TY - JOUR
AU - Kula, Robert, R
AB - Abstract Cerambycidae provide important ecological services in forests yet cause economic damage when they infest living trees. Parasitoids can regulate woodborer populations, providing considerable control of pest cerambycids. Identifying parasitoids of native cerambycids may be useful in managing cerambycid outbreaks and aid in new-association biocontrol of exotic invasive cerambycids. We investigated Cerambycidae and associated hymenopteran parasitoid communities infesting Acer rubrum, Pinus virginiana, and Carya tomentosa from a forest in Delaware from 2005 to 2012. Cerambycid abundance, diversity, and richness, as well as parasitoid abundance, were measured by collecting trees in different conditions: felled, girdled, and naturally infested. Effect of edge or interior red maple on cerambycid abundance, diversity, and richness was examined. Over 14,500 cerambycids of 56 species and 38 genera were collected during the 7-yr period. Eleven species represented 95% of all cerambycids collected. Treatment only affected red maple, showing increased cerambycid richness and diversity from naturally infested trees. Cerambycid richness and diversity were two times greater on hickory than other species when combining girdled and felled treatments. Over 19,000 parasitic Hymenoptera of 12 families emerged from woodborer-infested wood with >70% of individuals belonging to Braconidae. Thirteen known species, and two unknown species, of Braconidae were identified from a subsample of 495 specimens; Ontsira mellipes (Ashmead) (Hymenoptera: Braconidae) and Rhoptrocentrus piceus Marshall (Hymenoptera: Braconidae) were the most abundant. This study provides fundamental information on native parasitoids associated with Cerambycidae, including cerambycid larval host associations. Parasitoids identified herein should be investigated for potential adaptation to invasive Cerambycidae to benefit invasive woodborer management. Cerambycidae, Braconidae, new-association biocontrol, native species One of the largest and most diverse insect groups are the woodboring Cerambycidae. To date, >35,000 species of Cerambycidae have been described worldwide (Slipinski et al. 2011), with some 1,100 species endemic to North America (Yanega 1996, Svácha and Lawrence 2014). The phytophagous larvae develop within various types of woody plant tissues under conditions ranging from living to dead and decaying (Haack and Slansky 1987, Haack 2017). Conversely, most adults feed on pollen, foliage, twigs, and fungi (Butovitsch 1939, Duffy 1957, Linsley 1959), whereas others are nonfeeding, such as Prionus californicus Motschulsky and Callidiellum rufipenne (Motschulsky) (Haack 2017, Barbour et al. 2019). Cerambycids that develop in living hosts can have considerable economic impacts when they infest timber or wood products, urban trees, and fruit and nut trees (Solomon 1995). However, native species serve important ecological roles within forest ecosystems by decomposing woody debris (nutrient cycling; Grove 2002) and as resources for both vertebrate and invertebrate predators and parasitoids (Martin et al. 1951, Grove 2002). Moreover, the galleries created by larvae, accessible once adults emerge, serve as nesting sites and provide additional habitat for numerous arthropod species (Satoh et al. 2016, Sydenham et al. 2016). Yet, native Cerambycidae and woodboring species are threatened by the continued risk of displacement by invasive exotic forest pest species. For instance, Jennings et al. (2017) found significant shifts in cerambycid composition, diversity, and species richness associated with Fraxinus spp. from 2011 to 2014 in Maryland, resulting from the invasion of the exotic forest pest Agrilus planipennis Fairmaire (Coleoptera: Buprestidae). The natural history, biology, and taxonomy of adult North American Cerambycidae are well known (Hanks 1999, Langor et al. 2008) compared with immature stages. Yet information on host relationships is often complicated by the crepuscular behavior of adults and the cryptic nature of eggs and larvae (Linsley 1961). Comprehensive distribution data for adult cerambycids from various trapping methods are available for some areas of North America (e.g., Thomas et al. 2005, Hart et al. 2013), yet larval hosts are typically unknown or are unspecified (Haack 2017). Furthermore, larval hosts are often misidentified due to erroneous plant and insect identification, inaccurate interpretation of taxonomic changes, imprecise museum specimen labeling, and assumptions of host associations based on adult collection (Haack 2017). Surveys of adult cerambycids are implemented using passive sampling via the deployment of traps (e.g., cross vane panel, Malaise, pitfall, multiple funnel, and light traps) baited with sex pheromones or host volatiles that lure and capture active adults. Captured adults can be readily identified using diagnostic tools and information in a number of publications (e.g., Yanega 1996, Lingafelter 2007, LeBonte et al. 2013, Nearns et al. 2020). Conversely, larval surveying is labor intensive, requiring that trees are felled and entire boles and branches are split to observe for burrowing larvae. Although resources exist for larval identification, this stage is often more difficult to identify compared with adults. However, an alternative approach to larval sampling involves mechanically stressing living trees by removing bark sections (girdling) and leaving trees for colonization by adults attracted to these trees. These ‘trap trees’ can then be felled, sectioned, and stored to rear and identify emerging adult beetles. Thus, identification of cerambycid larvae is avoided, with the additional benefit of collecting other insects emerging from these bolts such as parasitoids associated with immature woodboring insect stages (e.g., Jennings et al. 2017, Duan et al. 2015b). Native parasitoid species can be particularly important in regulating insect pests of forest ecosystems and can provide considerable control of native pest species (Yang et al. 2014). In some cases, native parasitoid species have expanded their host ranges to include exotic invasive forest pest species. In turn, this can complement current management strategies for such pests (e.g., Duan et al. 2009, 2012). Cerambycid pests and their native natural enemies are particularly well studied in China. For example, the indigenous polyphagous predatory beetle Dastarcus helophoroides (Fairmaire) (Coleoptera: Bothrideridae) and various Sclerodermus spp. (Hymenoptera: Bethylidae) provide substantial control of woodboring Cerambycidae such as Anoplophora glabripennis (Motschulsky), Massicus raddei (Hope), and Monochamus alternatus Hope (Zhang and Yang 2006, Wei et al. 2009, Tang et al. 2012, Yang et al. 2014) in China. Although classical biological control can be used successfully against invasive exotic pests, sometimes effective host specific and coevolved natural enemies are absent or undiscovered, like those of A. glabripennis (Golec et al. 2018). Native parasitoids have been observed parasitizing novel exotic invasive hosts, and such occurrences have been documented with various invasive woodboring species of importance throughout different geographical areas. For instance, in North America, several native parasitic hymenopteran species, including Phasgnophora sulcata Westwood (Chalcididae), Atanycolus hicoriae Shenefelt, Atanycolus simplex (Cresson), Spathius floridanus Ashmead, and Leluthia astigma (Ashmead) (Braconidae), have been found parasitizing larvae of the invasive emerald ash borer (A. planipennis) at rates up to 60% in some locations (Bauer at al. 2005, 2008; Duan et al. 2009, 2012; Kula et al. 2010). Although parasitism rates of newly associated hosts are often less at the beginning of a new parasitoid–host association than those of their coevolved hosts, native natural enemies nevertheless contribute to biocontrol programs of invasive exotic insects (e.g., Duan et al. 2015b). Thus, identifying natural enemies associated with native woodborers may prove useful in controlling outbreaks of native beetles (via augmentation release) and may have potential as new-association biocontrol agents against exotic woodboring species (e.g., Hokkanen and Pimentel 1989). In addition, it is the best practice that before any natural enemy is imported, prerelease surveys are completed to identify potential native natural enemies that may have already adapted to the exotic pests. Interest in the biological control of invasive exotic pests will continue to grow as the potential for the establishment of non-native woodboring pests increases (Aukema et al. 2010). Equally important is enhancing