Integrating models to investigate critical phenological overlaps in complex ecological interactions: the mountain pine beetle-fungus symbiosis.

The fates of individual species are often tied to synchronization of phenology, however, few methods have been developed for integrating phenological models involving linked species. In this paper, we focus on mountain pine beetle (MPB, Dendroctonus ponderosae) and its two obligate mutualistic fungi, Grosmannia clavigera and Ophiostoma montium. Growth rates of all three partners are driven by temperature, and their idiosyncratic responses affect interactions at important life stage junctures. One critical phase for MPB-fungus symbiosis occurs just before dispersal of teneral (new) adult beetles, when fungi are acquired and transported in specialized structures (mycangia). Before dispersal, fungi must capture sufficient spatial resources within the tree to ensure contact with teneral adults and get packed into mycangia. Mycangial packing occurs at an unknown time during teneral feeding. We adapt thermal models predicting fungal growth and beetle development to predict overlap between the competing fungi and MPB teneral adult feeding windows and emergence. We consider a spectrum of mycangial packing strategies and describe them in terms of explicit functions with unknown parameters. Rates of growth are fixed by laboratory data, the unknown parameters describing various packing strategies, as well as the degree to which mycangial growth is slowed in woody tissues as compared to agar, are determined by maximum likelihood and two years of field observations. At the field location used, the most likely fungus acquisition strategy for MPB was packing mycangia just prior to emergence. Estimated model parameters suggested large differences in the relative growth rates of the two fungi in trees at the study site, with the most likely model estimating that G. clavigera grew approximately twenty-five times faster than O. montium under the bark, which is completely unexpected in comparison with observed fungal growth on agar.