Spatio-temporal analysis of Xyleborus glabratus (Coleoptera: Curculionidae [corrected] Scolytinae) invasion in eastern U.S. forests

The non-native redbay ambrosia beetle, Xyleborus glabratus Eichhoff (Coleoptera: Curculionidae: Scolytinae), has recently emerged as a significant pest of southeastern U.S. coastal forests. Specifically, a fungal symbiont (Raffaelea sp.) of X. glabratus has caused mortality of redbay (Persea borbonia) and sassafras (Sassafras albidum) trees in the region; several other Lauraceae species also seem susceptible. Although the range of X. glabratus continues to expand rapidly, little is known about the species' biology and behavior. In turn, there has been no broad-scale assessment of the threat it poses to eastern U.S. forests. To provide a basic information framework, we performed analyses exploiting relevant spatio-temporal data available for X. glabratus. First, we mapped the densities of redbay and sassafras from forest inventory data. Second, we used climate matching to delineate potential geographic limits for X. glabratus. Third, we used county infestation data to estimate the rate of spread and modeled spread through time, incorporating host density as a weighting factor. Our results suggest that (1) key areas with high concentrations of redbay have yet to be invaded, but some are immediately threatened; (2) climatic conditions may serve to constrain X. glabratus to the southeastern U.S. coastal region; and (3) if unchecked, X. glabratus may spread throughout the range of redbay in <40 yr. Disruption of anthropogenic, long-distance dispersal could reduce the likelihood of this outcome.     

Temporal analysis of sesquiterpene emissions from manuka and phoebe oil lures and efficacy for attraction of Xyleborus glabratus (Coleoptera: Curculionidae: Scolytinae)

Redbay ambrosia beetle, Xyleborus glabratus Eichhoff, is an exotic wood-borer that vectors the fungal agent (Raffaelea lauricola) responsible for laurel wilt. Laurel wilt has had severe impact on forest ecosystems in the southeastern United States, killing a large proportion of native Persea trees, particularly redbay (P. borbonia) and swampbay (P. palustris), and currently poses an economic threat to avocado (P. americana) in Florida. To control the spread of this lethal disease, effective attractants are needed for early detection of the vector. Two 12-wk field tests were conducted in Florida to evaluate efficacy and longevity of manuka and phoebe oil lures, and to relate captures of X. glabratus to release rates of putative sesquiterpene attractants. Two trap types were also evaluated, Lindgren funnel traps and sticky panel traps. To document lure emissions over time, a separate set of lures was aged outdoors for 12 wk and sampled periodically to quantify volatile sesquiterpenes using super-Q adsorbant and gas chromatography-mass spectroscopy analysis. Phoebe lures captured significantly more X. glabratus than manuka lures, and sticky traps captured more beetles than funnel traps. Phoebe lures captured X. glabratus for 10-12 wk, but field life of manuka lures was 2-3 wk. Emissions of alpha-copaene, alpha-humulene, and cadinene were consistently higher from phoebe lures, particularly during the 2-3 wk window when manuka lures lost efficacy, suggesting that these sesquiterpenes are primary kairomones used by host-seeking females. Results indicate that the current monitoring system is suboptimal for early detection of X. glabratus because of rapid depletion of sesquiterpenes from manuka lures.     

Taxonomic changes in palaeotropical Xyleborini (Coleoptera, Curculionidae, Scolytinae)

