A peer-reviewed open-access journal NeoBiota 84: 107—135 (2023) doi: 10.3897/neobiota.84.90829 te NeoB 10ta https:/ / neobi ota. pen soft. net Advancing research on alien species and biological invasions Jewels on the go: exotic buprestids around the world (Coleoptera, Buprestidae) Enrico Ruzzier', Robert A. Haack*, Gianfranco Curletti?, Alain Roques*, Mark G. Volkovitsh°®, Andrea Battisti! | Department of Agronomy, Food, Natural Resources, Animals and the Environment (DAFNAE), Viale dell’ Universita 16, Legnaro, 35020 Padova, Italy 2 USDA Forest Service, Northern Research Station, 3101 Discovery Drive, Suite E Lansing, Michigan, 48910, USA 3 Museo Civico di Storia Naturale, Parco Cascina Vigna, 10022 Carmagnola, Italia 4 INRAE- Zoologie Forestiere Centre de recherche, d'Orléans 2163 Avenue de la Pomme de Pin, CS 40001 ARDON 45075 Orléans, Cedex 2, France 5 Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, 199034 St. Petersburg, Russia Corresponding author: Enrico Ruzzier (enrico.ruzzier@unipd. it) Academic editor: Marc Kenis | Received 29 July 2022 | Accepted 10 March 2023 | Published 18 May 2023 Citation: Ruzzier E, Haack RA, Curletti G, Roques A, Volkovitsh MG, Battisti A (2023) Jewels on the go: exotic buprestids around the world (Coleoptera, Buprestidae). 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: 107-135. https://doi.org/10.3897/neobiota.84.90829 Abstract Buprestidae (Coleoptera: Buprestoidea) is one of the three wood-borer beetle groups of major phytosani- tary interest worldwide, together with Cerambycidae and Scolytinae (Curculionidae). As in other beetle families, some buprestid species have been unintentionally or intentionally introduced around the world, in some cases causing significant environmental and economic damage in the invaded territories. Despite the phytosanitary relevance of the Buprestidae, information regarding the identity of exotic buprestids, their biogeographic areas of origin, introduction pathways, and larval host plants, have remained scattered in the literature. Our objective was to summarize much of the existing knowledge on these topics in the present paper. Our analysis resulted in a list of 115 exotic buprestids worldwide, representing introduc- tions both within and between biogeographic realms and corresponding to less than 1% of the known buprestid species worldwide. Invasiveness does not seem to be linked to their larval host plant preferences, as introduced species utilize 158 plant genera in 70 plant families and are equally represented in all feeding guilds (monophagous, oligophagous, and polyphagous). As trade of plants or plant parts can serve as a pathway for future introductions, the information reported in this review can help in pest risk assessment. Keywords Biodiversity, exotic species, invasive alien species, jewel beetles Copyright Enrico Ruzzier et al. This is an open access article distributed under the terms of the CCO Public Domain Dedication. 108 Enrico Ruzzier et al. / NeoBiota 84: 107-135 (2023) Introduction Buprestidae Leach, 1815 (Coleoptera: Buprestoidea), commonly known as jewel bee- tles, include more than 15,000 described species distributed in all continents except Antarctica (Bellamy 2008). The family includes six subfamilies, namely Agrilinae, Bu- prestinae, Chrysochroinae, Galbellinae, Julodinae, and Polycestinae, (Bellamy 2003). All Buprestidae are phytophagous and generally oligophagous (i.e., associated with a single plant family) as both adults and larvae (Curletti 1994). Buprestid larvae de- velop in both living and dead plant tissues; most species are internal feeders, boring or mining in roots, stems, branches, and leaves of both woody plants and herbaceous plants (Bellamy and Volkovitsh 2005), while only Julodinae possess soil-dwelling lar- vae that feed externally on roots (Koliba¢ 2000). Many buprestids, especially the wood-boring species, select dead, dying, or stressed plants for oviposition (Chamorro et al. 2015); however, some species are capable of in- festing or even prefer healthy living hosts (Carlson and Knight 1969). ‘This last group can have an important economic impact on human activities because it includes pests in orchards and tree plantations (Bonsignore et al. 2008; Hashim et al. 2018; Dawadi et al. 2019). Furthermore, buprestids can have substantial negative impacts on the natural ecosystems during outbreaks (Coleman et al. 2012; Muilenburg and Herms 2012; Sallé et al. 2014; Vuts et al. 2016; Haack and Petrice 2019). The cryptic nature of most buprestid larvae, being hidden in woody tissues and, for some species, their slow larval development due to feeding in nutrient-poor xylem (Haack and Slansky 1987), has allowed multiple species to be transported in wood products and introduced to areas far from their place of origin. Much of this dispersal has been human-mediated and related to trade (Wu et al. 2017). One of the earliest ac- counts deals with the introduction of Chalcophora detrita detrita (Klug, 1829) from the Middle East to Southern Italy by the Etruscans or the Maritime Republics (from 1000 to 2000 years ago; Biagioni et al. 2015). However, since the end of the nineteenth cen- tury the introduction rate of exotic buprestids worldwide has substantially increased in similar fashion to many other invasive forest insects (Aukema et al. 2010; Chamorro et al. 2015; Hoebeke et al. 2017; Bozorov et al. 2018; Jendek et al. 2018; Roques et al. 2020; Volkovitsh et al. 2020). Buprestidae have taken advantage of globalization with the opening of new trade routes and the increase in the number and speed of movement of goods and people (PySek and Richardson 2010). In some cases, species such as Agrilus planipennis Fair- maire, 1888 (hosts: Chionanthus and Fraxinus [main host]), A. mali Matsumura, 1924 (hosts: Cydonia, Emmenopterys, Malus [main], Prunus, Pyrus, Sorbus), and Aphanisticus cochinchinae seminulum Obenberger, 1929 (hosts: Saccharum, Tripsacum) have become invasive, causing significant damage in urban and natural forests and agriculture, and often requiring significant investments for monitoring and control (Hespenheide 2007; Bauer et al. 2008; Jones et al. 2013; Volkovitsh et al. 2020). Consequently, Buprestidae is one of the Coleoptera families of major silvicultural interest worldwide (Maynard et al. 2004; Inghilesi et al. 2013; Haack