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The spruce beetle, Dendroctonus rufipennis, is an eruptive forest pest of significant economic and ecological importance. D. rufipennis has symbiotic associations with a number of microorganisms, especially the ophiostomatoid fungus Leptographium abietinum. The nature of this interaction is only partially understood. Additionally, mite and nematode associates can mediate bark beetle-fungal interactions, but this has not yet been studied for spruce beetles. In this study, we found eight mite species associated with spruce beetles: Tarsonemus ips, T. endophloeus, Histiogaster arborsignis, Dendrolaelaps quadrisetus, Proctolaelaps hytricoides, Trichouropoda alascae, T. n. sp. nr dalarenaensis, and Urobovella n. sp 767. The most prevalent species was H. arborsignis. In addition, 75% of beetles examined carried nematodes, with six species represented. These included a new species of Parasitorhabditis, Ektaphelenchus obtusus, Bursaphelenchus n. sp. 727, Aphelenchoides n. sp., Panagrolaimus sp., and Mykoletzkya ruminis. H. arborsignis showed strong feeding and oviposition preferences for L. abietinum among four fungal species tested in laboratory assays. Information on our attempts to culture the various nematode species collected from D. rufipennis is also provided. Bursaphelenchus were cultured from D. rufipennis nematangia plated on agar containing L. abietinum but not sterile agar. Thus, L. abietinum plays an important role in these gallery communities, affecting the tree-killing bark beetle, its phoretic mites, and nematodes. These data add to our understanding of bark beetle-microorganism interactions.

We report the Þrst study of gut-associated bacteria of bark beetles using both culture- dependent and culture-independent methods. These insects are major pests of pine trees but also contribute to important ecological functions such as nutrient cycling. We found members of the - and -Proteobacteria and Firmicutes in larvae of the southern pine beetle, Dendroctonus frontalis Zimmermann. Sequences from three larval guts were grouped into one to three operational taxonomic units (OTUs) at 3% difference among sequences. Communities in adult southern pine beetle guts consisted solely of members of the -Proteobacteria. These could be grouped into three to Þve OTUs at 3% difference between sequences. These gut communities have relatively low species richness, which may resect the specialization needed to exploit a nutrient-poor food source, colonize a chemically complex habitat, and maintain consistent associations with mutualistic fungi. However, there is considerable variation in gut microbiota composition among individual insects, suggesting the need for additional studies on sources of variation and potential substitutability among species performing similar functions.

We evaluated whether Dendroctonus frontalis Zimmermann populations were influenced by nontrophic interactions involving commensal mites, their mutualistic bluestain fungus Ophiostoma minus (Hedgc.) H. and P. Sydow, and beetle-mutualistic mycangial fungi. We tested for effects of delayed, nonlinear, or positive feedback from O. minus and mites on D. frontalis population growth. We predicted that (1) high mite densities have demographic consequences for beetles by influencing the prevalence of O. minus and antagonistic interactions between O. minus and mycangial fungi, and (2) inter-relations and abundances of mites and fungi differentially vary throughout the year in a seasonally variable climate. Surveys of D. frontalis populations revealed that temporal and spatial patterns in abundance of mites and their mutualistic fungus, O. minus were inversely related with beetle population growth. Negative demographic effects of O. minus on D. frontalis were nonlinear, only affecting beetle per capita reproduction when fungi colonized >35% of phloem habitat. Mite abundance was strongly correlated with O. minus and was an important driving force in promoting bluestain prevalence within trees. Spring abundances of mites and the prevalence of O. minus during D. frontalis infestation formation were strong predictors of beetle population decline later that year. The two mutualistic fungi associated with D. frontalis cycled seasonally but did not seem to influence beetle population dynamics.

Conifer resin and phloem tissue contain several phytochemical groups, composed primarily of monoterpenes, diterpene acids, and stilbene phenolics. The effects of monoterpenes and phenolics on stem-colonizing bark beetles and their associated microorganisms have been studied to some extent, but the roles of diterpene acids are largely unknown. Diterpene acids are known to have substantial feeding deterrent and growth inhibiting effects on a variety of insect groups and are known to inhibit a variety of fungi. We tested three diterpene acids present in red pine, Pinus resinosa ,a t various concentrations, on several life history components of the bark beetle Ips pini and the fungus Ophiostoma ips. No diterpene acid affected the host acceptance behavior or larval survival of Ips pini. In contrast, abietic acid and isopimaric acid strongly inhibited spore germination of O. ips, and abietic acid strongly inhibited mycelial growth. The levels of inhibition observed were higher than with any previous assays of monoterpenes or phenolics in this system. These results support the view that conifer defenses against bark beetleÐfungal complexes are multifaceted, with all three phytochemical groups being important to P. resinosa, but each with varying relative activity against the beetles and fungi.

