressitt, 1951</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus magnicollis (Fairmaire, 1888)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus pantherinus (Savenius, 1825)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus pekingensis Pic, 1939</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr></tbody></table>
<table><thead><tr><th>Subfamily</th><th>Tribe</th><th>Species</th><th>Origin</th><th>Europe</th><th>Asia</th><th>North America</th><th>The Caribbean</th><th>Australia</th><th>Total</th></tr></thead><tbody><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus rufilius Bates, 1884</td><td>Asia</td><td>0</td><td>27</td><td>0</td><td>0</td><td>0</td><td>27</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus rusticus (Linnaeus, 1758)</td><td>Europe/Asia</td><td>161</td><td>1</td><td>0</td><td>0</td><td>0</td><td>162</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus sagittatus (Germar, 1821)</td><td>North America</td><td>0</td><td>0</td><td>34</td><td>0</td><td>0</td><td>34</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td><strong>Xylotrechus stebbingi Gahan, 1906</strong></td><td>Asia</td><td>6089</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6054</td></tr><tr><td>Cerambycinae</td><td>Clytini</td><td>Xylotrechus undulatus (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>26</td><td>0</td><td>0</td><td>26</td></tr><tr><td>Cerambycinae</td><td>Deilini</td><td>Deilus fugax (Olivier, 1790)</td><td>Europe</td><td>87</td><td>0</td><td>0</td><td>0</td><td>0</td><td>87</td></tr><tr><td>Cerambycinae</td><td>Dryobiini</td><td>Dryobius sexnotatus Linsley, 1957</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Eburiini</td><td>Eburia dejeani Gahan, 1895</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Eburiini</td><td>Eburia octomaculata Chevrolat, 1862</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Eburiini</td><td>Eburia quadrigeminata (Say, 1827)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Elaphidiini</td><td>Anelaphus pumilus (Newman, 1840)</td><td>North America</td><td>0</td><td>0</td><td>531</td><td>0</td><td>0</td><td>531</td></tr><tr><td>Cerambycinae</td><td>Elaphidiini</td><td>Anelaphus villoosus (Fabricius, 1793)</td><td>North America</td><td>0</td><td>0</td><td>8</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Cerambycinae</td><td>Elaphidiini</td><td>Elaphidion mucronatum (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>110</td><td>0</td><td>0</td><td>110</td></tr><tr><td>Cerambycinae</td><td>Elaphidiini</td><td>Parelaphidion aspersum (Haldeman, 1847)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Elaphidiini</td><td>Parelaphidion incertum (Newman, 1840)</td><td>North America</td><td>0</td><td>0</td><td>4</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Cerambycinae</td><td>Graciliini</td><td>Axinopalpis gracilis (Krynicki, 1832)</td><td>Europe</td><td>8</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Cerambycinae</td><td>Graciliini</td><td>Gracilia minuta (Fabricius, 1781)</td><td>Europe</td><td>12</td><td>0</td><td>0</td><td>0</td><td>0</td><td>12</td></tr><tr><td>Cerambycinae</td><td>Graciliini</td><td>Penichroa fasciata (Stephens, 1831)</td><td>Europe</td><td>41</td><td>0</td><td>0</td><td>0</td><td>0</td><td>41</td></tr><tr><td>Cerambycinae</td><td>Hesperophanini</td><td>Gnatholea eburifera Thomson, 1861</td><td>Asia</td><td>0</td><td>10</td><td>0</td><td>0</td><td>0</td><td>10</td></tr><tr><td>Cerambycinae</td><td>Hesperophanini</td><td>Hesperophanes sericeus (Fabricius, 1787)</td><td>Europe</td><td>8</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Cerambycinae</td><td>Hesperophanini</td><td>Sromatium auratum (Böber, 1793)</td><td>Europe</td><td>7</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td></tr><tr><td>Cerambycinae</td><td>Hesperophanini</td><td><strong>Trichoferus campestris</strong><br>(Faldermann, 1835)</td><td>Asia</td><td>45</td><td>12</td><td>0</td><td>0</td><td>0</td><td>57</td></tr><tr><td>Cerambycinae</td><td>Hesperophanini</td><td>Trichoferus fasciculatus (Faldermann, 1837)</td><td>Europe</td><td>135</td><td>0</td><td>0</td><td>0</td><td>0</td><td>135</td></tr><tr><td>Cerambycinae</td><td>Hesperophanini</td><td>Trichoferus guerryi (Pic, 1915)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Hesperophanini</td><td>Trichoferus holosericeus (Rossi, 1790)</td><td>Europe</td><td>187</td><td>0</td><td>0</td><td>0</td><td>0</td><td>187</td></tr><tr><td>Cerambycinae</td><td>Hesperophanini</td><td>Trichoferus pallidus (Olivier, 1790)</td><td>Europe</td><td>145</td><td>0</td><td>0</td><td>0</td><td>0</td><td>145</td></tr><tr><td>Cerambycinae</td><td>Hylotrupini</td><td>Hylotrupes bajulus (Linnaeus, 1758)</td><td>Europe</td><td>79</td><td>0</td><td>0</td><td>0</td><td>0</td><td>79</td></tr><tr><td>Cerambycinae</td><td>Molorchini</td><td>Dolocerus reichii Mulsant, 1862</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Molorchini</td><td>Molorchus bimaculatus Say, 1824</td><td>North America</td><td>0</td><td>0</td><td>122</td><td>0</td><td>0</td><td>122</td></tr><tr><td>Cerambycinae</td><td>Molorchini</td><td>Molorchus minor (Linnaeus, 1758)</td><td>Europe</td><td>15</td><td>0</td><td>0</td><td>0</td><td>0</td><td>15</td></tr><tr><td>Cerambycinae</td><td>Molorchini</td><td>Molorchus umbellatarum (Schreber, 1759)</td><td>Europe</td><td>55</td><td>0</td><td>0</td><td>0</td><td>0</td><td>55</td></tr><tr><td>Cerambycinae</td><td>Neoibidionini</td><td>Neocomps cylindricollis (Fabricius, 1798)</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Obriini</td><td>Obrium brunneum (Fabricius, 1793)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Obriini</td><td>Obrium cantharinum (Linnaeus, 1767)</td><td>Europe</td><td>44</td><td>0</td><td>0</td><td>0</td><td>0</td><td>44</td></tr><tr><td>Cerambycinae</td><td>Obriini</td><td>Obrium maculatum (Olivier, 1800)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Phoracanthini</td><td><strong>Cordylomera spinicornis (Fabricius, 1775)</strong></td><td>Africa</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Cerambycinae</td><td>Phoracanthini</td><td><strong>Phoracantha recurva Newman, 1840</strong></td><td>Australasia</td><td>8</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Cerambycinae</td><td>Phoracanthini</td><td><strong>Phoracantha semipunctata</strong><br>(Fabricius, 1775)</td><td>Australasia</td><td>11</td><td>0</td><td>0</td><td>0</td><td>0</td><td>11</td></tr><tr><td>Cerambycinae</td><td>Phoracanthini</td><td><em>Thoris</em> sp.</td><td>Australasia</td><td>0</td><td>0</td><td>0</td><td>2</td><td>0</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Psebiini</td><td>Nathrius brevipennis (Mulsant, 1839)</td><td>Europe</td><td>649</td><td>0</td><td>0</td><td>0</td><td>0</td><td>649</td></tr><tr><td>Cerambycinae</td><td>Pytheini</td><td>Certallum ebulinum (Linnaeus, 1767)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Rhopalophorini</td><td>Rhopalophora longipes (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Stenhomalini</td><td>Stenhomalus fenestratus White, 1855</td><td>Asia</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Stenoderini</td><td>Syllitus sp.</td><td>Australasia</td><td>0</td><td>0</td><td>0</td><td>2</td><td>0</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Stenopterini</td><td>Callimoxys sanguinicolis (Olivier, 1800)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Stenopterini</td><td>Callimus abdominalis (Olivier, 1800)</td><td>Europe</td><td>11</td><td>0</td><td>0</td><td>0</td><td>0</td><td>11</td></tr><tr><td>Cerambycinae</td><td>Stenopterini</td><td>Callimus angulatus (Schrank, 1789)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Stenopterini</td><td>Stenopterus ater (Linnaeus, 1767)</td><td>Europe</td><td>20</td><td>0</td><td>0</td><td>0</td><td>0</td><td>20</td></tr><tr><td>Cerambycinae</td><td>Stenopterini</td><td>Stenopterus rufus (Linnaeus, 1767)</td><td>Europe</td><td>83</td><td>0</td><td>0</td><td>0</td><td>0</td><td>83</td></tr><tr><td>Cerambycinae</td><td>Tillomorphini</td><td>Bonfilsia pejoti Chalumeau &amp; Touroult, 2004</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>1</td></tr></tbody></table>
<table><thead><tr><th>Subfamily</th><th>Tribe</th><th>Species</th><th>Origin</th><th>Europe</th><th>Asia</th><th>North America</th><th>The Caribbean</th><th>Australia</th><th>Total</th></tr></thead><tbody><tr><td>Cerambycinae</td><td>Tillomorphini</td><td>Euderces picipes (Fabricius, 1787)</td><td>North America</td><td>0</td><td>0</td><td>9</td><td>0</td><td>0</td><td>9</td></tr><tr><td>Cerambycinae</td><td>Tillomorphini</td><td>Euderces pini (Olivier, 1800)</td><td>North America</td><td>0</td><td>0</td><td>93</td><td>0</td><td>0</td><td>93</td></tr><tr><td>Cerambycinae</td><td>Tillomorphini</td><td>Gourbeyrella madininae<br>Chalumeau &amp; Touroul, 2004</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Trachyderini</td><td>Anoplistes halodendri (Pallas, 1773)</td><td>Asia</td><td>0</td><td>2</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Trachyderini</td><td>Dicelosternus corallinus Gahan, 1900</td><td>Asia</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Cerambycinae</td><td>Trachyderini</td><td>Purpuricenus budensis (Götz, 1783)</td><td>Europe</td><td>18</td><td>0</td><td>0</td><td>0</td><td>0</td><td>18</td></tr><tr><td>Cerambycinae</td><td>Trachyderini</td><td>Purpuricenus globulicollis Dejean, 1839</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Cerambycinae</td><td>Trachyderini</td><td>Purpuricenus kaehleri (Linnaeus, 1758)</td><td>Europe</td><td>261</td><td>0</td><td>0</td><td>0</td><td>0</td><td>261</td></tr><tr><td>Cerambycinae</td><td>Trachyderini</td><td>Purpuricenus lituratus Ganglbauer, 1887</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Cerambycinae</td><td>Trachyderini</td><td>Purpuricenus temminckii<br>(Guérin-Méneville, 1844)</td><td>Asia</td><td>0</td><td>10</td><td>0</td><td>0</td><td>0</td><td>10</td></tr><tr><td>Cerambycinae</td><td>Trachyderini</td><td>Amarysius altajensis (Laxmann, 1770)</td><td>Asia</td><td>0</td><td>20</td><td>0</td><td>0</td><td>0</td><td>20</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Acanthocinus aedilis (Linnaeus, 1758)</td><td>Europe/Asia</td><td>6</td><td>24</td><td>0</td><td>0</td><td>0</td><td>30</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Acanthocinus griseus (Fabricius, 1793)</td><td>Europe/Asia</td><td>114</td><td>106</td><td>0</td><td>0</td><td>0</td><td>220</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Acanthocinus pusillus (Kirby, 1837)</td><td>North America</td><td>0</td><td>0</td><td>21</td><td>0</td><td>0</td><td>21</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Amniscus similis (Gahan, 1895)</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>5</td><td>5</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Astyleiopus variegatus (Haldeman, 1847)</td><td>North America</td><td>0</td><td>0</td><td>11</td><td>0</td><td>0</td><td>11</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Astylidius parvus (LeConte, 1873)</td><td>North America</td><td>0</td><td>0</td><td>17</td><td>0</td><td>0</td><td>17</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Astylopsis macula (Say, 1827)</td><td>North America</td><td>0</td><td>0</td><td>47</td><td>0</td><td>0</td><td>47</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Astylopsis sexguttata (Say, 1827)</td><td>North America</td><td>0</td><td>0</td><td>19</td><td>0</td><td>0</td><td>19</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Astylopsis sp.</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Graphisurus despectus (LeConte, 1850)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Graphisurus fasciatus (Degeer, 1775)</td><td>North America</td><td>0</td><td>0</td><td>86</td><td>0</td><td>0</td><td>86</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Graphisurus triangulifer (Haldeman, 1847)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Hyperplatys maculatus Haldeman, 1847</td><td>North America</td><td>0</td><td>0</td><td>4</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Lagocheirus araneiformis insulorum<br>Dillon, 1957</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td><td>4</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Leiopus fallaciosus Holzschuh, 1993</td><td>Asia</td><td>0</td><td>5</td><td>0</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Leiopus femoratus Fairmaire, 1859</td><td>Europe</td><td>3461</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3461</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Leiopus linnei Wallin, Nylander &amp;<br>Kvamme, 2009</td><td>Europe</td><td>548</td><td>0</td><td>0</td><td>0</td><td>0</td><td>548</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Leiopus nebulosus (Linneus, 1758)</td><td>Europe</td><td>1473</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1473</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Leptostylus transversus (Gyllenhal, 1817)</td><td>North America</td><td>0</td><td>0</td><td>101</td><td>0</td><td>0</td><td>101</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Lepturges angulatus (LeConte, 1852)</td><td>North America</td><td>0</td><td>0</td><td>20</td><td>0</td><td>0</td><td>20</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Lepturges confluens (Haldeman, 1847)</td><td>North America</td><td>0</td><td>0</td><td>26</td><td>0</td><td>0</td><td>26</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Lepturges sp.</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Sternidius alpha (Say, 1827)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Sternidius punctatus (Haldeman, 1847)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Sternidius rusticus (LeConte, 1852)</td><td>North America</td><td>0</td><td>0</td><td>19</td><td>0</td><td>0</td><td>19</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Styloleptus posticalis (Gahan, 1895)</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>1</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Trypanidius spilmani Villiers, 1980</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>1</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Urgleptes cobbeni Gilmour, 1963</td><td>Caribbean</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td><td>1</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Urgleptes querci (Fitch, 1859)</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lamiinae</td><td>Acanthocinini</td><td>Urgleptes signatus (LeConte, 1852)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Acanthoderini</td><td>Aegomorphus clavipes (Schrank von<br>Paula, 1781)</td><td>Europe</td><td>1412</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1412</td></tr><tr><td>Lamiinae</td><td>Acanthoderini</td><td>Aegomorphus francottei Sama, 1994</td><td>Europe</td><td>181</td><td>0</td><td>0</td><td>0</td><td>0</td><td>181</td></tr><tr><td>Lamiinae</td><td>Acanthoderini</td><td>Aegomorphus krueperi (Kraatz, 1859)</td><td>Europe</td><td>7</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td></tr><tr><td>Lamiinae</td><td>Acanthoderini</td><td>Aegomorphus modestus (Blais, 1817)</td><td>North America</td><td>0</td><td>0</td><td>58</td><td>0</td><td>0</td><td>58</td></tr><tr><td>Lamiinae</td><td>Acanthoderini</td><td>Aegomorphus quadrigibbus (Say, 1831)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Acanthoderini</td><td>Oplosia cinerea (Mulsant, 1839)</td><td>Europe</td><td>63</td><td>0</td><td>0</td><td>0</td><td>0</td><td>63</td></tr><tr><td>Lamiinae</td><td>Acanthoderini</td><td>Oplosia nubila (LeConte, 1862)</td><td>North America</td><td>0</td><td>0</td><td>4</td><td>0</td><td>0</td><td>4</td></tr></tbody></table>
