Volume 10  Numbers 2 & 3  2009  SPECIAL knowledge - conservation - sustainability

Journal of Life on Earth

INVASIVE ALIEN SPECIES The importance of taxonomy in responses to invasive alien species H. Douglas1, P.T. Dang2, B.D. Gill1, J. Huber3, P.G. Mason2, D.J. Parker1, B.J. Sinclair1.

Abstract. Invasive alien species (IAS) cause major economic losses and threaten biodiversity worldwide. Taxonomic information is essential to initiating defence against IAS, including early detection and identification of new IAS outbreaks. We demonstrate how current IAS strategies (i.e. prevention of new invasions, early detection and rapid response to new invaders and management of established invaders) require strong taxonomic infrastructure. Examples are given from terrestrial molluscs (Gastropoda) and such as the Gypsy Moth, Lymantria dispar Linnaeus (Lepidoptera: Lymantriidae), disease vector flies of the families Culicidae and Calliphoridae (Diptera:), Sirex noctilio (Fabricius) (: Siricidae) woodwasps and Dendroctonus bark beetles and the cabbage seedpod weevil, Ceutorhynchus obstrictus (Marsham) (Coleoptera: Curculionidae), that demonstrate the role of taxonomy in IAS response. Further, we show through the example of the Emerald Ash Borer, Agrilus planipennis Fairmaire (Coleoptera: Buprestidae), how international co-operation is essential to identify initial outbreaks of IAS. Such international co-operation is especially important because few countries can fund comprehensive pest identification organizations alone.

Au t h o r s ’ Ad d r e s s e s : The most important role of taxonomists is to produce the 1 Entomology, Ottawa Plant & Seed Laboratories, Canadian Food Inspection Agency, Bldg. 18, 960 Carling Ave., Ottawa, Ontario, K1A names, descriptions and keys to taxa and the predictive 0C6, Canada classifications that enable inference about IAS. This 2 Agriculture and Agri-Food Canada, Research Centre, K.W. Neatby primary taxonomic literature, and the secondary biological Building, 960 Carling Ave., Ottawa, ON, K1A 0C6, CANADA and ecological literature, based on the names occurring 3 Canadian Forest Service, Natural Resources Canada, c/o AAFC in the primary literature and authoritative identifications Research Centre, K.W. Neatby Building, 960 Carling Ave., Ottawa, ON, K1A 0C6, Canada by competent taxonomists, are the primary benefits of taxonomy. Taxonomic revisions, catalogues and checklists Introduction of various taxa for particular regions or the world provide a Invasive alien species (IAS) cause major economic losses and species diversity baseline from which predictions about IAS ecosystem damage worldwide. In the USA alone introduced can be made. The primary taxonomy literature, which tests pests cost agriculture about $13.5 billion per year and taxonomic and phylogenetic hypotheses using morphological, forestry about $2.1 billion (Pimentel et al. 2005). IAS were molecular sequence and other biological and ecological data identified as the primary cause for endangerment for about is scattered in thousands of publications worldwide. DNA half of all endangered species in USA (Wilcove et al. 1998). barcode data represent a powerful new tool for species Recent increases in world trade increase the likelihood of new hypothesis testing, but sound scientific conclusions require IAS introductions and effective quarantine actions against taxonomic investigation involving multiple evidence sources them require strong taxonomic knowledge of native and exotic (e.g. Schmidt and Sperling 2008). More comprehensive, biota (Lyal et al. 2008). Taxonomic information is essential synthetic works such as generic revisions containing illustrated for initiating defence against IAS by providing rapid, accurate identification keys for countries, biogeographical regions or, identification of IAS new to an area. Taxonomy is integral to all even the world, are required, but fewer are produced because stages of IAS response including prevention of new invasions, of the time required to gather and synthesize the specimens early detection of and rapid response to new invaders and and information for study. These primary sources provide management of established invaders. Once invaders arrive information for all secondary taxonomic works, such as and are detected and identified, further taxonomic research is catalogues, checklists, distribution maps and, eventually usually needed for short and long term response. Here, research (as tertiary works), taxonomic or biological treatments of on ecological interactions among the pest, host