Forest expansion and open vegetation responses during the past 14 ka at Zhagaer Co on the eastern Tibetan Plateau

Forest expansion and retreat are key drivers of alpine ecosystem dynamics, yet it remains unclear how forest expansion shapes alpine plant communities and whether alpine assemblages that re-emerge after forest decline resemble those of the Late Glacial. Here, we integrate sedimentary ancient DNA (sedaDNA) and pollen records from Zhagaer Co on the eastern Tibetan Plateau to reconstruct vegetation changes over the past [~]14 ka. Using Picea abundance as a proxy for forest cover, we apply constrained ordination of sedaDNA-inferred plant communities to identify plant taxa as "winners" and "losers" of forest expansion and compare the composition of loser taxa between the late Glacial and late Holocene cold-open phases. Our results show that forest expansion during the early to mid-Holocene favoured woody and forest-margin taxa (e.g. Rhododendron, Salicaceae), while suppressing alpine forbs and graminoids (e.g. Carex, Thalictrum), consistent with patterns expected under ecological filtering. However, late Holocene reopening did not fully restore a late Glacial-like alpine community; instead, it was characterised by a stronger contribution of alpine meadow and shrub taxa. This difference may reflect contrasting environmental backgrounds, including higher atmospheric CO2 levels during the late Holocene, ecological legacies of prior forest expansion, and increasing human influence. These findings suggest that forest expansion may lead to long-term restructuring of alpine plant assemblages, and that late Holocene cooling did not simply restore late Glacial alpine communities but instead produced a distinct alpine ecosystem state. Together, these results highlight the long-term legacy of treeline dynamics in shaping alpine ecosystem trajectories.