Dietary benefits of fungal associates to an eruptive herbivore: potential implications of multiple associates on host population dynamics

We used the mountain pine beetle (Dendroctonus ponderosae Hopkins) and its two fungal associates, Grosmannia clavigera and Ophiostoma montium, to study potential nutritional benefits of fungi to bark beetles. We tested for potential effects of feeding on phloem colonized by fungi on beetle performance in field and laboratory studies. The fungi increased nitrogen levels in the phloem of attacked trees by 40%, indicating that it may be an important source of dietary nitrogen for mountain pine beetles. However, nitrogen levels of phloem inoculated with fungi in the laboratory were similar to uncolonized phloem, indicating that the fungi may redistribute nitrogen from the sapwood to the phloem rather than increase absolute levels of nitrogen. Beetles emerging from attacked trees carrying G. clavigera were larger than beetles carrying O. montium, which in turn were larger than beetles lacking fungi. Results of experimental laboratory studies varied, likely because of differences in the growth and sporulation of fungi under artificial conditions. Results indicate that the two fungi may offer complementary benefits to the mountain pine beetle because larvae preferentially fed on phloem colonized by both fungi together over phloem colonized by one fungus or uncolonized phloem. Teneral adults preemergence fed on spores in pupal chambers when they were produced and consumed little phloem before emerging. Teneral adults mined extensively in the phloem before emerging when spores were not produced in the pupal chamber. Our results provide evidence for a nutritional role of fungi in the diet of bark beetles and show that multiple associates may differentially affect beetle performance, which could have important implications for bark beetle population dynamics.     

Ophiostoma species (Ascomycetes: Ophiostomatales) associated with bark beetles (Coleoptera: Scolytinae) colonizing Pinus radiata in northern Spain

Bark beetles (Coleoptera: Scolytinae) are known to be associated with fungi, especially species of Ophiostoma sensu lato and Ceratocystis. However, very little is known about these fungi in Spain. In this study, we examined the fungi associated with 13 bark beetle species and one weevil (Coleoptera: Entiminae) infesting Pinus radiata in the Basque Country of northern Spain. This study included an examination of 1323 bark beetles or their galleries in P. radiata. Isolations yielded a total of 920 cultures, which included 16 species of Ophiostoma sensu lato or their asexual states. These 16 species included 69 associations between fungi and bark beetles and weevils that have not previously been recorded. The most commonly encountered fungal associates of the bark beetles were Ophiostoma ips, Leptographium guttulatum, Ophiostoma stenoceras, and Ophiostoma piceae. In most cases, the niche of colonization had a significant effect on the abundance and composition of colonizing fungi. This confirms that resource overlap between species is reduced by partial spatial segregation. Interaction between niche and time seldom had a significant effect, which suggests that spatial colonization patterns are rarely flexible throughout timber degradation. The differences in common associates among the bark beetle species could be linked to the different niches that these beetles occupy.     

Phloemophagous and xylophagous insects, their parasitoids, predators and inquilines in the branches of the most important oak species in Serbia

Altogether 26 species of phloemophagous and xylophagous insects, 47 species of parasitoids, 14 species of predators and 7 species of inquilines were identified on the branches of Quercus cerris, Q. frainetto, Q. petraea and Q. robur, diameter 3–15 cm, at 24 sites in Serbia over the period 1992–1996. The greatest number of the identified species were taken from Q. petraea branches (66), followed by Q. cerris (49), Q. frainetto (48) and Q. robur (43). Among the identified phloemophagous and xylophagous insects, the most frequent and the most abundant species were Scolytus intricatus, Agrilus angustulus and Xylotrechus antilope. The most frequent and the most abundant parasitoid was Ecphylus silesiacus. In some samples, the species Entedon ergias, Cheiropachus quadrum and Dendrosoter protuberans were also among the more abundant parasitoids. The most significant predator was the species Tilloidea unifasciata, and the most significant inquilines were the species Poecilothrips albopictus and Asynapta pectoralis.     

