European vegetation during representative “warm” and “cold” intervals of stage-3 was inferred from pollen analytical data. The inferred vegetation differs in character and spatial pattern from that of both fully glacial and fully interglacial conditions and exhibits contrasts between warm and cold intervals, consistent with other evidence for stage-3 palaeoenvironmental fluctuations. European vegetation thus appears to have been an integral component of millennial environmental fluctuations during stage-3; vegetation responded to this scale of environmental change and through feedback mechanisms may have had effects upon the environment. The pollen-inferred vegetation was compared with vegetation simulated using the BIOME 3.5 vegetation model for climatic conditions simulated using a regional climate model (RegCM2) nested within a coupled global climate and vegetation model (GENESIS-BIOME). Despite some discrepancies in detail, both approaches capture the principal features of the present vegetation of Europe. The simulated vegetation for stage-3 differs markedly from that inferred from pollen analytical data, implying substantial discrepancy between the simulated climate and that actually prevailing. Sensitivity analyses indicate that the simulated climate is too warm and probably has too short a winter season. These discrepancies may reflect incorrect specification of sea surface temperature or sea-ice conditions and may be exacerbated by vegetation–climate feedback in the coupled global model.