Presence and Diversity of <Emphasis Type="Italic">Streptomyces</Emphasis> in <Emphasis Type="Italic">Dendroctonus</Emphasis> and Sympatric Bark Beetle Galleries Across North America

Recent studies have revealed several examples of intimate associations between insects and Actinobacteria, including the Southern Pine Beetle Dendroctonus frontalis and the Spruce Beetle Dendroctonus rufipennis. Here, we surveyed Streptomyces Actinobacteria co-occurring with 10 species of Dendroctonus bark beetles across the United States, using both phylogenetic and community ecology approaches. From these 10 species, and 19 other scolytine beetles that occur in the same trees, we obtained 154 Streptomyces-like isolates and generated 16S sequences from 134 of those. Confirmed 16S sequences of Streptomyces were binned into 36 distinct strains using a threshold of 0.2% sequence divergence. The 16S rDNA phylogeny of all isolates does not correlate with the distribution of strains among beetle species, localities, or parts of the beetles or their galleries. However, we identified three Streptomyces strains occurring repeatedly on Dendroctonus beetles and in their galleries. Identity of these isolates was corroborated using a house-keeping gene sequence (efTu). These strains are not confined to a certain species of beetle, locality, or part of the beetle or their galleries. However, their role as residents in the woodboring insect niche is supported by the repeated association of their 16S and efTu from across the continent, and also having been reported in studies of other subcortical insects.     

Ophiostomatoid fungi isolated from Japanese red pine and their relationships with bark beetles

We isolated ophiostomatoid fungi from bark beetles infesting Pinus densiflora and their galleries at 24 sites in Japan. Twenty-one ophiostomatoid fungi, including species of Ophiostoma, Grosmannia, Ceratocystiopsis, Leptographium, and Pesotum, were identified. Among these, 11 species were either newly recorded in Japan or were previously undescribed species. Some of these fungal species were isolated from several bark beetles, but other species were isolated from only a particular beetle species. Thus, it is suggested that some ophiostomatoid fungi have specific relationships with particular beetle species. In addition, fungus-beetle biplots from redundancy analysis (RDA) summarizing the effects of beetle ecological characteristics suggested that the association patterns between bark beetles and the associated fungi seemed to be related to the niches occupied by the beetles.     

Colonization of disturbed Scots pine trees by bark‐ and wood‐boring beetles

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<Emphasis Type="Italic">Geosmithia</Emphasis> Fungi are Highly Diverse and Consistent Bark Beetle Associates: Evidence from their Community Structure in Temperate Europe

Geosmithia spp. (Ascomycota: Hypocreales) are little-studied, dry-spored fungi that occur in galleries built by many phloeophagous bark beetles. This study mapped the distribution and environmental preferences of Geosmithia species occurring in galleries of temperate European bark beetles. One hundred seven host tree samples of 16 tree species infested with 23 subcortical insect species were collected from across Europe during the years 1997–2005. Over 600 Geosmithia isolates from the beetles were sorted into 17 operational taxonomic units (OTUs) based on their phenotype similarity and phylogeny of internal transcribed spacer (ITS) region of rDNA (ITS1-5.8S-ITS2). The OTUs represent six known species and eight undescribed taxa. Ninety-two samples infested with subcortical insects were characterized by the presence/absence of OTUs and the similarity among the samples was evaluated. Geographically distant populations of the same beetle species host relatively uniform Geosmithia communities across large geographic areas (ranging from southern Bulgaria to the Czech Republic). This suggests effective dispersal of Geosmithia spp. by bark beetles. Clustering of similar samples in ordination analysis is correlated predominantly with the isolation source (bark beetles and their respective feeding plant), but not with their geographical origin. The composition of the Geosmithia OTU community of each bark beetle species depends on the degree of isolation of the species’ niches. Thus, Geosmithia communities associated with regularly co-occurring bark beetle species are highly similar. The similarity decreases with decreasing frequency of beetle species’ co-occurrence, a pattern resembling that of entomochoric ophiostomatoid fungi. These findings suggest that: 1) communities of Geosmithia spp. are vector-specific; 2) at least in some cases, the association between Geosmithia OTUs and bark beetles may have been very stable and symbioses are likely to be a fundamental factor in the speciation of Geosmithia fungi; and 3) that even nonsticky spores of Geosmithia are suitable for maintaining an insect–fungus association, contrary to previous hypotheses. An erratum to this article can be found at     