and ecosystem species or genera that affect humans, ecosystems or particular are all indispensable in planning initial defensive strategies and commodities e.g., wheat or cattle. integrated control measures such as biological control. Such taxonomic tools and research capability are essential “Taxonomy is an information science responsible for erecting to all stages of IAS response. This includes not only initial formal classifications and maintaining names that make detection and identification of IAS, but also response and possible the storage, retrieval, organization and communication control, which requires understanding the ecology of invaders of billions of facts about millions of species” (Wheeler 2009). in their native and introduced ranges. Such understanding is This practical statement was part of his two part definition of needed to determine whether and how control measures should taxonomy (the other part treated the underlying evolutionary be initiated. Possible responses include localized intensive theories and the taxonomic conventions). Since Linnaeus, 250 eradication efforts, quarantine measures and biological control. years of taxonomic research have provided a large knowledge Here we provide examples of how taxonomy has contributed base about the diversity of living organisms from which to preventing, detecting or correcting short and long term useful, accurate predictions can be made about the possible response to outbreaks of Gastropoda (snails) and Lepidoptera, origins of various IAS. Diptera, Coleoptera and Hymenoptera. Further, we show how

92 TROPICAL CONSERVANCY international co-operation was essential to initially identify commodities to allow, and from where. With a comprehensive the Emerald Ash Borer beetle, Agrilus planipennis Fairmaire set of sound PRAs, countries would be better able to protect (Coleoptera: Buprestidae). Such international co-operation themselves against imports of commodities infested with IAS. is especially important because all insect collections have limited content, few countries can fund comprehensive pest Biological control from a forestry perspective identification organizations, and even in those countries the One further point is that IAS would be more common and number of taxonomists is declining (Haas and Häuser 2005; destructive without the insect and spider predators or Grant 2009). parasitoids (especially Hymenoptera and Diptera) that control them. Hymenoptera, in particular, can be specific and useful in Taxonomic names of accurately defined species are necessary to biological control, but implementation is hampered by a lack access all published information on species and when combined of taxonomic knowledge (Grissell 1999; Huber 2009). The with biological information allow further syntheses that provide taxonomy of predators and parasitoid groups is as important others, including quarantine officials, decision makers and non- as that of groups containing potential IAS. taxonomists, with needed information about invasive species. Examples of such synthetic works for IAS in forests are: Classical biological control (moving natural enemies to new Hendrickson (2002) for Canada, Moore (2005) and FAO (2009) areas where they do not occur) has been extremely important for the world. From a forestry perspective, the examples below for controlling IAS in forests. Many exotic sawflies and wood show the importance of taxonomy in determining what IAS wasps (Hymenoptera: Symphyta) have been reduced to minor are and what groups are likely to cause future problems. What or insignificant pest status by introduced parasitoids (e.g., taxonomy cannot usually do is predict specific details, such as ichneumon or chalcid wasps) or pathogens (e.g. nematodes, which particular species will become IAS, and where and when viruses), as shown by examples from Canada (McGugan and that might happen. However, Pest Risk Assessments (PRA) Coppel 1962; Langor et al. 2002) and New Zealand (Nuttall based on sound taxonomy can often predict which species are 1989). However, before biological control agents are deliberately Gypsy Moth, likely to be invasive pests once introduced beyond their native introduced into a new area they need to be correctly identified, Lymantria dispar L., Male, female and ranges. Such PRAs are part of the information that regulatory necessitating collaboration with taxonomists. Taxonomists, in caterpillar (Courtesy organizations use to make decisions about which import turn, may obtain useful biological information from biological of NRCan).

