@article {9447,
	title = {Assemblage of Hymenoptera arriving at logs colonized by Ips pini (Coleoptera: Curculionidae: Scolytinae) and its microbial symbionts in western Montana},
	journal = {Canadian Entomologist},
	volume = {141},
	number = {2},
	year = {2009},
	month = {March-April},
	pages = {172-199},
	abstract = {Colonization of a tree by bark beetles and their symbionts creates a new habitat for a diverse assemblage of arthropods, including competing herbivores, xylophages, fungivores, saprophages, predators, and parasitoids. Understanding these assemblages is important for evaluating, nontarget effects of various management tactics and for subsequently evaluating how changes in climate, the presence of invasive species. and altered forestry practices and land-use tenure may affect biodiversity. We characterized the assemblage of hymenopterans attracted to logs of ponderosa pine (Pinus ponderosa C. Lawson (Pinaceae)) colonized by the bark beetle Ips pini (Say) and its microbial symbionts. In one experiment, the composition and relative abundances, of species arriving at hosts colonized by I. pini, and possible sources of attraction, were determined. Treatments consisted of a loo containing I. pini with its natural complement of microorganisms, a log alone, and a blank control. A second experiment was carried out to determine whether or not Hymenoptera were attracted to microbial symbionts of I. pini. Treatments consisted of a blank control, a to, alone, a lop containing L pini with its natural complement of microrganisms, either Ophiostoma ips, Burk-holderia sp., or Pichia scolyti. and a log inoculated with a combination of these three microorganisms. Over 2 years, 5163 Hymenoptera were captured, of which over 98\% were parasitoids. Braconidae, Platygastridae, Encyrtidae. Pteromalidae, and Ichneumonidae were the most abundant. Seven known species of bark beetle parasitoids (all Pteromalidae) were captured. However. parasitoids of Diptera, Lepidoptera, Hymenoptera. and non-wood-boring Coleoptera were also common. Nineteen species showed preferential attraction to host plants infested with I. pini and its complement of microorganisms, host plants inoculated with I. pini microbial symbionts, or host plants alone. Interestingly, many of these species were parasitoids of phytophagous, fungivorous, and saprophytic insects rather than of bark beetles themselves. These results Suggest that a diverse assemblage of natural enemies that attack various feeding guilds within a common habitat exploit common olfactory cues.},
	keywords = {Ips pini},
	isbn = {0008-347X},
	author = {Boone, Celia K. and Six, Diana L. and Krauth, Steven J. and Raffa, Kenneth F.}
}
@article {9448,
	title = {The enemy of my enemy is still my enemy: competitors add to predator load of a tree-killing bark beetle},
	journal = {Agricultural and Forest Entomology},
	volume = {10},
	number = {4},
	year = {2008},
	month = {November},
	pages = {411-421},
	type = {Article},
	abstract = {1 The mountain pine beetle Dendroctonus ponderosae is a major tree-killing bark beetle in North America. We evaluated how the subsequent arrival of a competing bark beetle Ips pini influences the arrival of predators and their impact on both species.2 The predators Temnochila chlorodia and Enoclerus sphegeus were strongly attracted to pheromones of D. ponderosae. By contrast, Enoclerus lecontei was mostly attracted to I. pini pheromones. The host compound myrcene synergized attraction of both D. ponderosae and E. sphegeus to the pheromone of D. ponderosae. However, it inhibited attraction of both I. pini and E. lecontei to I. pini{\textquoteright}s pheromone.3 Dendroctonus ponderosae were more attracted to trees than logs treated with its pheromones, whereas I. pini were more attracted to logs than trees treated with its pheromones. Some 78\% of T. chlorodia were captured at hosts baited with D. ponderosae pheromones, whereas 83\% of E. lecontei were captured at hosts baited with I. pini pheromones. We characterized the sequence of arrival to live trees baited with pheromones of D. ponderosae as: D. ponderosae, T. chlorodia, E. sphegeus, I. pini, E. lecontei.4 Various combinations of I. pini and predators were added to logs colonized by D. ponderosae in the above sequence of arrival observed in live trees baited with D. ponderosae aggregation pheromones. Ips pini reduced D. ponderosae adult brood production. However, the combination of I. pini and E. lecontei did not raise D. ponderosae brood production above that observed with only I. pini present. Similarly, the combination of I. pini and T. chlorodia did not reduce D. ponderosae brood production below that observed with I. pini alone. By contrast, the combination of I. pini, T. chlorodia and E. lecontei caused more brood loss to D. ponderosae than I. pini alone.5 Enoclerus lecontei did not reduce brood production by T. chlorodia, whereas T. chlorodia substantially reduced brood production by E. lecontei.6 Secondary bark beetles that exploit the resource created by primary tree-killing species exert negative effects through both competition and increased predator load. Implications to the population dynamics, ecology and evolution of tree-killing bark beetles are discussed.},
