Non-native, invasive species pose one of the greatest threats to the health, diversity, and productivity of forests in North America (Wheeler and Hoebeke 2017). The introduction and establishment of non-native species through raw wood products, solid wood packaging material, or live plants has already had profound ecological effects on forests across the continent, including virtual eradication of several key species such as American chestnut [Castanea dentata (Marsh.) Borkh.] and several species of ash (Fraxinus) trees, and the economic costs to federal, state, provincial, and local governments and to private citizens have exceeded hundreds of millions of dollars (Aukema et al. 2011). Several non-native insects and pathogens—e.g., Asian longhorned beetle [Coleoptera: Cerambycidae: Anoplophora glabripennis (Motschulsky)], emerald ash borer (Coleoptera: Buprestidae: Agrilus planipennis Fairmaire), hemlock woolly adelgid [Hemiptera: Adelgidae: Adelges tsugae (Annand)], and sudden oak death (Pythiales: Phytophthora ramorum Werres, de Cock & Man in’t Veld)—have caused the decline and mortality of millions of trees in both forest and urban landscapes (Haack et al. 2010, Herms and McCullough 2014, Meentemeyer et al. 2007). Global trade (Fig. 1) has increased dramatically during the past 30 years in both volume and speed, resulting in an accelerated movement and establishment of non-native forest insects (Roques 2010, Meurisse et al. 2018). Exclusion, or preventing the introduction of non-native species, is the most effective first line of defense; however, species frequently escape detection and become established (National Invasive Species Council 2008). The National Invasive Species Council’s National Management Plan (2008) states that early detection, rapid assessment, and rapid response (EDRR) is a critical second line of defense and provides the greatest opportunity for eradication and cost-effective management. An effective EDRR program increases the likelihood that invasive species will be found, contained, and eradicated before they become widely established. Early detection and rapid response programs have focused on many different organisms, including, but not limited to, invasive plants, aquatic invertebrates, and several forest insects and diseases (National Invasive Species Council 2016). Container ship with cargo and solid wood packaging material potentially infested with invasive beetles. Bark and ambrosia beetles (Curculionidae: Scolytinae) are some of the most important insects that affect forests in North America and across the globe. Native species, such as the mountain pine beetle (Dendroctonus ponderosae Hopkins) and the southern pine beetle (Dendroctonus frontalis Zimmermann), have killed trees on millions of acres in western and southern North America, respectively (Clarke et al. 2016). Non-native species, such as the redbay ambrosia beetle (Xyleborus glabratus Eichhoff; Fig. 2a) and the polyphagous and Kuroshio shothole borers (Euwallacea spp.; Fig. 2b) have also had severe effects on forests and in urban settings in the southeastern U.S. and California, respectively (Ploetz et al. 2017, Eskalen et al. 2013). In addition, Scolytinae are the most commonly intercepted taxa of beetles at U.S. ports of entry. From 1984 through 2008, more than 8,000 interceptions of scolytine beetles from 85 different countries were reported at U.S. ports (Haack and Rabaglia 2013). In North America, more than 60 species of non-native scolytine species have become established in the U.S. (Haack and Rabaglia 2013, Smith et al. 2017, Gomez et al. 2018). Redbay (Persia borbonia) mortality from laurel wilt fungus vectored by redbay ambrosia beetle. Tree mortality caused by shothole borers (Euwallacea spp.) in a riparian area in southern California. The previous introduction and establishment of non-native bark beetles such as Scolytus multistriatus (Marsham) and Tomicus piniperda (L.) went undetected for many years (Chapman 1910, Haack and Kucera 1993), but the development of semiochemical lures and traps in the 1990s made a national EDRR program for non-native bark and ambrosia beetles feasible. In 2001, the USDA Forest Service, in coordination with the USDA Animal and Plant Health Inspection Service (APHIS), the National Plant Board, and the National Association of State Foresters began a pilot project for the early detection of and rapid response to non-native bark and ambrosia beetles (Rabaglia et al. 2008). In 2007, the Forest Service began national implementation of the EDRR project across the United States, which filled a gap among the other EDRR programs to survey urban/suburban woodlands for non-native bark and ambrosia beetles. This program provided a needed layer of surveillance for potentially invasive non-native beetles that may have eluded detection at ports and escaped into nearby natural areas. This paper summarizes the protocols, species collected, and accomplishments of the project from 2007 through 2016. This program resulted in the collection of >840,000 specimens, the discovery of three non-native species new to the United States, revised distribution records for 149 native and 28 non-native species, and observations of taxon-specific patterns in semiochemical lure response. The selection of target species for this EDRR project was based on interception data at U.S. ports of entry (Haack 2001), the potential damage a species might cause in U.S. forests, and the availability of effective traps and lures for a species. The species targeted (Fig. 3) were: Hylurgops palliatus (Gyllenhal) Hylurgus ligniperda (Fabricius) Tomicus minor (Hartig) Tomicus piniperda (Linnaeus) Pityogenes chalcographus (Linnaeus) Orthotomicus erosus (Wollaston) Ips sexdentatus (Boerner) Ips typographus (Linnaeus) Trypodendron domesticum (Linnaeus) Xyleborus species a–h. Representative EDRR target species: a) Hylurgops palliatus; b) Hylurgus