Following the recent reclassification of the Palaeotropic xyleborine genera (Hulcr and Cognato in press), additional species are transferred to correct genera or synonymized based on analysis of their morphological characters. The following species are given new combinations: Debus amphicranoides (Hagedorn), comb. n., Debus birmanus (Eggers, 1930), comb. n., Debus dolosus (Blandford, 1896), comb. n., Debus eximius (Schedl, 1970), comb. n., Debus interponens (Schedl, 1954), comb. n., Debus robustipennis (Schedl, 1954), comb. n., Debus spinatus (Eggers, 1923), comb. n., Microperus alpha (Beeson, 1929), comb. n., Microperus corporaali (Eggers), comb. n., Microperus eucalyptica (Schedl, 1938), comb. n., Microperus nugax (Schedl, 1939), comb. n., Pseudowebbia percorthylus (Schedl, 1935), comb. n., Truncaudum circumcinctus (Schedl, 1941), comb. n.THE FOLLOWING SPECIES ARE SYNONYMIZED: Arixyleborus hirtipennis (Eggers), syn. n., with Arixyleborus puberulus (Blandford); Coptoborus palmeri (Hopkins), syn. n., with Debus emarginatus (Eichhoff); Coptoborus terminaliae (Hopkins), syn. n., with Debus emarginatus (Eichhoff); Cyclorhipidion polyodon (Eggers), syn. n., with Truncaudum agnatum (Eggers); Euwallacea artelaevis (Schedl), syn. n., with Planiculus bicolor (Blandford); Xyleborinus perminutissimus (Schedl), syn. n., with Xyleborinus perpusillus (Eggers); Xyleborus exesus Blandford, syn. n., with Debus emarginatus (Eichhoff); Xyleborus fulvulus (Schedl), syn. n., with Microperus corporaali (Eggers); Xyleborus marginicollis (Schedl), syn. n., with Diuncus justus (Schedl); Xyleborus shoreae Stebbing, syn. n., with Debus fallax (Eichhoff).THE FOLLOWING SPECIES ARE GIVEN NEW STATUS: Streptocranus superbus (Schedl, 1951), restored name; Webbia divisus Browne, 1972, restored name; Webbia penicillatus (Hagedorn, 1910), restored name. Genus Taphrodasus Wood (1980) is declared not valid.     

Bark and Ambrosia Beetles (Coleoptera,Curculionidae: Scolytinae) of Northwest Russia: History of the Study,Composition and Genesis of the Fauna

Entomological Review - An essay of the fauna of bark beetles (Coleoptera: Curculionidae: Scolytinae) in five federal subjects of Northwest Russia is presented. The species composition of scolytids...     

Small Spruce Bark Beetle Ips amitinus (Eichhoff, 1872) (Coleoptera,Curculionidae: Scolytinae): a New Alien Species in West Siberia

The small spruce bark beetle Ips amitinus is a widespread species in many European countries that has been actively spreading into Northern Europe in the recent decades. In Russia, I. amitinus is present in the western, northwestern, and northern regions of the European part, with a tendency for range expansion. The species was first recognized in West Siberia in 2019 by characteristic morphological features and molecular genetic analysis. This bark beetle is abundant on Pinus sibirica in Siberian pine forests located near settlements within Tomsk and Kemerovo provinces, and is also sporadically found on the Siberian spruce Picea obovata. It colonizes the upper trunk and branches of standing and windfall trees. In the outbreak foci this bark beetle causes catastrophic drying of Siberian pines, starting from the crown top. This pattern of tree drying was noted for the first time near settlements in Yashkinsky District of Kemerovo Province in 2014, and now outbreak foci of I. amitinus exist in all the Siberian pine forests in this district. The population growth of I. amitinus was probably facilitated by dry and hot summer weather in the southeast of West Siberia during the last decade, in 2011 and 2012, and also by heavy winter snowfalls leaving numerous snapped tree branches which are easily colonized by the pest. In Tomsk Province, the most active outbreak focus of I. amitinus appeared in 2018 in the Siberian pine forest near Luchanovo and Ipatovo, following an outbreak of the Siberian moth Dendrolimus sibiricus. The invasion of I. amitinus in Siberia may increase the degradation rates not only of the gene-reserve Siberian pine forests but also of other dark coniferous stands.

     

Alimentary canal and reproductive tract of Hypothenemus hampei (Ferrari) (Coleoptera: Curculionidae, Scolytinae)

The alimentary canal and the reproductive tract of males and females of Hypothenemus hampei Ferrari are described. The alimentary canal of H. hampei showed the crop with several spine-like structures and the midgut with few gastric caeca. We evidenced for the first time that adult females need to feed on coffee in order to produce viable eggs before and during oviposition period. The presence of air bubbles inside the anterior midgut may be due to starvation rather than the response of the environment during flying as previously reported. Two ovaries and the same structures and arrangements common to individuals of the Curculionidae beetles composed the female reproductive system. The male reproductive tract showed a significant difference with respect to other Curculionidae and even other Scolytinae species as it showed a sclerotized aedeagus with several pore-like structures in the terminal portion where sperm is released during mating.     

