\u3ci\u3eDendroctonus Valens\u3c/i\u3e and \u3ci\u3eHylastes Porculus\u3c/i\u3e (Coleoptera: Scolytidae): Vectors of Pathogenic Fungi (Ophiostomatales) Associated With Red Pine Decline Disease.

A study was conducted to determine whether Dendroctonus valens and Hylastes porculus could vector their commonly associated fungi to red pine. Field collected adult D. valens transmitted Leptographium terebrantis, Leptographium procerum and Ophiostoma ips into 45%, 30%, and 5%, respectively of the wounded red pine roots onto which they were caged. Field collected H. porculus transmitted L. terebrantis, L. procerum and O. ips into 55%, 40%, and 5%, respectively, of the wounded red pine roots onto which beetles were caged. None of the control roots, which were mechanically wounded only, were found to contain O. ips, whereas only one control root contained L. terebrantis and only one control root contained L. procerum. This work demonstrates that D. valens and H. porculus can vector their associated Leptographium fungi to red pine trees and that these organisms are likely involved in red pine decline disease

Competitive Interactions among Symbiotic Fungi of the Southern Pine Beetle

The southern pine beetle, a damaging pest of conifers, is intimately linked to three symbiotic fungi. Two fungi, Ceratocystiopsis ranaculosus and Entomocorticium sp. A, are transported within specialized structures (mycangia) in the beetle exoskeleton and are mutualists of the beetle. A third fungus, Ophiostoma minus, is transported externally on the beetle exoskeleton (phoretically) and is an antagonist of the beetle. This study examined competitive interactions among these three fungi. The results of de Wit replacement series and primary and secondary resource capture assays with these fungi provide strong evidence for differential competition between the phoretic and mycangial fungi. O. minus was the most able to capture both uncolonized and colonized resources. Entomocorticium sp. A and C. ranaculosus, although equal to one another in compet- itive abilities, differed in their ability to compete with O. minus. Entomocorticium sp. A was able to maintain space free of O. minus to a much greater degree than was C. ranaculosus. The outcome of such competitive interactions may have significant impacts on the biology of this ecologically and economically important beetle

Effects of Feeding by Two Folivorous Arthropods on Susceptibility of Hybrid Poplar Clones to a Foliar Pathogen

We investigated variation in folivore-induced effects on subsequent plant suitability to a foliar pathogen. We used a leaf disk assay to expose three clones of hybrid poplar, NC11382, NE332 and NM6, to colonization by a leaf spot pathogen, Septoria musiva. Undamaged leaf disks of NE332 were the most resistant to S. musiva, followed by NM6 and NC11382, respectively. To test the effects of prior herbivory on subsequent susceptibility to this fungal pathogen, we inoculated S. musiva on leaf disks taken from leaves which had been exposed to feeding by Tetranychus mites or cottonwood leaf beetles. Prior activity by mites and cottonwood leaf beetle affected the subsequent susceptibility of clones NC 11382 and NE332 to S. musiva

Fungus Cultivation by Ambrosia Beetles: Behavior and Laboratory Breeding Success in Three Xyleborine Species

Fungus cultivation by ambrosia beetles is one of the four independently evolved cases of agriculture known in animals. Such cultivation is most advanced in the highly social subtribe Xyleborina (Scolytinae), which is characterized by haplodiploidy and extreme levels of inbreeding. Despite their ubiquity in forests worldwide, the behavior of these beetles remains poorly understood. This may be in part because of their cryptic life habits within the wood of trees. Here we present data obtained by varying a laboratory breeding technique based on artificial medium inside glass tubes, which enables behavioral observations. We studied species of the three most widespread genera of Xyleborina in the temperate zone: Xyleborus, Xyleborinus, and Xylosandrus. We raised several generations of each species with good breeding success in two types of media. The proportion of females of Xyleborinus saxesenii Ratzeburg producing offspring within 40 d depended significantly on founder female origin, which shows a transgenerational effect. Labor-intensive microbial sterilization techniques did not increase females' breeding success relative to a group of females shortly treated with ethanol. Gallery productivity measured as the mean number of mature offspring produced after 40 d varied between species and was weakly affected by the type of medium used and foundress origin (field or laboratory) in X. saxesenii, whereas different preparation and sterilization techniques of the beetles had no effect. Behavioral observations showed the time course of different reproductive stages and enabled to obtain detailed behavioral information in all species studied. We propose that the laboratory techniques we describe here are suited for extensive studies of sociality and modes of agriculture in the xyleborine ambrosia beetles, which may yield important insights into the evolution of fungal agriculture and advanced social organizatio