ligniperda; c) Ips sexdentatus; d) Ips typographus; e) Orthotomicus erosus; f) Pityogenes chalcographus; g) Tomicus piniperda; h) Trypodendron domesticum. From 2007 through 2009, following the pilot project and based on available funding, the goal was to survey approximately one third of the states every year, so that all 50 states would be surveyed every three years. By the end of 2009, 46 states had participated in the survey, with only Connecticut, Iowa, North Dakota, and Oklahoma not having been surveyed (Oklahoma and Connecticut were surveyed in 2011 and 2012, respectively). The U.S. Territories of Puerto Rico and Guam also were surveyed (in 2013 and 2014, respectively) to detect any invasive bark and ambrosia beetles established in these areas that could potentially spread to other areas. By 2016, all states except Iowa and North Dakota had been surveyed at least once (Table 1). States and territories participating in EDRR survey, 2007–2016. States and territories participating in EDRR survey, 2007–2016. In 2010, with a change in funding, the project shifted from surveying every state every three years to identifying high-risk states and targeting these states annually. Criteria for identifying high-risk states included the number of interceptions at ports of entry within a state, the number of non-native species established in a state, the amount of forest land in a state, and transportation corridors within a state. Based on these criteria, it was decided that five states (California, Florida, Georgia, New York, and Texas) would be surveyed every year; a second tier of 10 states would be surveyed every 3–5 years; and the remaining states would be surveyed every 5–7 years or as funding permitted. During the first three years of the project, state forestry or agriculture department partners in each state selected nine high-risk sites within the state; from 2010 through 2016, state partners selected 12 high-risk sites per year. High-risk sites were identified as locations at which potentially infested solid wood packaging material (such as dunnage, crating, and pallets) or other items were imported, stored, or recycled. These sites were selected in coordination with state Cooperative Agriculture Pest Survey (CAPS) committees, which often conducted APHIS-supported exotic wood borer surveys at similar high-risk sites and ports of entry. The CAPS survey sites were placed close to or within these sites (Fig. 4), whereas the EDRR survey traps were placed in wooded areas near these high-risk locations in fragmented forests or urban forests and parks (Fig. 5). In most states, because there was coordination between these surveys, sites selected for each survey were often in different locations to maximize coverage. Typical APHIS Cooperative Agriculture Pest Survey trap site. Typical EDRR trap site near wood recycling facility. During the entire survey, three 8- or 12-unit black Lindgren funnel traps were used at each survey location (Fig. 6). In most states, the wet trap option was used; the trap cup was filled with propylene glycol antifreeze to prevent escape of specimens and result in better preservation. Each trap was baited with one of the following three lures or lure combinations: Ultra-high release (UHR) ethanol lure only (general attractant for woodboring insects in hardwood and some coniferous hosts) UHR α-pinene and UHR ethanol lures together (general attractants for woodboring insects in coniferous hosts) Three-component exotic ips lure of ipsdienol, cis-verbenol, and methyl-butenol (more specific for conifer-feeding exotic bark beetles e.g., Hylurgus ligniperda, Ips typographus, and Orthotomicus erosus) Lindgren funnel traps used in EDRR survey. In 2009, manuka oil was added as a fourth lure at each site in Hawaii. Manuka oil was shown to be an effective lure for Xyleborus glabratus in the southeastern U.S. (Hanula and Sullivan 2008) and for the emerald ash borer (Crook et al. 2008); thus, the sesquiterpenes in the lure were potential attractants for other hardwood-infesting species. In Guam in 2014, ethanol + cubeb oil (a more readily available essential oil with a sesquiterpene profile similar to manuka oil [Hanula and Sullivan 2013]) was used instead of exotic ips lure (the pine-infesting Ips species were determined to be a low risk in Guam) as a potential attractant for hardwood-infesting ambrosia beetles. Trapping began in each state based on local phenology and knowledge of early emergence of bark and ambrosia beetles. From south to north, trapping usually coincided with bud break, when most bark and ambrosia beetles began emerging after overwintering (although a few early fliers may have been missed), and began in late February to early May. From 2007 through 2009, states trapped for a total of 20 weeks. Beginning in 2010, when the number of sites per state increased from nine to twelve, the trapping period changed to 12 weeks. During the pilot project and the first years of implementation, trap data showed that the majority of both bark and ambrosia beetles were captured during the first 12-week period, declining considerably thereafter. For the entire survey, traps were checked every two weeks, when samples were collected and antifreeze was changed. Every four weeks, traps were re-baited with fresh lures. Once per month, unsorted samples and samples that had been screened for all bark and ambrosia beetles were sent to regional taxonomists (Fig. 7). All bark and ambrosia beetles were identified to the most specific taxon possible using the most recently published taxonomic keys (e.g., Rabaglia et al. 2006; Fig. 8). A more extensive literature search and examination of type specimens from worldwide collections followed the discovery of potentially new non-native species. Occurrences of species were entered into an online database. The database, developed by the Forest Health Assessment and Applied Sciences