<table><thead><tr><th>Subfamily</th><th>Tribe</th><th>Species</th><th>Origin</th><th>Europe</th><th>Asia</th><th>North America</th><th>The Caribbean</th><th>Australia</th><th>Total</th></tr></thead><tbody><tr><td>Lamiinae</td><td>Agapanthiini</td><td>Agapanthia cardui (Linnaeus, 1767)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Agapanthiini</td><td>Agapanthia villosoviridescens (Degeer, 1775)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Apomecynini</td><td>Apomecyna saltator (Fabricius, 1787)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Ceroplesini</td><td>Moechotypa diphysis (Pascoe, 1871)</td><td>Asia</td><td>0</td><td>2</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lamiinae</td><td>Ceroplesini</td><td>Thysia wallichii tonkinensis (Kreische, 1924)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Desmiphorini</td><td>Anaesthetis testacea (Fabricius, 1781)</td><td>Europe</td><td>17</td><td>0</td><td>0</td><td>0</td><td>0</td><td>17</td></tr><tr><td>Lamiinae</td><td>Desmiphorini</td><td>Deroplia genei (Aragona, 1830)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Desmiphorini</td><td>Deroplia troberti (Mulsant, 1843)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Desmiphorini</td><td>Eupogonius pauper LeConte, 1852</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Desmiphorini</td><td>Eupogonius tomentosus (Haldeman, 1847)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Desmiphorini</td><td>Psenocerus supernotatus (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>9</td><td>0</td><td>0</td><td>9</td></tr><tr><td>Lamiinae</td><td>Dorcaschematini</td><td>Dorcaschema cinereum (Olivier, 1800)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Dorcaschematini</td><td>Olenecamptus bilobus (Fabricius, 1801)</td><td>Asia</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Exocentrini</td><td>Exocentrus adspersus Mulsant, 1846</td><td>Europe</td><td>5</td><td>0</td><td>0</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Lamiinae</td><td>Exocentrini</td><td>Exocentrus lusitanus (Linnaeus, 1767)</td><td>Europe</td><td>29</td><td>0</td><td>0</td><td>0</td><td>0</td><td>29</td></tr><tr><td>Lamiinae</td><td>Exocentrini</td><td>Exocentrus punctipennis Mulsant &amp; Guillebeau, 1856</td><td>Europe</td><td>28</td><td>0</td><td>0</td><td>0</td><td>0</td><td>28</td></tr><tr><td>Lamiinae</td><td>Lamiini</td><td>Lamiimimus gottschei Kolbe, 1886</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Lamiini</td><td>Pharsalia subgemmata (Thomson, 1857)</td><td>Asia</td><td>0</td><td>375</td><td>0</td><td>0</td><td>0</td><td>375</td></tr><tr><td>Lamiinae</td><td>Mesosini</td><td>Mesosa curculionoides (Linnaeus, 1761)</td><td>Europe</td><td>37</td><td>0</td><td>0</td><td>0</td><td>0</td><td>37</td></tr><tr><td>Lamiinae</td><td>Mesosini</td><td>Mesosa myops (Dalman, 1817)</td><td>Asia</td><td>0</td><td>29</td><td>0</td><td>0</td><td>0</td><td>29</td></tr><tr><td>Lamiinae</td><td>Mesosini</td><td>Mesosa nebulosa (Fabricius, 1781)</td><td>Europe</td><td>132</td><td>0</td><td>0</td><td>0</td><td>0</td><td>132</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Anoplophora beryllina (Hope, 1840)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Anoplophora chinensis (Forster, 1771)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Anoplophora glabripennis (Motschulsky, 1854)</td><td>Asia</td><td>0</td><td>9</td><td>0</td><td>0</td><td>0</td><td>9</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Microgoes oculatus (LeConte, 1862)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus alternatus Hope, 1842</td><td>Asia</td><td>0</td><td>1246</td><td>0</td><td>0</td><td>0</td><td>1246</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus bimaculatus Gahan, 1888</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus carolinensis (Olivier, 1797)</td><td>North America</td><td>0</td><td>0</td><td>77</td><td>0</td><td>0</td><td>77</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus galloprovincialis (Olivier, 1800)</td><td>Europe/Asia</td><td>6209</td><td>87</td><td>0</td><td>0</td><td>0</td><td>6296</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus maculosus Haldeman, 1847</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus notatus (Drury, 1773)</td><td>North America</td><td>0</td><td>0</td><td>256</td><td>0</td><td>0</td><td>256</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus saltuarius Gebler, 1830</td><td>Asia/Europe</td><td>13</td><td>985</td><td>0</td><td>0</td><td>0</td><td>998</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus sartor (Fabricius, 1787)</td><td>Europe</td><td>20</td><td>0</td><td>0</td><td>0</td><td>0</td><td>20</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus sartor urusovii (Fischer von Waldheim, 1806)</td><td>Asia/Europe</td><td>1</td><td>41</td><td>0</td><td>0</td><td>0</td><td>42</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus scutellatus (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>216</td><td>0</td><td>0</td><td>216</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus sutor (Linnaeus, 1758)</td><td>Europe/Asia</td><td>30</td><td>22</td><td>0</td><td>0</td><td>0</td><td>52</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Monochamus sutor longulus Pic, 1898</td><td>Asia</td><td>0</td><td>22</td><td>0</td><td>0</td><td>0</td><td>22</td></tr><tr><td>Lamiinae</td><td>Monochamini</td><td>Uraecha angusta (Pascoe, 1857)</td><td>Asia</td><td>0</td><td>15</td><td>0</td><td>0</td><td>0</td><td>15</td></tr><tr><td>Lamiinae</td><td>Obereini</td><td>Oberea linearis (Linnaeus, 1761)</td><td>Europe</td><td>8</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Lamiinae</td><td>Parmenini</td><td>Mesolita sp.</td><td>Australasia</td><td>0</td><td>0</td><td>0</td><td>3</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Parmenini</td><td>Parmena balteus (Linnaeus, 1767)</td><td>Europe</td><td>7</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td></tr><tr><td>Lamiinae</td><td>Parmenini</td><td>Parmena unifasciata (Rossi, 1790)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Phytoeciini</td><td>Phytoecia pustulata (Schrank von Paula, 1776)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Phytoeciini</td><td>Phytoecia nigricornis (Fabricius, 1782)</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lamiinae</td><td>Pogonocherini</td><td>Pogonocherus caroli Mulsant, 1862</td><td>Europe</td><td>5</td><td>0</td><td>0</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Lamiinae</td><td>Pogonocherini</td><td>Pogonocherus decoratus Fairmaire, 1855</td><td>Europe</td><td>139</td><td>0</td><td>0</td><td>0</td><td>0</td><td>139</td></tr><tr><td>Lamiinae</td><td>Pogonocherini</td><td>Pogonocherus fasciculatus (Degeer, 1775)</td><td>Europe</td><td>16</td><td>1</td><td>0</td><td>0</td><td>0</td><td>17</td></tr><tr><td>Lamiinae</td><td>Pogonocherini</td><td>Pogonocherus hispidulus (Piller &amp; Mitterpacher, 1783)</td><td>Europe</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr></tbody></table>
<table><thead><tr><th>Subfamily</th><th>Tribe</th><th>Species</th><th>Origin</th><th>Europe</th><th>Asia</th><th>North America</th><th>The Caribbean</th><th>Australia</th><th>Total</th></tr></thead><tbody><tr><td>Lamiinae</td><td>Pogonocherini</td><td>Pogonocherus hispidus (Linnaeus, 1758)</td><td>Europe</td><td>55</td><td>0</td><td>0</td><td>0</td><td>0</td><td>55</td></tr><tr><td>Lamiinae</td><td>Pogonocherini</td><td>Pogonocherus mixtus Haldeman, 1847</td><td>North America</td><td>0</td><td>0</td><td>8</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Lamiinae</td><td>Pogonocherini</td><td>Pogonocherus ovatus (Goeze, 1777)</td><td>Europe</td><td>19</td><td>0</td><td>0</td><td>0</td><td>0</td><td>19</td></tr><tr><td>Lamiinae</td><td>Pogonocherini</td><td>Pogonocherus penicillatus LeConte, 1850</td><td>North America</td><td>0</td><td>0</td><td>11</td><td>0</td><td>0</td><td>11</td></tr><tr><td>Lamiinae</td><td>Pogonocherini</td><td>Pogonocherus perroudi Mulsant, 1839</td><td>Europe</td><td>127</td><td>0</td><td>0</td><td>0</td><td>0</td><td>127</td></tr><tr><td>Lamiinae</td><td>Pteroplini</td><td>Niphona picticornis Mulsant, 1839</td><td>Europe</td><td>127</td><td>0</td><td>0</td><td>0</td><td>0</td><td>127</td></tr><tr><td>Lamiinae</td><td>Pteroplini</td><td>Sthenias gracilicornis Gressitt, 1937</td><td>Europe</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Menesia bipunctata (Zoubkoff, 1829)</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Panaglenea fortunei (Saunders, 1853)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Saperda alberti Plavilstshikov, 1915</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Saperda hosokawai Hasegawa, 2017</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Saperda octopunctata (Scopoli, 1772)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Saperda perforata (Pallas, 1773)</td><td>Europe</td><td>21</td><td>0</td><td>0</td><td>0</td><td>0</td><td>21</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Saperda populnea (Linnaeus, 1758)</td><td>Europe</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Saperda scalaris (Linnaeus, 1758)</td><td>Europe</td><td>24</td><td>0</td><td>0</td><td>0</td><td>0</td><td>24</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Stenostola dubia (Laicharting, 1784)</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lamiinae</td><td>Saperdini</td><td>Stenostola ferrea (Schrank von Paula, 1776)</td><td>Europe</td><td>28</td><td>0</td><td>0</td><td>0</td><td>0</td><td>28</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Alosterma tabacicolor (Degeer, 1775)</td><td>Europe</td><td>9</td><td>0</td><td>0</td><td>0</td><td>0</td><td>9</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Anastrangalia dubia (Scopoli, 1763)</td><td>Europe</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Anastrangalia reyi (Heyden, 1889)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Anastrangalia sanguinolenta (Linnaeus 1761)</td><td>Europe</td><td>13</td><td>0</td><td>0</td><td>0</td><td>0</td><td>13</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Anastrangalia scotodes continentalis (Plavilstshikov, 1936)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Anoplodera rufipes (Schaller, 1783)</td><td>Europe</td><td>5</td><td>0</td><td>0</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Anoplodera sexguttata (Fabricius, 1775)</td><td>Europe</td><td>9</td><td>0</td><td>0</td><td>0</td><td>0</td><td>9</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Brachyleptura brevis (Kirby, 1837)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Brachyleptura circumdata (Olivier, 1800)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Brachyleptura rubrica (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Leptura thoracica Creutzer, 1799</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Leptura aethiops Poda von Neuhaus, 1761</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Leptura aurulenta Fabricius, 1793</td><td>Europe</td><td>7</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Leptura quadrifasciata Linnaeus, 1758</td><td>Europe</td><td>9</td><td>0</td><td>0</td><td>0</td><td>0</td><td>9</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Neoalosterna capitata (Newman, 1841)</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Pachytodes erraticus (Dalman, 1817)</td><td>Europe</td><td>232</td><td>0</td><td>0</td><td>0</td><td>0</td><td>232</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Paracorymbia fulva (Degeer, 1775)</td><td>Europe</td><td>8</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Paracorymbia hybrida (Rey, 1885)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Pedostrangalia revestita (Linnaeus, 1767)</td><td>Europe</td><td>12</td><td>0</td><td>0</td><td>0</td><td>0</td><td>12</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Pseudovadonia livida (Fabricius, 1776)</td><td>Europe</td><td>5</td><td>0</td><td>0</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Rutpela maculata (Poda von Neuhaus, 1761)</td><td>Europe</td><td>74</td><td>0</td><td>0</td><td>0</td><td>0</td><td>74</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stenurella nigra (Linnaeus 1758)</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stenurella bifasciata (Müller, 1776)</td><td>Europe</td><td>16</td><td>0</td><td>0</td><td>0</td><td>0</td><td>16</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stenurella septempunctata (Fabricius, 1793)</td><td>Europe</td><td>5</td><td>0</td><td>0</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stenurella melanura (Linnaeus 1758)</td><td>Europe</td><td>33</td><td>0</td><td>0</td><td>0</td><td>0</td><td>33</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stictoleptura canadensis (Olivier, 1800)</td><td>North America</td><td>0</td><td>0</td><td>8</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stictoleptura cordigera (Fueßlins, 1775)</td><td>Europe</td><td>203</td><td>0</td><td>0</td><td>0</td><td>0</td><td>203</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stictoleptura erythroptera (Hagenbach, 1822)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stictoleptura fontenayi (Mulsant, 1839)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stictoleptura maculicornis (Degeer, 1775)</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stictoleptura rubra (Linnaeus, 1758)</td><td>Europe/asia</td><td>11</td><td>1</td><td>0</td><td>0</td><td>0</td><td>12</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stictoleptura scutellata (Fabricius, 1781)</td><td>Europe</td><td>29</td><td>0</td><td>0</td><td>0</td><td>0</td><td>29</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stictoleptura succedanea (Lewis, 1879)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Stictoleptura trisignata (Fairmaire, 1852)</td><td>Europe</td><td>7</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Strangalepta abbreviata (Germar, 1823)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr></tbody></table>