B I O D I V E R S I T Y 1 0 ( 2 & 3 ) 2 0 0 9 93 control workers helping them resolve taxonomic problems, known taxonomically, thus providing an easily accessible especially distinguishing cryptic species (Huber et al. 2001). information source on their systematics, hosts and biology. In contrast, almost nothing has been done with biological g y p s y m o t h control using parasitoids, the emphasis being placed instead Lymantria dispar Linnaeus (Lepidoptera: Lymantriidae) is an on competitors, predators and pathogens (Safranyik et al. important invasive pest in eastern North American deciduous 2002). Research on biological control organisms could reduce forests, and was considered by Lowe et al. (2004) to be among the risk that these Dendroctonus species pose. the 100 most invasive species worldwide. Ferguson (1978) revised the 46 North American species of Lymantriidae. He w o o d w a s p s reported 30% as pests, possibly the highest percentage for The family Siricidae (Hymenoptera), commonly called any group of insects, and found that all species in the tribe woodwasps, occurs only in the Northern Hemisphere except for Lymantriini in the Nearctic region were introduced from the one genus in central Africa. Sirex noctilio (Fabricius) is a pest Palearctic region. In contrast, he estimated that Afro-Eurasia of pine trees (Pinus spp.) that is native to the Palearctic region. has as many as 150 native Lymantria Hübner (Lymantriini) Several species of introduced pines grown in the Southern species. Pogue and Schaefer (2007) taxonomically treated Hemisphere are thus at risk to any Sirex spp. introduced from 31 species of Lymantria from Asia, where the genus is most the Northern Hemisphere. Sirex noctilio destroyed Pinus diverse, and considered all potentially invasive. radiata plantations in New Zealand and elsewhere until biocontrol agents were discovered and introduced. Five of the Despite there being no native Lymantria in the Nearctic ten biological Hymenoptera control agents established and region, 26 species of native Nearctic parasitoids and 17 insect spread, effectively reducing populations of S. noctilio to low predators have been found feeding on Gypsy Moth (Griffiths levels (Nuttall 1989). A nematode was also found but arrived and Quednau 1984), complementing the effects of nine naturally inside its Sirex host instead of being deliberately parasitoid species introduced from Europe. Gypsy Moth is introduced. All the parasitic wasp biocontrol agents obviously also attacked by a bacterium, Bacillus thuringiensis Berliner had to be accurately identified by taxonomists before being serovar kurstaki (Bacillaceae), two introduced pathogens: a released in New Zealand. North America is the latest region virus, nuclear polyhedrosis virus and a fungus, Entomophaga into which S. noctilio was accidentally introduced. In response, maimaiga Humber, Shimazu and Soper (Zygomycetes: a taxonomist is now revising the species of the entire family Entomophthoraceae). All agents, whether native or introduced, Siricidae for the New World, so that this or any other alien initially required identification and description by taxonomists. siricid species can be distinguished from all the native ones. The effects of the introduced parasitoids and pathogens on native biodiversity are largely unknown but examples of negative From a conservation perspective, Pinus species are exotic effects have been found. At least one parasitic fly, Compsilura to the southern hemisphere, so the establishment of pine concinnata (Meigen) (Diptera: Tachinidae), introduced from plantations there initially increased biodiversity but may Europe to control the Browntail Moth, Euproctis chrysorrhoea also have decreased it by displacing native species. Thus, an Linnaeus (Lepidoptera: Lymantriidae), has a wide host range unintentionally introduced pest of the introduced pines could and has been blamed for reducing numbers of non-target hosts be seen as a beneficial agent favouring native plant species including native North American Saturniidae (Lepidoptera) by reducing the deliberately introduced species considered (Boettner et al. 2000; Kellog et al. 2003). In this case releasing useful to humans. Intentional introduction of biological biological control organisms without sufficient knowledge of control agents of S. noctilio ultimately re-established the the biological characteristics and potential of the parasitoid desired situation from a forest resource perspective, i.e., species damaged native biodiversity. the continued presence of exotic pines in plantations in the Southern Hemisphere. Depending on one’s viewpoint any b a r k b e e t l e s (c o l e o p t e r a : curculionidae : s c o l y t i n a e ) of the introduced species could be viewed as deleterious or There are 19 species of Dendroctonus Erichson (Coleoptera: beneficial to the biodiversity of an area. In the New Zealand Curculionidae) bark beetles in the Nearctic region and only example, the introduced biocontrol agents of S. noctilio are 2 native to the Palaearctic. Dendroctonus species include perceived by most as beneficial to humans, regardless of their major forest pests, as shown by the over 50 pages of literature effect in maintaining intentionally introduced alien species references in the world catalog of Scolytinae (Wood and that may reduce the native biodiversity of the country. Bright 1992 and supplements up to 1999). Several are potential IAS, and D. valens LeConte (the Red Turpentine It is preferable that taxonomic syntheses of genera containing Beetle) is already an IAS in China, introduced accidentally potential IAS precede introductions, but such syntheses may in the 1980’s from North America. Governments in Eurasia be initiated in response to an IAS arriving in a new region, and North Africa clearly need to prevent Dendroctonus or as is being done for Sirex in North America. Regardless, it is other Scolytinae (e.g., Ips DeGeer species) from entering and clear that the ability of quarantine officers to recognize actual establishing in their countries. Comprehensive taxonomic and or potential IAS is seriously compromised without taxonomic bibliographic compilations on Scolytinae by two taxonomists, work to provide the necessary background knowledge in a S.L. Wood and D. E. Bright, Jr., have made Scolytinae well- useful form, e.g., identification keys, catalogues.