et al. 2014; MacQuarrie et al. 2020). Exotic Buprestidae around the world 109 Given this condition, great efforts have been made in the last few decades to iden- tify the main entry pathways, and to develop and implement early detection programs, effective monitoring strategies, and new tools for species identification (Meurisse et al. 2019; Poland and Rassati 2019). To date, however, little has been summarized about the main patterns of buprestid introductions worldwide, their taxonomic affinities, and their biogeographic origins. The purpose of this article is to provide a comprehensive review of natural and human-assisted translocation of buprestid species among and within various biogeo- graphic realms, describe the contribution of each realm and buprestid subfamily to this exchange of species, and provide the first comprehensive list of all introduced Buprestidae worldwide from the mid-1800s to present. Furthermore, a list of host plant associations at the genus and family level is provided, with an indication of the host range of each buprestid species. Our general aim is to provide information that can be used in pest risk assessment and invasion ecology. Methods In order to compile and then review the literature on exotic Buprestidae, we performed reiterated research in Google Scholar through the use of keywords such as “Bupresti- dae,” “introduced,” “exotic,” and “alien” and then integrated with the Boolean op- erators AND, OR, NOT and the use of “” for specific word combinations. We also obtained a considerable amount of literature that was not available in Google Scholar thanks to the support of many colleagues and buprestid specialists. Screening of the literature collected was done following the PRISMA approach and only the papers retained are cited in the Suppl. material 1 and were used for the analysis (Moher et al. 2009). The resulting reference library included papers in Chinese, English, French, German, and Italian. In the analysis, we considered only those publications where buprestids were identi- fied to species or subspecies level, and for those records published between 1850 and December 2020. In the taxonomic discussion, we did not consider the rank of subgenus. In particular, the non-native status of a given species was evaluated for its consistency throughout the reviewed literature; in case there was only a single reference publication and in the absence of any further information, the non-native status of a species was considered as valid. For each species included in the present research, we considered the most recent and comprehensive publication highlighting and explaining the non-native status as a key reference. For those buprestid species for which the literature was limited, we referred to the original faunistic record published. A full list of the Buprestidae spe- cies, associated with the reference literature, is provided in Suppl. material 1. Where the origin of a given taxon could not be assigned to a single biogeographic region, every possible area of origin was considered. The world’s biogeographic areas considered in this paper generally follow the interpretation and categorization pro- vided by Labl and Labl (2016). 110 Enrico Ruzzier et al. / NeoBiota 84: 107-135 (2023) At times it was difficult to know if an insect was firmly established in a new area or was simply intercepted at a port of entry, because papers varied in terminology and de- tail. In our dataset, when considering the species status, we have generally adopted the following categorization: A) Neonative: species native to a continent but introduced into regions other than the native ones either through natural spread indirectly favored by human activities (climate change, habitat change) or through accidental human- mediated introductions; B) Established: non-native species that sustain self-replacing populations over several life cycles (inclusive of single specimens collected in the wild away from potential entry points); C) Invasive: a non-native species established in natural or semi-natural ecosystems or habitat, which has impact and threatens native biological diversity; D) Intercepted: insects detected during inspection procedures or similar situations where no reproducing population is known to occur; E) Intention- ally introduced: species that have been actively introduced in areas other than their native range with a specific purpose, such as biological control of invasive plants; F) Unclear: all species for which the status is unclear (e.g., apparently extinct adventive populations, species described in areas where that specific genus does not occur, species record vague without any specific detail, mislabeling and misidentification). Data collected were organized in an Excel spreadsheet including the following information, organized by columns: subfamily, tribe, genus, species (full name plus author), biogeographic region of origin, biogeographic region of detection, status, and host plants. Detection region and host plant were associated with a specific column called references, which included all relevant information used to recover the data. Each species could have multiple entries (rows) in cases of multiple introduction events in different biogeographic areas, or in situations where the origin of the species was not reducible to a single biogeographic region. In the case of single introductions of widely distributed species in which it was clear the biogeographic region of origin of the insects, we considered only the record for that specific region. The taxonomy of plant genera and families used in the paper is based on the information available on the “Plants of The World Online” database (https://powo.science.kew.org/). Analyses and graphics were realized using the R software (version 4.1.2). Host plant preference was defined in the categories: monophagous (for buprestids feeding only on plant species of the same genus), oligophagous (buprestids feeding on different plant genera within the same host family), polyphagous (buprestids feeding on plant species from different host families). Results Faunistic part Our literature review identified 162 events of buprestid introductions among and within biogeographic regions that involved 115 distinct taxa (Suppl. material 1). The taxa included 44 species in the subfamily Agrilinae (tribes