our understanding of parasitoid and cerambycid host associations, as well as improving our knowledge of the habitat requirements of these taxa. Such information is critical for assessing local cerambycid and parasitoid communities and for improving conservation tactics for environments that support particular species. Moreover, this information can assist in predicting the potential impacts of exotic invasive woodboring species on native insect communities (e.g., Gandhi and Herms 2010, Jennings et al. 2017). We report Cerambycidae and their associated parasitoid communities (with an emphasis on Braconidae) emerging from red maple (Acer rubrum L.), Virginia pine (Pinus virginiana Mill.), and mockernut hickory [Carya tomentosa (Lam. ex Poir) Nuttall)] from a fragmented forest (Blackbird State Forest) in northern Delaware from 2005 to 2012. Our main goal was to characterize parasitic Hymenoptera (Braconidae) emerging from woodborer-infested trees to initiate future investigation into their ability to form new associations with exotic invasive Cerambycidae, thereby potentially contributing to invasive woodborer management via new-association biological control (e.g., Hokkanen and Pimentel 1989). Also, we were interested in cerambycid communities and the effect of tree species and survey method on their species abundance, richness, and diversity. To this end, we used three different sampling methods (felling, girdling, and collecting naturally infested trees) to examine Cerambycidae infesting Virginia pine, mockernut hickory, and red maple, as well as their associated parasitic hymenopteran communities, over a 7-yr period. First, we examined the effect of sampling method (treatments) and tree species on the relative abundance (%), richness, and diversity of cerambycids emerging from each tree species from 2005 to 2007. Then, from 2007 to 2012, we mechanically felled red maple trees from the forest edge and interior and subsequently determined the effect of red maple location on cerambycid abundance, richness, diversity, and differences between species occurring from the edge and interior. For hymenopteran parasitoids, we summarize the families, numbers of individuals, and their relative abundance (%) reared from cerambycid-infested Virginia pine, mockernut hickory, and red maple bolts from 2005 to 2012. Species-level identifications were limited to Braconidae. For this group, we provide a list of species and their relative abundance (%) collected from each tree species between 2005 and 2008. Materials and Methods Field Location Surveys were conducted from 2005 to 2012 in Blackbird State Forest in Townsend, DE. Blackbird State Forest is a mixed-deciduous forest (totaling >2,400 ha) consisting of A. rubrum, Liquidambar styraciflua L., Quercus bicolor Willd., and Nyssa sylvatica Marshall in the lowlands; A. rubrum, P. virginiana, Ilex opaca Aiton, L. styraciflua, and N. sylvatica in the understory; and A. rubrum, Carya glabra (Mill.) Sweet, C. tomentosa, Liriodendron tulipifera L., Quercus alba L., Q. rubra L., and P. virginiana dominating the overstory. Red maple (A. rubrum), mockernut hickory (C. tomentosa), and Virginia pine (P. virginiana) were the three most dominant tree species in the forest (E.A., personal observation). Trees were sampled along three trails, from 39.33035 N to 39.348701 N, −75.66582 E to −75.733737 E, and were within 3,000 m of the trails. Sampling for Cerambycidae and Braconidae The sampled tree diameter near breast height ranged from 8.9 to 18.5 cm for red maples, 13.1 to 39.2 cm for mockernut hickory, and 15.4 to 39.7 cm for Virginia pine. During 2005 and 2007, our major aim was to determine the possible effect of the three dominant tree species on species richness and diversity of cerambycids and to examine associated parasitoid taxa. All tree species were sampled simultaneously using two different methods to mechanically induce stress: 1) girdled tree and 2) felled tree. We girdled trees by removing the bark and outer xylem of a healthy tree in a single contiguous band (5 cm height, 2 cm depth) encircling each tree using a drawknife. Conversely, for felled trees, healthy trees were cut down using a chainsaw in early spring (April of each year) and left in the field to be colonized by woodboring beetles and associated parasitoids through the spring, summer, and fall months. That winter or the following spring (November—March of each year), girdled trees were felled, and together with the felled treatment, all tree trunks and large branches (average diameter = 13.4 ± 0.18) were cut into 50- to 56-cm-long bolts and inspected for signs of colonization. This included observations of frass expulsion, oviposition scars, and woodpecker feeding. Bolts were subsequently returned to the laboratory. However, signs of attack in mockernut hickory were unclear externally; thus, inspection of mockernut hickory bolts was modified whereby 5 cm2 sections of bark at 0.5-m increments were removed and inspected for colonization and developing cerambycid and parasitoid larvae and pupae. During these same years (2005–2007), naturally infested red maple, mockernut hickory, and Virginia pine trees that were wind felled or still standing and were near the mechanically stressed trees were also felled, sectioned, and brought back to the laboratory. This was completed to assess differences in the diversity, richness, and composition of cerambycids between naturally infested and mechanically stressed trees. From 2006 to 2008, a total of 688 and 590 mockernut hickory and Virginia pine bolts were collected, respectively, whereas 3,272 red maple bolts were collected between 2006 and 2013 (Table 1). Table 1. Summary of collection year, numbers of trees and bolts, and surface area of collected bolts from each tree species Tree species Years of collection Number of trees sampled Number of bolts collected Mean (± SE) Surface area per bolt (m2)a Virginia pine 2006–2008 47 590 0.357 ± 0.013 Mockernut hickory 2006–2008 60 688 0.319 ± 0.021 Red maple 2006–2013 772 3,272 0.310 ± 0.011 Tree species Years of collection Number of trees sampled Number of bolts collected Mean (± SE) Surface area per bolt (m2)a Virginia pine 2006–2008 47 590 0.357 ± 0.013 Mockernut hickory 2006–2008 60 688 0.319 ± 0.021 Red maple 2006–2013 772 3,272 0.310 ± 0.011 aSurface area (mean ± SE) was estimated using the formula of a cylinder (bolt length × pi × bolt diameter). Open in new tab Table 1. Summary of collection year, numbers of trees and bolts, and surface area of collected bolts from each tree species Tree species Years of collection Number of trees sampled Number of bolts collected Mean (± SE) Surface area per bolt (m2)a Virginia pine 2006–2008 47 590 0.357 ± 0.013 Mockernut hickory 2006–2008 60 688 0.319 ± 0.021 Red maple 2006–2013 772 3,272 0.310 ± 0.011 Tree species Years of collection Number of trees sampled Number of bolts collected Mean (± SE) Surface area per bolt (m2)a Virginia pine 2006–2008 47 590 0.357 ± 0.013 Mockernut hickory 2006–2008 60 688 0.319 ± 0.021 Red maple 2006–2013 772 3,272 0.310 ± 0.011 aSurface area (mean ± SE) was estimated using the formula of a cylinder (bolt length × pi × bolt diameter). Open in new tab Effect of Tree Location on Cerambycid Diversity and Richness To determine the possible effect of tree location (edge vs interior) on cerambycid abundance (%), richness, and diversity, 6–17 healthy red maple trees from the edge and interior of the forest were randomly selected and felled each year from 2007 to 2012 (except 2009). Trees on the edge were <20 m from the tree line or forest edge, whereas interior trees were ≥40 m from the forest edge. The distance between trees sampled from either the edge or interior ranged from 6.4 to 16.9 m. As described previously, all trees were felled in the spring of each year and left in the field. Trees were cut into 50- to 56-cm sections, collected over the following winter, and brought back to the laboratory. In total, 102 edge and 102 interior red maple trees were sampled between 2007 and 2012. Collection and Identification of Emerged Insects After infested bolts were retrieved from the field, they were individually placed inside cylindrical (length = 61 cm, diameter = 30 cm) cardboard Sonotubes (Sonoco, Hartsville, SC). Tubes containing bolts were held in an outdoor insectary at USDA-ARS-BIIRU (Newark, DE) under ambient temperature and light condition until the following spring when parasitoids and beetles emerged. Wood screws were partially screwed into bolts at 90° angles along the length of each infested bolt, thereby preventing the bolts from lying directly on the inner surface of tubes and preventing beetle and parasitoid emergence. There