Southern pine beetle (Coleoptera: Curculionidae) and its associated insect community: similarities and key differences between northeastern and southeastern pine forests.

We examine consequences of climate-induced range expansion on community composition and diversity within trees attacked by the southern pine beetle (Dendroctonus frontalis Zimmermann). At the northernmost limit of the southern pine beetle range where populations have persisted for multiple years (currently Long Island, NY), we collected and reared bark samples and placed emergence traps on southern pine beetle-attacked pitch pine, Pinus rigida Mill. (Pinales: Pinaceae). From these samples, we quantified southern pine beetle gallery length and emergence as well as the diversity and abundance of all associated insects including known and suspected competitors, predators, and parasitoids. We compared our results to that of historic sampling data (1975-1997) in the core of southern pine beetle's range in the southern United States. Key community members were present in both the northern and southern regions; composition and relative abundances differed markedly. A key predator, the clerid beetle Thanasimus dubius (Fabricius) (Coleoptera: Cleridae), was present in similar densities in both regions. Southern pine beetle infested a greater proportion of the length of the tree bole in the North. This increased tree utilization may be a consequence of a lack of resource competition by Ips De Geer (Coleoptera: Curculionidae) engraver beetles, which we found only in very low abundance in the northern sites. We discuss the implications of these results in the context of southern pine beetle range expansion. Continued study of the southern pine beetle community and temporal southern pine beetle dynamics in the North will add to our current knowledge base and aid preservation of rare and ecologically valuable pine barrens of New England.

Firewood Transport as a Vector of Forest Pest Dispersal in North America: A Scoping Review.

Native and nonnative insects and diseases can result in detrimental impacts to trees and forests, including the loss of economic resources and ecosystem services. Increases in globalization and changing human behaviors have created new anthropogenic pathways for long distance pest dispersal. In North America, literature suggests that once a forest or tree pest is established, the movement of firewood by the general public for recreational or home heating purposes is one of the primary pathways for its dispersal. Understanding human perceptions and behaviors is essential to inform the most effective strategies for modifying firewood and pest dispersal by humans. This scoping review seeks to assess trends and gaps in the existing literature, as well as patterns in behavior related to forest pest dispersal through firewood movement in North America. We identified 76 documents that addressed this topic to which we applied inclusion and exclusion criteria to select articles for further analysis. Twenty-five articles met the inclusion criteria and were categorized based on five identified themes: 1) insect incidence in firewood, 2) insect dispersal via firewood, 3) recreational firewood movement, 4) firewood treatments, and 5) behavior and rule compliance. The selected articles show trends that suggest that firewood movement presents a risk for forest insect dispersal, but that behavior can be modified, and compliance, monitoring, and treatments should be strengthened. This scoping review found limited research about western United States, Mexico, and Canada, various insect species and other organisms, regulation and management, awareness, and behavioral dimensions of firewood movement.

Correction: Dodds, K.J.; DiGirolomo, M.F. Effect of Cleaning Multiple-Funnel Traps on Captures of Bark and Woodboring Beetles in Northeastern United States. Insects 2020, 11, 702.

The authors wish to make the following corrections to this paper [...].

Effect of Cleaning Multiple-Funnel Traps on Captures of Bark and Woodboring Beetles in Northeastern United States.

Two experiments were conducted in mixed hardwood-conifer forests in the northeastern United States to test the effects of cleaning surfactant and non-surfactant treated multiple-funnel traps used to catch bark and woodboring beetles. Large amounts of pollen and other debris often form a crust on the interior of traps (personal observations). Such surface deposits may provide footholds for beetles to escape capture in traps. In one experiment, we tested cleaned surfactant and non-surfactant traps against non-cleaned surfactant and non-surfactant traps. In a second experiment, we tested field cleaning of modified multiple-funnel traps as an alternative to substituting clean traps on each collection visit. There was no effect of surfactant treated traps, cleaned or not, on total beetles or individual bark beetle species captured. However, in situ cleaned traps were statistically better at capturing total beetles, total bark beetles, and several bark beetle species than non-cleaned control traps. Surfactant-treated non-modified traps and cleaned modified traps had higher species richness and abundance than other treatments at the site level. Our results suggest that cleaning traps to remove accumulated pollen and debris may be helpful for some species but would have limited benefit for broad-scale trapping of bark and woodboring beetles in northeastern forests.

Sap Beetles (Coleoptera: Nitidulidae) in Oak Forests of Two Northeastern States: A Comparison of Trapping Methods and Monitoring for Phoretic Fungi.