<table><thead><tr><th>Subfamily</th><th>Tribe</th><th>Species</th><th>Origin</th><th>Europe</th><th>Asia</th><th>North America</th><th>The Caribbean</th><th>Australia</th><th>Total</th></tr></thead><tbody><tr><td>Lepturinae</td><td>Lepturini</td><td>Strangalia attenuata (Linnaeus 1758)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Strangalia luteicornis (Fabricius, 1775)</td><td>North America</td><td>0</td><td>0</td><td>4</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Strophiona nitens (Forster, 1771)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Trachysida mutabilis (Newman, 1841)</td><td>North America</td><td>0</td><td>0</td><td>4</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Trigonarthris proxima (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Trigonarthris subpubescens (Kirby, 1837)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Typocerus lunulatus (Swederus, 1787)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Typocerus velutinus (Olivier, 1800)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Lepturini</td><td>Vadonia unipunctata (Fabricius, 1787)</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Oxymirini</td><td>Anthophylax cyaneus (Haldeman, 1848)</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Oxymirini</td><td>Anthophylax viridis LeConte, 1850</td><td>North America</td><td>0</td><td>0</td><td>6</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Lepturinae</td><td>Oxymirini</td><td>Oxymirus cursor (Linnaeus, 1758)</td><td>Europe</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Anisorus quercus (Götz, 1783)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Brachyta interrogationis (Linnaeus, 1758)</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Carilia virginea (Linnaeus, 1758)</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Carilia virginea thalassina (Schrank von Paula, 1781)</td><td>Asia</td><td>0</td><td>14</td><td>0</td><td>0</td><td>0</td><td>14</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Centrodera decolorata (Harris, 1838)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Cortodera femorata (Fabricius, 1787)</td><td>Europe</td><td>11</td><td>0</td><td>0</td><td>0</td><td>0</td><td>11</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Cortodera flavimana (Waltl, 1838)</td><td>Europe</td><td>8</td><td>0</td><td>0</td><td>0</td><td>0</td><td>8</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Cortodera humenalis (Schaller, 1783)</td><td>Europe</td><td>99</td><td>0</td><td>0</td><td>0</td><td>0</td><td>99</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Dinoptera collaris (Linnaeus, 1758)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Acmaeops marginatus (Fabricius, 1781)</td><td>Europe/asia</td><td>3</td><td>11</td><td>0</td><td>0</td><td>0</td><td>14</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Acmaeops pratensis (Laicharting, 1784)</td><td>Europe</td><td>10</td><td>0</td><td>0</td><td>0</td><td>0</td><td>10</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Acmaeops proteus (Kirby, 1837)</td><td>North America</td><td>0</td><td>0</td><td>14</td><td>0</td><td>0</td><td>14</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Acmaeops septentrionis (C G Thomson, 1866)</td><td>Europe/asia</td><td>24</td><td>28</td><td>0</td><td>0</td><td>0</td><td>52</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Acmaeops smaragdulus (Fabricius, 1793)</td><td>Europe</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Evodinellus borealis (Gyllenhal, 1827)</td><td>Asia</td><td>0</td><td>2</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Gaurotes cyanipennis (Say, 1824)</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Grammoptera abdominalis (Stephens, 1831)</td><td>Europe</td><td>31</td><td>0</td><td>0</td><td>0</td><td>0</td><td>31</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Grammoptera ruficornis (Fabricius, 1781)</td><td>Europe</td><td>266</td><td>0</td><td>0</td><td>0</td><td>0</td><td>266</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Grammoptera ustulata (Schaller, 1783)</td><td>Europe</td><td>56</td><td>0</td><td>0</td><td>0</td><td>0</td><td>56</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Pachyta mediofasciata Pic 1936</td><td>Asia</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Pachyta quadrimaculata (Linnaeus, 1758)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Paragaurotes usuriensis (Blessig, 1873)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Pidonia lurida (Fabricius, 1792)</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Pseudosieversia japonica (Ohbayashi, 1937)</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Rhagium bifasciatum Fabricius, 1775</td><td>Europe</td><td>20</td><td>0</td><td>0</td><td>0</td><td>0</td><td>20</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Rhagium inquisitor (Linnaeus, 1758)</td><td>Holarctic</td><td>524</td><td>5</td><td>110</td><td>0</td><td>0</td><td>639</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Rhagium japonicum Bates, 1884</td><td>Asia</td><td>0</td><td>21</td><td>0</td><td>0</td><td>0</td><td>21</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Rhagium mordax (Degeer, 1775)</td><td>Europe</td><td>41</td><td>0</td><td>0</td><td>0</td><td>0</td><td>41</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Rhagium rugipenne Reitter, 1898</td><td>Asia</td><td>0</td><td>4</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Rhagium sycophanta (Schrank von Paula, 1781)</td><td>Europe</td><td>32</td><td>0</td><td>0</td><td>0</td><td>0</td><td>32</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Stenocorus cinnamopterus (Randall, 1838)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Lepturinae</td><td>Rhagiini</td><td>Stenocorus meridianus (Linnaeus, 1758)</td><td>Europe</td><td>71</td><td>0</td><td>0</td><td>0</td><td>0</td><td>71</td></tr><tr><td>Necydalinae</td><td>Necydalini</td><td>Necydalis major Linnaeus 1758</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Necydalinae</td><td>Necydalini</td><td>Necydalis ulmi (Chevrolat, 1838)</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Parandrinae</td><td>Parandrin</td><td>Neandra brunnea (Fabricius, 1798)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Prioninae</td><td>Aegosomatini</td><td>Aegosoma scabricorne (Scopoli, 1763)</td><td>Europe</td><td>33</td><td>0</td><td>0</td><td>0</td><td>0</td><td>33</td></tr><tr><td>Prioninae</td><td>Macrotomini</td><td>Prinobius myardi Mulsant, 1842</td><td>Europe</td><td>3</td><td>0</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Prioninae</td><td>Meroscelisini</td><td>Tragosoma harrisii LeConte, 1851</td><td>North America</td><td>0</td><td>0</td><td>236</td><td>0</td><td>0</td><td>236</td></tr><tr><td>Prioninae</td><td>Prionini</td><td>Dorysthenes sternalis (Fairmaire, 1902)</td><td>Asia</td><td>0</td><td>25</td><td>0</td><td>0</td><td>0</td><td>25</td></tr></tbody></table>
<table><thead><tr><th>Subfamily</th><th>Tribe</th><th>Species</th><th>Origin</th><th>Europe</th><th>Asia</th><th>North America</th><th>The Caribbean</th><th>Australia</th><th>Total</th></tr></thead><tbody><tr><td>Prioninae</td><td>Prionini</td><td>Dorysthenes paradoxus (Faldermann, 1833)</td><td>Asia</td><td>0</td><td>22</td><td>0</td><td>0</td><td>0</td><td>22</td></tr><tr><td>Prioninae</td><td>Prionini</td><td>Dorysthenes sp.</td><td>Asia</td><td>0</td><td>2</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Prioninae</td><td>Prionini</td><td>Mesoprionus besikanus (Fairmaire, 1855)</td><td>Europe</td><td>46</td><td>0</td><td>0</td><td>0</td><td>0</td><td>46</td></tr><tr><td>Prioninae</td><td>Prionini</td><td>Orthosoma brunneum (Forster, 1771)</td><td>North America</td><td>0</td><td>0</td><td>1</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Prioninae</td><td>Prionini</td><td>Prionus coriarius (Linnaeus, 1758)</td><td>Europe</td><td>4112</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4112</td></tr><tr><td>Prioninae</td><td>Prionini</td><td>Prionus insularis Motschulsky, 1857</td><td>Asia</td><td>0</td><td>241</td><td>0</td><td>0</td><td>0</td><td>241</td></tr><tr><td>Prioninae</td><td>Prionini</td><td>Prionus laticollis (Drury, 1773)</td><td>North America</td><td>0</td><td>0</td><td>3</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Prioninae</td><td>Prionini</td><td>Prionus sp.</td><td>Asia</td><td>0</td><td>1</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Spondylidinae</td><td>Anisarthrini</td><td>Alocerus moesiacus (Frivaldszky, 1837)</td><td>Europe</td><td>4</td><td>0</td><td>0</td><td>0</td><td>0</td><td>4</td></tr><tr><td>Spondylidinae</td><td>Anisarthrini</td><td>Anisarthron barbipes (Schrank von Paula, 1781)</td><td>Europe</td><td>19</td><td>0</td><td>0</td><td>0</td><td>0</td><td>19</td></tr><tr><td>Spondylidinae</td><td>Asemini</td><td>Arhopalus ferus (Mulsant, 1839)</td><td>Europe</td><td>338</td><td>0</td><td>0</td><td>0</td><td>0</td><td>338</td></tr><tr><td>Spondylidinae</td><td>Asemini</td><td>Arhopalus rusticus (Linnaeus, 1758)</td><td>Europe/Asia</td><td>4264</td><td>702</td><td>5</td><td>0</td><td>0</td><td>4971</td></tr><tr><td>Spondylidinae</td><td>Asemini</td><td>Asemum amurensis Kraatz, 1879</td><td>Asia</td><td>0</td><td>5</td><td>0</td><td>0</td><td>0</td><td>5</td></tr><tr><td>Spondylidinae</td><td>Asemini</td><td>Asemum striatum (Linnaeus, 1758)</td><td>Holarctic</td><td>21</td><td>181</td><td>289</td><td>0</td><td>0</td><td>491</td></tr><tr><td>Spondylidinae</td><td>Asemini</td><td>Asemum tenuicorne Kraatz, 1879</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Spondylidinae</td><td>Asemini</td><td>Cephalalbus oberthueri Sharp, 1905</td><td>Asia</td><td>0</td><td>14</td><td>0</td><td>0</td><td>0</td><td>14</td></tr><tr><td>Spondylidinae</td><td>Asemini</td><td>Cephalalbus sp.</td><td>Asia</td><td>0</td><td>3</td><td>0</td><td>0</td><td>0</td><td>3</td></tr><tr><td>Spondylidinae</td><td>Asemini</td><td>Cephalalbus unicolor (Gahan, 1906)</td><td>Asia</td><td>0</td><td>15</td><td>0</td><td>0</td><td>0</td><td>15</td></tr><tr><td>Spondylidinae</td><td>Asemini</td><td>Cephalocrius syriacus (Reitter, 1895)</td><td>Europe</td><td>2024</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2024</td></tr><tr><td>Spondylidinae</td><td>Nothorhinini</td><td>Nothorhina punctata (Fabricius, 1798)</td><td>Europe</td><td>2</td><td>0</td><td>0</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Spondylidinae</td><td>Saphanini</td><td>Oxypleurus nodieri Mulsant, 1839</td><td>Europe</td><td>25</td><td>0</td><td>0</td><td>0</td><td>0</td><td>25</td></tr><tr><td>Spondylidinae</td><td>Spondylidini</td><td>Spondylis buprestoides (Linnaeus, 1758)</td><td>Europe</td><td>2149</td><td>8</td><td>0</td><td>0</td><td>0</td><td>2157</td></tr><tr><td>Spondylidinae</td><td>Tetropiini</td><td>Tetropium castaneum (Linnaeus, 1758)</td><td>Europe</td><td>53</td><td>8</td><td>0</td><td>0</td><td>0</td><td>61</td></tr><tr><td>Spondylidinae</td><td>Tetropiini</td><td>Tetropium cinnamopterum Kirby, 1837</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Spondylidinae</td><td>Tetropiini</td><td>Tetropium fuscum (Fabricius, 1787)</td><td>Europe</td><td>100</td><td>0</td><td>0</td><td>0</td><td>0</td><td>100</td></tr><tr><td>Spondylidinae</td><td>Tetropiini</td><td>Tetropium gabrieli Weise, 1905</td><td>Europe</td><td>166</td><td>0</td><td>0</td><td>0</td><td>0</td><td>166</td></tr><tr><td>Spondylidinae</td><td>Tetropiini</td><td>Tetropium schwarzianum Casey, 1891</td><td>North America</td><td>0</td><td>0</td><td>2</td><td>0</td><td>0</td><td>2</td></tr><tr><td>Spondylidinae</td><td>Tetropiini</td><td>Tetrops praeustus (Linnaeus, 1758)</td><td>Europe</td><td>7</td><td>0</td><td>0</td><td>0</td><td>0</td><td>7</td></tr><tr><td>Spondylidinae</td><td>Tetropiini</td><td>Tetrops starkii Chevrolat, 1859</td><td>Europe</td><td>23</td><td>0</td><td>0</td><td>0</td><td>0</td><td>23</td></tr><tr><td>Disteniidae</td><td>Disteniini</td><td>Elytrimitatrix undata (Fabricius, 1775)</td><td>North America</td><td>0</td><td>0</td><td>6</td><td>0</td><td>0</td><td>6</td></tr><tr><td>Vesperidae</td><td>Vesperini</td><td>Vesperus conicicollis Fairmaire &amp; Coquerel, 1866</td><td>Europe</td><td>1</td><td>0</td><td>0</td><td>0</td><td>0</td><td>1</td></tr><tr><td>Vesperidae</td><td>Vesperini</td><td>Vesperus strepens (Fabricius, 1793)</td><td>Europe</td><td>6</td><td>0</td><td>0</td><td>0</td><td>0</td><td>6</td></tr></tbody></table>

