Mutualism is not restricted to tree‐killing bark beetles and fungi: the ecological stoichiometry of secondary bark beetles, fungi, and a scavenger

Abstract

1. All bark beetles are in symbiosis with fungi. Although obligate mutualisms with fungi are common with tree‐killing bark beetles (primaries), fungi associated with non‐tree‐killing bark beetles (secondaries) are usually dismissed as commensals.2. Using an ecological stoichiometric approach, we show secondaries are also involved in nutrition‐based mutualisms, some of which appear obligate, and that differences in symbiont provisioning efficiency have a potent effect on beetle carbon (C): nitrogen (N): phosphorus (P) ratios.3. Some secondary beetles have high P contents and require efficient P provisioning via fungi, while others have low P contents that may allow them to exploit less efficient fungi or a broader range of species with variable efficiencies. A co‐occurring scavenger that feeds on nutrient‐poor bark beetle frass (excrement/boring residues) exhibited the lowest phosphorus content yet recorded for an invertebrate.4. Our results generally support the growth‐rate hypothesis that posits differences in C:P and N:P ratios in consumers are due to differential allocation of P to P‐rich RNA to support growth. However, while the beetle species that accumulated the most biomass was considerably enriched in P and that with the least biomass was P‐poor, one beetle species that was P‐rich was also small possibly due to limitation by an element other than P.5. Our results indicate that fungi are important to a broader range of bark beetles than previously recognised. Additional research is needed to describe how these various symbioses influence forest ecosystems via differential effects of fungi on host beetle fitness.