Metabarcoding of fungal communities associated with bark beetles

Many species of fungi are closely allied with bark beetles, including many tree pathogens, but their species richness and patterns of distribution remain largely unknown. We established a protocol for metabarcoding of fungal communities directly from total genomic DNA extracted from individual beetles, showing that the ITS3/4 primer pair selectively amplifies the fungal ITS. Using three specimens of bark beetle from different species, we assess the fungal diversity associated with these specimens and the repeatability of these estimates in PCRs conducted with different primer tags. The combined replicates produced 727 fungal Operational Taxonomic Units (OTUs) for the specimen of Hylastes ater, 435 OTUs for Tomicus piniperda, and 294 OTUs for Trypodendron lineatum, while individual PCR reactions produced on average only 229, 54, and 31 OTUs for the three specimens, respectively. Yet, communities from PCR replicates were very similar in pairwise comparisons, in particular when considering species abundance, but differed greatly among the three beetle specimens. Different primer tags or the inclusion of amplicons in separate libraries did not impact the species composition. The ITS2 sequences were identified with the Lowest Common Ancestor approach and correspond to diverse lineages of fungi, including Ophiostomaceae and Leotiomycetes widely found to be tree pathogens. We conclude that Illumina MiSeq metabarcoding reliably captures fungal diversity associated with bark beetles, although numerous PCR replicates are recommended for an exhaustive sample. Direct PCR from beetle DNA extractions provides a rapid method for future surveys of fungal species diversity and their associations with bark beetles and environmental variables.     

Biosystematics of the straw itch mite with special reference to nomenclature and dermatology

• 1 Cross-mating of various populations of Pyemotes obtained from laboratory insect cultures demonstrated that the straw itch mite is a single, widely dispersed species, different from P.ventricosus, which was probably associated with the furniture beetle.
• 2 P.ventricosus should not be the approved scientific name for the straw itch mite. The correct name for the straw itch mite is Pyemotes tritici.
• 3 Various Pyemotes species differ in preferred habitats, hosts, and toxicity to man. Whereas P.tritici commonly attacks grain insects and is highly toxic to man, other species attack wood boring insects and may cause little or no irritation. P.scolyti and P.parviscolyti, which attack bark beetles, cause no observable symptoms.
• 4 It is suggested that before Pyemotes are used as biological control agents against bark beetles, it is important to identify the species correctly.

     

Ophiostoma arduennense sp. nov. (Ophiostomatales, Ascomycota) from Fagus sylvatica in southern Belgium

Ophiostoma arduennense sp. nov. is described from several cultures isolated from Fagus sylvatica in southern Belgium. The species is mainly characterized by globose perithecia with small button-like bases ornamented with brown hyphal hairs of variable length and, long cylindrical necks ending in ostiolate hyphae. It is homothallic with small reniform ascospores and no apparent anamorph. It is closely associated with the ambrosia beetles Xyloterus domesticus and X. signatus. Its phylogenetic relationships within Ophiostoma are discussed and the species is compared with other Ophiostoma species associated with European beech or other broad-leaved trees in Europe, especially species of the O. quercus–O. piceae complex. The species is responsible for a dark brown staining of the sapwood and its role in the decline of beech in Southern Belgium is also discussed.     