In Europe, a total of 192 cerambycid species were trapped, of which seven were non-natives (three Clytini: the North American *N. a. acuminatus* and the Asian *X. chinensis* and *Xylotrechus stebbingi* Gahan; three Phoracanthini: the African *Cordylomera spinicornis* (F.) and the Australasian *Phoracantha recurva* Newman and *P. semipunctata* (F.); and one Hesperophanini: the Asian *T. campestris*). The captures amounted to about 20% of the total European cerambycid fauna (955 species, including apterous species, according to Vitali and Schmitt 2017). In North America, a total of 97 species were captured, including two non-native species originating from Europe: the *Aseminii Arhopalus rusticus* (L.) and the *Callidiini P. testaceus*. In contrast, no non-native species were trapped in Asia (95 total species), the Caribbean (12 total species; i.e. 18% of the 65 species known in Martinique; Touroult and Poirier 2021) and Australia. Three species with Holarctic distribution (the *Aseminii Asemum striatum* [L.], the *Callidiini Callidium aeneum* [Degeer] and the *Rhagiini Rhagium inquisitor* [L.]) were trapped in Europe, Asia and North America, where-
as nine species with northern Palaearctic distribution were captured in both Europe and Asia (the Clytini *Xylotrechus rusticus* [L.], the Acanthocinini *Acanthocinus griseus* [F.], the Monochamini *M. galloprovincialis* and congeners *M. saltuarius* [Say] and *M. sartor urusso-vii* [Fischer von Waldheim], the Lepturini *Stictoleptura rubra* [L.], the Rhagiini congeners *Acmaeops marginatus* [F.] and *A. septentrionis* [C. G. Thomson] and the Asemini *A. rusticus*).