Team Forest Service, Forest Health as a for trap trap collection and These data were with the collection and other for species were the of the EDRR project was to on of the target species, it that a better was to all bark and ambrosia beetles to the most specific taxonomic The for this is of the trapping be on the EDRR at of in EDRR trap of identifying beetles in EDRR trap The of this project is the most rapid response possible to of non-native bark or ambrosia beetle species. EDRR collection and were to between collection and were with APHIS and states so that after and of a non-native bark or ambrosia beetle species new to North America, the or a a rapid risk could be conducted to the species a economic or ecological were established to the area in which the species This is to of or surveys for non-native species are to on funding, taxonomic and available The volume and of specimens through traps and such as used in this project, was this project targeted 10 non-native species, EDRR taxonomists the of identifying all captured The most important of these was detection of new non-native species that would have been only the target species were For taxonomists identified Scolytus for the first in North America a species that had been in the U.S. for many years but had undetected or had been included of recently or non-native and a better of the of non-native and native 2016, more than new state records were for a total of 149 species, including 28 non-native species (Table This is not only for North American scolytine but also provides to states and the of potential non-native or New state records for species collected in EDRR survey traps from 2007 through 2016. in non-native species established in North New state records for species collected in EDRR survey traps from 2007 through 2016. in non-native species established in North than specimens of bark and ambrosia beetles were collected and identified from states, Puerto and Guam the 10 years of this survey. of these specimens were identified to species, and were identified to these approximately to the and two that to the United States, the survey captured specimens of approximately species of the approximately species that in the U.S. The more species captured were to the semiochemical attractants and (e.g., Ips species, such as were collected species were by 10 or specimens in traps during the to the interception by the funnel traps than by The most species collected made of the specimens collected (Table non-native species already established in the U.S. of the most species and of all specimens of the species collected were non-native species established in the U.S. The three most species collected in and of the and all non-native species with to This established non-native species ambrosia in traps may be a of the species in the forests surveyed or a of these non-native species for may be to there is a for ethanol by invasive species this be used to potential also may be possible that the of non-native species captured in traps is an that they are native species and be to this potential of invasive species. most species in EDRR 2007–2016. in non-native species established in North most species in EDRR 2007–2016. in non-native species established in North The of EDRR have been the early detection of species new to North America, a rapid of potential and a rapid response. 2007, three species new to North America have been in EDRR traps (Fig. in and in (Rabaglia et al. in in 2010 et al. and in in 2010 et al. 2013). was in the pilot project, these three new non-native species were and the was with state agriculture and forestry and with and surveys to the distribution of the new species were conducted in the as the first detection or in the following year. In each trapping that these beetles were established several or across a eradication was not feasible. of ambrosia such as the three species 2007, is species in most hardwood and to undetected and to spread from a few new to United States in EDRR a) Scolytus b) c) d) Xyleborus e) Xyleborus f) is that not of the species during the years of EDRR were species in the U.S. for before These species were with the of surveys such as EDRR and CAPS in the late 1990s and early Scolytus is a of a species. was first in in and within two years had been from western states et al. of several collections specimens that had been collected before and as early as in New other species, it is that the EDRR surveys were the first to detect In the of specimens were first collected in during 2010, EDRR and CAPS surveys that had in in 2007 et al. 2011). A for from the 2010 collection that the introduction was limited to a few in the first of detection, the species was in three different more than the more than new state records during the 10 years of this survey, were for 28 non-native species already established in the U.S. (Table for these species is important not only to in the but also to of spread and Xyleborus the of the fungus & which the laurel was first in EDRR traps in in (Rabaglia et al. after which it spread through or movement to North and non-native ambrosia was first collected in in in in and and in in et al. it has been collected in new states through EDRR trapping (Table EDRR surveys have also the movement of Xyleborus in in and Rabaglia it has been in EDRR traps in and species to spread from the of into For first in was in new states during EDRR surveys and is from the and et al. The of data for all species identified and all states participating in EDRR from 2007 through be at 2010, there have been of species new to North America in EDRR is possible that all non-native species established in the states surveyed before 2010 have been and that any new will be from for a more effective rapid response. is also possible that the implementation of protocols, such as for and of the risk of wood have the number of woodboring insects