Late Pleistocene shrub expansion preceded megafauna turnover and extinctions in eastern Beringia

The collapse of the steppe-tundra biome (mammoth steppe) at the end of the Pleistocene is used as an important example of topdown ecosystem cascades, where human hunting of keystone species led to profound changes in vegetation across high latitudes in the Northern Hemisphere. Alternatively, it is argued that this biome transformation occurred through a bottom-up process, where climate-driven expansion of shrub tundra (Betula, Salix spp.) replaced the steppe-tundra vegetation that grazing megafauna taxa relied on. In eastern Beringia, these differing hypotheses remain largely untested, in part because the precise timing and spatial pattern of Late Pleistocene shrub expansion remains poorly resolved. This uncertainty is caused by chronological ambiguity in many lake sediment records, which typically rely on radiocarbon (14C) dates from bulk sediment or aquatic macrofossils-materials that are known to overestimate the age of sediment layers. Here, we reexamine Late Pleistocene pollen records for which 14C dating of terrestrial macrofossils is available and augment these data with 14C dates from arctic ground-squirrel middens and plant macrofossils. Comparing these paleovegetation data with a database of published 14C dates from megafauna remains, we find the postglacial expansion of shrub tundra preceded the regional extinctions of horse (Equus spp.) and mammoth (Mammuthus primigenius) and began during a period when the frequency of 14C dates indicates large grazers were abundant. These results are not consistent with a model of top-down ecosystem cascades and support the hypothesis that climate-driven habitat loss preceded and contributed to turnover in mammal communities.