Mites are the most common vectors of the fungus Gondwanamyces proteae in Protea infructescences

Entomochoric spore dispersal is well-documented for most ophiostomatoid fungal genera, most of which are associated with bark or ambrosia beetles. Gondwanamyces spp. are unusual members of this group that were first discovered in the flower heads of the primitive angiosperm genus Protea, that is mostly restricted to the Cape Floristic region of Africa. In this study, we present the discovery of the vectors of Gondwanamyces proteae in Protea repens infructescences, which were identified using PCR, direct isolation, and light microscopy. Gondwanamyces proteae DNA and ascospores were identified on diverse lineages of arthropods including beetles (Euderes lineicolis and Genuchus hottentottus), bugs (Oxycarenus maculates), a psocopteran species and five mite (Acari) species. Based on isolation frequency, however, a mite species in the genus Trichouropoda appears to be the most common vector of G. proteae. Gondwanamyces spores were frequently observed within pit mycangia at the base of the legs of these mites. Manipulative experiments demonstrated the ability of mites to carry viable G. proteae spores whilst in transit on the beetle G. hottentottus and that these mites are able to transfer G. proteae spores to uncolonised substrates in vitro. Interestingly, this same mite species has also been implicated as vector of Ophiostoma spores on P. repens and belongs to the same genus of mites that vector Ophiostoma spp. associated with pine-infesting bark beetles in the Northern Hemisphere.     

Effects of learning experience on behaviour of the generalist parasitoid Sclerodermus pupariae to novel hosts

Massicus raddei Blessig (Coleoptera: Cerambycidae), also referred to as the oak long‐horned beetle (OLB), is a non‐natural host for the generalist parasitoid Sclerodermus pupariae Yang et Yao (Hymenoptera: Bethylidae). To determine whether this generalist parasitoid might be a suitable agent for the control of OLB, the adaptive learning experience of adult female parasitoids to OLB larvae was investigated in the laboratory. A Y‐tube olfactometer bioassay was used to examine the effects of adaptive learning experience on the foraging ability of parasitoids for OLB larvae. The results indicated that parasitoids were significantly attracted by the volatiles of ash bark, Fraxinus velutina, with emerald ash borer (EAB), Agrilus planipennis Fairmaire (Coleoptera: Buprestidae) larvae and larval frass, after exposure to ash bark mixed with EAB larval frass (learning condition A). In contrast, after exposure to oak bark, Quercus liaotungensis, mixed with OLB larval frass (learning condition C), parasitoids showed significant preference for the volatiles of oak bark with OLB larvae and larval frass. On the basis of the results of no‐choice tests, we found that parasitoids exposed to learning condition C had greater paralysis efficiency and higher OLB larvae parasitism rates than those exposed to learning condition A or no experience. Furthermore, parasitoids fed on OLB larvae in learning condition C had significantly greater paralysis efficiency and higher OLB larvae parasitism rates than other parasitoids tested. Parasitoids fed on EAB larvae in learning condition A had the lowest paralysis efficiency and OLB larvae parasitism rates among the parasitoids tested. These findings suggested that adaptive learning significantly enhanced the ability of a generalist parasitoid to utilize a novel host. This may provide a new approach to controlling non‐natural hosts using generalist parasitoids.     

Discovery of fungus-mite mutualism in a unique niche

The floral heads (infructescences) of South African Protea L. represent a most unusual niche for fungi of the economically important genus Ophiostoma Syd. and P. Syd. emend. Z.W. de Beer et al. Current consensus holds that most members of Ophiostoma are vectored by tree-infesting bark beetles. However, it has recently been suggested that mites, phoretic on these bark beetles, may play a central role in the dispersal of Ophiostoma. No bark beetles are known from Protea. Therefore, identifying the vectors of Ophiostoma in Protea infructescences would independently evaluate the role of various arthropods in the dispersal of Ophiostoma. Infructescence-colonizing arthropods were tested for the presence of Ophiostoma DNA using polymerase chain reaction (PCR) and for reproductive propagules by isolation on agar plates. PCR tests revealed that few insects carried Ophiostoma DNA. In contrast, various mites (Proctolaelaps vandenbergi Ryke, two species of Tarsonemus Canestrini and Fonzago, and one Trichouropoda Berlese species) frequently carried Ophiostoma propagules. DNA sequence comparisons for 28S ribosomal DNA confirmed the presence of O. splendens G. J. Marais and M. J. Wingf., O. palmiculminatum Roets et al., and O. phasma Roets et al. on these mites. Two apparently undescribed species of Ophiostoma were also identified. Light and scanning electron microscopy revealed specialized structures in Trichouropoda and one Tarsonemus sp. that frequently contained Ophiostoma spores. The Trichouropoda sp. was able to complete its life cycle on a diet consisting solely of its identified phoretic Ophiostoma spp. This study provides compelling evidence that mites are the primary vectors of infructescence-associated Ophiostoma spp. in South Africa.     