94 TROPICAL CONSERVANCY Biological control from and the Habrocytus sp. was Stenomalina gracilis (Walker), an agriculture perspective but new surveys determined that of the three species only S. Taxonomy plays a key role in biological control of gracilis had established (Gibson et al. 2006b). Furthermore, agricultural pests. The example of Cabbage Seedpod more extensive taxonomic review of the parasitoids associated Weevil, Ceutorhynchus obstrictus (Marsham) (Coleoptera: with Cabbage Seedpod Weevil in North America revealed Curculionidae), in Canada illustrates how sound taxonomic that most species had been misidentified and included several research can explain perceived failure, clarify taxonomic species not previously known to be associated with Cabbage names and enhance understanding of biodiversity. This Seedpod Weevil (Gibson et al. 2005, 2006a & b). It was invasive alien species, native to Europe, was first recorded determined that all previous identifications of T. perfectus in North America in 1931 at the port city of Vancouver in North America were misidentifications of T. lucidus and (McLeod 1962), and has since spread to other parts of those of M. morys were misidentifications of an undescribed western and eastern North America (Baker 1936; Hagen species, subsequently described as M. moryoides Gibson 1946; Crowell 1952; Walz 1957; Anonymous 1977; Dolinski (Gibson et al. 2005). Collaborative taxonomic research also 1979; Boyd and Lentz 1994; Buntin et al. 1995; Cárcamo et resulted in description of a new parasitoid species associated al. 2001; Dosdall et al. 2002; Brodeur et al. 2001; Mason et with cabbage seedpod weevil in North America (Gibson et al. 2004). The cabbage seedpod weevil is a serious pest of al. 2005), development of new keys for the identification canola and rapeseed (Brassica napus L. and Brassica rapa of Trichomalus Thomson spp. (Muller et al. 2007) and L.) (Brassicaceae) in North America (McCaffrey 1992; Mesopolobus Westwood spp. (Bauer et al. 2007) associated Buntin et al. 1995; Cárcamo et al. 2001; Dosdall et al. 2002; with Ceutorhynchus species of European origin, and an Kuhlmann et al. 2002; Mason et al. 2004), dispersing at a illustrated key to the parasitoids of the cabbage seedpod rate of approximately 55 km per year (Dosdall et al. 2002) weevil in North America (Gibson et al. 2006b). Furthermore, and it is expected to eventually spread throughout the entire new parasitoid-host associations were determined (Gibson et canola-growing region of western Canada. Under climate al. 2005; Gillespie et al. 2006; Bauer et al. 2007; Dosdall et change scenarios, it is predicted that an average temperature al. 2007; Muller et al. 2007; Mason et al. unpublished). increase of 3oC would result in the expansion of the range of C. obstrictus into the northern areas of the Canadian prairies Parasitism levels of the Cabbage Seedpod Weevil remain low (Olfert and Weiss 2006). (<10%) even in British Columbia where the pest has been present since the 1930’s. Furthermore, abundance of the parasitoid Soon after Cabbage Seedpod Weevil was discovered, several species varies from region to region. It is clear that native parasitoids (Hymenoptera: Chalcidoidea) were reared from parasitoids have not adapted and are unlikely to adapt to Cabbage this host and identified as various taxa, including the European Seedpod Weevil and regulate pest populations. Thus, importation species Trichomalus perfectus (Walker) and Mesopolobus of parasitoid species important in regulating Cabbage Seedpod morys (Walker) (). These parasitoid species were Weevil in the area of origin, such as T. perfectus or M. morys, thought to have been introduced accidentally along with the remains the best option to control Cabbage Seedpod Weevil. pest (see review by Gibson et al. 2005). A classical biological However, native North American parasitoid species that attack control release against Cabbage Seedpod Weevil was also made this invasive alien species threaten native Ceutorhynchus weevils in 1949 in British Columbia with the release of three species that may also be hosts. High populations of Cabbage Seedpod of Pteromalidae (Hymenoptera) that were then identified as Weevil would provide a resource that would enable increases Trichomalus fasciatus (Thompson) [= Trichomalus lucidus in numbers of native parasitoids that could increase mortality of (Walker)], Xenocrepis pura Mayr [= Mesopolobus morys native Ceutorhynchus weevils, a parasitoid spillback (see Kelly (Walker)] and Habrocytus sp. (McLeod 1962). et al. 2009). The weevil’s invasion of the canola growing regions of In clarifying the status