Synchronous rise and fall of bark beetle and parasitoid populations in windthrow areas

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A major symbiont shift supports a major niche shift in a clade of tree-killing bark beetles

1. One small clade of bark beetles, out of thousands of species worldwide, has shifted from using phloem to using a combination of phloem and outer bark, or to completely using outer bark. 2. The shift to outer bark has been accompanied by a shift to non-typical bark beetle mutualist fungi in Entomocorticium (Basidiomycota) and Ceratocystiopsis (Ascomycota). 3. This study compared the growth and metabolic capabilities of fungi associated with a nearly phloem-independent species, Dendroctonus brevicomis, with those of mutualist fungi of Dendroctonus ponderosae, a completely phloem-colonising beetle in a sister clade associated with more typical Ascomycota in Grosmannia and Ophiostoma. 4. Only the basidiomycete associated with D. brevicomis could degrade cellulose and lignin, whereas both the ascomycete and basidiomycete could grow in outer bark. Ascomycetes associated with D. ponderosae could not degrade cellulose or lignin or grow in outer bark. 5. Beetles and fungi in these mutualisms may best be considered as co-niche constructors. For niche construction, one organism must modify a resource in a way that enhances its fitness while also influencing fitness of other organisms using the resource. Here, the beetles kill the tree, transport the fungi into the tree, and modify the woody substrate for use by the fungi. They have also evolved mycangia to ensure vertical dissemination of the fungi. In turn, the fungi modify tree tissues and provision nutrients to the host and have evolved traits that support their acquisition and transport by the beetle host in mycangia.     

The first fossil record of mites of the family Pyemotidae (Acari: Heterostigmata), with description of a new species of the genus <Emphasis Type="Italic">Pyemotes</Emphasis> from Rovno amber

The first fossil mite of the family Pyemotidae (Acari: Heterostigmata) is recorded. Pyemotes primus sp. nov. is described from the Late Eocene of the Rovno amber from a syninclusion with the bark beetles Taphramites rovnoensis Petrov et Perkovsky, 2008. In the presence of well developed longitudinal striation on hysterosomal tergites, the new species is similar to species from scolyti group, which is characterized by parasitoidism on immature stages of bark beetles and phoresy on adult stages. P. primus differs from the closely related species P. dryas (Vitzthum, 1923), P. parviscolyti Cross et Moser, 1971, P. johnmoseri Khaustov, 2004, and P. mandelshtami Khaustov, 1998 in the subequal setae h ₁ and h ₂.     

Short‐term effect of selection cutting in boreal balsam fir forest on cerambycid and scolytid beetles

The ecology and management of boreal forest ecosystems are drawing greater attention worldwide as their importance is being increasingly recognized for carbon sequestration or for harbouring the world's largest remaining intact forests. Selection cuts have been introduced as a more socially acceptable silvicultural method to improve the maintenance of habitat structure and functions as they mimic aspects of boreal forest succession dynamics. Many studies have shown that selection cutting helps maintaining arthropod communities in mature forests, but few have examined the increased risks of damage by bark‐ and wood‐boring insects in boreal forests of eastern North America. We used multidirectional flight‐interception traps to quantify the response of these beetles to 25 and 40% selection cutting in a balsam fir–white birch forest of Québec, Canada. The abundance and species number of both cerambycid and scolytid beetles were 5–6 times larger in selectively cut stands than in controls the year following treatments. Analyses revealed that bark‐ and wood‐boring beetles’ response was mostly associated with increased canopy openness in selectively cut stands (and sun‐exposed locations within them) and residual tree injuries caused by harvesting operations. These conditions attracted beetles such as Trypodendron lineatum (Scolytinae) and Rhagium inquisitor (Cerambycidae), two species known for their ability to attack weakened, dying and dead hosts. Most species were more abundant in selection cuts, except for Evodinus m. monticola (Cerambycidae) whose abundance was strongly reduced after treatment. Some beetles can have detrimental effects on residual trees and thus could reduce timber value, but most species found in treated stands do not represent a high risk for healthy trees. Thus, selection cuts do not seem favourable to the establishment of tree‐killing beetles. However, as they were found more active/abundant after selection cutting, it would be wise to further study their population dynamics over mid‐ and long‐term periods, along with the ecological and economic implications associated with this silvicultural treatment.     