Agrilini, Aphanisti- cini, Coraebini, and Tracheini) (Table 1), 51 species of Buprestinae (tribes Anthaxiini, Exotic Buprestidae around the world 111 Buprestini, Chrysobothrini, Curidini, Melanophilini, and Nascionini) (Table 2), 16 species of Chrysochroinae (Chalchophorini, Chrysochroini, Dicercini, Sphenopterini, Paraleptodemini, and Poecilonotini) (Table 3), and 6 species of Polycestinae (tribes Acmaeoderini, Polycestini, Prospherini and Ptosimini) (Table 4). No species of the subfamilies Galbellinae and Julodinae were recorded as introduced. The revision of all published records revealed that the buprestid species involved in introductions either within or between biogeographical realms constitute only 0.76% of all known bupres- tid species worldwide. The analysis showed that the introduction of exotic Buprestidae included all bio- geographic realms (with the obvious exclusion of the Antarctic realm), including in- troductions both among and within the realms (Fig. 1). In addition, our analysis re- vealed that the Nearctic and Palearctic bioregions contributed the greatest number of introduced species (90 in total) and also the most distinct introduction events (72.4% combined). ‘The realm that was the source for the highest number of buprestids intro- duced elsewhere was the Palearctic, with 52 out of approximately 2,500 native species (2.1%), followed by the Nearctic (38 out of ~800; 4.8%), the Indomalayan (13 out of ~2,800; 0.5%), the Neotropical (13 out of ~3,700; 0.4%), the Australasian (10 out of ~1,600; 0.6%), the Afrotropical (7 out of ~3,800; 0.2%), and the Oceanian (2 out of ~70; 2.9%). The analyses between the number of buprestid species per biogeographic realm and the number of species introduced elsewhere from each realm did not show any significant statistical relation (t = -0.10389, df = 5, p-value = 0.9213). Palearctic and Nearctic were the two regions with the highest number of intro- duced species (Fig. 1) but, despite somewhat similar environments, climate, and flora, there were substantial differences in the patterns of inter- and intra-biogeographic realm introductions (Table 5). In the case of intra-realm introductions, Palearctic and Nearctic showed a similar number of species (23 vs 20) despite the fact that the genera contributing to more than 50% of total introductions were, at least in part, different: Agrilus (9 species) and Buprestis (4 species) in the Palearctic, and Chrysobothris (9) and Agrilus (6) in the Nearctic. By contrast, when considering introductions between the two realms, it was possi- ble to observe a strong imbalance with 9 exotic species recorded in the Palearctic com- pared with 25 in the Nearctic. Furthermore, Agrilinae represented the majority of the exotic buprestids in the Nearctic, while Buprestinae were dominant in the Palearctic. With respect to all buprestid species considered introduced worldwide, we found 41 cases where the species were considered established, 43 cases as interceptions at en- try points, 32 cases where the status was unclear, and 22 cases of neonative species. We also classified 13 introductions where the species became invasive, and 6 cases where species were intentionally introduced. For the 41 cases of establishment, Buprestinae was the most represented subfamily, with 23 records subdivided among the genera Anthaxia (1 species), Buprestis (8 species), Belionota (1 species), Chrysobothris (6 species), and Trachypteris (1 species). Agrilinae accounted for 14 establishments, represented by 10 species of Agrilus, 1 Diphucrania, and 2 Trachys. The subfamilies Chrysochroinae and Polycestinae were involved in only a limited number of establishments, i.e., 1 Steraspis, 1 Prospheres and 2. Acmaeodera. 112 Table |. Subfamily Agrilinae: species list, biogeographic realms concerned, status, and larval host plants. Enrico Ruzzier et al. / NeoBiota 84: 107—135 (2023) * species confused with Agrilus coxalis Waterhouse, 1889 in the literature. Species Biogeographic realm Status Plant host genera origin introduction Agrilus angustulus (Iliger, 1803) Palearctic Palearctic Unclear Corylus, Ostrya (Betulaceae); Fagus, Castanea, Quercus (Fagaceae) Agrilus anxius Gory, 1841 Nearctic Nearctic Neonative Betula (Betulaceae) Agrilus auriventris Saunders, 1873 Australasian, | Oceanian Invasive Citrus (Rutaceae) Indomalayan Agrilus auroguttatus Schaeffer, 1905* Nearctic Invasive Quercus (Fagaceae) Agrilus bilineatus (Weber, 1801) Palearctic Established Castanea, Quercus (Fagaceae) Agrilus biguttatus (Fabricius, 1776) Palearctic Australasian | Intercepted | Fagus, Castanea, Quercus (Fagaceae); Tilia (Malvaceae); Populus (Salicaceae); Ulmus (Ulmaceae) Agrilus cavatus Chevrolat, 1838 Nearctic Neotropical Unclear Rhus (Anacardiaceae); Acaciella (Fabaceae) Agrilus convexicollis Redtenbacher, 1849 Palearctic Palearctic Neonative Euonymus (Celastraceae); Philadelphus (Hydrangeaceae); Fraxinus, Ligustrum, Olea, Syringa (Oleaceae) Agrilus cuprescens (Ménétriés, 1832) Palearctic Nearctic Established Rosa, Rubus (Rosaceae) Agrilus cyanenoniger Saunders, 1873 Palearctic Palearctic Neonative | Croton (Euphorbiaceae); Quercus (Fagaceae) Agrilus cyanescens (Ratzeburg, 1837) Palearctic Palearctic, Unclear, Lonicera, Symphoricarpos (Caprifoliaceae); Nearctic Established Rhamnus (Rhamnaceae) Agrilus derasofasciatus Lacordaire, 1835 Nearctic Non-native Vitis (Vitaceae) Agrilus difficilis Gory, 1841 Nearctic Established Gleditsia (Fabaceae); Zanthoxylum (Rutaceae) Agrilus extraneus Fisher, 1933 Oceanian Established Argemone (Papaveraceae) Agrilus fleischeri Obenberger, 1925 Nearctic Intercepted Populus, Salix (Salicaceae) Agrilus furcillatus Chevrolat, 1835 Nearctic, Nearctic Intercepted Pinus (Pinaceae); Zea (Poaceae); Coffea Neotropical (Rubiaceae); Salix (Salicaceae) Agrilus graminis Kiesenwetter, 1857 Palearctic Palearctic Neonative | Alnus, Corylus, Ostrya (Betulaceae); Euonymus (Celesteraceae); Castanea, Quercus (Fagaceae); Acer (Sapindaceae); Viburnum (Viburnaceae) Agrilus hyperici (Creutzer, 1799) Palearctic | Australasian, | Intentionally Hypericum (Hypericaceae) Nearctic introduced Agrilus kaluganus Obenberger, 1940 Palearctic Palearctic Neonative Corylus (Betulaceae) Agrilus livens Kerremans, 1892 Indomalayan | Palearctic Unclear Citrus (Rutaceae) Agrilus mali Matsumura, 1924 Palearctic Palearctic Neonative