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Subfossil pollen and plant macrofossil data derived from 14 C-dated sediment profiles can provide quantitative information on glacial and interglacial climates. The data allow climate variables related to growingseason warmth, winter cold, and plant-available moisture to be reconstructed. 123Clim Dyn (2011) 37:775-802 DOI 10.1007 surface-pollen assemblages are shown to be accurate and unbiased. Reconstructed LGM and MH climate anomaly patterns are coherent, consistent between variables, and robust with respect to the choice of technique. They support a conceptual model of the controls of Late Quaternary climate change whereby the first-order effects of orbital variations and greenhouse forcing on the seasonal cycle of temperature are predictably modified by responses of the atmospheric circulation and surface energy balance.

The spatio-temporal pattern of peak Holocene warmth (Holocene thermal maximum, HTM) is traced over 140 sites across the Western Hemisphere of the Arctic (0-180 W; north of B60 N). Paleoclimate inferences based on a wide variety of proxy indicators provide clear evidence for warmer-than-present conditions at 120 of these sites. At the 16 terrestrial sites where quantitative estimates have been obtained, local HTM temperatures (primarily summer estimates) were on average 1.670.8 C higher than present (approximate average of the 20th century), but the warming was time-transgressive across the western Arctic. As the precession-driven summer insolation anomaly peaked 12-10 ka (thousands of calendar years ago), warming was concentrated in northwest North America, while cool conditions lingered in the northeast. Alaska and northwest Canada experienced the HTM between ca 11 and 9 ka, about 4000 yr prior to the HTM in northeast Canada. The delayed warming in Quebec and Labrador was linked to the residual Laurentide Ice Sheet, which chilled the region through its impact on surface energy balance and ocean circulation. The lingering ice also attests to the inherent asymmetry of atmospheric and oceanic circulation that predisposes the region to glaciation and modulates the pattern of climatic change. The spatial asymmetry of warming during the HTM resembles the pattern of warming observed in the Arctic over the last several decades. Although the two warmings are described at different temporal scales, and the HTM was additionally affected by the residual Laurentide ice, the similarities suggest there might be a preferred mode of variability in the atmospheric circulation that generates a recurrent pattern of warming under positive radiative forcing. Unlike the HTM, however, future warming will not be counterbalanced by the cooling effect of a residual North American ice sheet. r ARTICLE IN PRESS

This paper integrates recent efforts to map the distribution of biomes for the late Quaternary with the detailed evidence that plant species have responded individualistically to climate change at millennial timescales. Using a fossil-pollen data set of over 700 sites, we review late-Quaternary vegetation history in northern and eastern North America across levels of ecological organization from individual taxa to biomes, and apply the insights gained from this review to critically examine the biome maps generated from the pollen data. Higher-order features of the vegetation (e.g., plant associations, physiognomy) emerge from individualistic responses of plant taxa to climate change, and different representations of vegetation history reveal different aspects of vegetation dynamics. Vegetation distribution and composition were relatively stable during full-glacial times (21 000-17 000 yr BP) [calendar years] and during the mid-to late Holocene (7000-500 yr BP), but changed rapidly during the late-glacial period and early Holocene (16 000-8 000 yr BP) and after 500 yr BP. Shifts in plant taxon distributions were characterized by individualistic changes in population abundances and ranges and included large east-west shifts in distribution in addition to the northward redistribution of most taxa. Modern associations such as Fagus-Tsuga and Picea-Alnus-Betula date to the early Holocene, whereas other associations common to the late-glacial period (e.g., Picea-Cyperaceae-Fraxinus-Ostrya/ Carpinus) no longer exist. Biomes are dynamic entities that have changed in distribution, composition, and structure over time. The late-Pleistocene suite of biomes is distinct from those that grew during the Holocene. The pollen-based biome reconstructions are able to capture the major features of late-Quaternary vegetation but downplay the magnitude and variety of vegetational responses to climate change by (1) limiting apparent land-cover change to ecotones, (2) masking internal variations in biome composition, and (3) obscuring the range shifts and changes in abundance among individual taxa. The compositional and structural differences between full-glacial and recent biomes of the same type are similar to or greater than the spatial heterogeneity in the composition and structure of present-day biomes. This spatial and temporal heterogeneity allows biome maps to accommodate individualistic behavior among species but masks climatically important variations in taxonomic composition as well as structural differences between modern biomes and their ancient counterparts.