Symbioses among bark beetles and their fungal and mite associates involve c
omplex, multi-level interactions. Dendroctonus frontalis attacks and kills
southern pines, introducing fungi into the tree. Ophiostoma minus, may init
ially aid beetles in killing trees, but later this "bluestain" fungus becom
es an antagonist, competing with larvae for host phloem. Two additional fun
gi, Entomocorticium sp. A and Ceratocystiopsis ranaculosus are carried with
in a specialized mycangium and inoculated into phloem where they are fed up
on by beetle larvae. The beetle also vectors several species of phoretic mi
tes which transport spores of O. minus and C. ranaculosus and can complete
their life cycle upon these two fungi. A similar interaction occurs between
Ips avulsus and its fungal (Entomocorticium spp.) and mite (Elattoma benne
tti) symbionts. Larval beetles feed on fungus as do the mites. Female mites
feed on fungus growing within beetle galleries, swelling to many times the
ir normal size. Larval mites develop and mate within the females who burst
open, giving birth to reproductively mature females. These relationships ma
y be best understood by considering the manner in which symbioses vary with
time and resources, and the degree to which other species may affect inter
actions among symbionts.