Stable isotopes and radiocarbon assess variable importance of plants and fungi in diets of arctic ground squirrels

Arctic ground squirrels (Urocitellus parryii) rely primarily on dietary protein derived from plants to fuel gluconeogenesis during hibernation, yet fungal sporocarps may be an important, yet overlooked, protein source. Fungivory levels depend on sporocarp productivity, which varies with the dominant plant species and is higher on acidic than on non-acidic soils. To test whether these factors altered fungal consumption, we used stable isotopes to investigate arctic ground squirrel diets at two sites in northern Alaska, Toolik (primarily moist acidic tundra) and Atigun (primarily moist non-acidic tundra). Radiocarbon estimates can also indicate fungivory levels because ectomycorrhizal fungi assimilate soil-derived organic nitrogen whose C-14 levels are higher than current photosynthesis. We measured radiocarbon in hair and delta C-13 and delta N-15 in hair, feces, ectomycorrhizal sporocarps, graminoids, and dicots. Feces were higher in delta C-13 and delta N-15 at Toolik than at Atigun, and fecal delta N-15 increased in August at Toolik, coincident with sporocarp production and fungal spores in feces. Mixing models indicated that graminoids contributed 64%, dicots 35%, and sporocarps 1% to Atigun hair protein, whereas graminoids contributed 37%, dicots 16%, and sporocarps 47% to Toolik hair protein. Acidic soils appeared to correlate with higher sporocarp production and fungivory at Toolik than at Atigun. Atigun hair resembled atmospheric CO2 in C-14, whereas Toolik hair had higher C-14, consistent with greater fungal consumption at Toolik. Late-season sporocarps may be a key protein source for some squirrels and may provide an integrated signal of the soil organic nitrogen assimilated by ectomycorrhizal fungi.