Early Wisconsinan (MIS 4) Arctic ground squirrel middens and a squirrel-eye-view of the mammoth-steppe

Fossil arctic ground squirrel (Spermophilus parryii) middens were recovered from ice-rich loess sediments in association with Sheep Creek-Klondike and Dominion Creek tephras (ca 80 ka) exposed in westcentral Yukon. These middens provide plant and insect macrofossil evidence for a steppe-tundra ecosystem during the Early Wisconsinan (MIS 4) glacial interval. Midden plant and insect macrofossil
data are compared with those previously published for Late Wisconsinan middens dating to ca 25-29 14C ka BP (MIS 3/2) from the region. Although multivariate statistical comparisons suggest differences between the relative abundances of plant macrofossils, the co-occurrence of steppe-tundra
plants and insects (e.g., Elymus trachycaulus, Kobresia myosuroides, Artemisia frigida, Phlox hoodii, Connatichela artemisiae) provides evidence for successive reestablishment of the zonal steppe-tundra habitats during cold stages of the Late Pleistocene. Arctic ground squirrels were well adapted to the cold, arid climates, steppe-tundra vegetation and well-drained loessal soils that characterize cold stages of Late Pleistocene Beringia. These glacial conditions enabled arctic ground squirrel populations to expand their range to the interior regions of Alaska and Yukon, including the Klondike, where they are absent
today. Arctic ground squirrels have endured numerous Quaternary climate oscillations by retracting populations to disjunct “interglacial refugia” during warm interglacial periods (e.g., south-facing steppe slopes, well-drained arctic and alpine tundra areas) and expanding their distribution across the mammoth-steppe biome during cold, arid glacial intervals.