Cydonia, Malus, Prunus, Pyrus, Sorbus (Rosaceae); Emmenopterys (Rubiaceae) Agrilus nicolanus Obenberger, 1924 Palearctic Neonative Quercus (Fagaceae); Ulmus (Ulmaceae) Agrilus occipitalis (Eschscholtz, 1822) Australasian, | Oceanian Invasive Citrus (Rutaceae) Indomalayan, Palearctic Agrilus pilosovittatus Saunders, 1873 Palearctic Nearctic Established Wisteria (Fabaceae) Agrilus planipennis Fairmaire, 1888 Palearctic Nearctic, Invasive, Chionanthus, Fraxinus (Oleaceae) Palearctic Neonative Agrilus prionurus Chevrolat, 1838 Nearctic Neonative Chionanthus (Oleaceae); Sapindus (Sapindaceae) Agrilus pulchellus Bland, 1865 Nearctic Nearctic Intercepted Chrysothamnus sp., Erigeron (Asteraceae); Amsinkia (Boraginaceae); Celtis (Cannabaceae); Quercus (Fagaceae); Sphaeralcea (Malvaceae); Allionia, Boerhavia (Nyctaginaceae) Agrilus ribesi Schaefer, 1946 Nearctic Invasive Ribes (Grossulariaceae) Agrilus sinuatus (Olivier, 1790) Palearctic Nearctic Established Crataegus, Malus, Prunus, Pyrus, Sorbus (Rosaceae) Agrilus smaragdifrons Gang|bauer, 1890 Palearctic Nearctic Established Ailanthus (Simaroubaceae) Agrilus sulcicollis Lacordaire, 1835 Palearctic Nearctic Established Fagus, Castanea, Quercus (Fagaceae) Agrilus subrobustus Saunders, 1873 Indomalayan, Nearctic Established Albizia (Fabaceae) Palearctic Aphanisticus antennatus Saunders, 1873 Indomalayan,} Unclear Not available Neotropical Exotic Buprestidae around the world LS Species Biogeographic realm Status Plant host genera origin introduction Aphanisticus cochinchinae seminulum Indomalayan | Nearctic, Invasive Saccharum, Tripsacum (Poaceae) Obenberger, 1929 Neotropical, ane Oceanian Coraebus andrewesi Obenberger, 1922 Indomalayan, | Neotropical Unclear Not available Palearctic Coraebus rubi (Linnaeus, 1767) Palearctic Neonative Rosa, Rubus (Rosaceae) Coraebus undatus (Fabricius, 1787) Palearctic Palearctic | Intercepted Diospyros (Ebenaceae); Castanea, Fagus, Quercus (Fagaceae) Diphucrania viridipurpurea Carter, 1924 | Australasian | Palearctic | Established Not available Hylaeogena jureceki Obenberger, 1941 Neotropical | Afrotropical, | Intentionally Dolichandra (Bignoniaceae) Australasian | introduced Leiopleura carbonata (LeConte, 1860) Neotropical Unclear Not available Leiopleura otero (Fisher, 1935) Neotropical Unclear Not available Lius poseidon Napp, 1972 Neotropical | Oceanian | Intentionally} Miconia, Chaetogastra (Melastomataceae) introduced Trachys minutus (Linnaeus, 1758) Palearctic Nearctic Established Corylus (Betulaceae); Sorbus (Rosaceae); Salix (Salicaceae), Ulmus (Ulmaceae) Trachys troglodytiformis Obenberger, 1918] Palearctic Nearctic Established Althea, Hibiscus, Malva (Malvaceae) Table 2. Subfamily Buprestinae: species list, biogeographic realms concerned, status, and larval host plants. Species Fon ode Gory © Laporte 1) Anthaxia laticeps Abeille de Perrin, 1900 Biogeographic realm introduction Palearctic Status Neonative Plant host genera Picea, Pinus (Pinaceae) Pinus (Pinaceae) Palearctic Palearctic Anthaxia proteus Saunders, 1873 Anthaxia salicis (Fabricius, 1776) Palearctic Unclear Nearctic Established Pinus (Pinaceae) Castanea, Quercus (Fagaceae); Salix (Salicaceae); Acer (Sapindaceae) Cobosina willineri (Cobos, 1972) Neotropical | Neotropical Neonative Not available Buprestis apricans Herbst, 1801 Nearctic Neotropical Established Pinus (Pinaceae) Nearctic Buprestis aurulenta Linnaeus, 1767 Australasian, Neotropical, Palearctic, Oceanian Intercepted, Established, Unclear, Established Thuja, Juniperus (Cupressaceae); Abies, Picea, Pinus, Pseudotsuga (Pinaceae) Palearctic Nearctic Palearctic Buprestis dalmatina Mannerheim, 1837 Buprestis decora Fabricius, 1775 Buprestis haemorrhoidalis Herbst, 1780 Nearctic, Palearctic Neotropical, Palearctic Afrotropical, Australasian, Nearctic, Neotropical, Palearctic Intercepted Neonative Established Unclear, Intercepted, Established, Unclear, Unclear Pinus (Pinaceae) Pinus (Pinaceae) Callitris (Cupressaceae); Abies, Picea, Pinus (Pinaceae) Bruprestis humeralis Klug, 1829 Palearctic Palearctic Neonative Pinus (Pinaceae) Buprestis lineata Fabricius, 1781 Nearctic Australasian, Nearctic, Neotropical, Palearctic Intercepted, Neonative, Established, Unclear Pinus (Pinaceae) Nearctic Neotropical Established Buprestis maculativentris Say, 1825 Buprestis maculipennis Gory, 1841 Buprestis novemmaculata Linnaeus, 1767 | _ Palearctic Buprestis salisburyensis Herbst, 1801 Nearctic Australasian Afrotropical, Indomalayan, Nearctic, Neotropical, Palearctic Nearctic Intercepted Unclear, Unclear, Intercepted, Established, Established Established Abies, Picea, Pinus (Pinaceae) Taxodium (Cupressaceae); Pinus, Tsuga (Pinaceae) Larix, Picea, Pinus (Pinaceae) Pinus (Pinaceae) 114 Species Trachykele blondeli Marseul, 1865 Belionota prasina (Thunberg, 1789) Merimna atrata (Gory & Laporte, 1837) | Australasian Biogeographic realm Status introduction Nearctic Australasian, | Intercepted, Palearctic Non-native Australasian, Afrotropical, Established, Intercepted, Established, Established, Intercepted Indomalayan | Australasian, Nearctic, Neotropical Palearctic Oceanian Intercepted Enrico Ruzzier et al. / NeoBiota 84: 107-135 (2023) Plant host genera Calocedrus, Chamaecyparis, Cupressus, Juniperus, Thuja (Cupressaceae) Anacardium, Mangifera (Anacardiaceae); Delonix, Pithecellobium (Fabaceae); Casuarina (Casuarinaceae); Hopea (Dipterocarpaceae); Ceiba (Malvaceae) Eucalyptus (Myrtaceae) Chrysobothris adelpha Gemminger & Harold, 1869 Chrysobothris acutipennis Chevrolat, 1835 | Nearctic, Neotropical Established Neotropical Chrysobothris affinis (Fabricius, 1794) Nearctic Oceanian Intercepted Palearctic Australasian Intercepted Prospis (Fabaceae); Carya (Juglandaceae); Amelanchier (Rosaceae) Ebenopsis, Leucaena (Fabaceae) Pistacia (Anacardiaceae); Alnus, Betula, Carpinus, Corylus, Ostrya (Betulaceae); Cornus (Cornaceae); Arbutus (Ericaceae); Cercis, Gleditsia, Robinia (Fabaceae); Castanea, Fagus, Quercus (Fagaceae); Punica (Lythraceae); /uglans (Juglandaceae); Tilia (Malvaceae); Ficus, Morus (Moraceae); Eucalyptus (Myrtaceae); Fraxinus (Oleaceae); Cedrus (Pinaceae); Platanus (Platanaceae); Crataegus, Malus, Prunus, Pyrus, Rosa, Sorbus (Rosaceae); Populus, Salix (Salicaceae); Acer (Sapindaceae); Ulmus (Ulmaceae) Chrysobothris analis LeConte, 1860 Chrysobothris cavifrons Deyrolle, 1864 