The Polyphagous Bark Beetle Hypothenemus eruditus (Westwood, 1836) (Coleoptera,Curculionidae, Scolytinae): the First Record on Buxus Trees in Russia

Entomological Review - The formation of Hypothenemus eruditus foci is described on a tree of Buxus sempervirens ssp. colchica which died as the result of damage caused by larvae of Neoglyphodes...     

First Record of the Four-Eyed Fir Bark Beetle Polygraphus proximus Blandford, 1894 (Coleoptera,Curculionidae: Scolytinae) from Irkutsk Province,Russia

Entomological Review - Polygraphus proximus Blandford, 1894 is recorded for the first time for Irkutsk Province of Russia. The species was collected in the vicinity of Utulik Village in Slyudyansky...     

Five new species of Triotemnus (Coleoptera, Curculionidae, Scolytinae) from Morocco and Yemen

Fivenew species of the genus Triotemnus from Morocco and Yemen are described. Triotemnus is a new genus of Scolytinae for the Yemen region. External morphology of the new species and all morphologically related species of the genus were studied. While the new species from Morocco are morphologically similar to the known species from the corresponding region, all three newly described species from Yemen, mainly two of them living in Socotra, are morphologically very different from all other known species of the genus. Geographical distribution and the probability of endemicity are discussed.     

Phylogeography of the bark beetle Dendroctonus mexicanus Hopkins (Coleoptera: Curculionidae: Scolytinae)

Dendroctonus mexicanus is polyphagous within the Pinus genus and has a wide geographical distribution in Mexico and Guatemala. We examined the pattern of genetic variation across the range of this species to explore its demographic history and its phylogeographic pattern. Analysis of the mtDNA sequences of 173 individuals from 25 Mexican populations allowed to us identify 53 geographically structured haplotypes. High haplotype and low nucleotide diversities and Tajima’s D indicate that D. mexicanus experienced rapid population expansion during its dispersal across mountain systems within its current range. The nested clade phylogeographic analysis indicates that the phylogeographic pattern of D. mexicanus is explained by continuous dispersion among lineages from the Sierra Madre Occidental, the Sierra Madre Oriental and the Trans-Mexican Volcanic Belt. However, we also observed isolation events among haplotypes from the Cofre de Perote/Trans-Mexican Volcanic Belt/Sierra Madre Oriental and the Trans-Mexican Volcanic Belt/Sierra Madre del Sur, which is consistent with the present conformation of mountain systems in Mexico and the emergence of geographical barriers during the Pleistocene.     

Mating Behavior of the Coffee Berry Borer, Hypothenemus hampei (Ferrari) (Coleoptera: Curculionidae: Scolytinae)

The mating behavior of the coffee berry borer, Hypothenemus hampei (Ferrari), was observed under laboratory conditions. Pairs of adult virgin male and female beetles were placed in the wells of a polystyrene microtiter plate, one pair per well. The mating activity of each pair was recorded for 24?h. The mating behavior of the coffee berry borer was similar to that of other Scolytinae and was clearly divided into precopulatory, copulatory and postcopulatory phases. The beetles started to mate within a few hours of emergence. Repeated mating occurred during the 24-hour period and increased in frequency with age. However, we cannot address multiple matings in H. hampei, since we did not simulate the female-biased sex ratio of this species and the experimental design did not allow females to avoid additional mating attempts by males. In addition, further studies are necessary that focus on the effectiveness of sperm transmission and direct and indirect effects of multiple matings on the H. hampei females and their offspring.     

