ABSTRACT
Fungal and mite associates may drive changes in bark beetle populations, and mechanisms constraining beetle irruptions may be hidden in endemic populations. We characterized common fungi of endemic-level Jeffrey pine beetle (JPB) in western USA and analyzed their dissemination by JPB (maxillae and fecal pellet) and fungivorous mites to identify if endogenous regulation drove the population. We hypothesized that: (1) as in near-endemic mountain pine beetle populations, JPB's mutualistic fungus would either be less abundant in endemic than in non-endemic populations or that another fungus may be more prevalent; (2) JPB primarily transports its mutualistic fungus, while its fungivorous mites primarily transport another fungus, and (3) based on the prevalence of yeasts in bark beetle symbioses, that a mutualistic interaction with blue-stain fungi present in that system may exist. Grosmannia clavigera was the most frequent JPB symbiont; however, the new here reported antagonist, Ophiostoma minus, was second in frequency. As hypothesized, JPB mostly carried its mutualist fungus while another fungus (i.e., antagonistic) was mainly carried by mites, but no fungal transport was obligate. Furthermore, we found a novel mutualistic interaction between the yeast Kuraishia molischiana and G. clavigera which fostered a growth advantage at temperatures associated with beetle colonization.
ABSTRACT
Differences in defensive traits of tree species may predict why some conifers are susceptible to bark beetle-fungal complexes and others are not. A symbiotic fungus (Leptographium abietinum (Peck) M.J. Wingf.) associated with the tree-killing bark beetle (Dendroctonus rufipennis Kirby) is phytopathogenic to host trees and may hasten tree decline during colonization by beetles, but defense responses of mature trees to the fungus have not been experimentally examined. To test the hypothesis that interspecific variation in spruce resistance is explained by defense traits we compared constitutive (bark thickness and constitutive resin ducts) and induced defenses (resin flow, monoterpene composition, concentration, phloem lesion formation and traumatic resin ducts) between two sympatric spruces: Engelmann spruce (Picea engelmannii Parry ex Engelm.-a susceptible host) and blue spruce (Picea pungens Engelm.-a resistant host) in response to fungal inoculation. Four central findings emerged: (i) blue spruce has thicker outer bark and thinner phloem than Engelmann spruce, which may restrict fungal access to phloem and result in less beetle-available resource overall; (ii) both spruce species induce monoterpenes in response to inoculation but blue spruce has higher constitutive monoterpene levels, induces monoterpenes more rapidly, and induces higher concentrations over a period of time consistent with spruce beetle attack duration; (iii) Engelmann and blue spruce differed in the monoterpenes they upregulated in response to fungal inoculation: blue spruce upregulated α-pinene, terpinolene and γ-terpinene, but Engelmann spruce upregulated 3-carene and linalool; and (iv) blue spruce has a higher frequency of constitutive resin ducts and produces more traumatic resin ducts in annual growth increments than Engelmann spruce, though Engelmann spruce produces more resin following aseptic wounding or fungal inoculation. These findings suggest that higher constitutive resin duct densities and monoterpene concentrations, as well as the ability to rapidly induce specific monoterpenes in response to L. abietinum inoculation, are phenotypic traits associated with hosts resistant to spruce beetle colonization.