was 1- to 4-cm gap between the bolt and the inner surface of the tubes, allowing air to freely circulate around the bolt. Prior to inserting the bolt, the interior surfaces of the tubes were sterilized with 5% bleach solution. Each tube was sealed on either end with tight-fitting plastic lids; however, one side of the lid was modified with a collection cup that captured emerging adult beetles and parasitoids (Fig. 1). Fig. 1. Open in new tabDownload slide Example of Sonotubes modified with collection cups used to store tree bolts and capture emerging insects. Fig. 1. Open in new tabDownload slide Example of Sonotubes modified with collection cups used to store tree bolts and capture emerging insects. Emergence cups were checked Monday–Friday weekly for the presence of insects, and all emerged insects were recorded. All herbivores, including cerambycids, buprestids, and other wood borers, along with parasitic Hymenoptera, were preserved for identification. Because no buprestids emerged from mockernut hickory and few species were found on Virginia pine and red maple, they were not analyzed. Analyses of species richness and diversity focused on Cerambycidae. All cerambycids were identified morphologically to species using Yanega (1996). A subsample of hymenopteran parasitoids (Braconidae; n = 495) was sent to the USDA-ARS Systematic Entomology Laboratory (Washington, DC) for identification by R.R.K. These braconids were identified to genus using Wharton et al. (1997) and then sorted into morphospecies. Morphospecies were identified to species, when possible, using keys and diagnostic information as indicated in Yu et al. (2016) and their identities confirmed through comparison with authoritatively identified specimens in the Smithsonian Institution National Museum of Natural History, Washington, DC (USNM). Voucher specimens have been deposited at both the USNM and USDA-ARS-BIIRU. We were most interested in identifying braconid parasitoids emerging from woodborer-infested wood; therefore, identification of parasitoid species focused on this group. Data Analysis We summarized data on the cerambycid and hymenopteran parasitoid taxa, including their relative abundance, from 2006 to 2008 for Virginia pine and mockernut hickory and from 2006 to 2013 for red maple. We then calculated the species richness (S represents number of cerambycid species in each tree sample) and Shannon’s diversity index (H=∑Pi|ln(Pi)|) ⁠, where Pi is the proportion of individuals of one species divided by the total number of species found in a sample for cerambycids emerging from each tree species. The effect of sampling method (girdled, felled, and naturally infested) and tree species on both the richness and diversity of cerambycids was analyzed using independent two-way analysis of variance. After no difference was detected between mechanical treatments (girdled and felled) in the richness and diversity within each tree species, we pooled the data from these treatments to compare overall differences in cerambycid richness and diversity between mockernut hickory, Virginia pine, and red maple based on samples from 2006 to 2008. To determine the effect of red maple location (edge vs interior) on the species richness and diversity of cerambycids, we analyzed data collected from 2008 to 2013 using a mixed linear model, where the year was considered a random factor and location a fixed factor. Prior to analyses, data were checked for the normality of residuals and homoscedasticity using Shapiro’s and Bartlett’s tests. Mean values among different treatments were separated using Tukey’s Honestly Significant Difference test. All analyses were conducted using JMP Pro 13.1.1 statistical software (Sall et al. 2017). Results Insect Communities Associated with Sampled Trees In total, 98,843 individual insects comprising 5 orders and 55 families were collected from Virginia pine, mockernut hickory, and red maple bolts from 2006 to 2013 (Fig. 2; Supp Tables 1–3 [online only]). Cerambycidae and parasitic Hymenoptera represented over a third of all insect specimens collected (n = 33,972). This included 14,562 Cerambycidae consisting of 38 genera and 56 species, and 19,410 parasitic Hymenoptera comprising 12 families with >70% (n = 14,563) of the specimens belonging to Braconidae. Fig. 2. Open in new tabDownload slide Relative abundance of the most common insect families (>1% of total number collected) emerging from Virginia pine (2006 to 2008), mockernut hickory (2006 to 2008), and red maple (2006 to 2013) bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware, USA). Fig. 2. Open in new tabDownload slide Relative abundance of the most common insect families (>1% of total number collected) emerging from Virginia pine (2006 to 2008), mockernut hickory (2006 to 2008), and red maple (2006 to 2013) bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware, USA). Overall, Cerambycidae comprised >50% (n = 6,431) of the specimens collected from mockernut hickory among the families represented, though their abundance was considerably less on Virginia pine (14%, n = 299) and red maple (9%, n = 8,007; Fig. 2). Across all survey methods, years, and tree species, nine species comprised >90% of all Cerambycidae collected: Neoclytus acuminatus (Fabricius) (31%, n = 4,544), Neoclytus mucronatus (Fabricius) (21%, n = 3,064), Saperda discoidea Fabricius (13%, n = 1,881), Xylotrechus colonus (Fabricius) (8%, n = 1,174), Elaphidion mucronatum (Say) (7%, n = 1,067), Graphisurus fasciatus (DeGeer) (5%, n = 755), Curius dentatus Newman (3%, n = 494), Euderces pini (Olivier) (3%, n = 479), and Monochamus sp. (1%, n =191). Specimens of Braconidae represented nearly a fifth of the specimens for all families collected from all tree species over all sampling years; their abundance was highest on mockernut hickory (22%, n = 2,571) followed by Virginia pine (19%, n = 415) and red maple (14%, n = 11,577; Fig. 3; Supp Tables 1–3 [online only]). Fig. 3. Open in new tabDownload slide Relative abundance of all parasitic hymenopteran families emerging from woodborer-infested Virginia pine (2006 to 2008), mockernut hickory (2006 to 2008), and red maple (2006 to 2013) bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware, USA). Numbers above bars indicate total number of individual specimens collected from each tree species. Fig. 3. Open in new tabDownload slide Relative abundance of all parasitic hymenopteran families emerging from woodborer-infested Virginia pine (2006 to 2008), mockernut hickory (2006 to 2008), and red maple (2006 to 2013) bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware, USA). Numbers above bars indicate total number of individual specimens collected from each tree species. Cerambycidae Abundance, Richness, and Diversity from Virginia Pine, Mockernut Hickory, and Red Maple Virginia Pine In total, 12 cerambycid species (n = 245 individuals) were collected from Virginia pine bolts between 2006 and 2008, with six species unique to this tree species (Table 2). One species, Monochamus sp., accounted for 75% (n = 184) of all species collected from Virginia pine (Table 2). The aforementioned species combined with the abundances of Monochamus carolinensis (Olivier) and Astylopsis sexguttata (Say) comprised over 90% (n = 222) of all individuals collected from Virginia pine (Table 2). There was no difference between girdled, felled, and naturally infested treatments on cerambycid richness and diversity (Table 5). Table 2. Cerambycidae species emerging from Virginia pine bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2008 in order of abundance. Year Cerambycidae species Total Relative abundance (%) 2006 2007 2008 Monochamus sp. 184 75.1 138 45 1 Monochamus carolinensis (Olivier) 23 9.39 0 0 23 Astylopsis sexguttata (Say)a 15 6.12 0 14 1 Astylopsis collaris (Haldeman) 7 2.86 2 4 1 Neoclytus mucronatus (Fabricius) 5 2.04 5 0 0 Xylotrechus colonus (Fabricius) 4 1.63 3 1 0 Asemum striatum (Linnaeus)a 2 0.82 0 2 0 Acanthocinus obsoletus (Olivier)a 1 0.41 0 1 0 Neoclytus sp. 1 0.41 1 0 0 Neoclytus acuminatus (Fabricius) 1 0.41 1 0 0 Rhagium inquisitor (Linnaeus)a 1 0.41 0 1 0 Xylotrechus s. sagittatus (Germar)a 1 0.41 0 1 0 Total 245 150 69 26 Year Cerambycidae species Total Relative abundance (%) 2006 2007 2008 Monochamus sp. 184 75.1 138 45 1 Monochamus carolinensis (Olivier) 23 9.39 0 0 23 Astylopsis sexguttata (Say)a 15 6.12 0 14 1 Astylopsis collaris (Haldeman) 7 2.86 2 4 1 Neoclytus mucronatus (Fabricius) 5 2.04 5 0 0 Xylotrechus colonus (Fabricius) 4 1.63 3 1 0 Asemum striatum (Linnaeus)a 2 0.82 0 2 0 Acanthocinus obsoletus (Olivier)a 1 0.41 0 1 0 Neoclytus sp. 1 0.41 1 0 0 Neoclytus acuminatus (Fabricius) 1 0.41 1 0 0 Rhagium