Nearctic Nearctic Established Australasian Palearctic Intercepted Rhus (Anacardiaceae); Celtis (Cannabaceae); Diospyros (Ebenaceae); Cercis, Ebenopsis, Haematoxylum, Leucaena, Mimosa, Parkinsonia, Prosopis (Fabaceae); Carya, Juglans (Juglandaceae); Coccoloba (Polygonaceae); Prunus (Rosaceae); Citrus (Rutaceae); Sapindus (Sapindaceae); Ulmus (Ulmaceae) Not available Chrysobothris cerceripraeda Westcott & Thomas, 2015 Nearctic Nearctic Unclear Not available Chrysobothris chrysonota Deyrolle, 1864 | Australasian Chrysobothris costata Kerremans, 1895 Chrysobothris costifrons Waterhouse, 1887 Palearctic Intercepted Oceanian Oceanian Invasive Nearctic Nearctic Neonative Not available Intsia (Fabaceae); Citrus (Rutaceae) Quercus (Fagaceae) Chrysobothris dorsata (Fabricius, 1787) Chrysobothris ellyptica Deyrolle, 1864 Chrysobothris femorata (Olivier, 1790) Afrotropical, | Palearctic Unclear Palearctic Palearctic Intercepted Nearctic Australasian, Intercepted Oceanian, Palearctic Acacia, Ceratonia (Fabaceae) Not available Liquidambar (Altingiaceae); Carpinus (Betulaceae); Celtis (Cannabaceae); Diospyros (Ebenaceae); Cercis (Fabaceae); Castanea, Quercus (Fagaceae); Carya, Juglans (Juglandaceae); Tilia (Malvaceae); Fraxinus (Oleaceae); Platanus (Platanaceae); Amelanchier, Crategus, Cydonia, Malus, Prunus, Sorbus (Rosaceae); Populus, Salix (Salicaceae); Acer (Sapindaceae); Ulmus (Ulmaceae) Chrysobothris igniventris Reitter, 1895 Palearctic Nearctic Intercepted Larix, Pinus (Pinaceae) Chrysobothris indica Castelnau & Gory, 1837 Established Oceanian Indomalayan Terminalia (Combrentaceae); Shorea (Dipterocarpaceae); Acacia (Fabaceae); Myristica (Myristicaceae); Mimusops (Sapotaceae) Chrysobothris knulli Nelson, 1975 Nearctic Nearctic Established Acacia (Fabaceae) Exotic Buprestidae around the world 115 Species Biogeographic realm Status Plant host genera introduction Chrysobothris mali Horn, 1886 Nearctic Nearctic Intercepted | Alnus, Betula, Corylus (Betulaceae); Arbutus, Arctostaphylos (Ericaceae); Pickeringia, Prosopis, Wisteria (Fabaceae); Fagus, Quercus (Fagaceae); Ribes (Grossulariaceae); Juglans (Juglandaceae); Persea (Lauraceae); Liriodendron (Magnioliaceae); Ficus (Moraceae); Eucalyptus (Myrtaceae); Platanus (Platanaceae); Ceanothus, Rhamnus (Rhamnaceae); Adenostoma, Cercocarpus, Cotoneaster, Crataegus, Cydonia, Malus, Oemleria, Photinia, Prunus, Pyracantha, Pyrus, Rhaphiolepis, Rosa, Rubus, Sorbus (Rosaceae); Populus, Salix (Salicaceae); Acers, Aesculus (Sapindaceae); Ulmus (Ulmaceae) Chrysobothris octocola LeConte, 1858 Nearctic Oceanian Established | Acacia, Parkinsonia, Prosopis (Fabaceae); Prunus (Rosaceae); Salix (Salicaceae) Chrysobothris pupureoplagiata Nearctic Nearctic Intercepted | Canotia sp. (Celasteraceae), Psorothamnus Scheaffer, 1904 (Fabaceae) Chrysobothris quadriimpressa Gory & Nearctic Nearctic Neonative Liquidambar (Altaginaceae); Quercus Laporte, 1837 (Fagaceae); /uglans (Juglandaceae); Sapindus (Sapindaceae) Chrysobothris rotundicollis Gory & Neotropical Unclear Ebenopsis (Fabaceae); Larix, Pinus Laporte, 1837 (Pinaceae) Chrysobothris rugosiceps Melsheimer, 1845] Nearctic Nearctic Neonative Castanea, Quercus (Fagaceae) Chrysobothris sexpunctata, Fabricius 1801] Neotropical | Neotropical Established Not available Chrysobothris superba Deyrolle, 1864 Australasian Palearctic Intercepted Not available Chrysobothris tranquebarica Neotropical Nearctic Unclear Casuarina (Casuarinaceae); Conocarpus (Combrentaceae); Cassia (Fabaceae); Pinus (Pinaceae); Rhizophora (Rhizophoraceae) Chrysobothris trinervia (Kirby, 1837) Nearctic Nearctic Intercepted | Larix, Picea, Pinus, Pseudotsuga (Pinaceae) Australasian (Gmelin, 1790) Anilara hoscheki Obenberger, 1916 Palearctic Intercepted Not available Melanophila consupta LeConte, 1857 Oceanian Non-native Calocedrus (Cupressaceae); Eucalyptus (Myrtaceae); Pinus Pseudotsuga (Pinaceae) Phaenops cyanea (Fabricius, 1775) Palearctic Nearctic Intercepted Abies, Larix, Pinus (Pinaceae) Phaenops drummondi (Kirby, 1837) Nearctic Nearctic, Intercepted Abies, Cedrus, Larix, Picea, Pseudotsuga Palearctic (Pinaceae) Trachypteris picta decostigma Palearctic Neotropical Established Populus, Salix (Salicaceae) (Fabricius, 1787) Nascio vetusta (Boisduval, 1835) Australasian | Australasian Intercepted Eucalyptus, Metrosideros (Myrtaceae); Xanthorrhoea (Asphodelaceae) With respect to the 43 cases where the buprestids were apparently only intercepted, the Buprestinae had the highest number of interceptions worldwide (28), which included 24 species. Ihe most commonly intercepted genus was Chrysobothris (14 species), followed by Buprestis (6 species). There were 6 cases of intercepted Agrilinae, involving 4 species of Agrilus and 1 Coraebus. For both Chrysochroinae and Polycestinae there were multiple single species interceptions. For 28 species among Agrilinae, Buprestinae, Chrysochroinae and Polycestinae it was not possible to assign their status to any of the existing categories; therefore, they were classified as “unclear.” We recognize that many more species of Bu- prestidae have been intercepted at ports throughout the world, but in almost all cases these datasets are not available to the public and therefore could not be considered in our paper. 116 Enrico Ruzzier et al. / NeoBiota 84: 107—135 (2023) Table 3. Subfamily Chrysochroinae: species list, biogeographic realms concerned, status, and larval host plants. Species Status Plant host genera origin introduction Chalcophora angulicollis (LeConte, 1857) Nearctic Nearctic, Unclear Abies, Pinus, Pseudotsuga (Pinaceae) Palearctic Chalcophora japonica (Gory, 1840) Intercepted Pinus (Pinaceae) Chalcophora virginiensis (Drury, 1770) Nearctic | Neotropical, Unclear Taxodium (Cupressaceae); Pinus (Pinaceae) Palearctic Cyphogastra foveicollis (Boisduval, 1835) Australasian | Palearctic | Intercepted Not available Dicerca moesta (Fabricius, 1794) Palearctic Nearctic, Intercepted, Abies, Pinus, Picea (Pinaceae) Palearctic Unclear Dicerca furcata (Thunberg, 1787) Intercepted Betula (Betulaceae) Dicerca tuberculata (Laporte & Gory, 1837)} Nearctic Neotropical | Non-native Tsuga (Pinaceae) Euchroma gigantea (Linnaeus, 1758) Neotropical | Neotropical Unclear Ceiba, Pachira, Pseudobombax (Malvaceae) Lampetis bahamica Fisher, 1925 Neotropical | Neotropical | Intercepted Not available Lampetis