Three species were notably abundant with captures exceeding > 5,000, including the European native *P. testaceus* (which was also trapped in the USA as a non-native species), the Palaearctic *M. galloprovincialis* (trapped in Europe and Northern China) and the Asian *X. stebbingi* which has invaded Europe. Sixteen species were represented by more than 1,000 specimens, 58 species by more than 100 specimens (Fig. 3) and 84 species by more than 50 individuals. In contrast, 109 cerambycid species from the total of 374 species were represented by only one or two specimens. More than 1,000 individuals were caught for four of the Callidiini species, of which three were native European species (*P. testaceus* – 15,126 individuals, 41 of which were trapped in North America where they have been introduced; *Pyrrhidium sanguineum* [L.]- 4,388 individuals and *Phymatodes alni* [L.]- 2,295 individuals), along with the North American *Phymatodes amoenus* (Say) (3,100 individuals).

Some invasive species were trapped in both their native range and in invaded regions (Fig. 4). The Chinese Clytini *X. chinensis* was captured in its native range around Beijing as well as in the invaded European areas in Spain, Greece (Crete) and southern France. The European species *P. testaceus* (Callidiini) and *A. rusticus* (Asemini) were

<figure><img src="image_4.png" /><figcaption>**Figure 3.** Cerambycid species with more than 100 individuals captured, ranked by decreasing order and showing the continent of capture.</figcaption></figure>
<figure><img src="image_5.png" /><figcaption>**Figure 4.** Cerambycid species trapped in both their native range (circles) and invaded range (triangles). Each species is shown by a different colour.</figcaption></figure>

trapped in large numbers in their native Europe, but also as non-native species in North America. Conversely, the North American Clytini *N. a. acuminatus* was captured in its native range in the USA, but also in the invaded areas of Italy. A noticeable anomaly was the Himalayan Clytini *X. stebbingi*, which was caught in large numbers in southern Europe (6,089 specimens) where it is invasive, but not at all in the traps deployed in Asia. The African Phoracanthini *C. spinicornis* was regularly trapped within European ports-of-entry, but not in nearby woody areas.

