Responses of Native and Non-native Bark and Ambrosia Beetles (Coleoptera: Curculionidae: Scolytinae) to Different Chemical Attractants: Insights From the USDA Forest Service Early Detection and Rapid Response Program Data Analysis.

More than 60 non-native bark and ambrosia beetle species (Coleoptera: Curculionidae: Scolytinae) are established in North America and several have had severe negative impacts on ecosystems. Non-native scolytines can introduce fungi which may cause vascular wilts and compete with native fungi and lead to reductions in native species through host reduction. The Early Detection Rapid Response (EDRR) program was created by the USDA Forest Service in 2007 to detect non-native bark and ambrosia beetles and provide a baseline for tracking populations over time. This program has led to new collection records and increased communication among agencies to delimit non-native scolytine populations and perform appropriate management. Although insect responses to different lure types vary, it is unknown how different lures compare in attracting bark and ambrosia beetles. Our goal was to examine how lure combinations used in the EDRR program affect captures of bark and ambrosia beetle communities and to determine the most effective combination of lures for targeting non-native scolytines. The highest proportion of non-native scolytines was captured with ethanol, as was the greatest total number of species, and the most diverse beetle community. Traps with Ips (Coleoptera: Curculionidae) lures captured the highest proportion of native scolytines but the lowest total number of total species and was also the least diverse. Communities of scolytines differed significantly among lures, states, and years. While ethanol is an appropriate lure for generalist trapping and targeting a wide range of non-native bark and ambrosia beetles, more targeted lures are needed for monitoring certain species of non-natives.

The bark and ambrosia beetles (Coleoptera: Curculionidae: Scolytinae and Platypodinae) of American Samoa.

A survey of five of the islands of American Samoa was conducted from 2016-2018 utilizing multi-funnel traps baited with ethanol and quercivorol (attractants for xyleborine ambrosia beetles). Specimens of Scolytinae and Platypodinae from this survey, as well as specimens in the American Samoa Community College Collection were identified. A total of 53 species of Scolytinae and two species of Platypodinae are reported. Fourteen species of Scolytinae and one species of Platypodinae are reported as new to American Samoa. The following new synonymy and new combinations are proposed: Xyleborinus artelineatus (Beeson, 1929) (=Xyleborinus cinctipennis (Schedl, 1980) syn. nov.; Ambrosiophilus wilderi (Beeson, 1929) comb. nov. from Ambrosiodmus; Beaverium swezeyi (Beeson, 1929) comb. nov. from Xyleborus; and Debus fallaxoides (Schedl, 1955) comb. nov. from Xyleborus. The distribution on the islands of American Samoa and taxonomic history of each species is provided.

Establishment of a non-native xyleborine ambrosia beetle, Xyleborus monographus (Fabricius) (Coleoptera: Curculionidae: Scolytinae), new to North America in California.

Specimens of an ambrosia beetle, Xyleborus monographus (Fabricius), were found infesting oak trees in California. This is the first record of this species established in North America. Based on collection information, this species most likely has been established in the Napa County area for several years. A modified key to Xyleborus in North America, and diagnosis of the species is provided.

North American Xyleborini north of Mexico: a review and key to genera and species (Coleoptera, Curculionidae, Scolytinae).

Bark and ambrosia beetles (Scolytinae) are the most successful group of invasive wood borers worldwide, and the most invasive among them are species in the tribe Xyleborini. This haplodiploid, highly inbred, fungus-farming group is represented by 30 non-native species in North America, of which at least five are serious pests. The few identification resources for Xyleborini that exist are becoming outdated due to new species arrivals and nomenclatural changes. Here we present a new comprehensive key to Xyleborini currently known from the continental United States. Compared to the previous key, the following species have been added to the North American fauna: Ambrosiodmus minor (Stebbing), Ambrosiophilus nodulosus (Eggers), Anisandrus maiche Kurentsov, Coptoborus pseudotenuis (Schedl), Cyclorhipidion fukiense (Eggers), Dryocoetoides reticulatus Atkinson, Dryoxylon onoharaense (Murayama), Euwallacea interjectus (Blandford), Xyleborinus andrewesi (Blandford), Xyleborinus artestriatus (Eichhoff), Xyleborinus octiesdentatus (Murayama), Xyleborus bispinatus Eichhoff, Xyleborus seriatus Blandford, Xyleborus spinulosus Blandford, and Xylosandrus amputatus (Blandford).