corruscans (Carter, 1924) Australasian | Australasian Unclear Not available Lampetis fastuosa (Fabricius, 1775) Unclear Areca (Arecaceae); Acacia (Fabaceae); Eucalyptus (Myrtaceae); Tectona (Lamiaceae) Lamprodila festiva (Linnaeus, 1767) Palearctic Palearctic | Neonative Callitris, Chamaecyparis, Cupressus, Juniperus, Platycladus, Tetraclinis, Thuja (Cupressaceae); Ziziphus (Rhamnaceae); Tamarix (Tamaricaceae) Lamprodila vivata (Lewis, 1893) Intercepted Cryptomeria, Chamaecyparis, Juniperus (Cupressaceae) Sphenoptera jugoslavica Obenberger, 1926 Intentionally Centaurea (Asteraceae) introduced Steraspis squamosa (Klug, 1829) Afrotropical,| Palearctic | Established, Tamarix (Tamaricaceae) Palearctic Neonative Table 4. Subfamily Polycestinae: species list, biogeographic realms concerned, status, and larval host plants. Species Biogeographic realm Status Plant host genera origin | introduction Acmaeodera bipunctata Palearctic | Palearctic Neonative Euphorbia (Euphorbiaceae); Juniperus, Thuja (Olivier, 1790) (Cupressaceae); Ficus (Moraceae); Abies, Cedrus, Larix, Pinus (Pinaceae) Acmaeodera flavomarginata| Nearctic, | Neotropical Established Acacia, Prosopis (Fabaceae); Diospyros (Ebenaceae) (Gray, 1832) Neotropical Acmaeodera neoneglecta Nearctic Fisher, 1949 Prospheres aurantiopictus (Laporte & Gory, 1837) Ptosima undecimmaculata (Herbst, 1784) Palearctic Nearctic Nearctic Intercepted Acacia, Ebenopsis, Leucaena, Prosopis, Sophora (Fabaceae); Carya (Juglandaceae); Ulmus (Ulmaceae) Australasian | Australasian Established Araucaria (Araucariaceae); Pinus (Pinaceae) Intercepted Mangifera (Anacardiaceae); Ceratonia (Fabaceae), Crataegus, Malus, Prunus, Pyrus (Rosaceae); Citrus (Rutaceae); Vitis (Vitaceae) Among all the taxa investigated, 22 species were considered as neonatives. ‘There were 10 Agrilinae (9 Agrilus and 1 Coraebus); 9 Buprestinae (2 Anthaxia, 1 Cobosina, 3 Buprestis, and 3 Chrysobothris); 2 Chrysochroinae (1 Steraspis and 1 Lamprodila), and 1 Polycestinae (1 Acmaeodera). Neonative species were recorded almost exclu- sively in the Northern Hemisphere, with 15 species in the Palearctic and 6 in the Exotic Buprestidae around the world 117 ! Afro ; goanta” 'ropica, Figure |. World map illustrating the number of introduced species of Buprestidae within and between biogeographic realms (above) and graphical representation of the exchanges (below), with the thickness of the arrows directly proportional to the number of introduction events. The length of the colored arc of each realm corresponds to the total number of introduced species, either in or out. 118 Enrico Ruzzier et al. / NeoBiota 84: 107-135 (2023) Table 5. Comparison between buprestid introductions within and between the Nearctic and Palearctic realms, with details on the number of species within each genus. within Palearctic within Nearctic Palearctic to Nearctic Nearctic to Palearctic 9 Agrilus 9 Chrysobothris 12 Agrilus (one species 3 Buprestis 4 Buprestis 6 Agrilus intentionally introduced) 2 Chalcophora 3 Anthaxia 2 Buprestis 3 Buprestis 1 Agrilus 2 Coraebus 1 Acmaeodera 2 Trachys 1 Chrysobothris 1 Acmaeodera 1 Chalcophora 1 Anthaxia 1 Phaenops 1 Chrysobothris 1 Phaenops 1 Chalcophora 1 Trachykele 1 Dicerca 1 Chrysobothris 1 Steraspis 1 Dicerca 1 Lamprodila 1 Lamprodila 1 Phaenops 1 Ptosima 1 Sphenoptera (intentionally introduced) Nearctic realm. Agrilus was the most represented genus in the Palearctic with 7 species, while Chrysobothris was the most represented genus in the Nearctic with 3 species. A single species of Cobosina was the only example of a neonative taxon in the Neotropic realm. All 13 cases of invasive buprestids are species of Agrilinae and Buprestinae. These species became invasive once introduced to the Nearctic, Oceanian and Neotropical realms. There were 6 species of invasive Agrilinae (5 Agrilus and 1 Aphanistichus), and only two invasive Buprestinae in the genera Belionota and Chrysobothris. Six cases of intentionally introduced taxa were found, representing 4 species in the genera Agrilus (Agrilini), Sphenoptera (Sphenopterini), Hylaeogena and Lius (Tracheini). These species were introduced into the Nearctic, Afrotropical, and Australasian realms. Larval host plants The analysis of larval host plants for all Buprestidae introduced worldwide identified 158 different plant genera within 70 families (3 Gymnosperms and 67 Angiosperms), with only a few introduced buprestids without host information (Tables 1-4). The exotic Buprestidae included sets of species with wide variation in the range of their larval hosts, varying from highly polyphagous on non-phylogenetically related plant families to monophagous on a single plant genus. Larval host specificity (i.e., monophagous, oligophagous and polyphagous) of introduced Buprestidae is equally distributed among the subfamilies (Kruskal-Wallis chi-squared = 1.2007, df = 2, p- value = 0.5486) (Table 6). The larval host families most represented were Pinaceae (60 host records), Rosaceae (52), Fabaceae (49), Fagaceae (36), and Cupressaceae (24), which together accounted for 52% of all host records (Table 7). Considering introductions within and between biogeographic realms, it emerged that the most common genera of host plants varied greatly among world biogeographic realms, both in abundance and diversity (Table 8). Exotic Buprestidae around the world 119 Table 6. Number of introduced species with different levels of larval host-use specialization by bupres- tid subfamilies. Monophagous Agrilinae 13 Buprestinae 11 Chrysochroinae Polycestinae Total 29 Oligophagous Polyphagous Unknown 9 17 e: 11 20 8 3 3 0 6 0 24 46 16 Table 7. Summary table of the main plant families and genera exploited as larval host plants by in- troduced Buprestidae by subfamily. Numbers between parenthesis refers to the number of records, not distinct species. Plant Genera Pinus (27), Abies (8), Picea (8), Plant Families Pinaceae (60) Larix (7) Rosaceae (52) Prunus (9), Matus (7), Sorbus (7), Pyrus (5) Fabaceae (49) Acacia (9), Prosopis (6), Ebenopsis (4), Leucaena (3) Fagaceae (35) Quercus (18), Castanea (11), Fagus (6) Cupressaceae (23) Juniperus (5), Thuja (4) Betulaceae (18) Corylus (6), Betula (4), Alnus (3) Salicaceae (16) Salix (9), Populus (7) Buprestid subfamilies Buprestinae (45), Chrysochroinae (9), Polycestinae (4), Agrilinae (1) Buprestinae (30), Agrilinae (15), Polycestinae (7) Buprestinae (31), Polycestinae (14), Agrilinae (20), Buprestinae (13), Polycestinae (2) Chrysochroinae (11), Buprestinae (10), Polycestinae (2) Buprestinae (9), Agrilinae (8), Chrysochroinae (1) Buprestinae (10), Agrilinae (5), Polycestinae (1) Buprestid genera Buprestis (21), Chrysobothris (10), Phaenops (8), Chalcophora (5) Chrysobothris (30), Agrilus (10), Ptosima (4), Acmaeoderella (3) Chrysobothris (29), Acmaeodera (7), Acmaeoderella (6), Agrilus (3) Agrilus (17), Chrysobothris (11), Coraebus (3) Lamprodila (10), Trachykele (5), Buprestis (2), Acmaeodera (2) Chrysobothris (9), Agrilus (7) Chrysobothris (7), Agrilus (4), Trachypteris (2) Discussion The low introduction rate, 0.76% compared for example to the 2.17% out of ~ 6000 taxa of Curculionidae Scolytinae (Lantschner et al. 2020), indicates a general low propensity for Buprestidae to be introduced by humans, either directly or indirectly. In support of this contention is the high number of single buprestid introductions (i.e, one species introduced only once and only in a single biogeographic realm), with respect to the total number of introduction events. In addition, the invasiveness does not seem to be linked to larval host plant preferences, as introduced species are included in all feeding guilds (monophagous, oligophagous, and polyphagous). The genera Agrilus (Agrilinae: Agrilini), Buprestis (Buprestinae: Buprestini), and Chrysobothris (Buprestinae: Chrysobothrini) would seem to be more predisposed to introduction events than other genera, possibly owing to both their morphological and biological traits. Agrilus are generally small in size and univoltine (Solomon 1995; Chamorro et al. 2015). They infest mostly live plants and signs of their presence are difficult to detect prior to adult emergence and host dieback. Therefore, several Agrilus species have likely been moved over time through trade of live plants, such as orna- mentals or nursery stock, as well as through domestic and international movements 120 Enrico Ruzzier et al. / NeoBiota 84: 107-135 (2023) Table 8. Summary table of the most common plant genera exploited as larval host plants by buprestid species introduced either within or between biogeographic realms. Origin — Introduction realm Afrotropical — Palearctic Australasian — Australasian Australasian — Oceanian Australasian — Palearctic Indomalayan — Afrotropical Indomalayan — Australasian Indomalayan — Palearctic Indomalayan — Nearctic Indomalayan — Neotropical Indomalayan — Oceanian Nearctic — Australasian Nearctic — Nearctic Nearctic — Oceanian Nearctic — Palearctic Nearctic — Neotropical Neotropical — Afrotropical Neotropical — Australasian Neotropical — Nearctic Neotropical — Neotropical Neotropical — Oceanian Palearctic — Afrotropical Palearctic — Australasian Palearctic — Indomalayan Palearctic — Nearctic Palearctic — Neotropical Palearctic — Oceanian Palearctic — Palearctic Oceanian — Oceanian Most common larval host plant genera exploited by those species with a narrow host range Angiosperms: Acacia, Ceratonia, Tamarix Angiosperms: Eucalyptus Angiosperms: Citrus Angiosperms: Anacardium, Casuarina, Ceiba, Delonix, Hopea, Mangifera, Pithecellobium Angiosperms: Anacardium, Casuarina, Ceiba, Delonix, Hopea, Mangifera, Pithecellobium Angiosperms: Anacardium, Casuarina, Ceiba, Delonix, Hopea, Mangifera, Pithecellobium Angiosperms: Citrus Angiosperms: Albizia, Anacardium, Casuarina, Ceiba, Delonix, Hopea, Mangifera, Pithecellobium, Saccharum, Tripsacum Angiosperms: Anacardium, Casuarina, Ceiba, Delonix, Hopea, Mangifera, Pithecellobium, Saccharum, Tripsacum Angiosperms: Citrus Gymnosperms: Pinus Angiosperms: Acacia, Juglans, Prosopis, Sapindus, Ulmus Gymnosperms: Pinus, Pseudotsuga Angiosperms: Amelanchier, Carya, Prosopis, Prunus, Salix Gymnosperms: Pinus, Pseudotsuga Gymnosperms: Abies, Pinus, Pseudotsuga Gymnosperms: Pinus Angiosperms: Dolichandra Angiosperms: Dolichandra Gymnosperms: Pinus Angiosperms: Acacia, Ceiba, Diospyros, Ebenopsis, Leucaena, Pachira, Prosopis, Pseudobombax Angiosperms: Miconia, Tibouchina Gymnosperms: Picea, Pinus Angiosperms: Castanea, Fagus, Populus, Quercus, Tilia Ulmus Gymnosperms: Larix, Picea, Pinus Angiosperms: Salix Gymnosperms: Abies, Larix, Picea, Pinus Gymnosperms: Picea, Pinus Angiosperms: Citrus Angiosperms: Castanea, Quercus Gymnosperms: Abies, Picea, Pinus Angiosperms: Argemone, Citrus, Intsia of recently cut logs and manufactured wood products, especially when not debarked. The example of the emerald ash borer, A. planipennis, is remarkable in the number of pathways (e.g., logs, firewood, nursery stock) by which it has moved in North America (Herms and McCullough 2014; Haack et al. 2015). By contrast to Agrilus, most Buprestis and Chrysobothris species have longer lar- val developmental periods; they can infest both living, stressed, and dead plants; and they typically tunnel in host xylem, including both sapwood and heartwood (Solomon 1995; Evans et al. 2004). As a consequence of this multi-year develop- mental period deep inside wood, infestations are generally difficult to detect until adult emergence. Although most species oviposit in bark cracks or under the bark, a few species can oviposit directly on exposed wood (xylem). Moreover, once lar- vae have entered the xylem, the presence of bark is no longer required. Therefore, Exotic Buprestidae around the world 121 introductions of these species can result from movement of logs and milled wood products either with or without bark. Given the relatively low number of exotic buprestids investigated and the hetero- geneity of the sources consulted, it has not been possible to delineate an exact tempo- ral trend for worldwide buprestid introductions, although it seems evident that most species were likely introduced before the 1970s, with very few ever intercepted during port surveys. This condition likely reflects the lack of strict phytosanitary regulations in the early 1900s (Eschen et al. 2015). In addition, international trade among European countries and their overseas colonies likely facilitated the movement of some species early on, as well as later during the two world wars. Examples come from Buprestis au- rulenta Linnaeus, 1767 and Buprestis novemmaculata Linnaeus, 1767, two species in- troduced in all biogeographic realms edging the Atlantic