Abundance and dynamics of filamentous fungi in the complex ambrosia gardens of the primitively eusocial beetle Xyleborinus saxeseniiRatzeburg (Coleoptera: Curculionidae, Scolytinae)

Insect fungus gardens consist of a community of interacting microorganisms that can have either beneficial or detrimental effects to the farmers. In contrast to fungus-farming ants and termites, the fungal communities of ambrosia beetles and the effects of particular fungal species on the farmers are largely unknown. Here, we used a laboratory rearing technique for studying the filamentous fungal garden community of the ambrosia beetle, Xyleborinus saxesenii, which cultivates fungi in tunnels excavated within dead trees. Raffaelea sulfurea and Fusicolla acetilerea were transmitted in spore-carrying organs by gallery founding females and established first in new gardens. Raffaelea sulfurea had positive effects on egg-laying and larval numbers. Over time, four other fungal species emerged in the gardens. Prevalence of one of them, Paecilomyces variotii, correlated negatively with larval numbers and can be harmful to adults by forming biofilms on their bodies. It also comprised the main portion of garden material removed from galleries by adults. Our data suggest that two mutualistic, several commensalistic and one to two pathogenic filamentous fungi are associated with X. saxesenii. Fungal diversity in gardens of ambrosia beetles appears to be much lower than that in gardens of fungus-culturing ants, which seems to result from essential differences in substrates and behaviour

Effects of Feeding by Two Folivorous Arthropods on Susceptibility of Hybrid Poplar Clones to a Foliar Pathogen

We investigated variation in folivore-induced effects on subsequent plant suitability to a foliar pathogen. We used a leaf disk assay to expose three clones of hybrid poplar, NC11382, NE332 and NM6, to colonization by a leaf spot pathogen, Septoria musiva. Undamaged leaf disks of NE332 were the most resistant to S. musiva, followed by NM6 and NC11382, respectively. To test the effects of prior herbivory on subsequent susceptibility to this fungal pathogen, we inoculated S. musiva on leaf disks taken from leaves which had been exposed to feeding by Tetranychus mites or cottonwood leaf beetles. Prior activity by mites and cottonwood leaf beetle affected the subsequent susceptibility of clones NC 11382 and NE332 to S. musiva

Antagonisms, mutualisms and commensalisms affect outbreak dynamics of the southern pine beetle

Feedback from community interactions involving mutualisms are a rarely explored mechanism for generating complex population dynamics. We examined the effects of two linked mutualisms on the population dynamics of a beetle that exhibits outbreak dynamics. One mutualism involves an obligate association between the bark beetle, Dendroctonus frontalis and two mycangial fungi. The second mutualism involves Tarsonemus mites that are phoretic on D. frontalis ( commensal ), and a blue-staining fungus, Ophiostoma minus. The presence of O. minus reduces beetle larval survival ( antagonistic ) by outcompeting beetle-mutualistic fungi within trees yet supports mite populations by acting as a nutritional mutualist. These linked interactions potentially create an interaction system with the form of an endogenous negative feedback loop. We address four hypotheses: (1) Direct negative feedback: Beetles directly increase the abundance of O. minus, which reduces per capita reproduction of beetles. (2) Indirect negative feedback: Beetles indirectly increase mite abundance, which increases O. minus, which decreases beetle reproduction. (3) The effect of O. minus on beetles depends on mites, but mite abundance is independent of beetle abundance. (4) The effect of O. minus on beetles is independent of beetle and mite abundance. High Tarsonemus and O. minus abundances were strongly correlated with the decline and eventual local extinction of beetle populations. Manipulation experiments revealed strong negative effects of O. minus on beetles, but falsified the hypothesis that horizontal transmission of O. minus generates negative feedback. Surveys of beetle populations revealed that reproductive rates of Tarsonemus, O. minus, and beetles covaried in a manner consistent with strong indirect interactions between organisms. Co-occurrence of mutualisms embedded within a community may have stabilizing effects if both mutualisms limit each other. However, delays and/ or non-linearities in the interaction systems may result in large population fluctuations. © Springer-Verlag 2005

Diversity and evolution of entomocorticium (Russulales, peniophoraceae), a genus of bark beetle mutualists derived from free-living, wood rotting peniophora