Nematodes associated with Ips cembrae (Coleoptera: Curculionidae): comparison of generations,sexes and sampling methods

A study of nematodes associated with the large larch bark beetle Ips cembrae (Heer 1836) was carried out at three locations in the Czech Republic. The proportion of beetles infested by endoparasitic nematodes (representatives of genera Contortylenchus, Parasitylenchus, Cryptaphelenchus and Parasitorhabditis) ranged from 29.9 to 50.9%. Significant differences were determined in nematode infestation levels among locations, generations and sampling methods. No differences were found in infestation rates between males and females. The percentage of bark beetles with phoretic nematodes ranged from 18 to 42.9%. Phoretic nematodes directly found under elytra, on wings and between body segments of the bark beetles belong to the genus Micoletzkya. However, adults and juveniles of other two phoretic species Laimaphelenchus penardi and Bursaphelenchus sp. were found in the gallery frass of I. cembrae. Infestation by phoretic nematodes positively correlated with the presence of mites under elytra.     

Ophiostomatoid fungi including two new fungal species associated with pine root-feeding beetles in northern Spain

Many bark beetles live in a symbiosis with ophiostomatoid fungi but very little is known regarding these fungi in Spain. In this study, we considered the fungi associated with nine bark beetle species and one weevil infesting two native tree species (Pinus sylvestris and Pinus nigra) and one non-native (Pinus radiata) in Cantabria (Northern Spain). This included examination of 239 bark beetles or their galleries. Isolations yielded a total of 110 cultures that included 11 fungal species (five species of Leptographium sensu lato including Leptographium absconditum sp. nov., five species of Ophiostoma sensu lato including Ophiostoma cantabriense sp. nov, and one species of Graphilbum). The most commonly encountered fungal associates of the bark beetles were Grosmannia olivacea, Leptographium procerum, and Ophiostoma canum. The aggressiveness of the collected fungal species was evaluated using inoculations on two-year-old P. radiata seedlings. Leptographium wingfieldii, Leptographium guttulatum, and Ophiostoma ips were the only species capable of causing significant lesions.     

Two Ophiostoma species associated with bark beetles in wave-regenerated Abies veitchii forests in Japan

Two species of Ophiostoma were isolated from four bark beetles (i.e., Cryphalus montatus, C. piceae, Dryocoetes hectographus, and Polygraphus proximus) infesting Abies veitchii and from their galleries in the wave-regenerated forests in the central part of the main island (Honshu) of Japan. One of them is described here as a new species, Ophiostoma subalpinum, and the other is identified as O. davidsonii, newly reported in Japan. Ophiostoma subalpinum is characterized by short ostiolar hyphae, oblong or allantoid ascospores enclosed in a thin, hyaline sheath, and a Pesotum anamorph. Received: June 4, 2001 / Accepted: December 25, 2001     

Fungi Vectored by the Bark Beetle Ips typographus Following Hibernation Under the Bark of Standing Trees and in the Forest Litter

The bark beetle Ips typographus has different hibernation environments, under the bark of standing trees or in the forest litter, which is likely to affect the beetle-associated fungal flora. We isolated fungi from beetles, standing I. typographus-attacked trees, and forest litter below the attacked trees. Fungal identification was done using cultural and molecular methods. The results of the two methods in detecting fungal species were compared. Fungal communities associated with I. typographus differed considerably depending on the hibernation environment. In addition to seven taxa of known ophiostomoid I. typographus-associated fungi, we detected 18 ascomycetes and anamorphic fungi, five wood-decaying basidomycetes, 11 yeasts, and four zygomycetes. Of those, 14 fungal taxa were detected exclusively from beetles that hibernated under bark, and six taxa were detected exclusively from beetles hibernating in forest litter. The spruce pathogen, Ceratocystis polonica, was detected occasionally in bark, while another spruce pathogen, Grosmannia europhioides, was detected more often from beetles hibernating under the bark as compared to litter. The identification method had a significant impact on which taxa were detected. Rapidly growing fungal taxa, e.g. Penicillium, Trichoderma, and Ophiostoma, dominated pure culture isolations; while yeasts dominated the communities detected using molecular methods. The study also demonstrated low frequencies of tree pathogenic fungi carried by I. typographus during its outbreaks and that the beetle does not require them to successfully attack and kill trees.     