inquisitor (Linnaeus)a 1 0.41 0 1 0 Xylotrechus s. sagittatus (Germar)a 1 0.41 0 1 0 Total 245 150 69 26 aSpecies only found from Virginia pine. Open in new tab Table 2. Cerambycidae species emerging from Virginia pine bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2008 in order of abundance. Year Cerambycidae species Total Relative abundance (%) 2006 2007 2008 Monochamus sp. 184 75.1 138 45 1 Monochamus carolinensis (Olivier) 23 9.39 0 0 23 Astylopsis sexguttata (Say)a 15 6.12 0 14 1 Astylopsis collaris (Haldeman) 7 2.86 2 4 1 Neoclytus mucronatus (Fabricius) 5 2.04 5 0 0 Xylotrechus colonus (Fabricius) 4 1.63 3 1 0 Asemum striatum (Linnaeus)a 2 0.82 0 2 0 Acanthocinus obsoletus (Olivier)a 1 0.41 0 1 0 Neoclytus sp. 1 0.41 1 0 0 Neoclytus acuminatus (Fabricius) 1 0.41 1 0 0 Rhagium inquisitor (Linnaeus)a 1 0.41 0 1 0 Xylotrechus s. sagittatus (Germar)a 1 0.41 0 1 0 Total 245 150 69 26 Year Cerambycidae species Total Relative abundance (%) 2006 2007 2008 Monochamus sp. 184 75.1 138 45 1 Monochamus carolinensis (Olivier) 23 9.39 0 0 23 Astylopsis sexguttata (Say)a 15 6.12 0 14 1 Astylopsis collaris (Haldeman) 7 2.86 2 4 1 Neoclytus mucronatus (Fabricius) 5 2.04 5 0 0 Xylotrechus colonus (Fabricius) 4 1.63 3 1 0 Asemum striatum (Linnaeus)a 2 0.82 0 2 0 Acanthocinus obsoletus (Olivier)a 1 0.41 0 1 0 Neoclytus sp. 1 0.41 1 0 0 Neoclytus acuminatus (Fabricius) 1 0.41 1 0 0 Rhagium inquisitor (Linnaeus)a 1 0.41 0 1 0 Xylotrechus s. sagittatus (Germar)a 1 0.41 0 1 0 Total 245 150 69 26 aSpecies only found from Virginia pine. Open in new tab Mockernut Hickory From 2006 to 2008, 6,352 cerambycid beetles were collected from mockernut hickory bolts representing 24 species of which 9 were unique to this tree species (Table 3). Three species, N. mucronatus, S. discoidea, and X. colonus, accounted for nearly 96% (n = 6,094) of all individuals collected from mockernut hickory (Table 4). Treatment of trees (felled or girdled) had no effect on cerambycid richness or diversity (Table 4). Table 3. Cerambycidae species emerging from mockernut hickory bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2008 in order of abundance Year Cerambycidae species Total Relative abundance (%) 2006 2007 2008 Neoclytus mucronatus (Fabricius) 3,052 48.1 1,777 487 788 Saperda discoidea Fabricius 1,880 29.6 345 1,478 57 Xylotrechus colonus (Fabricius) 1,162 18.3 870 281 11 Stenosphenus notatus (Olivier)a 56 0.88 0 8 48 Saperda lateralis Fabriciusa 55 0.87 0 55 0 Graphisurus fasciatus (DeGeer) 43 0.68 0 41 2 Neoclytus acuminatus (Fabricius) 37 0.58 5 2 30 Megacyllene caryae (Gahan)a 34 0.54 0 34 0 Monochamus sp. 7 0.11 7 0 0 Heterachthes pallidus Haldemana 6 0.09 0 0 6 Neoclytus sp. 6 0.09 6 0 0 Curius dentatus Newman 2 0.03 0 2 0 Graphisurus despectus (Le Conte)a 2 0.03 0 2 0 Liopinus mimeticus (Casey) 2 0.03 2 0 0 Achryson surinamum (Linnaeus)a 1 0.02 0 1 0 Astylopsis collaris (Haldeman) 1 0.02 0 1 0 Astylopsis macula (Say) 1 0.02 0 1 0 Dorcaschema cinereum (Olivier)a 1 0.02 0 1 0 Euderces pini (Olivier) 1 0.02 0 1 0 Lepturges confluens (Haldeman)a 1 0.02 0 0 1 Lepturges pictus (Le Conte)a 1 0.02 0 0 1 Tilloclytus geminatus (Haldeman) 1 0.02 0 0 1 Total 6,352 3,012 2,395 945 Year Cerambycidae species Total Relative abundance (%) 2006 2007 2008 Neoclytus mucronatus (Fabricius) 3,052 48.1 1,777 487 788 Saperda discoidea Fabricius 1,880 29.6 345 1,478 57 Xylotrechus colonus (Fabricius) 1,162 18.3 870 281 11 Stenosphenus notatus (Olivier)a 56 0.88 0 8 48 Saperda lateralis Fabriciusa 55 0.87 0 55 0 Graphisurus fasciatus (DeGeer) 43 0.68 0 41 2 Neoclytus acuminatus (Fabricius) 37 0.58 5 2 30 Megacyllene caryae (Gahan)a 34 0.54 0 34 0 Monochamus sp. 7 0.11 7 0 0 Heterachthes pallidus Haldemana 6 0.09 0 0 6 Neoclytus sp. 6 0.09 6 0 0 Curius dentatus Newman 2 0.03 0 2 0 Graphisurus despectus (Le Conte)a 2 0.03 0 2 0 Liopinus mimeticus (Casey) 2 0.03 2 0 0 Achryson surinamum (Linnaeus)a 1 0.02 0 1 0 Astylopsis collaris (Haldeman) 1 0.02 0 1 0 Astylopsis macula (Say) 1 0.02 0 1 0 Dorcaschema cinereum (Olivier)a 1 0.02 0 1 0 Euderces pini (Olivier) 1 0.02 0 1 0 Lepturges confluens (Haldeman)a 1 0.02 0 0 1 Lepturges pictus (Le Conte)a 1 0.02 0 0 1 Tilloclytus geminatus (Haldeman) 1 0.02 0 0 1 Total 6,352 3,012 2,395 945 aSpecies only found from mockernut hickory. Open in new tab Table 3. Cerambycidae species emerging from mockernut hickory bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2008 in order of abundance Year Cerambycidae species Total Relative abundance (%) 2006 2007 2008 Neoclytus mucronatus (Fabricius) 3,052 48.1 1,777 487 788 Saperda discoidea Fabricius 1,880 29.6 345 1,478 57 Xylotrechus colonus (Fabricius) 1,162 18.3 870 281 11 Stenosphenus notatus (Olivier)a 56 0.88 0 8 48 Saperda lateralis Fabriciusa 55 0.87 0 55 0 Graphisurus fasciatus (DeGeer) 43 0.68 0 41 2 Neoclytus acuminatus (Fabricius) 37 0.58 5 2 30 Megacyllene caryae (Gahan)a 34 0.54 0 34 0 Monochamus sp. 7 0.11 7 0 0 Heterachthes pallidus Haldemana 6 0.09 0 0 6 Neoclytus sp. 6 0.09 6 0 0 Curius dentatus Newman 2 0.03 0 2 0 Graphisurus despectus (Le Conte)a 2 0.03 0 2 0 Liopinus mimeticus (Casey) 2 0.03 2 0 0 Achryson surinamum (Linnaeus)a 1 0.02 0 1 0 Astylopsis collaris (Haldeman) 1 0.02 0 1 0 Astylopsis macula (Say) 1 0.02 0 1 0 Dorcaschema cinereum (Olivier)a 1 0.02 0 1 0 Euderces pini (Olivier) 1 0.02 0 1 0 Lepturges confluens (Haldeman)a 1 0.02 0 0 1 Lepturges pictus (Le Conte)a 1 0.02 0 0 1 Tilloclytus geminatus (Haldeman) 1 0.02 0 0 1 Total 6,352 3,012 2,395 945 Year Cerambycidae species Total Relative abundance (%) 2006 2007 2008 Neoclytus mucronatus (Fabricius) 3,052 48.1 1,777 487 788 Saperda discoidea Fabricius 1,880 29.6 345 1,478 57 Xylotrechus colonus (Fabricius) 1,162 18.3 870 281 11 Stenosphenus notatus (Olivier)a 56 0.88 0 8 48 Saperda lateralis Fabriciusa 55 0.87 0 55 0 Graphisurus fasciatus (DeGeer) 43 0.68 0 41 2 Neoclytus acuminatus (Fabricius) 37 0.58 5 2 30 Megacyllene caryae (Gahan)a 34 0.54 0 34 0 Monochamus sp. 7 0.11 7 0 0 Heterachthes pallidus Haldemana 6 0.09 0 0 6 Neoclytus sp. 6 0.09 6 0 0 Curius dentatus Newman 2 0.03 0 2 0 Graphisurus despectus (Le Conte)a 2 0.03 0 2 0 Liopinus mimeticus (Casey) 2 0.03 2 0 0 Achryson surinamum (Linnaeus)a 1 0.02 0 1 0 Astylopsis collaris (Haldeman) 1 0.02 0 1 0 Astylopsis macula (Say) 1 0.02 0 1 0 Dorcaschema cinereum (Olivier)a 1 0.02 0 1 0 Euderces pini (Olivier) 1 0.02 0 1 0 Lepturges confluens (Haldeman)a 1 0.02 0 0 1 Lepturges pictus (Le Conte)a 1 0.02 0 0 1 Tilloclytus geminatus (Haldeman) 1 0.02 0 0 1 Total 6,352 3,012 2,395 945 aSpecies only found from mockernut hickory. Open in new tab Table 4. Cerambycidae emerging from red maple bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2013 in order of abundance Cerambycidae species Total Relative abundance (%) Year 2006 2007 2008 2009 2010 2011 2012 2013 Neoclytus acuminatus (Fabricius) 4,506 56.6 63 181 552 355 155 1,622 677 901 Elaphidion mucronatum (Say)a 1,067 13.4 0 2 26 0 11 555 444 29 Graphisurus fasciatus (DeGeer) 712 8.94 0 6 61 92 16 337 165 35 Curius dentatus Newman 492 6.18 2 11 11 30 3 124 123 188 Euderces pini (Olivier)b 478 6.00 0 291 164 3 9 4 4 3 Astylopsis macula (Say) 218 2.74 3 22 11 18 4 85 58 17 Molorchus bimaculatus celti Knull 103 1.29 0 77 0 0 9 0 17 0 Tilloclytus geminatus (Haldeman)b 85 1.07 0 33 1 2 38 10 1 0 Liopinus mimeticus (Casey) 82 1.03 0 0 0 0 1 57 19 5 Urgleptes querci (Fitch)a 65 0.82 0 56 0 0 2 7 0 0 Hyperplatys aspersa (Say)a 39 0.49 0 7 3 2 3 17 1 6 Leptostylus asperatus (Haldeman)a 35 0.44 0 0 7 4 0 22 2 0 Parandra b. brunnea Fabriciusa,c 10 0.13 0 0 0 0 0 0 10 0 Aegomorphus modestus (Blais)a 9 0.11 1 0 1 0 0 3 0 4 Xylotrechus colonus (Fabricius) 8 0.10 1 1 2 1 0 2 1 0 Neoclytus mucronatus (Fabricius) 7 0.09 2 1 4 0 0 0 0 0 Molorchus bimaculatus bimaculatus Saya 6 0.08 0 0 0 0 2 3 0 1 Anelaphus pumilus (Newman)a,c 5 0.06 0 2 0 0 0 2 0 1 Molorchus bimaculatus Saya 4 0.05 0 0 0 0 1 0 0 3 Anelaphus villosus (Fabricius)a,c 3 0.04 0 0 0 1 2 0 0 0 Trachysida mutabilis (Newman)a 3 0.04 0 3 0 0 0 0 0 0 Astylopsis arcuata (LeConte)a 2 0.03 0 0 2 0 0 0 0 0 Astylopsis collaris (Haldeman)b 2 0.03 0 1 1 0 0 0 0 0 Bellamira scalaris (Say)a 2 0.03 0 0 0 0 2 0 0 0 Ecyrus dasycerus (Say)a,c 2 0.03 0 0 0 0 1 1 0 0 Neoalosterna capitata (Newman)a,b 2 0.03 0 0 0 0 0 0 0 2 Neoclytus jouteli Davis 2 0.03 0 0 0 0 1 1 0 0 Parelaphidion aspersum (Haldeman)a 2 0.03 0 0 0 2 0 0 0 0 Strangalia acuminata (Olivier)a 2 0.03 0 1 0 1 0 0 0 0 Acanthocinus sp.a 1 0.01 1 0 0 0 0 0 0 0 Anelaphus parallelus (Newman)a 1 0.01 0 1 0 0 0 0 0 0 Brachyleptura rubrica (Say)a 1 0.01 0 0 0 0 1 0 0 0 Cyrtinus pygmaeus (Haldeman) 1 0.01 0 1 0 0 0 0 0 0 Ecyrus d. dasycerus (Say)a 1 0.01 0 1 0 0 0 0 0 0 Knulliana cincta (Drury)a,b 1 0.01 0 0 0 0 0 0 0 1 Stenelytrana emarginata (Fabricius)a 1 0.01 0 0 0 0 1 0 0 0 Pogonocherus mixtus Haldemana 1 0.01 0 1 0 0 0 0 0 0 Typocerus velutinus (Olivier)a 1 0.01 0 0 0 0 1 0 0 0 Xylotrechus aceris Fishera 1 0.01 0 0 0 0 0 0 0 1 Total 7,963 73 699 2,854 2,520 2,273 4,863 3,534 1,197 Cerambycidae species Total Relative abundance (%) Year 2006 2007 2008 2009 2010 2011 2012 2013 Neoclytus acuminatus (Fabricius) 4,506 56.6 63 181 552 355 155 1,622 677 901 Elaphidion mucronatum (Say)a 1,067 13.4 0 2 26 0 11 555 444 29 Graphisurus fasciatus (DeGeer) 712 8.94 0 6 61 92 16 337 165 35 Curius dentatus Newman 492 6.18 2 11 11 30 3 124 123 188 Euderces pini (Olivier)b 478 6.00 0 291 164 3 9 4 4 3 Astylopsis macula (Say) 218 2.74 3 22 11 18 4 85 58 17 Molorchus bimaculatus celti Knull 103 1.29 0 77 0 0 9 0 17 0 Tilloclytus geminatus (Haldeman)b 85 1.07 0 33 1 2 38 10 1 0 Liopinus mimeticus (Casey) 82 1.03 0 0 0 0 1 57 19 5 Urgleptes querci (Fitch)a 65 0.82 0 56 0 0 2 7 0 0 Hyperplatys aspersa (Say)a 39 0.49 0 7 3 2 3 17 1 6 Leptostylus asperatus (Haldeman)a 35 0.44 0 0 7 4 0 22 2 0 Parandra b. brunnea Fabriciusa,c 10 0.13 0 0 0 0 0 0 10 0 Aegomorphus modestus (Blais)a 9 0.11 1 0 1 0 0 3 0 4 Xylotrechus colonus (Fabricius) 8 0.10 1 1 2 1 0 2 1 0 Neoclytus mucronatus (Fabricius) 7 0.09 2 1 4 0 0 0 0 0 Molorchus bimaculatus bimaculatus Saya 6 0.08 0 0 0 0 2 3 0 1 Anelaphus pumilus (Newman)a,c 5 0.06 0 2 0 0 0 2 0 1 Molorchus bimaculatus Saya 4 0.05 0 0 0 0 1 0 0 3 Anelaphus villosus (Fabricius)a,c 3 0.04 0 0 0 1 2 0 0 0 Trachysida mutabilis (Newman)a 3 0.04 0 3 0 0 0 0 0 0 Astylopsis arcuata (LeConte)a 2 0.03 0 0 2 0 0 0 0 0 Astylopsis collaris (Haldeman)b 2 0.03 0 1 1 0 0 0 0 0 Bellamira scalaris (Say)a 2 0.03 0 0 0 0 2 0 0 0 Ecyrus dasycerus (Say)a,c 2 0.03 0 0 0 0 1 1 0 0 Neoalosterna capitata (Newman)a,b 2 0.03 0 0 0 0 0 0 0 2 Neoclytus jouteli Davis 2 0.03 0 0 0 0 1 1 0 0 Parelaphidion aspersum (Haldeman)a 2 0.03 0 0 0 2 0 0 0 0 Strangalia acuminata (Olivier)a 2 0.03 0 1 0 1 0 0 0 0 Acanthocinus sp.a 1 0.01 1 0 0 0 0 0 0 0 Anelaphus parallelus (Newman)a 1 0.01 0 1 0 0 0 0 0 0 Brachyleptura rubrica (Say)a 1 0.01 0 0 0 0 1 0 0 0 Cyrtinus pygmaeus (Haldeman) 1 0.01 0 1 0 0 0 0 0 0 Ecyrus d. dasycerus (Say)a 1 0.01 0 1 0 0 0 0 0 0 Knulliana cincta (Drury)a,b 1 0.01 0 0 0 0 0 0 0 1 Stenelytrana emarginata (Fabricius)a 1 0.01 0 0 0 0 1 0 0 0 Pogonocherus mixtus Haldemana 1 0.01 0 1 0 0 0 0 0 0 Typocerus velutinus (Olivier)a 1 0.01 0 0 0 0 1 0 0 0 Xylotrechus aceris Fishera 1 0.01 0 0 0 0 0 0 0 1 Total 7,963 73 699 2,854 2,520 2,273 4,863 3,534 1,197 aSpecies only found from red maple. bSpecies only reared from bolts collected on forest edge. cSpecies only reared from bolts collected in forest interior. Open in new tab Table 4. Cerambycidae emerging from red maple bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2013 in order of abundance Cerambycidae species Total Relative abundance (%) Year 2006 2007 2008 2009 2010 2011 2012 2013 Neoclytus acuminatus (Fabricius) 4,506 56.6 63 181 552 355 155 1,622 677 901 Elaphidion mucronatum (Say)a 1,067 13.4 0 2 26 0 11 555 444 29 Graphisurus fasciatus (DeGeer) 712 8.94 0 6 61 92 16 337 165 35 Curius dentatus Newman 492 6.18 2 11 11 30 3 124 123 188 Euderces pini (Olivier)b 478 6.00 0 291 164 3 9 4 4 3 Astylopsis macula (Say) 218 2.74 3 22 11 18 4 85 58 17 Molorchus bimaculatus celti Knull 103 1.29 0 77 0 0 9 0 17 0 Tilloclytus geminatus (Haldeman)b 85 1.07 0 33 1 2 38 10 1 0 Liopinus mimeticus (Casey) 82 1.03 0 0 0 0 1 57 19 5 Urgleptes querci (Fitch)a 65 0.82 0 56 0 0 2 7 0 0 Hyperplatys aspersa (Say)a 39 0.49 0 7 3 2 3 17 1 6 Leptostylus asperatus (Haldeman)a 35 0.44 0 0 7 4 0 22 2 0 Parandra b. brunnea Fabriciusa,c 10 0.13 0 0 0 0 0 0 10 0 Aegomorphus modestus (Blais)a 9 0.11 1 0 1 0 0 3 0 4 Xylotrechus colonus (Fabricius) 8 0.10 1 1 2 1 0 2 1 0 Neoclytus mucronatus (Fabricius) 7 0.09 2 1 4 0 0 0 0 0 Molorchus bimaculatus bimaculatus Saya 6 0.08 0 0 0 0 2 3 0 1 Anelaphus pumilus (Newman)a,c 5 0.06 0 2 0 0 0 2 0 1 Molorchus bimaculatus Saya 4 0.05 0 0 0 0 1 0 0 3 Anelaphus villosus (Fabricius)a,c 3 0.04 0 0 0 1 2 0 0 0 Trachysida mutabilis (Newman)a 3 0.04 0 3 0 0 0 0 0 0 Astylopsis arcuata (LeConte)a 2 0.03 0 0 2 0 0 0 0 0 Astylopsis collaris (Haldeman)b 2 0.03 0 1 1 0 0 0 0 0 Bellamira scalaris (Say)a 2 0.03 0 0 0 0 2 0 0 0 Ecyrus dasycerus (Say)a,c 2 0.03 0 0 0 0 1 1 0 0 Neoalosterna capitata (Newman)a,b 2 0.03 0 0 0 0 0 0 0 2 Neoclytus jouteli Davis 2 0.03 0 0 0 0 1 1 0 0 Parelaphidion aspersum (Haldeman)a 2 0.03 0 0 0 2 0 0 0 0 Strangalia acuminata (Olivier)a 2 0.03 0 1 0 1 0 0 0 0 Acanthocinus sp.a 1 0.01 1 0 0 0 0 0 0 0 Anelaphus parallelus (Newman)a 1 0.01 0 1 0 0 0 0 0 0 Brachyleptura rubrica (Say)a 1 0.01 0 0 0 0 1 0 0 0 Cyrtinus pygmaeus (Haldeman) 1 0.01 0 1 0 0 0 0 0 0 Ecyrus d. dasycerus (Say)a 1 0.01 0 1 0 0 0 0 0 0 Knulliana cincta (Drury)a,b 1 0.01 0 0 0 0 0 0 0 1 Stenelytrana emarginata (Fabricius)a 1 0.01 0 0 0 0 1 0 0 0 Pogonocherus mixtus Haldemana 1 0.01 0 1 0 0 0 0 0 0 Typocerus velutinus (Olivier)a 1 0.01 0 0 0 0 1 0 0 0 Xylotrechus aceris Fishera 1 0.01 0 0 0 0 0 0 0 1 Total 7,963 73 699 2,854 2,520 2,273 4,863 3,534 1,197 Cerambycidae species Total Relative abundance (%) Year 2006 2007 2008 2009 2010 2011 2012 2013 Neoclytus acuminatus (Fabricius) 4,506 56.6 63 181 552 355 155 1,622 677 901 Elaphidion mucronatum (Say)a 1,067 13.4 0 2 26 0 11 555 444 29 Graphisurus fasciatus (DeGeer) 712 8.94 0 6 61 92 16 337 165 35 Curius dentatus Newman 492 6.18 2 11 11 30 3 124 123 188 Euderces pini (Olivier)b 478 6.00 0 291 164 3 9 4 4 3 Astylopsis macula (Say) 218 2.74 3 22 11 18 4 85 58 17 Molorchus bimaculatus celti Knull 103 1.29 0 77 0 0 9 0 17 0 Tilloclytus geminatus (Haldeman)b 85 1.07 0 33 1 2 38 10 1 0 Liopinus mimeticus (Casey) 82 1.03 0 0 0 0 1 57 19 5 Urgleptes querci (Fitch)a 65 0.82 0 56 0 0 2 7 0 0 Hyperplatys aspersa (Say)a 39 0.49 0 7 3 2 3 17 1 6 Leptostylus asperatus (Haldeman)a 35 0.44 0 0 7 4 0 22 2 0 Parandra b. brunnea Fabriciusa,c 10 0.13 0 0 0 0 0 0 10 0 Aegomorphus modestus (Blais)a 9 0.11 1 0 1 0 0 3 0 4 Xylotrechus colonus (Fabricius) 8 0.10 1 1 2 1 0 2 1 0 Neoclytus mucronatus (Fabricius) 7 0.09 2 1 4 0 0 0 0 0 Molorchus bimaculatus bimaculatus Saya 6 0.08 0 0 0 0 2 3 0 1 Anelaphus pumilus (Newman)a,c 5 0.06 0 2 0 0 0 2 0 1 Molorchus bimaculatus Saya 4 0.05 0 0 0 0 1 0 0 3 Anelaphus villosus (Fabricius)a,c 3 0.04 0 0 0 1 2 0 0 0 Trachysida mutabilis (Newman)a 3 0.04 0 3 0 0 0 0 0 0 Astylopsis arcuata (LeConte)a 2 0.03 0 0 2 0 0 0 0 0 Astylopsis collaris (Haldeman)b 2 0.03 0 1 1 0 0 0 0 0 Bellamira scalaris (Say)a 2 0.03 0 0 0 0 2 0 0 0 Ecyrus dasycerus (Say)a,c 2 0.03 0 0 0 0 1 1 0 0 Neoalosterna capitata (Newman)a,b 2 0.03 0 0 0 0 0 0 0 2 Neoclytus jouteli Davis 2 0.03 0 0 0 0 1 1 0 0 Parelaphidion aspersum (Haldeman)a 2 0.03 0 0 0 2 0 0 0 0 Strangalia acuminata (Olivier)a 2 0.03 0 1 0 1 0 0 0 0 Acanthocinus sp.a 1 0.01 1 0 0 0 0 0 0 0 Anelaphus parallelus (Newman)a 1 0.01 0 1 0 0 0 0 0 0 Brachyleptura rubrica (Say)a 1 0.01 0 0 0 0 1 0 0 0 Cyrtinus pygmaeus (Haldeman) 1 0.01 0 1 0 0 0 0 0 0 Ecyrus d. dasycerus (Say)a 1 0.01 0 1 0 0 0 0 0 0 Knulliana cincta (Drury)a,b 1 0.01 0 0 0 0 0 0 0 1 Stenelytrana emarginata (Fabricius)a 1 0.01 0 0 0 0 1 0 0 0 Pogonocherus mixtus Haldemana 1 0.01 0 1 0 0 0 0 0 0 Typocerus velutinus (Olivier)a 1 0.01 0 0 0 0 1 0 0 0 Xylotrechus aceris Fishera 1 0.01 0 0 0 0 0 0 0 1 Total 7,963 73 699 2,854 2,520 2,273 4,863 3,534 1,197 aSpecies only found from red maple. bSpecies only reared from bolts collected on forest edge. cSpecies only reared from bolts collected in forest interior. Open in new tab Red Maple In total, 7,963 individuals comprising 39 species were collected from red maple bolts across all survey methods from 2006 to 2013 (Table 4). Of these species, 25 were unique to red maple (Table 4). Neoclytus acuminatus was the most abundant species (n = 4,506) accounting for approximately 58% of the total specimens collected (Table 4). The abundance of N. acuminatus combined with that of E. mucronatum (13%, n = 1,067), G. fasciatus (9%, n = 712), C. dentatus (6%, n = 492), E. pini (6%), and A. macula (Say) (3%, n = 218) represented nearly 94% (n = 7,576; Table 4) of the cerambycid specimens collected from red maple. Naturally infested red maple trees supported significantly higher cerambycid richness and diversity compared with felled and girdled trees (Table 5). Table 5. Effect of tree species and sampling method on Cerambycidae species richness and diversity from 2006 to 2008 Tree species Treatment n Cerambycidae Richnessa Diversity Virginia pine Naturally infested 6 1.17 ± 0.31a 0.22 ± 0.14a Felled 20 1.75 ± 0.16a 0.34 ± 0.07a Girdled 8 1.50 ± 0.63a 