Ocean, including Azores and Canary Islands, two important bridgeheads in the trade routes between Europe and the Americas (Steckley 1972; Crosby 1984; de Avilez Rocha 2019). Similarly, sugar cane cultivation is associated with the worldwide spread of Aphanisticus cochinchinae seminulum Obenberger, 1929 (Zack et al. 2009). In more recent times, many examples of intracontinental spread of buprestids have been reported, especially for certain species of Agrilus, Anthaxia, and Chryso- bothris (Westcott 2005; Fagerstrém et al. 2009; Izzillo 2013; Orlova-Bienkowskaja and Volkovitsh 2015; Westcott et al. 2018; Curletti and Ranghino 2020). Rapid in- tracontinental spread probably reflects greater connectivity among trading partners as well as increased speed of transport, especially in the European Union and North America. Range expansion of some neonative species has apparently resulted from human-caused climate and environmental changes, such as for Agrilus graminis Kiesen- wetter, 1857; Agrilus nicolanus Obenberger, 1924; Buprestis dalmatina Mannerheim, 1837; Lamprodila festiva (Linnaeus, 1767). In the USA, the southward and westward spread of the native birch specialist Agrilus anxius Gory, 1841 has been attributed to the widespread planting of ornamental birch trees in many areas outside the native range of North American birch species (Muilenburg and Herms 2012). It is interesting to note that most neonatives have caused little damage, although there are a few exceptions often associated with the inadvertent movement of infested live plants. For example, the introduction of Agrilus planipennis from Eastern Asia to the Moscow area resulted in severe mortality of ash (Fraxinus) trees in European Russia (Orlova-Bienkowskaja 2014); however, it is also plausible that Agrilus planipennis could have been introduced in Moscow on ash nursery stock imported from North America (Haack et al. 2015). Another example is Lamprodila festiva (Linnaeus, 1767), a southern European — circum-Mediterranean species, which has expanded its distribution north- ward and eastward, benefiting from extensive plantings of its host plants (Cupressaceae) as ornamental plants in private and public gardens (Nitzu et al. 2016; Rabl et al. 2017; Volkovitsh and Karpun 2017; Ruicanescu and Stoica 2019). Similarly, Agrilus mali Matsumura, 1924, an eastern Palearctic species, has taken advantage of expanding cul- tivation of Rosaceae fruit trees and patches of natural forest as a springboard to spread westward in the Palearctic (Volkovitsh et al. 2020; Zhang et al. 2021; Lu et al. 2022). 122 Enrico Ruzzier et al. / NeoBiota 84: 107-135 (2023) Only four buprestid species have been intentionally introduced as biological control agents against invasive weeds in North America, South Africa, and Austral- ia. Sphenoptera jugoslavica Obenberger, 1926 has been intentionally introduced and successfully established in the western USA where it is used to control the invasive plant Centaurea diffusa Lam. (Asteraceae) (Lang et al. 1998); Agrilus hyperici (Creutzer, 1799) was introduced in the USA and Australia where it provides efficient control of invasive Hypericum species (Hypericaceae); while Hylaeogena jureceki Obenberger, 1941 was introduced and established with different rates of success in South Africa and Australia to control the invasive plant Dolichandra unguis-cati (L.) L.G.Lohmann (Bignoniaceae) (King et al. 2011; Snow and Dhileepan 2014). The Neotropical Lius poseidon Napp, 1972 was instead intentionally introduced to Hawai'i to control the invasive Miconia crenata (Vahl) Michelang (Melastomataceae); however, in Hawai’i the species naturally became a biocontrol agent of another invasive plant Chaetogastra herbacea (DC.) PJ.EGuim. & Michelang. (Melastomataceae) (Culliney and Nagamine 2000; Conant and Hirayama 2001; Conant et al. 2013). Conclusion The family Buprestidae is highly diverse with a global distribution defined by multiple abiotic and biotic factors, including human-mediated introductions. Although some biological and ecological traits, such as apparent obligate outbreeding and obligate maturation feeding for all buprestids, can serve as barriers to successful establishment, the opening of new continental and intercontinental trade routes as well as the ever-in- creasing volume and types of goods and plants traded increases the risk of future intro- ductions or passive diffusion of more buprestid species. With respect to climate change and the widespread practice of introducing exotic plants for ornamental, agricultural, and forestry purposes around the world, it will be important to identify possible new introduction pathways for exotic Buprestidae along with pest risk assessments. In this regard, more research is needed on buprestid taxonomy and ecology, together with training and funding of more buprestid specialists. ‘Ihe development of new technolo- gies for rapid species identification, either morphological or molecular, would be very useful for the management of this important group of plant pests, which are becoming of increasing economic importance worldwide. Acknowledgements The present paper resulted from activities framed into the HOMED project (HOlistic Management of Emerging forest pests and Diseases), which was funded from the Eu- ropean Union's Horizon 2020 research and innovation programme under grant agree- ment no. 771271. The authors thank Eduard Jendek (Faculty of Forestry and Wood Sciences, Czech University of Life Sciences) for providing important literature useful for this contribution. Exotic Buprestidae around the world 125 References Alfieri A (1976) The Coleoptera of Egypt. Mémoires de la Société Entomologique d’Egypte 5: 1-361. 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Entomologia Generalis 41(3): 257-266. https://doi.org/10.1127/ entomologia/2021/0974 Zhu H-B, Gao Z-X, Han Z-C, Lu J, Chen J-D (2008) The jewel beetles (Buprestidae) inter- cepted on timbers from Papua New Guinea. Chinese Bulletin of Entomology 45: 647-650. Supplementary material | Systematic list of all Coleoptera Buprestidae introduced around the world be- tween 1850 and 2020 Authors: Enrico Ruzzier, Robert A. Haack, Mark G. Volkovitsh Data type: table (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 (ODDbL) 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.84.90829.suppl1