into the United States (Haack et al. The of and of this survey it among bark and ambrosia beetle surveys in the United of this and extensive better the movement of non-native species, the for and that may affect EDRR data have already been used to patterns of bark and ambrosia beetles in the United States et al. and between and non-native scolytine beetle establishment et al. 2011). The EDRR survey also increased of the distribution of native species of bark and ambrosia beetles in the U.S. of the records are for species that may be for a state, but for which records have been e.g., in and For collection records from EDRR the of native Scolytus was than with specimens collected in and and (Table the detection and of native species was not a of this survey, this be used to the potential that non-native species may have on native species. of the non-native established in North America are ambrosia beetles et al. 2018). all of these species these are not usually to the in some such as Xyleborus the is and has caused extensive mortality to North American et al. 2007). The EDRR program provided funding to the of species established in the United States et al. et al. et al. as as some species that could potentially become established in the U.S. et al. 2017, et al. 2017). the of laurel most of these non-native are not beetle are not and in native and non-native are species that are not may have ecological by et al. identified a new wood fungus with the non-native ambrosia beetle minor which is to wood in the southeastern U.S. this project, new species of with bark or ambrosia beetles have been et al. et al. 2017). This provides a better of and target potential high-risk species that may be The lures for this survey targeted high-risk species, such as some Ips species (e.g., Ips and other more high-risk such as ambrosia beetles. Based on the of specimens, and species these lures were effective in a of species in different were trapped during the survey, and some may have been trapped by many species and showed patterns with a response to specific lures. this was not a with and and traps were in different areas with different each year, based on the number of trap sites (more than per for 10 some be made of taxa to the different lures (Table ambrosia beetles usually showed a for traps and Rabaglia however, some and species were by the of species, which and were not by the of α-pinene to but to ethanol by the was by the of α-pinene (Table a and response to ethanol by Xyleborus which was by the of α-pinene (Table most Ips species were to the exotic ips lure Ips typographus used in the however, Ips was more to ethanol + α-pinene (Table The exotic ips lure used ipsdienol, which is not a of as the species of were most to ethanol + but the species were most to exotic ips lure (Table and was the response of to exotic ips because this species on et al. to exotic ips lure may an response to Ips the of and et al. 2017). The patterns of and response in this survey be used to selection of lures and target different of specimens in EDRR survey traps baited with different and 2007–2016. of specimens in EDRR survey traps baited with different and 2007–2016. of the of the National Invasive Species Council’s National Management Plan (2008) is to and the to and to invasive Bark and ambrosia beetles are not to from the and which are the of the non-native beetles in the United States (Haack and Rabaglia 2013). keys and collections not for these From the of the EDRR it was that there was a for taxonomic and keys to potentially non-native species in was provided to and keys and 2014, Smith and and is in to keys for the worldwide species of A of this project has been to the national for of bark and ambrosia beetles. In this has been through for EDRR from a of land and These have published keys and taxonomic (e.g., Rabaglia but they also the development of and to of of this (such as commonly trapped species and non-native e.g., Xyleborus for and taxonomists are not Scolytinae this survey more than traps in more than locations across states, the of and the Territories of Puerto Rico and Guam (Fig. the was to the potential for invasive species the United States participating in this project were to high-risk sites such as wooded areas near ports of urban or this project trapping may of trap locations to within a state, or of all traps in a in a high-risk area of a state. response to the lures used in the project were effective across many different different lures will be used to target specific high-risk high-risk species may a response to other for Xyleborus glabratus response to ethanol most but is more to sesquiterpene et al. et al. 2016). Beginning in 2017, some of these new lures were added to the EDRR trap locations in the United States, 2007–2016. Non-native including bark and ambrosia have had a on North American forest the early detection of many of these species has to be a EDRR provides the opportunity for the and cost-effective of these potentially invasive species. 10 have of U.S. bark and ambrosia beetle and provided a of the and of species in the U.S. also have identified potential areas for such as the selection of lures and development of trapping and this program it is that any non-native species will be recently species that be for and effective response. This project would not have been possible the and taxonomic of many federal, state, and the regional Forest Service surveys with the many state, and collected and specimens, and entered into the database. this project would not have been The was developed by and by the Forest Service, Forest Health Assessment and Applied Sciences with this important of the also Smith State for taxonomic of and for extensive on and Texas) for with species and to and of the to EDRR also and for to the National at the to with of non-native species.
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