Efforts to describe the complex relationships between bark beetles and the ophiosto- matoid (stain) fungi they transport have largely resulted in a dichotomous classiÞcation. These symbioses have been viewed as either mutualistic (i.e., fungi help bark beetles colonize living trees by overcoming tree defenses or by providing nutrients after colonization in return for transport to a host) or antagonistic (i.e., fungi compete for a limited resource and reduce brood development with no apparent beneÞt to the beetle). We investigated several components ofone beetleÐf ungus interaction. SpeciÞcally, we addressed whether beetle entry into, and development within, a host tree vary with the degree ofcolonization by ophiostomatoid fungi. Ips pini (Say) transports several species ofophiostomatoid fungi, the most common being Ophiostoma ips (Rumbold) Nannfeldt, in the process ofcolonizing its host, Pinus resinosa Aitman. We introduced this fungus 0, 3, 7, and 10 d before beetle entry to characterize its effects on I. pini colonization and development. This sequence allowed quantiÞcation of temporal effects and comparison of results with other systems. Fungal growth was greatest when inoculated before beetle colonization. Fungal colonization reduced beetle entry into logs, but increased brood production. Mate capture was not signiÞcantly affected by fungal growth. The beneÞts imparted by O. ips to its beetle vector during brood development are compared with results from other systems. This difference may in part be related to the exploitation of highly stressed and dead trees, rather than vigorous hosts, by I. pini.

Dendroctonus frontalis Zimmerman is a major disturbance agent in American pine forests, but attack preferences for various host species, and their relative suitability for reproduction, are poorly known. We studied patterns of beetle attack and reproduction during an infestation of stands containing Virginia pine and loblolly pine. Nearly all Virginia pine were attacked and killed, whereas a third of the loblolly pine escaped attack. Among attacked trees, the density of landings and attacks on Virginia pine was 56–106% higher than on loblolly pine at one site, whereas it was similar between species at another site. Paradoxically, D. frontalis preferred the host that was least suitable for reproduction: mean ± SE = 0.89 ± 0.33 versus 4.65 ± 1.40 progeny/attack in Virginia pine versus loblolly pine. Poor reproduction in Virginia pine was attributable to increased adult mortality, decreased oviposition, and decreased larval survival. Phloem thickness and nitrogen content were similar between the two...

Temperature has strong effects on metabolic processes of individuals and demographics of populations, but effects on ecological communities are not well known. Many economically and ecologically important pest species have obligate associations with other organisms; therefore, effects of temperature on these species might be mediated by strong interactions. The southern pine beetle (Dendroctonus frontalis Zimmermann) harbors a rich community of phoretic mites and fungi that are linked by many strong direct and indirect interactions, providing multiple pathways for temperature to affect the system. We tested the effects of temperature on this community by manipulating communities within naturally infested sections of pine trees. Direct effects of temperature on component species were conspicuous and sometimes predictable based on single-species physiology, but there were also strong indirect effects of temperature via alteration of species interactions that could not have been predicted based on autecological temperature responses. Climatic variation, including directional warming, will likely influence ecological systems through direct physiological effects as well as indirect effects through species interactions.

Symbiosis is receiving increased attention among all aspects of biology because of the unifying themes it helps construct across ecological, evolutionary, developmental, semiochemical, and pest management theory. Insects show a vast array of symbiotic relationships with a wide diversity of microorganisms. These relationships may confer a variety of benefits to the host (macrosymbiont), such as direct or indirect nutrition, ability to counter the defenses of plant or animal hosts, protection from natural enemies, improved development and reproduction, and communication. Benefits to the microsymbiont (including a broad range of fungi, bacteria, mites, nematodes, etc.) often include transport, protection from antagonists, and protection from environmental extremes. Symbiotic relationships may be mutualistic, commensal, competitive, or parasitic. In many cases, individual relationships may include both beneficial and detrimental effects to each partner during various phases of their life histories or as environmental conditions change. The outcomes of insect-microbial interactions are often strongly mediated by other symbionts and by features of the external and internal environment. These outcomes can also have important effects on human well being and environmental quality, by affecting agriculture, human health, natural resources, and the impacts of invasive species. We argue that, for many systems, our understanding of symbiotic relationships will advance most rapidly where context dependency and multipartite membership are integrated into existing conceptual frameworks. Furthermore, the contribution of entomological studies to overall symbiosis theory will be greatest where preoccupation with strict definitions and artificial boundaries is minimized, and integration of emerging molecular and quantitative techniques is maximized. We highlight symbiotic relations involving bark beetles to illustrate examples of the above trends.

The spruce beetle, Dendroctonus rufipennis, is an eruptive forest pest of significant economic and ecological importance. D. rufipennis has symbiotic associations with a number of microorganisms, especially the ophiostomatoid fungus Leptographium abietinum. The nature of this interaction is only partially understood. Additionally, mite and nematode associates can mediate bark beetle-fungal interactions, but this has not yet been studied for spruce beetles. In this study, we found eight mite species associated with spruce beetles: Tarsonemus ips, T. endophloeus, Histiogaster arborsignis, Dendrolaelaps quadrisetus, Proctolaelaps hytricoides, Trichouropoda alascae, T. n. sp. nr dalarenaensis, and Urobovella n. sp 767. The most prevalent species was H. arborsignis. In addition, 75% of beetles examined carried nematodes, with six species represented. These included a new species of Parasitorhabditis, Ektaphelenchus obtusus, Bursaphelenchus n. sp. 727, Aphelenchoides n. sp., Panagrolaimus sp., and Mykoletzkya ruminis. H. arborsignis showed strong feeding and oviposition preferences for L. abietinum among four fungal species tested in laboratory assays. Information on our attempts to culture the various nematode species collected from D. rufipennis is also provided. Bursaphelenchus were cultured from D. rufipennis nematangia plated on agar containing L. abietinum but not sterile agar. Thus, L. abietinum plays an important role in these gallery communities, affecting the tree-killing bark beetle, its phoretic mites, and nematodes. These data add to our understanding of bark beetle-microorganism interactions.