Simultaneous captures of non-target Coleopteran species were mostly bark and ambrosia beetles (> 100,000 individuals), which are not yet identified to species, but also predators in the family Cleridae, essentially *Clerus mutillarius* Fabricius, 1775 (> 5,000 individuals) and *Thanasimus* spp. (> 2,000 individuals) and Trogossitidae (*Temnoscheila* spp.; > 500 individuals).

## Additional captures resulting from the 10-pheromone blend

The addition of trichoferone and (E)-2-cis-6,7-epoxynonenal to the 8-pheromone blend in France and China in 2019 onwards, did not significantly change the previous trapping spectrum of the 8-pheromone blend (paired *t*-test; *P* = 0.750). However, the 10-pheromone blend resulted in trapping large numbers of four *Trichoferus* species, including the Asian *T. campestris* in its invasive range in Europe and native range in China (Table 3). In addition, large numbers of *Aromia bungii* were trapped in their native Chinese range. Only a few specimens of these five species had been previously trapped using the 8-pheromone blend.
Discussion

Capturing 376 species of cerambycid beetles from eight different subfamilies and 60 tribes on different continents, with 84 species captured in numbers greater than 50 individuals, clearly demonstrates the potential of the multi-pheromone lure to constitute an effective tool for the detection of ‘unexpected’ cerambycid invaders that are accidentally translocated outside their native ranges. Our hypothesis regarding the generic effectiveness of the blend was based on the evolutionary conservatism observed in many cerambycid pheromone structures. Pheromone constituents of the blend composition are shared by phylogenetically-related species on different continents (cf. references in Table 2). Therefore, their combination was expected to simultaneously attract multiple species of different tribes and subfamilies. These expected generic effects were largely supported for the targeted subfamilies, namely the Cerambycinae, Lamiinae, Spondylidinae and Prioninae.

The best represented tribe was Clytini (Cerambycinae). A total of 64 species were trapped overall, including catches in Asia (27 spp.), Europe (22 spp.) and North America (15 spp.). Two of these species were captured in both the native and invaded continents (*X. chinensis* - Asia/Europe, *N. a. acuminatus* - North America/Europe). This richness probably resulted from the presence in the blend of C6-ketol (3-hydroxyhexan-2-one) and *anti*-2,3-hexanediol. Both are known to be male-emitted attractants for a number of species in this tribe (Millar and Hanks 2017; Imrei et al. 2021). Furthermore, using these two compounds in a similar multipheromone blend in tropical China, Wickham et al. (2021) captured 26 Clytini species of which only four were in common with the present study (*Rhaphuma anongi* Gressit & Rondon, *Rhaphuma laosica* Gressit & Rondon, *Xylotrechus buqueti* [Laporte de Castelnau & Gory] and *X. chinensis*), thus suggesting an even larger potential of the blend. The Asian Clytini *X. stebbingi*, a native of the Himalayas (India, Pakistan), provides an illustrative example of the potential of detection of ‘unexpected’ cerambycid invaders, with a total of 6,089 specimens captured in the invaded countries of southern Europe. The species was trapped each year at more than 50 sites in southern France, Greece, Italy, Spain and Switzerland, but also in nurseries near Paris far from the invaded areas. However, it has apparently not invaded China. Somewhat surprisingly, the pheromone blend of this abundant invasive species has not yet been identified, although racemic 3-hydroxyhexan-2-one was suggested to be a key component of its pheromone (Rasati et al. 2021) and a number of other *Xylotrechus* species have pheromones comprised of 3-hydroxyalkan-2-ones, 2-hydroxyalkan-3-ones and/or 2,3-alkanediols (Millar and Hanks 2017). The richness in the trapped *Xylotrechus* species (19 spp.), a genus known to include a number of invasive species, is especially important for the early detection of new invaders.

In the same subfamily Cerambycinae, the tribe Callidiini was represented by 15 species trapped in Europe and five in North America, including a total of 11 species in the genus *Phymatodes* on the two continents. The very large number of captures (from ~ 2,000 to more than 15,000 individuals) of three *Phymatodes* species, two native to
Europe (*P. testaceus* and *P. alni*) and one from North America (*P. amoeneus*) and those of the closely-related European *P. sanguineum*, probably reflects the inclusion in the blend of both C6-ketol and 2-methylbutan-1-ol, known to be attractants for a number of *Phymatodes* spp. (Millar and Hanks 2017). Hanks et al. (2019) had previously trapped *P. testaceus* and *P. amoenus* in large numbers using these compounds, but also confirmed that the attraction to C6-ketol is antagonised by 2-methylbutan-1-ol for another species of *Phymatodes*, *P. aereus* (Newman) (Mitchell et al. 2011). Probably for the same reason, few *P. aereus* (14 individuals) were trapped during our study. A species of the Callidiini, *Callidium aeneum*, was represented by more than 200 individuals, but is another example of a species for which pheromones have not yet been identified. However, the congeners *C. antennatum hesperum* Casey and *C. pseudotsugae* Fisher are known to use C6-ketol along with semanopyrrole as their aggregation-sex pheromone blend (Millar et al. 2019). The C6-ketol compound is also likely to be a pheromone component for *Bethelium tillides* (Pascoe), a representative of another Cerambycinae tribe, Callidiopini, in Australia (Hayes et al. 2016), as indicated by our captures of *Bethelium* sp. in Australia and another Callidiopini, *C. flavus*, in Martinique.

In the subfamily Lamiinae, large numbers of individuals of 12 species of Monochamini in the genus *Monochamus* were trapped in Europe, Russia (Siberia), China and North America. This likely resulted from the inclusion in the blend of monochamol (2-[undecyloxy]-ethanol), known as a sex-aggregation pheromone for at least 14 *Monochamus* species in Europe (*M. galloprovincialis*; Pajares et al. 2010), North America (e.g. *M. carolinensis* [Olivier] and *M. scutellatus* [Say]; Millar and Hanks 2017) and Asia (*M. alternatus* Hope; Lee et al. 2018). The captures included species such as *M. sartor* (F.) and *M. sutor longulus* Pic for which no attractant had previously been recorded (but known for *M. sutor* [L.], Pajares et al. 2013). These results confirmed those obtained by Boone et al. (2018), who trapped six *Monochamus* species in North America and *M. alternatus* in Asia using monochamol. Given the importance of *Monochamus* beetles in vectoring the lethal pinewood nematode, the multilure blend would be useful for early detection of such invading species in ports and other high-risk sites. Additionally, another Monochamini in a different genus, *Uraecha angusta* (Pascoe), was trapped in China in high numbers, further indicating that monochamol is not restricted to the genus *Monochamus* Dejean.

In the subfamily Spondylidinae, Žunič-Kosi et al. (2019) recently showed that (S)-fuscumol, with geranylacetone as a minor component, is a sex-aggregation pheromone for a European representative of the Asemini tribe, *A. rusticus*. They also suggested that the fuscumol motif is probably shared more broadly in this subfamily. Our trapping of 15 spondylidine species supports this assumption, as the blend included fuscumol, fuscumol acetate and geranylacetone. Besides the ~ 5,000 trapped *A. rusticus* individuals, other European Asemini were captured in substantial numbers as well, including *Cephalocrius syriacus* Sharp (> 2,000 individuals), *Arhopalus ferus* (Mulsant) and *A. striatum* in Europe, but also other *Asemum* and *Cephalallus* species in China (Table 3). High numbers of captures were also noted for several species in other spondylidine tribes, such as *Spondylis buprestoides* (L.) (Spondylidini, > 2,000
individuals) in both Europe and China, for which no attractants are yet known and the alpine *Tetropium gabrieli* Weise (Tetropiini), for which Schroeder et al. (2021) recently showed an attraction to (E)-fuscumol.

In the subfamily Prioninae, the inclusion of prionic acid, originally identified as a female-produced sex pheromone of the North American species *Prionus californicus* Motschulsky (Rodstein et al. 2009), but also as an attractant for most, if not all, other North American species of *Prionus* (Barbour et al. 2011; Millar and Hanks 2017) and for the Asian Prionini *Dorysthenes granulosus* (Thomson) (Wickham et al. 2016a), resulted in substantial catches (> 4,000 specimens) of the European *Prionus coriarius* (L.). Additional captures of other Prionini, such as the Balkanic *Mesoprionus besikanus* (Fairmaire), the Asian *Prionus insularis* Motschulsky, and three Chinese species of *Dorysthenes* (*D. sternalis* [Fairmaire], *D. paradoxus* [Faldermann] and an, as yet, unidentified species), suggest a broad attractive spectrum for prionic acid in this tribe.