0.35 ± 0.18a F = 0.740 F = 0.245 P = 0.385 P = 0.784 Mockernut hickory Felled 20 3.70 ± 0.16a 0.87 ± 0.04a Girdled 26 3.69 ± 0.50a 0.69 ± 0.10a F = 0.002 F = 2.403 P = 0.989 P = 0.128 Red maple Naturally infested 36 2.78 ± 0.30a 0.69 ± 0.08a Felled 70 1.87 ± 0.16b 0.36 ± 0.05b Girdled 13 1.23 ± 0.23b 0.20 ± 0.09b F = 6.993 F = 10.286 P < 0.001 P < 0.001 Tree species Treatment n Cerambycidae Richnessa Diversity Virginia pine Naturally infested 6 1.17 ± 0.31a 0.22 ± 0.14a Felled 20 1.75 ± 0.16a 0.34 ± 0.07a Girdled 8 1.50 ± 0.63a 0.35 ± 0.18a F = 0.740 F = 0.245 P = 0.385 P = 0.784 Mockernut hickory Felled 20 3.70 ± 0.16a 0.87 ± 0.04a Girdled 26 3.69 ± 0.50a 0.69 ± 0.10a F = 0.002 F = 2.403 P = 0.989 P = 0.128 Red maple Naturally infested 36 2.78 ± 0.30a 0.69 ± 0.08a Felled 70 1.87 ± 0.16b 0.36 ± 0.05b Girdled 13 1.23 ± 0.23b 0.20 ± 0.09b F = 6.993 F = 10.286 P < 0.001 P < 0.001 aDifferent letters within columns indicate significant differences between treatments via Tukey’s Honestly Significant Difference test (P = 0.05). Open in new tab Table 5. Effect of tree species and sampling method on Cerambycidae species richness and diversity from 2006 to 2008 Tree species Treatment n Cerambycidae Richnessa Diversity Virginia pine Naturally infested 6 1.17 ± 0.31a 0.22 ± 0.14a Felled 20 1.75 ± 0.16a 0.34 ± 0.07a Girdled 8 1.50 ± 0.63a 0.35 ± 0.18a F = 0.740 F = 0.245 P = 0.385 P = 0.784 Mockernut hickory Felled 20 3.70 ± 0.16a 0.87 ± 0.04a Girdled 26 3.69 ± 0.50a 0.69 ± 0.10a F = 0.002 F = 2.403 P = 0.989 P = 0.128 Red maple Naturally infested 36 2.78 ± 0.30a 0.69 ± 0.08a Felled 70 1.87 ± 0.16b 0.36 ± 0.05b Girdled 13 1.23 ± 0.23b 0.20 ± 0.09b F = 6.993 F = 10.286 P < 0.001 P < 0.001 Tree species Treatment n Cerambycidae Richnessa Diversity Virginia pine Naturally infested 6 1.17 ± 0.31a 0.22 ± 0.14a Felled 20 1.75 ± 0.16a 0.34 ± 0.07a Girdled 8 1.50 ± 0.63a 0.35 ± 0.18a F = 0.740 F = 0.245 P = 0.385 P = 0.784 Mockernut hickory Felled 20 3.70 ± 0.16a 0.87 ± 0.04a Girdled 26 3.69 ± 0.50a 0.69 ± 0.10a F = 0.002 F = 2.403 P = 0.989 P = 0.128 Red maple Naturally infested 36 2.78 ± 0.30a 0.69 ± 0.08a Felled 70 1.87 ± 0.16b 0.36 ± 0.05b Girdled 13 1.23 ± 0.23b 0.20 ± 0.09b F = 6.993 F = 10.286 P < 0.001 P < 0.001 aDifferent letters within columns indicate significant differences between treatments via Tukey’s Honestly Significant Difference test (P = 0.05). Open in new tab Effects of Mechanical Stress Treatments on Cerambycidae Richness and Diversity Overall, mockernut hickory (felled and girdled treatments only) supported nearly two times the cerambycid richness (df = 2, 200; F = 25.05; P < 0.001) and diversity (df = 2, 200; F = 15.44; P < 0.0001; Fig. 4) than did the other stressed tree species (red maple and Virginia pine) from 2006 to 2008. Fig. 4. Open in new tabDownload slide Cerambycidae species richness (A) and diversity (B) from mechanically stressed (pooled girdled and felled treatments) trees only from 2008 to 2006. Different letters above bars indicate significant differences via Tukey’s Honestly Significant Difference test (P = 0.05). Fig. 4. Open in new tabDownload slide Cerambycidae species richness (A) and diversity (B) from mechanically stressed (pooled girdled and felled treatments) trees only from 2008 to 2006. Different letters above bars indicate significant differences via Tukey’s Honestly Significant Difference test (P = 0.05). Effect of Red Maple Location on Cerambycidae Richness and Diversity Red maple location (edge vs interior) had no effect on cerambycid richness (df = 1, 204; F = 0.187; P = 0.666) or diversity (df = 1, 204; F = 0.119; P = 0.731). Mean cerambycid richness and diversity from the forest edge was 2.5 ± 0.2 and 0.54 ± 0.05, respectively, whereas mean richness and diversity from the forest interior was 2.4 ± 0.2 and 0.52 ± 0.05, respectively. Several species were found only inhabiting red maple trees collected from either the forest edge or interior. For instance, Anelaphus pumilus (Newman), Anelaphus villosus (Fabricius), Ecyrus dasycerus (Say), and Parandra b. brunnea Fabricius larvae were only found from red maple trees harvested at the forest exterior (Table 4). Conversely, Astylopsis collaris (Haldeman), E. pini, Knulliana cincta (Drury), Neoalosterna capitata (Newman), and Tilloclytus geminatus (Haldeman) larvae were only found from red maples tree in the interior (Table 4). Braconidae Emerging from Woodborer-Infested Virginia Pine, Mockernut Hickory, and Red Maple More than 14,000 individual braconid specimens were collected. The last author (R.R.K.) examined a subsample of 495 specimens reared from cerambycid-infested Virginia pine, mockernut hickory, and red maple bolts (all sampling methods combined) from 2006 to 2008 and unequivocally determined 420 specimens to species or morphospecies. This resulted in 13 known species and two unknown species of Cenocoelius and Doryctes (Table 6). Across all tree species (n = 420), 77% of the braconid specimens belonged to either Ontsira mellipes (Ashmead [Hymenoptera: Braconidae]; n = 223) or Rhoptrocentrus piceus Marshall (Hymenoptera: Braconidae) (n = 100; Table 6). Relative abundances of braconid species varied between tree species. For instance, O. mellipes (n = 12), Spathius parvulus Matthews (n = 7), and Cenocoelius nigrisoma (Rohwer) (n = 4) comprised >88% (n = 26) of specimens identified from Virginia pine (Fig. 5), whereas >95% (n = 326) of braconid specimens from red maple were identified as O. mellipes (n = 210) and R. piceus (n = 100; Fig. 5). Conversely, Wroughtonia ligator (Say) (n = 16), Doryctes erythromelas (Brullé) (n = 16), S. floridanus (n = 13), Doryctes anatolikus Marsh (n = 12), and Doryctes fartus (Provancher) (n = 6) comprised nearly 93% (n = 68) of specimens identified from mockernut hickory (Fig. 5). Table 6. Braconidae emerging from woodborer-infested bolts (all tree species and sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2008 in order of abundance Braconidae species Total Relative abundance (%) Ontsira mellipes (Ashmead) 223 53.1 Rhoptrocentrus piceus Marshall 100 23.8 Wroughtonia ligator (Say) 20 4.76 Doryctes erythromelas (Brullé) 17 4.05 Spathius floridanus Ashmead 13 3.10 Doryctes anatolikus Marsh 12 2.86 Doryctes fartus (Provancher) 10 2.38 Spathius parvulus Matthews 7 1.67 Cenocoelius nigrisoma (Rohwer) 6 1.43 Doryctes rufipes (Provancher) 3 0.71 Cenocoelius sp. 2 0.48 Doryctes sp. 2 0.48 Ecphylus lepturgi Rohwer 2 0.48 Spathius laflammei Provancher 2 0.48 Atanycolus ulmicola (Viereck) 1 0.24 Total 420 Braconidae species Total Relative abundance (%) Ontsira mellipes (Ashmead) 223 53.1 Rhoptrocentrus piceus Marshall 100 23.8 Wroughtonia ligator (Say) 20 4.76 Doryctes erythromelas (Brullé) 17 4.05 Spathius floridanus Ashmead 13 3.10 Doryctes anatolikus Marsh 12 2.86 Doryctes fartus (Provancher) 10 2.38 Spathius parvulus Matthews 7 1.67 Cenocoelius nigrisoma (Rohwer) 6 1.43 Doryctes rufipes (Provancher) 3 0.71 Cenocoelius sp. 2 0.48 Doryctes sp. 2 0.48 Ecphylus lepturgi Rohwer 2 0.48 Spathius laflammei Provancher 2 0.48 Atanycolus ulmicola (Viereck) 1 0.24 Total 420 Open in new tab Table 6. Braconidae emerging from woodborer-infested bolts (all tree species and sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2008 in order of abundance Braconidae species Total Relative abundance (%) Ontsira mellipes (Ashmead) 223 53.1 Rhoptrocentrus piceus Marshall 100 23.8 Wroughtonia ligator (Say) 20 4.76 Doryctes erythromelas (Brullé) 17 4.05 Spathius floridanus Ashmead 13 3.10 Doryctes anatolikus Marsh 12 2.86 Doryctes fartus (Provancher) 10 2.38 Spathius parvulus Matthews 7 1.67 Cenocoelius nigrisoma (Rohwer) 6 1.43 Doryctes rufipes (Provancher) 3 0.71 Cenocoelius sp. 2 0.48 Doryctes sp. 2 0.48 Ecphylus lepturgi Rohwer 2 0.48 Spathius laflammei Provancher 2 0.48 Atanycolus ulmicola (Viereck) 1 0.24 Total 420 Braconidae species Total Relative abundance (%) Ontsira mellipes (Ashmead) 223 53.1 Rhoptrocentrus piceus Marshall 100 23.8 Wroughtonia ligator (Say) 20 4.76 Doryctes erythromelas (Brullé) 17 4.05 Spathius floridanus Ashmead 13 3.10 Doryctes anatolikus Marsh 12 2.86 Doryctes fartus (Provancher) 10 2.38 Spathius parvulus Matthews 7 1.67 Cenocoelius nigrisoma (Rohwer) 6 1.43 Doryctes rufipes (Provancher) 3 0.71 Cenocoelius sp. 2 0.48 Doryctes sp. 2 0.48 Ecphylus lepturgi Rohwer 2 0.48 Spathius laflammei Provancher 2 0.48 Atanycolus ulmicola (Viereck) 1 0.24 Total 420 Open in new tab Fig. 5. Open in new tabDownload slide Relative abundance of unequivocally (n = 420) identified Braconidae from a subsample (n = 495) obtained from woodborer-infested Virginia pine, mockernut hickory, and red maple bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2008. Fig. 5. Open in new