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Abstract Rising temperatures, increasing hydroclimate variability and intensifying disturbance regimes increase the risk of rapid ecosystem conversions. We can leverage multi‐proxy records of past ecosystem transformations to understand their causes and ecosystem vulnerability to rapid change.Prior to Euro‐American settlement, northern Indiana was a mosaic of prairie, oak‐dominated forests/woodlands and beech‐dominated hardwood forests. This heterogeneity, combined with well‐documented but poorly understood past beech population declines, make this region ideal for studying the drivers of ecosystem transformations.Here, we present a new record from Story Lake, IN, with proxies for vegetation composition (pollen), fire (charcoal) and beech intrinsic water use efficiency (δ¹³C of beech pollen; δ¹³C_beech). Multiple proxies from the same core enable clear establishment of lead–lag relationships. Additionally, δ¹³C_beechenables direct comparisons between beech population abundance and physiological responses to changing environments. We compare Story Lake to a nearby lake‐level reconstruction and to pollen records from nearby Pretty and Appleman Lakes and the distal Spicer Lake, to test hypotheses about synchrony and the spatial scale of governing processes.The 11.7 ka sediment record from Story Lake indicates multiple conversions between beech‐hardwood forest and oak forest/woodland. Beech pollen abundances rapidly increased between 7.5 and 7.1 ka, while oak declined. Oak abundances increased after 4.6 ka and remained high until 2.8 ka, indicating replacement of mesic forests by oak forest/woodland. At 2.8 ka, beech abundances rapidly increased, indicating mesic forest reestablishment. Beech and oak abundances correlate with charcoal accumulation rates but beech abundance is not correlated with δ¹³C_beech.Fluctuations in beech abundances are synchronous among Story, Appleman and Pretty Lakes, but asynchronous between Story and Spicer Lakes, suggesting regulation by local‐scale vegetation‐fire‐climate feedbacks and secondarily by regional‐scale drivers.Holocene forest composition and fire dynamics appear to be closely co‐regulated and may be affected by local to regional climate variations. The importance of extrinsic drivers and positive/negative feedbacks changes over time, with higher ecoclimate sensitivity before 2.8 ka and greater resilience afterwards.Synthesis: Overall, oak‐ and beech‐dominated ecosystems were highly dynamic over the Holocene, with multiple ecosystem conversions driven by shifting interactions among vegetation, hydroclimate and fire regime.