Despite the general efficiency of our blend, a number of species, especially those trapped with less than 50 individuals, are likely either random catches or were attracted by physical characteristics (e.g. trap shape and/or colour). Based on the previous results of Fan et al. (2019), who showed highly significant differences in the number of species trapped by multipheromone blends and unbaited traps, we did not deploy any unbaited control traps in this study which could help in clarifying this point. However, it is noteworthy that, very unexpectedly, we trapped a total of 79 lepturine species, including 49 in Europe, 12 in Asia and 18 in North America, although few pheromones are known in the subfamily Lepturinae and none of them, such as *cis*-vaccenyl acetate (Ray et al. 2011) or (R)-desmolactone (Ray et al. 2014), was included in the blend. Most of these species were caught in small numbers, consistent with random captures of individuals. Only three species (*Pachytodes erraticus* [Dalman], *Stictoleptura cordigera* [Fueßlins] and *Rutpela maculata* [Poda von Neuhaus]) out of the 44 captured in the tribe Lepturini, were represented by more than 50 individuals (Table 3). For these species, we can only speculate that the beetles were attracted to the trap colour or silhouette, rather than to the lure (see details in Cavaletto et al. 2021). Analogous results were obtained for another Lepturine tribe, the Rhaginii, where more than 600 specimens of a species considered as Holarctic, *R. inquisitor*, were caught in Europe, Asia and North America, together with the closely-related *R. japonicum* Bates trapped in China.

Attraction of these lepturines may also have been a result of the addition of high release rate ethanol and α-pinene lures to traps, rather than attraction to the blend of synthetic pheromones. Plant volatiles can, in some cases, effectively enhance the attraction of cerambycids to pheromone lures (e.g. for *Monochamus* species; Pajares et al. 2010). Indeed, plant volatiles alone, such as turpentine, have long been used as generic attractants for wood-boring insects, including some species of cerambycids, but are not as effective as pheromone-baited traps for target species. For example, Rassati et al. (2019) showed that traps deployed in Italy with a multi-lure including most of the compounds of our blend (C6-ketol, racemic 3-hydroxyoctan-2-one, *syn*-2,3-hexanediols, [*E/Z*]-fuscumol and [*E/Z*]-fuscumol acetate) caught more than twice the number of cerambycid species as ethanol-baited traps.
Our results also provide leads to possible pheromone structures in new species (see also above), building on the previously-articulated concept of “pheromone identification by proxy”, in which identification of pheromones for one species may provide leads for the identification of pheromones of related taxa (Millar et al. 2019). For the 16 species caught in numbers > 1,000 individuals, it is likely that the major component(s) of their pheromones were present in the blend. Pheromones or possible pheromones had been identified from only about half of these species (*M. alternatus*, *M. galloprovincialis* and *M. saltuarius*, *A. rusticus*, *P. amoenus* and *P. testaceus*, *P. sanguineum*, *P. coriarius*, *Xylotrechus antilope* [Schönherr]; Millar and Hanks 2017). Thus, our data represent the first possible leads to the pheromone structures for a number of cerambycine species in the tribe Clytini (*C. glabromaculatus*, > 1,000 individuals), three Lamiinae in the tribes Acanthocinini (*Leiopus femoratus* Fairmaire, > 3,000 individuals; *Leiopus nebulosus* [L.], > 1,000 individuals) and Acanthoderini (*Aegomorphus clavipes* [Schrank von Paula], > 1,000 individuals) and two Spondylidinae in the tribes Asemini (*C. syriacus*, > 2,000 individuals) and Spondylidini (*S. buprestoides*, > 2,000 individuals). It is likely that the same could be true for at least some of the remaining 46 species caught in numbers > 100 individuals (and maybe even > 50), but for which pheromones have not been formally identified. For instance, Wickham et al. (2021) considered that their substantial captures of the Clytini *R. anongi* and *R. laosica* in tropical China suggested the presence of pheromone components in the blend that they tested, which was similar to ours. Our results further support this assumption because we trapped ~ 100 *R. anongi* and > 25 *R. laosica* with traps deployed at a single site of southern China.

Trapping of some invasive species in both the native and invaded ranges revealed the potential of the multilure blend for detecting invaders. Some of these non-native species have been present for a long time in the invaded areas (e.g. the European *P. testaceus* and *A. rusticus* in North America and the North American *N. a. acuminatus* in Europe). However, the trapping of very recent invaders within and near ports-of-entry is noteworthy and is indicative of the sensitivity of the blend for early detection at low population levels. For example, the Chinese Clytini *X. chinensis* was captured in its native range around Beijing, as well as in all the scattered European areas it has invaded and established in relatively recently (2013 in Spain-Catalonia; 2017 in Greece-Crete island and 2018 in southern France-Port of Sète; https://gd.eppo.int/taxon/XYLOCH/distribution/ES). Similarly, when the 10-pheromone blend including trichoferone was deployed, the Chinese Hesperophanini *T. campestris* was trapped in both its native range in China and in the river port of Huningue (France), where this invasive species had not yet been recorded. Interestingly, despite its presumably low abundance, our trapping studies allowed us to follow the dispersal of this invading species from the port. For example, in 2019 and 2020, specimens were only detected in traps placed within the Huningue Port but, in 2021, the species was captured in traps placed within a 1 km-radius from the Port. Numerous catches of *X. stebbingi* in ports-of-entry and nurseries of northern France, far from the known invaded southern area of France, also highlighted the sensitivity of the blend for its detection.
## What possible improvements can be expected?

Is it possible and useful to continue increasing the number of pheromones included in the blend? The addition of trichoferone and the pheromone of *Aromia bungii* to the 8-pheromone blend in some field trials in France and China since 2020 resulted in relatively high numbers of captures of several *Trichoferus* species (three native European species and one native Chinese species invasive in Europe), as well as individuals of *A. bungii* in China, without reducing the trapping scope observed in nearby traps baited with the primary blend, especially the *cerambycine* *P. testaceus*. Millar et al. (2021) obtained analogous results in Pennsylvania, USA, where the responses of the major *cerambycid* species were not affected, except for the lamiine species *Sternidius alpha* (Say) whose catches were shut down by the addition of the pheromones of the two non-native species. Preliminary experiments (not detailed here) carried out in 2020 and 2021 in south-central France consisting of adding the sex-aggregation pheromone of *Rosalia alpina* (Linnaeus), an alkylated pyrone (Žunič-Kosi et al. 2017), to the 8-pheromone blend also resulted in captures of *R. alpina* without altering the *cerambycid* species richness. Tests of addition of the semanopyrrole structure (1-[1H-pyrrol-2-yl]-1,2-propanedione) could also be of interest, given that it occurs in pheromones of species from several continents. Wickham et al. (2016b) suggested that semanopyrrole may correspond to another well-conserved sex-aggregation pheromone motif within the subfamily *Cerambycinae*, being highly attractive in combination with C6-ketol for the Callidiini *Callidiellum villosulum* (Fairmaire) and as a single component for the Phoracanthini *Allotraeus asiaticus* (Schwarzer) in China. Silva et al. (2017) also identified semanopyrrole as a component of the sex-aggregation pheromone of two South American *Cerambycinae* in the tribe Elaphidiini, *Ambonus distinctus* (Newman) and *Ambonus electus* (Gahan). Recent work has also shown that 10-methyldodecanol, 11-methyltridecanol and their corresponding aldehydes may form another conserved structural motif, with examples of pheromones from South American (Silva et al. 2020) and North American *cerambycid* species (JGM and LMH, work in progress). As all these additional compounds exhibit chemical structures substantially different from the ones used in the 8-pheromone blend, it may be hypothesised that their addition will be unlikely to interfere with the attraction of species to the other eight pheromones.

For a more general approach of early detection of xylophagous invaders, targeting not only *cerambycids*, but also other groups, such as bark and ambrosia beetles (*Curculionidae*, *Scolytinae*), woodwasps (*Siricidae*) and jewel beetles (*Buprestidae*), represents a valuable opportunity. In fact, traps baited with some (e.g. Marchioro et al. 2020; Miller et al. 2022) or all (Cavaletto et al. 2020) the eight longhorn beetle pheromones used in this study, allowed us to catch a high number of other wood-boring beetle species and associated predators, especially when synergised by generic attractants, such as ethanol or α-pinene. The possibility of augmenting the blend with complementary attractants specific and/or generic to each of these groups is also of interest. Miller et al. (2016) already tested the combination of monochamol and α-pinene with ipsenol, a pheromone component of *Ips* bark beetles (*Curculionidae: Scolytinae*) and
did not observe any negative effect nor synergy on catches of Cerambycidae or on any associated species of bark beetles, weevils or bark beetle predators. However, some of the cerambycid pheromones used in the 8-pheromone blend have been shown to have either positive or negative effects on catches of bark and ambrosia beetles, depending on the species (Marchioro et al. 2020).

The position of the trap also has rather to be carefully managed. In our study, standardisation of trap position was not possible due to the different trapping locations (ports-of-entry, urban parks, forests) and the variety of environments amongst the countries included in the study. However, several recent studies have confirmed that trap position can have a considerable influence on the captures of cerambycid beetles, on a vertical gradient from the forest understorey up to the canopy (Wermelinger et al. 2007; Graham et al. 2012; Flaherty et al. 2019; Rassati et al. 2019; Miller et al. 2020; Wickham et al. 2021), as well as on a horizontal gradient between the forest edge and forest interior (Allison et al. 2019; Sweeney et al. 2020). For instance, *Monochamus maculosus* Haldeman and *M. scutellatus* were more abundant in the clearing adjacent to the forest than in the forest, whereas the contrary was observed for *M. carolinensis* and *M. titillator* (F.). As most of our field trials did not consider the upper canopy and were not deployed along forest edge-interior gradients, except for the specific 2019 experiment, it is possible that a number of species with specific types of behaviour related to these micro-environments were not trapped at all. Moreover, weather conditions during trapping periods, as well as the size of the local populations, were also likely to influence the captures.