First records of Cyclorhipidion fukiense (Eggers) (Coleoptera: Curculionidae: Scolytinae: Xyleborini), an ambrosia beetle native to Asia, in North America.

The Asian ambrosia beetle, Cyclorhipidion fukiense (Eggers) was detected for the first time in North America based on three specimens trapped in 2012 from three localities in South Carolina and two other specimens intercepted at the port of Savannah, Georgia, in 2010. The species is characterized, illustrated with high-resolution images, and compared with two other congeneric, adventive species (C. bodoanum and C. pelliculosum) presently established in eastern North America. Morphometric measurements are provided and a provisional key is presented to the species of Cyclorhipidion occurring in North America.

Bark and Ambrosia Beetles Show Different Invasion Patterns in the USA.

Non-native bark and ambrosia beetles represent a threat to forests worldwide. Their invasion patterns are, however, still unclear. Here we investigated first, if the spread of non-native bark and ambrosia beetles is a gradual or a discontinuous process; second, which are the main correlates of their community structure; third, whether those correlates correspond to those of native species. We used data on species distribution of non-native and native scolytines in the continental 48 USA states. These data were analyzed through a beta-diversity index, partitioned into species richness differences and species replacement, using Mantel correlograms and non-metric multidimensional scaling (NMDS) ordination for identifying spatial patterns, and regression on distance matrices to test the association of climate (temperature, rainfall), forest (cover area, composition), geographical (distance), and human-related (import) variables with ß-diversity components. For both non-native bark and ambrosia beetles, ß-diversity was mainly composed of species richness difference than species replacement. For non-native bark beetles, a discontinuous invasion process composed of long distance jumps or multiple introduction events was apparent. Species richness differences were primarily correlated with differences in import values while temperature was the main correlate of species replacement. For non-native ambrosia beetles, a more continuous invasion process was apparent, with the pool of non-native species arriving in the coastal areas that tended to be filtered as they spread to interior portions of the continental USA. Species richness differences were mainly correlated with differences in rainfall among states, while rainfall and temperature were the main correlates of species replacement. Our study suggests that the different ecology of bark and ambrosia beetles influences their invasion process in new environments. The lower dependency that bark beetles have on climate allowed them to potentially colonize more areas within the USA, while non-native ambrosia beetles, being dependent on rainfall, are typically filtered by the environment.

New Fungus-Insect Symbiosis: Culturing, Molecular, and Histological Methods Determine Saprophytic Polyporales Mutualists of Ambrosiodmus Ambrosia Beetles.

Ambrosia symbiosis is an obligate, farming-like mutualism between wood-boring beetles and fungi. It evolved at least 11 times and includes many notorious invasive pests. All ambrosia beetles studied to date cultivate ascomycotan fungi: early colonizers of recently killed trees with poor wood digestion. Beetles in the widespread genus Ambrosiodmus, however, colonize decayed wood. We characterized the mycosymbionts of three Ambrosiodmus species using quantitative culturing, high-throughput metabarcoding, and histology. We determined the fungi to be within the Polyporales, closely related to Flavodon flavus. Culture-independent sequencing of Ambrosiodmus minor mycangia revealed a single operational taxonomic unit identical to the sequences from the cultured Flavodon. Histological sectioning confirmed that Ambrosiodmus possessed preoral mycangia containing dimitic hyphae similar to cultured F. cf. flavus. The Ambrosiodmus-Flavodon symbiosis is unique in several aspects: it is the first reported association between an ambrosia beetle and a basidiomycotan fungus; the mycosymbiont grows as hyphae in the mycangia, not as budding pseudo-mycelium; and the mycosymbiont is a white-rot saprophyte rather than an early colonizer: a previously undocumented wood borer niche. Few fungi are capable of turning rotten wood into complete animal nutrition. Several thousand beetle-fungus symbioses remain unstudied and promise unknown and unexpected mycological diversity and enzymatic innovations.