	keywords = {Dendroctonus ponderosae, Ips pini},
	isbn = {1461-9555},
	author = {Boone, Celia K. and Six, Diana L. and Raffa, Kenneth F.}
}
@article {9449,
	title = {Parasitoids and dipteran predators exploit volatiles from microbial symbionts to locate bark beetles},
	journal = {Environmental Entomology},
	volume = {37},
	number = {1},
	year = {2008},
	month = {February},
	pages = {150-161},
	type = {Article},
	abstract = {Host location by parasitoids and dipteran predators of bark beetles is poorly understood. Unlike coleopteran predators that locate prey by orienting to prey pheromones, wasps and flies often attack life stages not present until after pheromone production ceases. Bark beetles have important microbial symbionts, which could provide sources of cues. We tested host trees, trees colonized by beetles and symbionts, and trees colonized by symbionts alone for attractiveness to hymenopteran parasitoids and dipteran predators. Field studies were conducted with Ips pini in Montana. Three pteromalid wasps were predominant. All were associated with the second and third instars of L pini. Heydenia unica was more attracted to logs colonized by either I. pini or the fungus Ophiostoma ips than logs alone or blank controls (screen with no log). Rhopalicus pulchripennis was more attracted to logs colonized by L pini than logs alone or blank controls. Dibrachys cavus was attracted to logs but did not distinguish whether or not they were colonized. Two dolichopodid predators were predominant. A Medetera species was more attracted to colonized than uncolonized logs and more attracted to logs than blank controls. It was also more attracted to logs colonized with the yeast Pichia scolyti than uncolonized logs, but attraction was less consistent. An unidentified dolichopodid was more attracted to logs colonized with L pini, O. ips, and the bacteria Burkholderia sp., than to uncolonized logs, It was also attracted to uncolonized logs. Its responses were less consistent and pronounced than H. unica. These results suggest some parasitoids and dipteran predators exploit microbial symbionts of bark beetles to locate hosts. Overall, specialists showed strong attraction to fungal cues, whereas generalists were more attracted by plant volatiles. These results also show how microbial symbionts can have conflicting effects on host fitness},
	keywords = {Ips pini},
	isbn = {0046-225X},
	author = {Boone, Celia K. and Six, Diana L. and Zheng, Yanbing and Raffa, Kenneth F.}
}
@article {8936,
	title = {Gene genealogies reveal cryptic species and host preferences for the pine fungal pathogen Grosmannia clavigera},
	journal = {Molecular Ecology},
	volume = {20},
	number = {12},
	year = {2011},
	month = {Jun},
	pages = {2581-2602},
	abstract = {Grosmannia clavigera is a fungal pathogen of pine forests in western North America and a symbiotic associate of two sister bark beetles: Dendroctonus ponderosae and D. jeffreyi. This fungus and its beetle associate D. ponderosae are expanding in large epidemics in western North America. Using the fungal genome sequence and gene annotations, we assessed whether fungal isolates from the two beetles inhabiting different species of pine in epidemic regions of western Canada and the USA, as well as in localized populations outside of the current epidemic, represent different genetic lineages. We characterized nucleotide variations in 67 genomic regions and selected 15 for the phylogenetic analysis. Using concordance of gene genealogies and distinct ecological characteristics, we identified two sibling phylogenetic species: Gc and Gs. Where the closely related Pinus ponderosa and P. jeffreyi are infested by localized populations of their respective beetles, Gc is present. In contrast, Gs is an exclusive associate of D. ponderosae mainly present on its primary host-tree P. contorta; however, in the current epidemic areas, it is also found in other pine species. These results suggest that the host-tree species and the beetle population dynamics may be important factors associated with the genetic divergence and diversity of fungal partners in the beetle-tree ecosystems. Gc represents the original G. clavigera holotype, and Gs should be described as a new species.},
	keywords = {Dendroctonus jeffreyi, Dendroctonus ponderosae},
	isbn = {0962-1083},
	author = {Alamouti, Sepideh M. and Wang, Vincent and DiGuistini, Scott and Six, Diana L. and Bohlmann, Joerg and Hamelin, Richard C. and Feau, Nicolas and Breuil, Colette}
}