tabDownload slide Relative abundance of unequivocally (n = 420) identified Braconidae from a subsample (n = 495) obtained from woodborer-infested Virginia pine, mockernut hickory, and red maple bolts (all sampling methods combined) collected from Blackbird State Forest (Delaware) from 2006 to 2008. Identification was equivocal for 75 of the 495 specimens examined; these were identified to the extent possible and consisted of Heterospilus, Spathius, Doryctes, and Atanycolus. Heterospilus comprised 54 of the specimens and were tentatively sorted into seven morphospecies collectively from red maple, mockernut hickory, and Virginia pine. An additional 13 specimens of Spathius are morphologically similar to S. floridanus (n = 6, from mockernut hickory and red maple) and S. parvulus (n = 7, from Virginia pine) but could not be placed in those species unequivocally. This was also the case for six specimens of Doryctes morphologically similar to D. fartus (n = 4) and D. erythromelas (n = 2) from mockernut hickory. Last, two specimens of Atanycolus reared from mockernut hickory are possibly A. simplex. Discussion Native Cerambycidae serve important ecological roles within forest ecosystems; however, they can also have extensive impacts on economically important wood products, urban and shade trees, and fruit and nut production (Solomon 1995). Moreover, the continued threat from invasive exotic Cerambycidae significantly affects both the tree species they infest and the arthropods and vertebrates that depend on these resources (e.g., Gandhi and Herms 2010). Thus, identifying larval cerambycid hosts and associated native natural enemies of these larvae may assist in improving approaches to managing outbreaks of native cerambycid species. Furthermore, enhancing our understanding of native natural enemies associated with cerambycids may aid in selecting native natural enemies that are effective in managing invasive exotic pests, particularly when they lack host specific and coevolved natural enemies. Studies have examined cerambycid diversity throughout the United States (Yanega 1996, Thomas et al. 2005, Lingafelter 2007, MacRae and Rice 2007, Hart et al. 2013), yet few have focused on the mid-Atlantic region particularly Delaware and the surrounding states of Pennsylvania, Maryland, and New Jersey. Many previous studies have used pheromones, host volatiles, or a combination of the three in baited flight intercept traps to capture active adult beetles. These methods are advantageous for passively capturing large numbers of adult Cerambycidae and assessing species presence and absence, numbers of individuals captured, phenology, and species richness and diversity (e.g., Hanks and Millar 2013, Hanks et al. 2014). However, effective attractants of many Cerambycidae are currently unknown, and this limits the utility of these traps to capture these species (Handley et al. 2015, Hanks and Millar 2016). Furthermore, of the synthetic pheromones that have been developed, cerambycid species may respond differently to varying concentrations of pheromones (Hanks et al. 2014). This suggests that pheromone-baited traps may under-represent the beetle species present, as well as their abundances in ecosystems where baited traps are located (Handley et al. 2015). Also, critical data on host relationships cannot be established from these methods, and consequently, cerambycid habitat requirements remain undetermined. Our approach, although labor and time intensive, may have improved our understanding of the cerambycid community by not influencing or biasing beetle species and numbers via lure and trap types or trap location, as has been evidenced by previous studies (McIntosh et al. 2001, Dodds et al. 2010, Handley et al. 2015). Through this approach we were able to provide precise host–habitat information on all Cerambycidae collected, as well as generating extensive data on other insect taxa associated with the sampled tree species. Moreover, data on associations of Cerambycidae, their natural enemies, and the other insect taxa identified serve as important factors in predicting the potential impacts from exotic invasive species that threaten red maple, mockernut hickory, and Virginia pine. The most abundant cerambycid species collected during this survey were similar to two previous investigations from Delaware and one from Pennsylvania despite their use of pheromone-, pheromone-plant volatile, or ethanol-baited flight interception traps. For example, in fragmented forest habitats in Delaware, Dunn et al. (2016) and Handley et al. (2015) collected large numbers of X. colonus, E. mucronatum, N. mucronatus, and G. fasciatus. Similarly, Hanks and Millar (2013) captured large numbers of N. a. acuminatus and X. colonus in several Pennsylvania counties. In contrast, none of the previous investigations captured S. discoidea (the third most abundant species collected in this study) in comparable numbers. Variation in the cerambycid species and numbers captured in this study and from those using flight intercept traps probably resulted from the differential attraction of beetle species to pheromones versus host plant volatiles, trap deployment times (night vs day) and dates, or disparities between tree species (hosts) present and wood types available (live, dead, dying, or decaying; Ohsawa 2008, Mitchell et al. 2015, Skabeikis et al. 2016, Wong et al. 2017). Whereas tree-stressing (felled vs girdled) treatments had no effect on cerambycid richness and diversity from Virginia pine and mockernut hickory, naturally infested red maple had nearly 1.5 times greater cerambycid richness and diversity compared with girdled and felled treatments. Many cerambycids vary in their wood type preference and feed on a range of woody tissues from living to dead, dying, or weakened in addition to stressed and standing or recently felled wood (Hanks 1999, Jonsell and Weslien 2003). Moreover, as wood conditions decline through tree death and decay, successional changes in cerambycid community composition occur (Gibb et al. 2006, Saint-Germain et al. 2007, Lee et al. 2014, Haack 2017). The most abundantly collected cerambycids, N. acuminatus, N. mucronatus, and X. colonus, are polyphagous and typically feed on weakened and dead trees; yet, they have also been observed on freshly felled boles (Linsley 1964, Yanega 1996, Ginzel and Hanks 2004, Haack 2017). Conversely, S. discoidea is primarily known to feed on living tree tissues (Haack 2017). Increased cerambycid richness and diversity on standing infested red maple compared with mechanically induced stressed treatments (girdled and felled) may have resulted from differences in the availability of standing infested trees compared with the more ephemeral mechanically damaged trees. Yet another possible variable contributing to increased cerambycid richness and diversity between treatments may have resulted from differences between volatile organic compounds (VOCs) emitted from mechanically damaged and naturally infested host plants. Long-distance host location in some cerambycids is mediated by host plant VOCs (e.g., Fettköther et al. 2000, Barata et al. 2002, Allison et al. 2004); however, different stressors such as girdling, felling, and insect herbivory can alter VOC composition and in turn affect insect attraction to stressed hosts (Crook et al. 2008, McCullough et al. 2009). The effect of environmental gradients on distribution patterns of cerambycids is limited and often is highly variable, particularly of perpendicular forest interior-edge gradients. For instance, Dodds (2011) examined the effect of trap placement between forest interior, edge, and clearings on cerambycid richness and abundance and observed higher abundance and richness from edge habitats compared with the interior and clearing, as well as a greater number of unique species captured along the forest edge and clearing. Conversely, Allison et al. (2018) observed cerambycid abundance to be highest in the interior and decreasing as the distance to the forest edge increased. In contrast to Dodds (2011) and Allison et al. (2018), we found no differences in either cerambycid richness or diversity between red maples harvested from forest edge or interior habitats. Criticisms of classical biological control have renewed interest in the potential contribution of native parasitoids to the management of invasive woodboring species. Although current management of woodborers such as A. planipennis has focused on classical biological control, considerable work has been done investigating native (resident) hymenopteran species attacking this beetle since its arrival in North America. For instance, field surveys in