A New Genus and Species of Scolytinae (Coleoptera: Curculionidae) from the Rovno Amber

A new genus, Klesovia gen. nov., with the species K. pubescens sp. nov. (Coleoptera: Curculionidae) are described from the Rovno amber. The new genus is closely related to Xylechinus (tribe Hylurgini) and distinguished by the wide flattened ovoid antennal club with rounded apex and the metepisternum covered with long thin sparse setae; it differs from other genera of Hylurgini in having a five-segmented funicle, four-segmented club, and large distance between the procoxa.     

Polyphyly of Xylosandrus Reitter inferred from nuclear and mitochondrial genes (Coleoptera: Curculionidae: Scolytinae)

The Xyleborina ambrosia beetle genus Xylosandrus contains 54 species, several of which are of economic importance. The monophyly of the genus was tested using a data set comprised of multiple gene loci: 28S rDNA; the mitochondrial gene cytochrome oxidase I (COI); and the nuclear genes arginine kinase (ArgK), rudimentary (CAD), and Elongation Factor 1α (EF-1α). The nuclear protein-coding genes CAD and ArgK were used for the first time in phylogenetics of Scolytinae. Analyses were performed using Parsimony and Bayesian optimality criteria. Our analyses included 43 specimens representing 15 Xylosandrus species and 20 species from Amasa, Anisandrus, Cnestus, Euwallacea and Xyleborus, and two species from the outgroup genus Coccotrypes. All analyses recovered a polyphyletic Xylosandrus. Several species of Xylosandrus were consistently placed in clades with the genera Anisandrus and Cnestus with high support values (100% bootstrap support). Among these, was the economically important invasive species X. mutilatus, which was consistently recovered as part of the “Cnestus” clade. In our analyses, both CAD and ArgK demonstrated phylogenetic utility across varying nodal depths. Despite the selection of genes with signals at complementary phylogenetic depths, the data set used herein did not resolve the phylogeny of Xylosandrus and related genera. Since the taxon sample available for molecular work represents only a fraction of Xylosandrus species, a complete revision that combines molecular and morphological data in a total evidence approach is recommended for the genus.     

New ichnotaxon Megascolytinus zherikhini (Coleoptera: Curculionidae: Scolytinae) from upper cretaceous deposits of mongolia

Gallery systems of Scolytinae on a petrified Araucaria trunk fragment from the Late Cretaceous of Gobi, Mongolia, similar to those of extant members of the supertribe Hylesininae, but exceptionally large (estimated body length of the beetle about 20 mm), are described as a new ichnotaxon, Megascolytinus zheriknini.     

Lack of genetic differentiation in aggressive and secondary bark beetles (Coleoptera: Curculionidae, Scolytinae) from Arizona

Bark beetles (Coleoptera: Curculionidae, Scolytinae) play an important role as disturbance agents in ponderosa pine (Pinus ponderosa Douglas ex Lawson) forests of Arizona. However, from 2001 to 2003, elevated bark beetle activity caused unprecedented levels of ponderosa pine mortality. A better understanding of the population structure of these species will facilitate analysis of their dispersal patterns and improve management strategies. Here, we use fluorescently labeled amplified fragment length polymorphism (fAFLP) analysis to resolve genetic variation among and within sampling locations in northcentral Arizona of Ips pini (Say), Dendroctonus brevicomis LeConte, and D. frontalis Zimmermann. We generated genetic fingerprints for >500 beetle specimens and analyzed genetic diversity. For all species, gene flow estimates among sampling locations were high, and significant population subdivision was not discernible across a large portion of ponderosa pine forests in Arizona. However, a weak relationship was detected with I. pini population structure and elevation. Because of the lack of genetic differentiation detected throughout the large study area, our findings suggest these insects are capable of long distance dispersal and exhibit a high degree of gene flow across a broad region. We conclude that our results are consistent with strong dispersal patterns and large population sizes of all three species.     