Another important point is the colour of the trap. Most traps used in the study were black multifunnel traps (1069 out of 1289; 83%). However, Cavaletto et al. (2021), using the same 8-pheromone blend, showed that trap colour had a considerable influence on cerambycid captures in both species' richness and abundance for several subfamilies, but in different ways. For example, black traps caught significantly fewer species of Cerambycinae than yellow ones, whereas for Lamiinae, both brown and red traps caught significantly more species than black traps and colours in the portion of the visible electromagnetic spectrum (yellow, green, blue) attracted higher numbers of lepturine species than did black traps. A relationship with adult beetle behaviour could be hypothesised. Cavaletto et al. (2021) observed that the number of species of flower-visiting cerambycids was significantly lower in black traps than in yellow, blue and green ones, whereas yellow and green traps were significantly less efficient than black traps for non-flower-visiting species. Therefore, systematic tests of traps of different colours in different world regions could significantly enhance the efficiency of trapping species which respond to the multipheromone blend. Data obtained during the present study will be later merged with those of specifically-designed experiments to test for the influence of trap colour.

The impact of such trappings on local insect biodiversity could be questioned. As all specimens from non-target Coleopteran groups have not been identified yet, we cannot exclude that a few species other than cerambycids, bark and ambrosia beetles and beetle predators (clerids, trogossitids) have also been trapped in significant
numbers (> 500 ind.). However, any trapping study, like our one, is necessarily limited
in scope by cost and logistical factors. Thus, unless trappings are intensively conducted
over a whole region or country, which is very unlikely, they are likely to affect local
biodiversity in a very limited way.

In conclusion, we are delivering a database of nearly 400 species which were trapped
during the course of our multiyear field trials with the multipheromone blend, and the
two hypotheses of our study are strongly supported. First, the trapping of a species in
significant numbers on a continent effectively increased the probability that it can be
detected upon arrival in other countries/continents, as shown by the species trapped in
large numbers in both native and invaded ranges, supporting hypothesis 1. Second, the
multipheromone blend was shown to be an effective generic attractant for multiple spe-
cies from several cerambycid subfamilies, including numerous species for which phero-
mones have not yet been identified, supporting hypothesis 2. In addition, some species,
such as the lepturine species caught in large numbers, were probably trapped because
of trap colour or the host plant lure, rather than as a result of the blend composition.
However, regardless of cues used by beetles, trapping of non-native species when they
arrive at ports-of-entry has the same value for phytosanitary officials. Antagonistic ef-
fects between compounds exist, but appear to be fairly limited and so should not com-
promise the overall detection potential. Finally, further advances in the effectiveness of
detection of cerambycids by multipheromone lures can be expected as parameters, such
as trap colour and height, are optimised and as the number of pheromone components
which are found to be conserved within and across related taxa and continents expands.

## Acknowledgements

We thank Filippo Giannone, Riccardo Poloni, Kate Van Rooyen, Chantelle Kostano-
wicz, Vincent Webster, Andrej Kapla, Matic Gabor, Mischa Giasson and Cory Hughes
for technical assistance in the lab and field and for species identification. Paige Payter,
Michigan State University (MSU), installed and monitored traps in Michigan and
Page Payter and Gary Parsons (MSU) identified the captured cerambycids. We are
indebted to Fréderic Delport, François-Xavier Saintonge, Jean-Baptiste Daubrée and
all colleagues of the “Santé des Forêts” Department (DSF) and local offices (SRAL)
of the French Ministry of Agriculture for the management of the traps in France.
Marie-Pierre Dufresne from Fredon Centre - Val de Loire and Sylvain Amiot from
the Direction Patrimoine végétal et Biodiversité of Tours-Métropole helped to settle
traps in the Val de Loire area, France. We are also grateful to Eddy Poirier and Nicolas
Moulin for the management of the traps in Martinique and to the forest health team
of Vaersa and the forest management service (SOGF) of the Generalitat Valenciana
(Spain). We thank very much Xing Zhong-Ping for his help in the trappings in Yun-
nan and Anastasia Knorre for helping us with field research in the State Nature Reserve
“Stolby” (Krasnoyarsk, Russia). We also want to thank Prof. Ana Paula Ramos for
enabling a connection with the administration of the municipalities of Lisbon and
Setúbal for the trappings in Portugal. We are indebted to Eng. Rui Simão and Eng. Ana Júlia Francisco in CM- Lisboa, Dr. António Nobre from the administration board of Lisbon harbour, Eng. Sérgio Gaspar from CM-Setúbal for allowing us to conduct this work in their municipalities. The municipality of L'Argentière la Bessée provided invaluable assistance for the management of the traps in the southern French Alps. We thank Robert Haack, Nicolas Meurisse and a third anonymous reviewer for their very helpful comments and suggestions on the manuscript.

This work was essentially supported by the HOMED project (HOlistic Manage-
ment of Emerging Forest Pests and Diseases) which received funding from the Euro-
pean Union's Horizon 2020 research and innovation programme under grant agree-
ment No. 771271 (https://homed-projecteu/). We also acknowledge funding from the
European project LIFE SAMFIX (SAving Mediterranean Forests from Invasions of Xy-
losandrus Beetles and associated Pathogenic Fungi, LIFE17 NAT/IT/000609, https://
wwwlifesamfixeu/), from the EUPHRESCO project MULTITRAP ("Multi-lure and
multi-trap surveillance for invasive tree pests"). Trappings in France were supported
from funding by the French Ministry of Agriculture under the PORTRAP project
"Test de l'efficacité de pièges génériques multicomposés pour la détection précoce d'in-
sectes exotiques xylophages dans les sites potentiels d'entrée sur le territoire national"
and by the Région Centre- Val de Loire under the CANOPEE project ("Biodiversité
des insectes des canopées de chênes dépérissants" - Project No. 2018-00124136). Trap-
ping in Austria was supported by the Austrian Federal Ministry for Agriculture, For-
stry, Regions and Water Management (Research Project 101183). Trapping in China
was partially supported by a grant from the French Embassy in Beijing under the
programme CaiYuanPei. LR, JS, YQL and YY are grateful for funding by the National
Key Research and Development Program of China (Grant No 2021YFC2600400) and
the National Natural Science Foundation of China (31770687). Research conducted
in Nova Scotia, Canada, was funded by the Pest Risk Management Program of Natu-
ral Resources Canada, Canadian Forest Service. Trapping in Michigan was supported
by a grant from the Michigan Department of Agriculture and Rural Development.
Research in Siberia (Russia) was supported by Sukachev Institute of Forest SB RAS
(the basic project, grant No 0287-2021-0011) [field collection] and the Russian Sci-
ence Foundation (grant No 22-16-00075) [species identification]. AMR and EKLF
are grateful for the Robert Borcer Endowment and the Undergraduate Research Fund
of Xavier University and USDA-APHIS cooperative agreement numbers AP19PPQS
and T00C082 and AP20PPQS and T00C173. MZ and CM acknowledge funding
by the New South Wales government and Hort Innovation project 16004 NSW DPI
component of the Macadamia Integrated Pest Management. Trials in Martinique were
part of a natural site inventory funded by the DEAL Martinique (French Ministry of
Environment). The work in Slovenia was financially supported by the Slovenian Re-
search Agency (Research Core Funding P1-0255). Trapping in England was supported
by the Department for Environment, Food and Rural Affairs (Defra). JGM and LMH
gratefully acknowledge support from United States Department of Agriculture, Ani-
mal and Plant Health Inspection Service (APHIS) grants 19- to 22-8130-1422-CA.
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**Supplementary material I**

**Total trapping network**

Authors: Alain Roques, Lili Ren, Davide Rassati, Juan Shi, Evgueni Akulov, Neil Audsley, Marie-Anne Auger-Rozenberg, Dimitrios Avtzis, Andrea Battisti, Richard Belanger, Alexis Bernard, Iris Bernadinelli, Manuela Branco, Giacomo Cavaletto, Christian Cocquempot, Mario Contarini, Béatrice Courtial, Claudine Courtin, Olivier Denux, Miloň Dvořák, Jian-ting Fan, Nina Feddern, Joseph Francese, Emily K. L. Franzen, André Garcia, Georgi Georgiev, Margarita Georgieva, Federica Giarruzzo, Martin Gossner, Louis Gross, Daniele Guarneri, Gernot Hoch, Doris Hölling, Mats Jonsell, Natalia Kirichenko, Antoon Loomans, You-qing Luo, Deborah McCullough, Craig Maddox, Emmanuelle Magnoux, Matteo Marchioro, Petr Martinek, Hugo Mas, Bruno Mériguet, Yong-zhi Pan, Régis Phélut, Patrick Pineau, Ann Marie Ray, Olivier Roques, Marie-Cécile Ruiz, Victor Sarto i Monteys, Stefano Speranza, Jiang-hua Sun, Jon D. Sweeney, Julien Touroult, Lionel Valladares, Loïs Veillat, Yuan Yuan, Myron P. Zalucki, Yunfan Zou, Alenka Žunič-Kosi, Lawrence M. Hanks, Jocelyn G. Millar

Data type: site description (excel document)

Copyright notice: This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0/). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.

Link: https://doi.org/10.3897/neobiota.84.91096.suppl1