Ethanol and (-)-alpha-Pinene: attractant kairomones for bark and ambrosia beetles in the southeastern US.

In 2002-2004, we examined the flight responses of 49 species of native and exotic bark and ambrosia beetles (Coleoptera: Scolytidae and Platypodidae) to traps baited with ethanol and/or (-)-alpha-pinene in the southeastern US. Eight field trials were conducted in mature pine stands in Alabama, Florida, Georgia, North Carolina, and South Carolina. Funnel traps baited with ethanol lures (release rate, about 0.6 g/day at 25-28 degrees C) were attractive to ten species of ambrosia beetles (Ambrosiodmus tachygraphus, Anisandrus sayi, Dryoxylon onoharaensum, Monarthrum mali, Xyleborinus saxesenii, Xyleborus affinis, Xyleborus ferrugineus, Xylosandrus compactus, Xylosandrus crassiusculus, and Xylosandrus germanus) and two species of bark beetles (Cryptocarenus heveae and Hypothenemus sp.). Traps baited with (-)-alpha-pinene lures (release rate, 2-6 g/day at 25-28 degrees C) were attractive to five bark beetle species (Dendroctonus terebrans, Hylastes porculus, Hylastes salebrosus, Hylastes tenuis, and Ips grandicollis) and one platypodid ambrosia beetle species (Myoplatypus flavicornis). Ethanol enhanced responses of some species (Xyleborus pubescens, H. porculus, H. salebrosus, H. tenuis, and Pityophthorus cariniceps) to traps baited with (-)-alpha-pinene in some locations. (-)-alpha-Pinene interrupted the response of some ambrosia beetle species to traps baited with ethanol, but only the response of D. onoharaensum was interrupted consistently at most locations. Of 23 species of ambrosia beetles captured in our field trials, nine were exotic and accounted for 70-97% of total catches of ambrosia beetles. Our results provide support for the continued use of separate traps baited with ethanol alone and ethanol with (-)-alpha-pinene to detect and monitor common bark and ambrosia beetles from the southeastern region of the US.

Biology and host associations of redbay ambrosia beetle (Coleoptera: Curculionidae: Scolytinae), exotic vector of laurel wilt killing redbay trees in the southeastern United States.

The redbay ambrosia beetle, Xyleborus glabratus Eichhoff (Coleoptera: Curculionidae: Scolytinae), and its fungal symbiont, Raffaelea sp., are new introductions to the southeastern United States responsible for the wilt of mature redbay, Persea borbonia (L.) Spreng., trees. In 2006 and 2007, we investigated the seasonal flight activity of X. glabratus, its host associations, and population levels at eight locations in South Carolina and Georgia where infestations ranged from very recent to at least several years old. Adults were active throughout the year with peak activity in early September. Brood development seems to take 50-60 d. Wood infested with beetles and infected with the Raffaelea sp. was similar in attraction to uninfested redbay wood, whereas both were more attractive than a nonhost species. Sassafras, Sassafras albidium (Nutt.) Nees, another species of Lauraceae, was not attractive to X. glabratus and very few beetle entrance holes were found in sassafras wood compared with redbay. Conversely, avocado, Persea americana Mill., was as attractive to X. glabratus as swampbay, P. palustris (Raf.) Sarg., and both were more attractive than the nonhost red maple, Acer rubrum L. However, avocado had relatively few entrance holes in the wood. In 2007, we compared X. glabratus populations in areas where all mature redbay have died to areas where infestations were very active and more recent. Trap catches of X. glabratus and numbers of entrance holes in trap bolts of redbay were correlated with the number of dead trees with leaves attached. Older infestations where mature host trees had been eliminated by the wilt had low numbers of beetles resulting in trap catches ranging from 0.04 to 0.12 beetles per trap per d compared with 4-7 beetles per trap per d in areas with numerous recently dead trees. Our results indicate beetle populations drop dramatically after suitable host material is gone and provide hope that management strategies can be developed to restore redbay trees. The lack of attraction of X. glabratus to sassafras suggests that spread of X. glabratus may slow once it is outside the range of redbay.