Brief Review of the Associations of Xylobiont Nematodes with Bark Beetles (Coleoptera,Curculionidae: Scolytinae)

Nematode pathogens cause wilt diseases in conifers and deciduous trees. The longhorn beetles (Coleoptera: Cerambycidae) and bark beetles (Coleoptera: Curculionidae: Scolytinae) act as nematode vectors spreading the invasive juvenile stages during their maturation feeding or during oviposition on the plant hosts. There are numerous reviews of nematodes associated with bark beetles on conifers, while little attention has been paid to the nematodes of deciduous trees. The development of Dutch elm disease and ash dieback is mainly caused by fungal pathogens transmitted by bark beetles; the latter act as vectors of not only fungi but also nematodes enclosed in nematangia under their elytra, and also in the tracheae and Malpighian canals. Apart from phytopathogenic nematodes, bark beetles transmit mycophagous and bacterivorous nematodes and own parasites of bark beetles. The ecological groups of nematodes associated with Scolytinae are reviewed; the known records of associations of nematodes with bark beetles are listed for coniferous host plants of Russia and neighboring countries; the world-wide list of these associations for deciduous plant hosts is given.

     

Phylogeography of spruce beetles (Dendroctonus rufipennis Kirby) (Curculionidae: Scolytinae) in North America

Tree-feeding insects that are widespread in north temperate regions are excellent models for studying how past glaciations have impacted differentiation and speciation. We used mitochondrial DNA (mtDNA) sequences and allele frequencies at nine microsatellite loci to examine genetic population structure across the current range of the spruce beetle (Dendroctonus rufipennis), an economically important insect in North America. Two major haplotype groups occur across northern North America, from Newfoundland to Alaska, on white spruce (Picea glauca), and a third distinctive haplotype group occurs throughout the Rocky Mountains on Engelmann spruce (Picea engelmannii). The two mtDNA lineages found in northern populations are 3-4% divergent from each other and from the lineages found in the Rocky Mountains. Analyses of microsatellite data also suggest the existence of major population groupings associated with different geographical regions. In the Pacific Northwest, concordant contact zones for genetically distinct populations of spruce beetles and their principal hosts appear to reflect recent secondary contact. Although we could detect no evidence of historical mtDNA gene flow between allopatric population groups, patterns of variation in the Pacific Northwest suggest recent hybridization and introgression. Together with the pollen record for spruce, they also suggest that beetles have spread from at least three glacial refugia. A minimum estimate of divergence time between the Rocky Mountain and northern populations was 1.7 Myr (million years), presumably reflecting the combined effects of isolation during multiple glacial cycles.     

Optimizing ethanol-baited traps for monitoring damaging ambrosia beetles (Coleoptera: Curculionidae, Scolytinae) in ornamental nurseries

The exotic ambrosia beetles Xylosandrus crassiusculus (Motschulsky) and Xylosandrus germanus (Blandford) (Coleoptera: Curculionidae: Scolytinae) are serious pests in ornamental tree nurseries. To optimize bottle-traps as a monitoring system for X. crassiusculus and X. germanus in nurseries, we tested whether increasing the rate of commercial ethanol lures improved captures or early detection of these species. Experiments were conducted in Ohio (2008 and 2009) and Virginia (2008), two states that have experienced significant damage from X. crassiusculus, X. germanus, or both. There were four treatments: no-lure (unbaited control), 1-ethanol lure, 2-ethanol lures and 1 + 1-ethanol lures (one lure in the trap and one suspended 0.5 m above the trap). Captures of X. crassiusculus and X. germanus were higher in all ethanol treatments than unbaited controls, and were generally higher in treatments with two lures versus one. There was no difference in beetle captures between the 2-lure and 1 + 1-lure treatments. First detection of X. crassiusculus and X. germanus occurred more consistently in the treatments with two lures than one lure. Xyleborinus saxesenii (Ratzeburg), Anisandrus sayi Hopkins, Hypothenemus dissimilis Zimmermann, and Hypothenemus eruditus Westwood were also more attracted to traps baited with ethanol than unbaited controls. X. saxesenii was captured in higher numbers in the treatments with two lures than one in Virginia but not in Ohio. There was no difference in captures of the other species among ethanol treatments. The current research shows that ethanol release rates influence sensitivity of traps for detecting emergence of overwintered ambrosia beetles.