Interactions betweenAulonium ruficorne [Coleoptera: Colydiidae] and other natural enemies of bark beetles [Coleoptera: Scolytidae]

The within tree distribution of some common natural enemies of bark beetles in pine plantations in Israel and some aspects of their feeding habits were studied with special emphasis on the potential impact on the predatorAulonium ruficorne Olivier. A total of 12 predators and 2 parasite species were found associated withA. ruficorne in the natural enemy complex of bark beetles on pines. No secondary parasites were detected. The anthocoridScoloposcelis pulchella (Zetterstedt) and the dipteranMedetera striata Parent were observed feeding on immature stages ofA. ruficorne in the absence of scolytids. The associated Coleoptea:Nemosema elongatum F.,Rhizophagus bipustulatus L.,Corticeus spp.,Plastysoma spp. andPlegaderus discisus Erickson are thought to compete withA. ruficorne on larvae and pupae of bark beetles when the latter are in limited quantities (especially in the lower section of the tree). The parasites, mainlyMetacolus unifasciatuss Forster andDendrosoter caenopachoides Ruschka are assumed to compete withA. ruficorne during the larval period in the smooth bark section of the stem. Competition might occur mainly during spring and fall. Deutonymphs of the miteIpiduropoda sellnicki were detected on the abdomen ofA. ruficorne adults. Larvae of the predator were rarely infected in the field by the bacteriaSerratia sp. while laboratory cultures suffered high rate of mortality caused by this pathogen.      

Responses of Tribolium castaneum to olfactory cues from cotton seeds,the fungi associated with cotton seeds,and cereals

We tested, in an olfactometer, whether or not Tribolium castaneum Herbst (Coleoptera: Tenebrionidae) responds preferentially to the volatiles that emanate from the fungi associated with cotton [Gossypium hirsutum L. (Malvaceae)] seed over those that emanate from cereals, because cereals are usually portrayed as the primary resources of these beetles. Pairwise comparisons were conducted between cotton seed, wheat (Triticum aestivum L.), and sorghum [Sorghum bicolor (L.) Moench] (both Poaceae); volatiles were tested from intact seeds and from both water and ethanol extracts. The results demonstrate that T. castaneum is attracted more strongly to cotton seeds with its lint contaminated with fungi, than to the conventional resources of this species (i.e., wheat and sorghum). Further tests prove that it is the fungus on the lint that produces the active volatiles, because the beetles did not respond to sterilized cotton lint (i.e., without the fungi typically associated with it when cotton seed is stored). Tests with five fungal cultures (each representing an unidentified species that was isolated from the field‐collected cotton lint) were variable across the cultures, with only one of them being significantly attractive to the beetles. The others were not attractive and one may even have repulsed the beetles. The results are consistent with the beetles having a strong ecological association with fungi and suggest it would be worth investigating the ecology of T. castaneum from this perspective.     

Differences in the fruit maturation stages at which oviposition occurs among insect seed predators feeding on the fruits of five dipterocarp tree species

The seeds of dipterocarp trees are the main food resources for many species of weevils, bark beetles and small moths; however, for most seed‐eating insects on dipterocarp tropical trees, seed utilization patterns remain poorly investigated. This study aimed to determine the fruit maturation stages at which eggs are laid by different insect seed predators feeding on the seeds or fruits of the following five dipterocarp species: Dipterocarpus globosus, Dryobalanops aromatica, Shorea beccariana, S. acuta and S. curtisii, which reproduced during the same period. We investigated the occurrence frequencies of the insect seed predators at various growth stages by collecting both unfallen and fallen fruit on several occasions during the period of seed/fruit maturation in a tropical rainforest in Borneo from September to December 2013. Weevils and bark beetles were the dominant insect seed predators of the five tree species. One or two weevil species of Alcidodes, Damnux and/or Nanophyes preyed on the seeds of each of the five tree species, and one bark beetle species, Coccotrypes gedeanus, preyed on the seeds of all five tree species. Many larvae, pupae and adults of each weevil species were found in pre‐dispersal (unfallen) fruit, whereas bark beetles at various growth stages were found in post‐dispersal (fallen) fruit. These results suggested that, among the dominant insect seed predators of the five dipterocarp species, weevil species oviposit on pre‐dispersal fruit and begin their larval growth before seed dispersal, whereas the oviposition and larval development of bark beetle species occurs in post‐dispersal fruit.