Michigan, Pennsylvania, Ohio, and Ontario, Canada have so far identified 12 resident parasitoid species in North American attacking and developing on A. planipennis (Bauer et al. 2005, 2008; Duan et al. 2009, 2015a; Kula et al. 2010; Lyons 2010). Although we were unable to determine parasitoid–host relationships during this study due to the concealed nature of woodboring Cerambycidae and their associated parasitoids, all Braconidae emerging from cerambycid-infested trees during this study belonged the subfamilies Braconinae, Cenocoelinae, Doryctinae, and Helconinae. Of these, the Braconinae are predominately idiobiont ectoparasitoids of concealed insect larvae, primarily in the orders Coleoptera and Lepidoptera but also Diptera and Hymenoptera (Quicke 2015). The Doryctinae are almost exclusively idiobiont ectoparasitoids of larvae in various insect orders but predominantly Coleoptera and frequently woodboring beetle larvae (Marsh 1997, Quicke 2015). Helconinae sensu stricto are solitary koinobiont endoparasitoids of woodboring beetles, particularly Cerambycidae (Sharkey 1997, Quicke 2015). Last, as far as is known, the Cenocoeliinae are solitary koinobiont endoparasitoids of concealed coleopteran larvae (Van Achterberg 1997, Quicke 2015). Many species in these braconid subfamilies play important roles in regulating native woodborers and have been used in biocontrol programs against invasive woodborers. For instance, the doryctines Rhoptrocentrus quercusi Yang and Chao, Doryctes petiolatus Shestakov, and Zombrus bicolor (Enderlein) are important native parasitoids of M. raddei, a considerable pest of Quercus spp. in China (Cao et al. 2015). Moreover, Syngaster lepidus Brullé and Jarra phoracantha Marsh and Austin (Doryctinae) are important natural enemies and classical biocontrol agents of the invasive eucalyptus borers Phoracantha semipunctata (Fabricius) and Phoracantha recurva Newman (Millar et al. 2002). The helconine, Helconidea dentator (Fabricius), and the cenocoeliine, Lestricus secalis (L.), are important parasitoids of woodboring cerambycids, such as those in Tetropium and Pogonocherus, that infest both living and dead conifers throughout Europe (Hilszczański 1998, Kenis and Hilszczanski 2004). Aside from D. fartus, for which host use is unknown, all species in Table 6 except S. parvulus have been reported previously as parasitoids of Cerambycidae; however, none of them have been reported from A. glabripennis or a congener. Doryctes fartus and S. parvulus have been reported as associated with pine, with the latter species reported previously from curculionids (Yu et al. 2016). The most abundant parasitoid in this survey, O. mellipes (Table 6), has been reported previously from an undetermined species of Callidium (Coleoptera: Cerambycidae) in Juniperus virginiana L. (Marsh 1966, Yu et al. 2016). The second most abundant parasitoid, R. piceus (Table 6), has been reported previously from cerambycids in six genera, as well beetles in three other families and hosts from one family each of Hymenoptera and Lepidoptera (Yu et al. 2016, host plant associations not listed). Three of the species in Table 6 are koinobiont endoparasitoids: W. ligator, C. nigrisoma, and Cenocoelius sp.; all other species are idiobiont ectoparasitoids, including S. parvulus. Notable among the life history of idiobiont ectoparasitoids is that their host is paralyzed and its development terminated upon oviposition; the immature stages of the parasitoid usually feed externally (Cornell and Hawkins 1993). In contrast, in koinobiont endoparasitoids, host development continues upon parasitoid oviposition, and the immature stages of the parasitoid usually feed internally. Therefore, koinobiont endoparasitoids must contend with host immune defense systems if they are to successfully emerge. Idiobiont species, on the other hand, typically do not require mechanisms to escape physiological host detection; thus, they are more capable of novel host exploration (Hawkins et al. 1992, Cornell and Hawkins 1993, Askew and Shaw 1986). This life-history trait has allowed idiobiont parasitoids to be successful in new-association approaches to managing exotic invasive woodboring pests (Daniel 1928, Steenburgh 1931, Allen et al. 1940, Hawkins et al. 1992, Cornell and Hawkins 1993). New-association biological control utilizes parasitoids with broad physiological host ranges that can acquire noncoevolved hosts (Hokkanen and Pimentel 1984) that fit within their ecological host range (Wiedenmann and Smith 1997). Because it is generally considered that exotic invasive pests are best managed by coevolved natural enemies, few efforts have focused on controlling such pests using new-association agents (Pimentel 1963) and in particular the use of native natural enemies against exotic pests in their newly invaded regions. Nevertheless, native natural enemies have been successfully implemented in new-association biological control programs. For instance, the North American braconid Macrocentrus ancylivora Rohwer (Hymenoptera), a dominant parasitoid attacking twig, stem, and fruit boring lepidopteran larvae, was found parasitizing and developing on the exotic invasive host Grapholita molesta (Busck) (Lepidoptera: Tortricidae), a newly destructive pest of peaches throughout northeastern North American (including Canada). This parasitoid was subsequently mass released against this exotic woodboring pest in New York, New Jersey, Pennsylvania, and Ontario, Canada (Daniel 1928, Driggers 1930, Steenburgh 1931, Garman and Brigham 1933, Allen et al. 1940, Brunson and Allen 1944). Thus, the use of endemic natural enemies to suppress exotic invasive pests may be useful when coevolved natural enemies in the pests’ area of endemism cannot be found. One such example of invasive woodborers lacking effective coevolved natural enemies is A. glabripennis, a significant threat to forest and urban and suburban trees throughout North America and Europe. Three natural enemies, D. helophoroides, Scleroderma guani Xiao and Wu (Hymenoptera: Bethylidae), and Iphiaulax imposter (Scopoli) (Hymenoptera: Braconidae), have been discovered that attack the larval stage of A. glabripennis in its area of endemism (Tang et al. 1996, Wang et al. 1999, Gao and Li 2001, Hu et al. 2009), but none provide effective management of A. glabripennis or of other Anoplophora species in Asia (Turgeon and Smith 2013). Moreover, these are all generalist natural enemies and thus are not suitable for A. glabripennis management outside of Asia, as they could have detrimental effects on native North American woodboring beetle populations (Gould et al. 2018). Interestingly, several of the native North American braconid species found during this survey formed new-association with A. glabripennis in laboratory settings, including O. mellipes, R. piceus, S. laflammei, and two unidentified species of Heterospilus and Atanycolus (Duan et al. 2015b). In conclusion, the present field study documented the occurrence and relative abundance of ecologically and phylogenetically related cerambycids infesting mockernut hickory, Virginia pine, and red maple along with hymenopteran parasitoids (particularly Braconidae) associated with the infestations. These results provide insights about the possibility of recruitment of native parasitoids to invasive cerambycid pests in the framework of new association biological control. Although associations between native North American parasitoids and A. glabripennis are currently unknown in nature, we propose that rearing woodborers and their associated parasitoids presents a unique opportunity to discover potentially effective new-association agents for exotic invasive woodborers. Using the methods presented in this study, it is possible to screen a variety of native natural enemies against exotic woodborers to assess their potential as either stand-alone agents or as supplemental agents to compliment current biological control strategies for established exotic invasive woodborers. Acknowledgments We thank Jim Dobson (Northern Regional Forester, Delaware Department of Agriculture, Forest Service, Blackbird State Forest) for permission to carry out experiments and fell trees at Blackbird State Forest. 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TI - Cerambycid Communities and Their Associated Hymenopteran Parasitoids From Major Hardwood Trees in Delaware: Implications for Biocontrol of Invasive Longhorned Beetles
JO - Environmental Entomology
DO - 10.1093/ee/nvz169
DA - 2020-04-14
UR - https://www.deepdyve.com/lp/oxford-university-press/cerambycid-communities-and-their-associated-hymenopteran-parasitoids-r7Q6bbloo9
SP - 1
VL - Advance Article
IS - 
DP - DeepDyve
ER -