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.     

Effect of nutrition of parasites with nectar of melliferous plants on parasitism of the elm bark beetles (Col., Scolytidae)

In the forest of Kljestevica, the elm is attacked by four species of elm bark beetles: Scolytus multistriatus (Marsh.), Scolytus scolytus (Fab.), Scolytus pygmaeus (Fab.) and Scolytus ensifer (Rich.) (Col., Scolytidae). Scolytus multistriatus and S. pygmaeus are the most numerous species. Elm bark beetle is parasitized by four species of parasites: Dendrosoter protuberans (Nees), Ecphylus silesiacus (Ratz.), Coeloides scolyticida (Wesm.) (Hym., Braconidae) and Entedon leucogramma (Ratz.) (Hym., Eulophidae). Ecphylus silesiacus and D. protuberans are the most numerous species. The nectar of mustard ( Sinapis alba L., Brassicaceae) flowers, sweet basil ( Stachys recta L., Lamiaceae) flowers and of wild carrot ( Daucus carota L., Umbelliferae), has an important effect on the number of parasites and on the parasitism of the elm bark beetles. Wild carrot is the most attractive plant to the parasites for additional nutrition, especially for D. protuberans . Mustard and sweet basil flowers are suitable for additional nutrition of E. silesiacus imagos. On meadow flowers growing spontaneously in forest clearings, the lowest number of parasite imagos was collected by an entomological catcher, and the lowest percentage of parasitized larvae and eggs of the elm bark beetle was recorded in elm control catch trunks that were not near the cultivated melliferous plants.     

Assessing the pathogenic effect ofFusarium, Geosmithia andOphiostoma fungi from broad-leaved trees

Phytopathogenic effect of Geosmithia pallida, G. langdonii, Ophiostoma grandicarpum, O. querci, two isolates of O. piceae, and two isolates of Fusarium solani was compared using plant growth test (stem and root length of garden cress plants seeded on mycelium-covered potato carrot agar); Ophiostoma spp. and F. solani were isolated from oak, Geosmithia spp. from galleries of Scolytus intricatus on beech. All fungi inhibited more the root elongation than that of stems. F. solani led to plant collapse after briefly stimulating the growth of stem and in one case also root. G. langdonii inhibited stem and root growth to 20% and led to plant collapse. G. pallida inhibited root growth to 25% whereas stem growth was almost unimpaired. Ophiostoma spp. reduced stem growth to approximately 60-80% and root growth to 25-60%. O. piceae and O. querci caused plant collapse after 15-20 d.     

Leptographium pruni, sp. nov. from bark beetle-infested Prunus jamasakura in Japan

Leptographium species are anamorphs of Ophiostoma, commonly isolated from conifer. There are, however, a small number of these fungi that have been collected from angiosperm hosts. In this study, we describe Leptographium pruni, sp. nov. isolated from the bark of Prunus jamasakura infested by the bark beetle Polygraphus ssiori. This new species is unusual in having a distinct Sporothrix synanamorph with ramoconidia. No evidence of a teleomorph was found, but a high level of tolerance to the antibiotic cycloheximide and the presence of a Sporothrix synanamorph suggest that L. pruni is an Ophiostoma anamorph. Analysis of sequence data for the domain 1 region of the LSUrDNA operon also supports the phylogenetic relationship of L. pruni with Ophiostoma. In addition, sequence data suggest that L. pruni is related to other species of Leptographium rather than Pesotum species with distinct Sporothrix synanamorphs.     

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.     