of the parasitoid complex associated Alberta, Saskatchewan, Ontario and Quebec in the late 1990’s with Cabbage Seedpod Weevil, taxonomy played a critical led to new surveys to identify the parasitoids present, what role in establishing the actual species that parasitize this pest in their impact on Cabbage Seedpod Weevil was, and whether North America, The correction of the parasitoid identifications the introduction of additional parasitoids from Europe would was made possible because of improved taxonomy of the taxa be warranted. In particular, the apparent ineffectiveness of T. in Europe in the 1960’s (particularly Graham’s 1969 revision perfectus as a biological control agent of the Cabbage Seedpod of Pteromalidae of Northwestern Europe) and deposition of Weevil in North America, although it is highly effective in authoritatively identified specimens of European species in the Europe (typically occurring at infection prevalence levels CNC which were then available for comparison. Furthermore, >50% and up to 90%) indicated the need for taxonomic this example demonstrates that initial identifications of IAS examination of the parasitoid complex associated with the pest/parasitoids are often misidentifications because of the Cabbage Seedpod Weevil. Resulting taxonomic study of immediate lack of taxonomic knowledge or comparative voucher specimens of the three species introduced into specimens. Therefore, initial identifications of the pest and British Columbia in 1949 revealed that the species introduced biological control agents should be reviewed as taxonomic as T. fasciatus was actually Trichomalus perfectus (Walker) knowledge improves in both the area of origin and the

B I O D I V E R S I T Y 1 0 ( 2 & 3 ) 2 0 0 9 95 region of introduction. The most recent taxonomic work also preserved voucher specimens quickly and efficiently across corrected the assumption that introduced European species several international borders. Once this pest had been identified, of parasitoids had failed for biological control of cabbage the published literature could be reviewed to summarize the seedpod weevil. Finally, our understanding of the diversity of known distribution and biology (Wei et al. 2004) and research parasitoids associated with ceutorhynchine weevils in North could begin to address additional questions on the biology America was improved. and to search for potential biological control agents (Liu et al. 2003; Bauer et al. 2008). Unfortunately, the rise of national Agrilus planipennis and the necessity for and international regulations that prevents the free exchange or international cooperation in taxonomy shipment of material among taxonomists threatens to undermine Invasive alien species pose a special challenge to taxonomists such cooperation, which imperils the flora and fauna that many and quarantine services because they can suddenly appear of these regulations were designed to protect. If Jendek had in port or industrial areas with no indication of their origins. not resided in Slovakia, the identification of EAB could have The first hurdle is to decide if the specimen represents an been problematic. While North Americans were struggling to undescribed native species, which can be a challenge, or if it identify EAB in 2002, at least one South Asian government is a described exotic species (a bigger challenge), or perhaps was busy passing a Biological Diversity Act in response to the an undescribed exotic species (the biggest challenge). Most United Nations Convention on Biological Diversity (CBD). The identification specialists are familiar with their local and aim of the CBD is to protect the genetic resources of sovereign regional faunas, or with a modest range of taxa on a global nations and to prevent the commercial exploitation of native basis. When an unknown species arrives from an unknown flora and fauna. While biological diversity acts are valuable area, identifiers are unlikely to be able to make an accurate tools to protect wildlife and reduce biopiracy, they can prevent identification due to a lack of appropriate literature and the legitimate exchange of specimens across borders, penalizing voucher specimens. The free exchange of information and both domestic and foreign scientists alike (Prathapan et al. specimens among international experts is critical for the rapid 2006, 2008). Canada and USA are not immune, as Canadian and accurate identification of new IAS. scientists, trying to ship dead insects to colleagues in the U.S. for identification, have had shipments stopped and been notified The discovery of Agrilus planipennis or the Emerald Ash Borer by