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Climate-driven changes in high-elevation forest distribution and reductions in snow and ice cover have major implications for ecosystems and global water security. In the Greater Yellowstone Ecosystem of the Rocky Mountains (United States), recent melting of a high-elevation (3,091 m asl) ice patch exposed a mature stand of whitebark pine (Pinus albicaulis) trees, located ~180 m in elevation above modern treeline, that date to the mid-Holocene (c. 5,950 to 5,440 cal y BP). Here, we used this subfossil wood record to develop tree-ring-based temperature estimates for the upper-elevation climate conditions that resulted in ancient forest establishment and growth and the subsequent regional ice-patch growth and downslope shift of treeline. Results suggest that mid-Holocene forest establishment and growth occurred under warm-season (May-Oct) mean temperatures of 6.2 °C (±0.2 °C), until a multicentury cooling anomaly suppressed temperatures below 5.8 °C, resulting in stand mortality by c. 5,440 y BP. Transient climate model simulations indicate that regional cooling was driven by changes in summer insolation and Northern Hemisphere volcanism. The initial cooling event was followed centuries later (c. 5,100 y BP) by sustained Icelandic volcanic eruptions that forced a centennial-scale 1.0 °C summer cooling anomaly and led to rapid ice-patch growth and preservation of the trees. With recent warming (c. 2000–2020 CE), warm-season temperatures now equal and will soon exceed those of the mid-Holocene period of high treeline. It is likely that perennial ice cover will again disappear from the region, and treeline may expand upslope so long as plant-available moisture and disturbance are not limiting. 

Growing season temperatures play a crucial role in controlling treeline elevation at regional to global scales. However, understanding of treeline dynamics in response to long-term changes in temperature is limited. In this study, we analyze pollen, plant macrofossils, and charcoal preserved in organic layers within a 10,400-year-old ice patch and in sediment from a 6000-year-old wetland located above present-day treeline in the Beartooth Mountains, Wyoming, to explore the relationship between Holocene climate variability and shifts in treeline elevation. Pollen data indicate a lower-than-present treeline between 9000 and 6200 cal yr BP during the warm, dry summer and cold winter conditions of the early Holocene. Increases in arboreal pollen at 6200 cal yr BP suggest an upslope treeline expansion when summers became cooler and wetter. A possible hiatus in the wetland record at ca. 4200–3000 cal yr BP suggests increased snow and ice cover at high elevations and a lowering of treeline. Treeline position continued to fluctuate with growing season warming and cooling during the late-Holocene. Periods of high fire activity correspond with times of increased woody cover at high elevations. The two records indicate that climate was an important driver of vegetation and treeline change during the Holocene. Early Holocene treeline was governed by moisture limitations, whereas late-Holocene treeline was sensitive to increases in growing season temperatures. Climate projections for the region suggest warmer temperatures could decrease effective growing season moisture at high elevations resulting in a reduction of treeline elevation.