Hymenopterous parasitoids of bark beetles [Scolytidae] in Israel: Host relation,host plant,abundance and seasonal history

Twenty six species of parasitoids were recorded from 15 species of bark beetles in Israel. Parasitoids were collected from infested cut branches of broadleaves and trap-trees of pine and cypress. Most of the parasitoids are oligophagous and range fromEcphylus caudatus, a monophagous species exclusive toHypoborus ficus, toHeydenia pretiosa, which develops on 11 species.Scleroderma domesticus, Agrilocida ferrieri andEupelmus nearurozonus are occasional parasitoids of bark beetles.Cephalonomia hypobori, Israelius carthami, S. domesticus, Entedon ergias, Cerocephala eccoptogasteri andRoptocerus xylophagorum are cryptoparasitoids and search for their host in the galleries. Known parasitoids are listed from 15 tree genera belonging to 8 botanical families. A typical range for the common parasitoids is presented with within-tree distribution of pine parasitoids. Relatively monophagous species are usually dominant parasitoids in a specific beetle population, whereas relatively polyphagous parasitoids vary in abundance among different host populations.Calosota spp., 2Eurytoma spp. andDendrosoter hartigii are usually present in small numbers. Information concerning the seasonal history of the parasitoids suggests that the major species may raise about 5 to 7 annual generations. Contribution from the Agricultural Research Organization, Bet Dagan, Israel N^o 1331-E, 1984 series     

Ophiostomatoid fungi associated with hardwood-infesting bark and ambrosia beetles in Poland: Taxonomic diversity and vector specificity

Fungi in the orders Ophiostomatales and Microascales (Ascomycota), often designated as ophiostomatoid fungi, are frequent associates of scolytine bark and ambrosia beetles that colonize hardwood and coniferous trees. Several species, e.g., Ophiostoma novo-ulmi, are economically damaging pathogens of trees. Because little is known regarding the ophiostomatoid fungi in Europe, we have explored the diversity of these fungi associated with hardwood-infesting beetles in Poland. This study aims to clarify the associations between fungi in the genera Ambrosiella, Graphium (Microascales), Graphilbum, Leptographium, Ophiostoma and Sporothrix (Ophiostomatales) and their beetle vectors in hardwood ecosystems. Samples associated with 18 bark and ambrosia beetle species were collected from 11 stands in Poland. Fungi were isolated from adult beetles and galleries. Isolates were identified based on morphology, DNA sequence comparisons for five gene regions (ITS, LSU, ßT, TEF 1-α, and CAL) and phylogenetic analyses. In total, 36 distinct taxa were identified, including 24 known and 12 currently unknown species. Several associations between fungi and bark and ambrosia beetles were recorded for the first time. In addition, associations between Dryocoetes alni, D. villosus, Hylesinus crenatus, Ernoporus tiliae, Pteleobius vittatus and ophiostomatoid fungi were reported for the first time, and Sporothrix eucastanea was reported for the first time outside of the USA. Among the species of Ophiostomatales, 14 species were in Ophiostoma s. l., two species were in Graphilbum, nine species were in Sporothrix, and seven species were in Leptographium s. l. Among the species of Microascales, three species were in Graphium, and one was in Ambrosiella. Twenty taxa were present on the beetles and in the galleries, twelve only on beetles, and four only in galleries. Bark and ambrosia beetles from hardwoods appear to be regular vectors, with ophiostomatoid fungi present in all the beetle species. Most ophiostomatoid species had a distinct level of vector/host specificity, although Ophiostoma quercus, the most frequently encountered species, also had the greatest range of beetle vectors and tree hosts. Plant pathogenic O. novo-ulmi was found mainly in association with elm-infesting bark beetles (Scolytus multistriatus, S. scolytus, and P. vittatus) and occasionally with H. crenatus on Fraxinus excelsior and with Scolytus intricatus on Quercus robur.     

Rapid-cooling and storage of plant nematodes at -140 degrees C

Low temperatures can assure the long-term or even indefinite preservation of important biological specimens. Nematode cryopreservation allows for the availability of large numbers of living nematodes at any one time, especially for experimental purposes. New isolates of Bursaphelenchus have recently been collected, including Bursaphelenchus eremus (Rühm) Goodey. This species was identified in north-central Italy on dying oak trees and from the bark beetle Scolytus intricatus Ratzeburg as dauer larvae. We therefore, sought to develop a cryopreservation technique for the long-term storage of all available Bursaphelenchus spp. The technique consists of a rapid-cooling protocol involving immersion in a liquid nitrogen bath before storage of the frozen samples in a mechanical freezer at -140 degrees C. The survival of nematodes subjected to this rapid-cooling protocol was higher than previously reported using slow-cooling methods and is suitable for several species of Bursaphelenchus and other phytoparasitic nematodes.     