the U.S. Fish and Wildlife Service that the genus and species (EAB) as it is called in North America is a good example that names for all specimens must be listed for the shipment to be highlights the importance of international cooperation among delivered. If this basic information was known, there would taxonomists in the battle against invasive alien species. EAB have been no reason to ship the specimens in the first place. In was first reared in North America from dying ash trees in the Canada, the Wild and Plant Protection and Regulation Detroit area of Michigan in May and June of 2002 (Haack of International and Interprovincial Act (WAPPRIITA) makes et al. 2002; Cappaert et al. 2005). Specimens were initially it an offence to violate any foreign law related to wildlife. identified as anAgrilus species by an entomologist at Michigan For those countries that declare all species of flora and fauna State University who suspected that it might be exotic because protected, possession of any specimens, including dead fruit it did not resemble any native species. Samples were sent to flies or cockroaches, becomes an offence. The implementation the United States Department of Agriculture and to leading of these well intentioned laws needs to be tempered with the buprestid taxonomists in the U.S. for further identification. realization that the free exchange of voucher specimens among A consensus emerged that it was likely an Asian Agrilus, but taxonomists is critical in protecting native flora and fauna from with no voucher specimens in North American collections it invasive alien species. was not possible to determine the actual species. An email with digital images was subsequently sent by Richard Westcott (Oregon Department of Agriculture) to Eduard Jendek (Slovak Taxonomy and terrestrial mollusc Academy of Sciences, Bratislava) who had been working on pests (Gastropoda) in Canada Asian Agrilus species for many years. Jendek made a tentative Molluscs are the second largest animal phylum on earth after identification of A. planipennis based on the email, which he the . The terrestrial snails and slugs of Canada confirmed once he received pinned specimens by mail (Haack (land snails hereafter) are poorly known. There have been few et al. 2002). After the Michigan specimens were identified, faunistic surveys of land snails across Canada and there are beetles from neighbouring Windsor, Ontario, submitted to probably many undescribed native species. Of the 200 species the Canadian Food Inspection Agency were forwarded to known to occur in Canada about 50 remain undescribed. Also, Westcott who confirmed the presence of this pest in Canada. 10–25% of Canada’s fauna are exotic species (Forsyth 2009, personal communication). In British Columbia, almost one- The identity of this new IAS was solved by Jendek who third of all land snail species are exotic (Forsyth 2004). The had previously been able to study the type specimen of A. biggest barrier to responding to these exotic species in Canada planipennis in the Muséum national d’Histoire Naturelle in is that there are no terrestrial malacologists in Canadian federal Paris and had published on it (Jendek 1994). The accurate or provincial laboratories. The Canadian Food Inspection identification of EAB in the U.S. and Canada required the Agency, who regulate land snail imports and gastropods as collaboration of international experts and the ability to send plant pests, often seek assistance outside their own diagnostic

96 TROPICAL CONSERVANCY service for mollusc interceptions. Canada has terrestrial mollusc a serious threat to the endemic passerine fauna (Fessl et al. collections at the Canadian Museum of Nature (Ottawa), The 2006; Wiedenfeld et al. 2007). Traditional taxonomy played Royal British Columbia Museum (Victoria), the Royal Ontario a key role in determining that P. downsi was not native to the Museum (Toronto) and several smaller museums. Because of islands and that it comprised a single invasive species (Sinclair its poorly known fauna, the lack of taxonomic researchers, unpublished data), which has been verified in recent genetic outdated literature, small collections and increasing global studies of this parasitic fly (Dudaniec et al. 2008). trade, Canada is vulnerable to new introductions of exotic terrestrial molluscs. However, recently there have been several Conclusions promising developments in diagnostic keys for terrestrial As demonstrated in the above examples, taxonomy is required molluscs. The Canadian Food Inspection Agency is funding at all stages of interaction with IAS insect and terrestrial mollusc a