Symbiosis between insects and fungi arose multiple times during the evolution of both groups, and some of the most biologically diverse and economically important are mutualisms in which the insects cultivate and feed on fungi. Among these are bark beetles, whose ascomycetous cultivars are better known and studied than their frequently-overlooked and poorly understood basidiomycetous partners. In this study, we propose five new species of Entomocorticium, fungal mutualists in the Russulales (Basidiomycota) that are mutualistic symbionts of scolytine beetles. We have isolated these fungi from the beetle mycangia, which are structures adapted for the selective storage and transportation of fungal mutualists. Herein, we present the most complete phylogeny of the closely related genera Entomocorticium and Peniophora and provide insights into how an insect-associated taxon (Entomocorticium) evolved from within a wood-decaying, wind-dispersed lineage (Peniophora). Our results indicate that following a transition from angiosperms to gymnosperms, fungal domestication by beetles facilitated the evolution and diversification of Entomocorticium. We additionally propose four new species: Entomocorticium fibulatum Araújo, Li & Hulcr, sp. nov.; E. belizense Araújo, Li & Hulcr, sp. nov.; E. perryae Araújo, Li & Hulcr, sp. nov.; and E. macrovesiculatum Araújo, Li, Six & Hulcr, sp. nov. Our findings highlight the fact that insect-fungi associations remain an understudied field and that these associations harbor a large reservoir of novel fungal species.Fil: Araújo, João P. M.. University of Florida; Estados Unidos. New York Botanical Garden; Estados UnidosFil: Li, You. University of Florida; Estados UnidosFil: Six, Diana. University of Montana; Estados UnidosFil: Rajchenberg, Mario. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Centro de Investigación y Extensión Forestal Andino Patagónico; ArgentinaFil: Smith, Matthew E.. University of Florida; Estados UnidosFil: Johnson, Andrew J.. University of Florida; Estados UnidosFil: Klepzig, Kier D.. University of Georgia; Estados UnidosFil: Crous, Pedro W.. Westerdijk Fungal Biodiversity Institute; Países BajosFil: Leal Dutra, Caio A.. Universidad de Copenhagen; DinamarcaFil: Skelton, James. The College of William and Mary; Estados UnidosFil: Adams, Sawyer N.. University of Florida; Estados UnidosFil: Hulcr, Jiri. University of Florida; Estados Unido

Diversity and evolution of entomocorticium (Russulales, peniophoraceae), a genus of bark beetle mutualists derived from free-living, wood rotting peniophora

Symbiosis between insects and fungi arose multiple times during the evolution of both groups, and some of the most biologically diverse and economically important are mutualisms in which the insects cultivate and feed on fungi. Among these are bark beetles, whose ascomycetous cultivars are better known and studied than their frequently-overlooked and poorly understood basidiomycetous partners. In this study, we propose five new species of Entomocorticium, fungal mutualists in the Russulales (Basidiomycota) that are mutualistic symbionts of scolytine beetles. We have isolated these fungi from the beetle mycangia, which are structures adapted for the selective storage and transportation of fungal mutualists. Herein, we present the most complete phylogeny of the closely related genera Entomocorticium and Peniophora and provide insights into how an insect-associated taxon (Entomocorticium) evolved from within a wood-decaying, wind-dispersed lineage (Peniophora). Our results indicate that following a transition from angiosperms to gymnosperms, fungal domestication by beetles facilitated the evolution and diversification of Entomocorticium. We additionally propose four new species: Entomocorticium fibulatum Araújo, Li & Hulcr, sp. nov.; E. belizense Araújo, Li & Hulcr, sp. nov.; E. perryae Araújo, Li & Hulcr, sp. nov.; and E. macrovesiculatum Araújo, Li, Six & Hulcr, sp. nov. Our findings highlight the fact that insect-fungi associations remain an understudied field and that these associations harbor a large reservoir of novel fungal species.Fil: Araújo, João P. M.. University of Florida; Estados Unidos. New York Botanical Garden; Estados UnidosFil: Li, You. University of Florida; Estados UnidosFil: Six, Diana. University of Montana; Estados UnidosFil: Rajchenberg, Mario. Consejo Nacional de Investigaciones Científicas y Técnicas; Argentina. Centro de Investigación y Extensión Forestal Andino Patagónico; ArgentinaFil: Smith, Matthew E.. University of Florida; Estados UnidosFil: Johnson, Andrew J.. University of Florida; Estados UnidosFil: Klepzig, Kier D.. University of Georgia; Estados UnidosFil: Crous, Pedro W.. Westerdijk Fungal Biodiversity Institute; Países BajosFil: Leal Dutra, Caio A.. Universidad de Copenhagen; DinamarcaFil: Skelton, James. The College of William and Mary; Estados UnidosFil: Adams, Sawyer N.. University of Florida; Estados UnidosFil: Hulcr, Jiri. University of Florida; Estados Unido