Ophiostoma gemellus and Sporothrix variecibatus from mites infesting Protea infructescences in South Africa

Ophiostoma (Ophiostomatales) represents a large genus of fungi mainly known from associations with bark beetles (Curculionidae: Scolytinae) infesting conifers in the northern hemisphere. Few southern hemisphere native species are known, and the five species that consistently occur in the infructescences of Protea spp. in South Africa are ecologically unusual. Little is known about the vectors of Ophiostoma spp. from Protea infructescences, however recent studies have considered the possible role of insects and mites in the distribution of these exceptional fungi. In this study we describe a new species of Ophiostoma and a new Sporothrix spp. with affinities to Ophiostoma, both initially isolated from mites associated with Protea spp. They are described as Ophiostoma gemellus sp. nov. and Sporothrix variecibatus sp. nov. based on their morphology and comparisons of DNA sequence data of the 28S ribosomal, beta-tubulin and internal transcribed spacer (ITS1, 5.8S, ITS2) regions. DNA sequences of S. variecibatus were identical to those of a Sporothrix isolate obtained from Eucalyptus leaf litter in the same area in which S. variecibatus occurs in Protea infructescences. Results of this study add evidence to the view that mites are the vectors of Ophiostoma spp. that colonize Protea infructescences. They also show that DNA sequence comparisons are likely to reveal additional cryptic species of Ophiostoma in this unusual niche.     

Species of Ophiostomataceae isolated from Picea abies infested by the bark beetle Ips typographic

Isolations of microorganisms were made from the sapwood of Norway spruce ( Picea abies ) (L.) Karst. infested by the spruce bark beetle Ips typographus L. The most important group of fungi isolated, the species of Ophiostomataceae, is treated. Ten species were isolated, of which Ophiostoma polonicum Siem. appears to be the most active invader of the sapwood. Three new species are described, Ophiostoma ainoae sp. nov., O. flexuosum sp. nov. and O. cucullatum sp. nov. and three new combinations are proposed, O. davidsonii (Olchow. & Reid.) comb. nov., O. europhioides (Wright & Cain) comb. nov. and O. sagmatospora (Wright & Cain) comb. nov.     

Large Shift in Symbiont Assemblage in the Invasive Red Turpentine Beetle

Changes in symbiont assemblages can affect the success and impact of invasive species, and may provide knowledge regarding the invasion histories of their vectors. Bark beetle symbioses are ideal systems to study changes in symbiont assemblages resulting from invasions. The red turpentine beetle (Dendroctonus valens) is a bark beetle species that recently invaded China from its native range in North America. It is associated with ophiostomatalean fungi in both locations, although the fungi have previously been well-surveyed only in China. We surveyed the ophiostomatalean fungi associated with D. valens in eastern and western North America, and identified the fungal species using multi-gene phylogenies. From the 307 collected isolates (147 in eastern North America and 160 in western North America), we identified 20 species: 11 in eastern North America and 13 in western North America. Four species were shared between eastern North America and western North America, one species (Ophiostoma floccosum) was shared between western North America and China, and three species (Grosmannia koreana, Leptographium procerum, and Ophiostoma abietinum) were shared between eastern North America and China. Ophiostoma floccosum and O. abietinum have worldwide distributions, and were rarely isolated from D. valens. However, G. koreana and L. procerum are primarily limited to Asia and North America respectively. Leptographium procerum, which is thought to be native to North America, represented >45% of the symbionts of D. valens in eastern North America and China, suggesting D. valens may have been introduced to China from eastern North America. These results are surprising, as previous population genetics studies on D. valens based on the cytochrome oxidase I gene have suggested that the insect was introduced into China from western North America.     

Chalcids of the subfamily Pteromalinae (Hymenoptera,Pteromalidae) as parasitoids of the bark beetles (Coleoptera,Scolytidae) in the fauna of Russia and adjacent territories

Eighteen species of ten genera of bark beetle parasitoids were found in the fauna of Russia and adjacent countries. Sixteen species of the parasitoids are recorded as new to the faunas of Belarus, Ukraine, Azerbaijan, Kazakhstan, Tajikistan, Uzbekistan, Kyrgyzstan, and regions of Russia. An annotated list of nineteen species of pteromalids, with data on the material examined, geographic distribution, and hosts is given. New hosts are reported for five species of the bark beetle ectoparasitoids. An original key to eighteen species and ten genera of Pteromalidae parasitizing bark beetles is provided.     

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

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