Abstract. We examined direct and indirect impacts of millennial-scale climate change on fire regimes in the south-central Brooks Range, Alaska, USA, using four lake sediment records and existing paleoclimate interpretations. New techniques were introduced to identify charcoal peaks semi-objectively and to detect statistical differences between fire regimes. Peaks in charcoal accumulation rates provided estimates of fire return intervals (FRIs), which were compared among vegetation zones identified by fossil pollen and stomata. Climatic warming between ca. 15 000-9000 yr BP (calendar years before Common Era [CE] 1950) coincided with shifts in vegetation from herb tundra to shrub tundra to deciduous woodlands, all novel species assemblages relative to modern vegetation. Two sites cover this period and show decreased FRIs with the transition from herb to Betula-dominated shrub tundra ca. 13 300-14 300 yr BP (FRI mean ¼ 144 yr; 95% CI ¼ 120-169 yr), when climate warmed but remained cooler than present. Although warming would have favored shorter FRIs in the shrub tundra, the shift to more continuous, flammable fuels relative to herb tundra was probably a more important cause of increased burning. Similarly, a vegetation shift to Populus-dominated deciduous woodlands overrode the influence of warmer-and drier-than-present summers, resulting in lower fire activity from ca. 10 300-8250 yr BP (FRI mean ¼ 251 yr; 95% CI ¼ 156-347 yr). Three sites record the mid-to-late Holocene, when climatic cooling and moistening allowed Picea glauca forest-tundra and P. mariana boreal forests to establish ca. 8000 and 5500 yr BP, respectively. FRIs in forest-tundra were either similar to or shorter than those in the deciduous woodlands (FRI mean range ¼ 131-238 yr). The addition of P. mariana ca. 5500 yr BP increased landscape flammability, overrode the effects of climatic cooling and moistening and resulted in lower FRIs (FRI mean ¼ 145 yr; 95% CI ¼ 130-163). Overall, shifts in fire regimes were strongly linked to changes in vegetation, which were responding to millennial-scale climate change. We conclude that shifts in vegetation can amplify or override the direct influence of climate change on fire regimes, when vegetation shifts significantly modify landscape flammability. Our findings emphasize the importance of biophysical feedbacks between climate, fire, and vegetation in determining the response of ecosystems to past, and by inference, future climate change.

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

Contents 38I.38II.Approaches for reconstructing refugia: strengths, limitations and recent advances39III.46IV.47V.48VI.4949References49 Summary Climate refugia, locations where taxa survive periods of regionally adverse climate, are thought to be critical for maintaining biodiversity through the glacial–interglacial climate changes of the Quaternary. A critical research need is to better integrate and reconcile the three major lines of evidence used to infer the existence of past refugia – fossil records, species distribution models and phylogeographic surveys – in order to characterize the complex spatiotemporal trajectories of species and populations in and out of refugia. Here we review the complementary strengths, limitations and new advances for these three approaches. We provide case studies to illustrate their combined application, and point the way towards new opportunities for synthesizing these disparate lines of evidence. Case studies with European beech, Qinghai spruce and Douglas‐fir illustrate how the combination of these three approaches successfully resolves complex species histories not attainable from any one approach. Promising new statistical techniques can capitalize on the strengths of each method and provide a robust quantitative reconstruction of species history. Studying past refugia can help identify contemporary refugia and clarify their conservation significance, in particular by elucidating the fine‐scale processes and the particular geographic locations that buffer species against rapidly changing climate.

Wildfire activity in boreal forests is anticipated to increase dramatically, with far-reaching ecological and socioeconomic consequences. Paleorecords are indispensible for elucidating boreal fire regime dynamics under changing climate, because fire return intervals and successional cycles in these ecosystems occur over decadal to centennial timescales. We present charcoal records from 14 lakes in the Yukon Flats of interior Alaska, one of the most flammable ecoregions of the boreal forest biome, to infer causes and consequences of fire regime change over the past 10,000 y. Strong correspondence between charcoal-inferred and observational fire records shows the fidelity of sedimentary charcoal records as archives of past fire regimes. Fire frequency and area burned increased ∼6,000–3,000 y ago, probably as a result of elevated landscape flammability associated with increased Picea mariana in the regional vegetation. During the Medieval Climate Anomaly (MCA; ∼1,000–500 cal B.P.), the period most similar to recent decades, warm and dry climatic conditions resulted in peak biomass burning, but severe fires favored less-flammable deciduous vegetation, such that fire frequency remained relatively stationary. These results suggest that boreal forests can sustain high-severity fire regimes for centuries under warm and dry conditions, with vegetation feedbacks modulating climate–fire linkages. The apparent limit to MCA burning has been surpassed by the regional fire regime of recent decades, which is characterized by exceptionally high fire frequency and biomass burning. This extreme combination suggests a transition to a unique regime of unprecedented fire activity. However, vegetation dynamics similar to feedbacks that occurred during the MCA may stabilize the fire regime, despite additional warming.

Forest fire occurrence is affected by multiple controls that operate at local to regional scales. At the spatial scale of forest stands, regional climatic controls may be obscured by local controls (e.g., stochastic ignitions, topography, and fuel loads), but the long-term role of such local controls is poorly understood. We report here stand-scale (<100 ha) fire histories of the past 5000 years based on the analysis of sediment charcoal at two lakes 11 km apart in southeastern British Columbia. The two lakes are today located in similar subalpine forests, and they likely have experienced the same late-Holocene climatic changes because of their close proximity. We evaluated two independent properties of fire history: (1) fire-interval distribution, a measure of the overall incidence of fire, and (2) fire synchroneity, a measure of the co-occurrence of fire (here, assessed at centennial to millennial time scales due to the resolution of sediment records). Fire-interval distributions differed between the sites prior to, but not after, 2500 yr before present. When the entire 5000-yr period is considered, no statistical synchrony between fire-episode dates existed between the two sites at any temporal scale, but for the last 2500 yr marginal levels of synchrony occurred at centennial scales. Each individual fire record exhibited little coherency with regional climate changes. In contrast, variations in the composite record (average of both sites) matched variations in climate evidenced by late-Holocene glacial advances. This was probably due to the increased sample size and spatial extent represented by the composite record (up to 200 ha) plus increased regional climatic variability over the last several millennia, which may have partially overridden local, non-climatic controls. We conclude that (1) over past millennia, neighboring stands with similar modern conditions may have experienced different fire intervals and asynchronous patterns in fire episodes, likely because local controls outweighed the synchronizing effect of climate; (2) the influence of climate on fire occurrence is more strongly expressed when climatic variability is relatively great; and (3) multiple records from a region are essential if climate-fire relations are to be reliably described.