field-guide to the native and introduced molluscs of eastern pests. Without adequate taxonomic infrastructure, detections Canada (to be published in 2009). There is also a recent and accurate identifications are delayed or not possible, resulting publication on the land snails of British Columbia (Forsyth in delayed or misguided responses and controls. Ideally, the 2004). These guides allow genus or species level identification systematics of pest groups and their natural agents of biological of most established and commonly intercepted land snail pests, control should be well understood before introduction, whether but without taxonomic support how can Canada protect itself accidental or intentional, as we demonstrate above. This is against invasive exotic snails and slugs? essential for timely detection, identification and management of IAS. Such studies should include ‘rare pests’ that do not Many terrestrial snails and slugs damage agriculture and normally cause measurable damage in their native ranges. forestry, the environment and human health (Forsyth 2004). This is because invasive species often have novel ecological Snails are known vectors of serious animal pathogens. Without characteristics or lack the parasitoids, predators and other control curated collections, faunistic surveys, libraries, taxonomists agents in their introduced range that naturally suppress their and identifiers, Canada will be unaware of new introductions populations in their area of origin. Efforts to understand insect and will only be capable of reaction to invasive species long faunas in this detail are only practicable through international after establishment. The CFIA regularly intercepts many collaboration among scientists. Exchange of specimens for non-indigenous snails but there is a limit to the effectiveness study could be facilitated by minor amendments to the existing of quarantine when so little is known of the Canadian fauna laws and regulations of various national governments. and those of its trading partners. Governments and universities would benefit from collaborating to support taxonomic In the absence of robust taxonomies of pest groups, good research in terrestrial malacology. The resulting survey of the taxonomic research capacity is essential to IAS response. Specimen terrestrial land snail biodiversity of Canada would make this collections, libraries and laboratories staffed with qualified country better prepared for new invasive introductions. Here, scientists, identifiers and technicians allow both provisional as elsewhere, detection of biological invasions requires a specimen identification even without a complete literature and the comprehensive literature and personnel who are able to use it. capacity to conduct taxonomic research. This research should be both proactive, to prepare for expected biological invasions, and Taxonomy and invasive alien Diptera emergency-based, in response to recent invasions or interceptions. Numerous invasive flies (Diptera) in the families Agromyzidae, The examples discussed above demonstrate the need for continued Anthomyiidae, Calliphoridae, Culicidae, Drosophilidae, taxonomic work in response to ongoing invasions and the costs of Muscidae, Phoridae, Psilidae, Sarcophagidae and Tephritidae implementing biological control without sufficient involvement of are known, but our attention is mostly focussed toward pests taxonomic researchers. and disease vectors. Detailed studies of the devastating effects of introduced vector mosquitoes and the diseases they transmit Acknowledgments have been well documented (Lounibos 2002). In addition, there We thank M. Damus, G. Gibson and an anonymous reviewer have been extensive investigations into the taxonomic genetic for their helpful comments. structure of disease vector mosquitoes (Collins & Paskewitz References 1996; Krzywinski & Besansky 2003; Scheffer 2005). The Anonymous. 1977. 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Bulletin of Entomological Research 97: 387-397. malaria and a suitable vector, Culex quinquefasciatus Say Bauer, L. H. Liu and J. Gould. 2008. Progress on biological control of (Culicidae), has been documented (van Riper et al. 1986). In emerald ash borer. In: Mastro, Victor; Lance, David; Reardon, Richard; Parra, Gregory, comps. Emerald ash borer research and development the Galápagos Islands the introduction of the nestling parasite meeting; 2007 October 23-24; Pittsburgh, PA. FHTET 2008-07. Philornis downsi Dodge & Aitken (Muscidae) is considered Morgantown, WV: U.S. Department of Agriculture, Forest Service,

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