Oak wilt is slowly expanding in the northeastern United States. Several nitidulid beetle species are known vectors of the fungus [Bretziella fagacearum (Bretz) Z. W. De Beer, Marinc., T. A. Duong, and M. J. Wingf (Microascales: Ceratocystidaceae)] that causes this disease, acquiring spores from fungal mats on infected trees and transmitting them to uninfected trees. Survey and fungal isolation from captured nitidulid beetles could be an important tool for detecting the presence of this disease in a geographic area not previously known to have oak wilt. In preparation for monitoring activities in such areas, two trapping studies were conducted in the northeastern United States: 1) trap test comparing the efficacy of wind-oriented pipe, multiple-funnel, and modified pitfall traps for nitidulids and 2) wet and dry collection cup comparison. Lures were a combination of nitidulid pheromones and fermenting liquid. Results support the use of multiple-funnel traps over the other two trap types, for both targeted species-specific surveys and community sampling. More total nitidulids, Colopterus truncatus (Randall), and Glischrochilus fasciatus (Olivier) were captured in wet collection cups compared with dry cups. Twenty-seven fungal species were isolated, none of which were B. fagacearum. Many fungi isolated from beetles were plant pathogens, indicating that in addition to the oak wilt fungus, sap beetles may contribute to the spread of other plant diseases.

Suitability of eastern pines for oviposition and survival of Sirex noctilio F.

The European woodwasp, Sirex noctilio F., is a pest of pines in many areas around the world. Since its introduction to North America, the distribution of S. noctilio overlaps with a known host (Pinus sylvestris) and hosts native to North America. Direct comparisons of suitability for oviposition and larval survival among these pines have not been made. We tested the relative suitability of four common pine species in northeastern North America (P. sylvestris, P. resinosa, P. banksiana, and P. strobus) as hosts for S. noctilio in a controlled, but in situ experiment. In a mixed pine forest in northern Ontario, we caged S. noctilio mating pairs on 10 freshly cut pine logs of each species, and estimated oviposition, counted adult S. noctilio (F1 generation) that emerged from logs, and calculated survivorship from egg to adult. Pinus sylvestris and P. resinosa were optimal hosts according to all three metrics of S. noctilio performance. Pinus strobus was a suitable larval host, but was not perceived as such by females, as evidenced by lower oviposition. Pinus banksiana was perceived as a suitable host by females, but was the least suitable larval host. Our results suggest that P. sylvestris and P. resinosa are more suitable hosts, at least in cut logs, than P. strobus and P. banksiana for S. noctilio in eastern North America.

Do Native Insects and Associated Fungi Limit Non-Native Woodwasp, Sirex noctilio, Survival in a Newly Invaded Environment?

Sirex noctilio F. (Hymenoptera: Siricidae) is an introduced pest of pines (Pinus spp.) in several countries in the Southern Hemisphere. Although S. noctilio is established in North America (first discovered in 2004), it has not been a destructive pest there so far, where forest communities more closely resemble those in its native Eurasian range­where it is not a pest. To investigate the influence of the existing community of associated insects (competitors + natural enemies) and fungi (vectored by insects) on S. noctilio survival in North America, we examined stage-specific mortality factors and their relative importance, generating life tables drawn from experimentally-manipulated and natural cohorts of Sirex spp. (mostly S. noctilio, but some native S. nigricornis F.). For both natural and experimentally-manipulated cohorts, factors which acted during the earliest Sirex life stages, most likely tree resistance and/or competition among fungal associates, were paramount in dictating woodwasp survival. Experimentally-manipulated life tables revealed that protection from the community of associates resulted in a significantly, and substantially larger (>15x) S. noctilio F1 generation than exposure to it. Seventy percent of generation mortality in the exposed cohort was due to tree resistance or unknown causes early in larval development, which could have included competition among other bark- or wood-inhabiting insects and/or their fungal associates. Only 46% of generation mortality in the protected cohort was due to tree resistance and/or unknown causes. Parasitoids, particularly endoparasitoids (Ibalia spp.), showed limited ability to control S. noctilio, and reduced the experimentally-established cohort by only 11%, and natural cohorts an average of 3.4%. The relative importance of tree resistance vs. competition with bark- and wood-borers in reducing S. noctilio survival remains unclear. Tree resistance and/or competition likely contribute more than natural enemies in maintaining the S. noctilio population in North America below damaging levels.

Type of intercept trap not important for capturing female Sirex noctilio and S. nigricornis (Hymenoptera: Siricidae) in North America.

Current detection tools for Sirex noctilio F. (Hymenoptera Siricidae) in North America are poor. To determine the importance of intercept trap type for capturing females of S. noctilio and its native congener, Sirex nigricornis F., in eastern North America, we report on seven trap comparison studies from different years and geographic locations. Among studies, total numbers of S. noctilio captured were low (mean of < or = 1 wasp per trap). Total numbers of S. nigricornis caught were generally greater, and ranged from a mean of 1-13 wasps per trap. Nearly all studies found no significant differences among intercept trap types in the number of woodwasps caught. For future studies, we recommend that either panel or 12-unit Lindgren funnel traps be used to catch S. noctilio or S. nigricornis in eastern North America.

Effects of Invasive Winter Moth Defoliation on Tree Radial Growth in Eastern Massachusetts, USA.

Winter moth, Operophtera brumata L. (Lepidoptera: Geometridae), has been defoliating hardwood trees in eastern Massachusetts since the 1990s. Native to Europe, winter moth has also been detected in Rhode Island, Connecticut, eastern Long Island (NY), New Hampshire, and Maine. Individual tree impacts of winter moth defoliation in New England are currently unknown. Using dendroecological techniques, this study related annual radial growth of individual host (Quercus spp. and Acer spp.) trees to detailed defoliation estimates. Winter moth defoliation was associated with up to a 47% reduction in annual radial growth of Quercus trees. Latewood production of Quercus was reduced by up to 67% in the same year as defoliation, while earlywood production was reduced by up to 24% in the year following defoliation. Winter moth defoliation was not a strong predictor of radial growth in Acer species. This study is the first to document impacts of novel invasions of winter moth into New England.

Trap lure blend of pine volatiles and bark beetle pheromones for Monochamus spp. (Coleoptera: Cerambycidae) in pine forests of Canada and the United States.

In 2007-2008, we examined the flight responses of Monochamus titillator (F.) complex [M. titillator, Monochamus carolinensis (Olivier), and any possible hybrids], Monochamus scutellatus (Say), Monochamus clamator (LeConte), Monochamus obtusus Casey, and Monochamus mutator LeConte (Coleoptera: Cerambycidae) to multiple-funnel traps baited with and without host volatiles and bark beetle pheromones. Experiments were conducted in mature pine (Pinus) stands in Alberta (Canada), and Arkansas, Arizona, California, Florida, Idaho, Michigan, Montana, New Hampshire, North Carolina, Ohio, Oregon, Tennessee, Utah, and Wisconsin (United States). At each location, traps were deployed in 10 replicate blocks of four traps per block. The trap treatments were: 1) blank control; 2) ipsenol and ipsdienol; 3) ethanol and alpha-pinene; and 4) a quaternary blend of ipsenol, ipsdienol, ethanol, and alpha-pinene. All five species or species complex of Monochamus preferred traps baited with the quaternary blend over all other treatments. The consistency of these results across such a large geographic area suggests that similar selection pressures may be acting on Monochamus spp. in pine forests, regardless of variation in stand composition and climatic conditions. Our results suggest that multiple-funnel traps baited with the quaternary blend ofipsenol, ipsdienol, ethanol, and alpha-pinene may be highly effective for monitoring various Monochamus spp. in pine forests of North America, and may have utility in trapping and detection programs in North America and overseas.

Colonization of Three Maple Species by Asian Longhorned Beetle, Anoplophora glabripennis, in Two Mixed-Hardwood Forest Stands.

Asian longhorned beetle (ALB), Anoplophora glabripennis (Motschulsky), is an invasive insect that has successfully established multiple times in North America. To investigate host colonization and reproductive success (exit holes/eggs), two ALB infested forest stands were sampled in central Massachusetts, USA. Infested Acer platanoides L., Acer rubrum L., and Acer saccharum Marsh. were felled, bucked into 1 m sections and dissected to determine indications of ALB infestations, such as presence of life stages or signs of damage on trees. ALB damage was also aged on a subset of trees to determine the earliest attacks on the three Acer species. In one stand, ALB oviposition was significantly higher on the native A. rubrum and A. saccharum than the exotic A. platanoides. In the second stand, ALB oviposition was significantly higher and cumulative reproductive success was higher on A. rubrum than A. platanoides or A. saccharum. An A. saccharum had the earliest signs of attack that occurred in 2006. Acer rubrum (2007) and A. platanoides (2010) were colonized shortly thereafter. Overall, ALB was more successful in A. rubrum, where adults emerged from 53% and 64% of trees in each stand, compared to A. platanoides (11% and 18%) or A. saccharum (14% and 9%).

Arboreal insects associated with herbicide-stressed Pinus resinosa and Pinus sylvestris used as Sirex noctilio trap trees in New York.

In September of 2004, Sirex noctilio F. (Hymenoptera: Siricidae) was detected in New York State and later found to be established over a larger area, including parts of southeastern Canada and the northeastern United States. A key component of S. noctilio detection and management plans in other parts of the world where S. noctilio has become established are chemically girdled trap trees. Trap tree usage in North America is confounded by the presence of diverse communities of organisms that inhabit dead and dying trees. We trapped a portion of the arboreal insect community arriving at Pinus resinosa Ait. and Pinus sylvestris L., trap trees girdled 3 mo before (April), one month before (June), and at S. noctilio flight (July) in central New York. Multiple-funnel traps attached to trap trees captured 30,031 individuals from 109 species of Scolytinae, Cerambycidae, and Siricidae. Ips pini (Say) and Ips grandicollis (Eichhoff) accounted for almost 50% of the scolytines captured at trap trees and were present on all girdling dates. Significantly more scolytines and cerambycids were captured on P. sylvestris compared with P. resinosa, but species richness of captured insects did not differ between the two trees. More total and conifer-inhabiting scolytines and cerambycids were captured in traps on trees girdled in April and June and higher observed species richness was found on trees girdled in April and controls. Results from this study suggest a large community of arboreal insects and associated organisms are attracted to chemically girdled trap trees and likely interact with S. noctilio.

Effects of habitat type and trap placement on captures of bark (Coleoptera: Scolytidae) and longhorned (Coleoptera: Cerambycidae) beetles in semiochemical-baited traps.

Two studies were conducted to determine the effect of habitat selection and trap placement on catches of Scolytidae and Cerambycidae (Coleoptera) in northeastern U.S. forests. In a nonreplicated case study, four habitat types--closed canopy hardwood stand, closed canopy conifer stand, a low-intensity thinned Pinus strobus L. stand, and a high-intensity thinned P. strobus stand--were surveyed using alpha-pinene, ethanol, ipsenol, ipsdienol, and lanierone. Average trap catches, species richness, and the number of unique species captured were all highest in at least one of the thinned habitats. A second experiment that was replicated evaluated the placement of traps in relation to habitat patches. Semiochemical-baited traps (alpha-pinene and ethanol) were placed under a closed canopy forest, along an edge, and in a clearing and tested for effects on Scolytidae and Cerambycidae trap catches. Abundance and species richness were generally higher in the closed canopy and edge placements compared with traps in the open area. The highest number of unique species were captured in the edge and clearing.

Trap type, lure placement, and habitat effects on Cerambycidae and Scolytinae (Coleoptera) catches in the Northeastern United States.

Semiochemical-based exotic species surveys targeting forest Coleoptera have gradually expanded in North America and elsewhere. Determining how various factors affect trap catches and increase species richness in traps is important for maximizing the efficacy of survey efforts. Studies were conducted in southern Maine and New Hampshire by using ethanol and alpha-pinene as lures to determine the influence of trap type, lure placement and size, and habitat type on catches of Scolytinae and Cerambycidae in coniferous forests. Three trap types (canopy malaise, intercept panel, and multiple-funnel), three lure placements/sizes (standard placement, above trap, and enlarged), and two habitat types (margins of clearcuts and shelterwood) were tested in three experiments. The three trap types performed equally well in terms of average number of species captured, but the canopy malaise caught more unique species than the other traps. In most cases, traps with lures placed above traps caught fewer beetles than lures hanging from the side of traps or with an expanded surface area. Generally, more insects were captured in shelterwood treatments versus the margins of clearcuts.

Test of nonhost angiosperm volatiles and verbenone to protect trap trees for Sirex noctilio (Hymenoptera: Siricidae) from attacks by bark beetles (Coleoptera: Scolytidae) in the northeastern United States.

Sirex noctilio F. (Hymenoptera: Siricidae) is an invasive woodwasp, currently established in northeastern North America. In other regions of the world, stressed trap trees are used to monitor populations of S. noctilio and to provide inoculation points for the biological control nematode Deladenus siricidicola Bedding. However, the operational use of trap trees for S. noctilio in North America may be compromised by the large community of native organisms that inhabit stressed and dying pine trees. Common bark beetles such as Ips pini (Say) and Ips grandicollis (Eichhoff) (Coleoptera: Scolytidae) could potentially compete with S. noctilio and associates for resources on trap trees, possibly reducing the efficacy of trap trees as habitats for the woodwasp. In an attempt to develop a technology to mitigate this potential issue, three common semiochemical interruptants--conophthorin, green leaf volatile mix, and verbenone--were tested for effectiveness in reducing arrivals of I. pini and I. grandicollis on trap trees treated with herbicides in northeastern United States. In addition, the effects of these compounds were determined independently with pheromone-baited multiple-funnel traps. None of the interruptants reduced numbers of I. pini or I. grandicollis either arriving on trap trees or caught in pheromone-baited traps. However, verbenone increased catches of I. grandicollis in traps baited with its pheromone, ipsenol. The mix of green leaf volatiles reduced catches of a native ambrosia beetle, Gnathotrichus materiarius (Fitch), whereas verbenone reduced trap catches of an exotic ambrosia beetle, Xylosandrus germanus (Blandford). Catches of X. germanus in traps adjacent to trap trees were enhanced with conophthorin.