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.

Abstract1. In northern Idaho (USA), more than 100 vascular plant species are disjunct >160 km from their main distribution along the Pacific Northwest coast. It remains unclear whether most species within this interior forest disjunction, including Tsuga mertensiana, survived the last glacial period in a north-Idaho refugium or whether these species colonized the region via long-distance dispersal during the Holocene.2. Sediment cores were extracted from three mid-to high-elevation lakes within T. mertensiana-dominated forests in the Northern Rocky Mountains of Idaho.Pollen and macrofossils were used to reconstruct forest composition, determine the timing of T. mertensiana establishment, examine the hypothesis that the region was a glacial refugium, and contrast how climate, competition and/or dispersal limitation have influenced its modern distribution.3. The modern distribution of T. mertensiana was analysed by constructing a range map and modelling the potential species distribution. The presence of outlier populations surrounding the Idaho disjunction along with broad areas of unoccupied suitable habitat indicates that the range of T. mertensiana is currently expanding. To assess the accuracy of T. mertensiana pollen at detecting its range limit, a network of pollen surface samples was used to analyse the probability of detecting T. mertensiana pollen as a function of distance from its geographical range limit. Consistent T. mertensiana pollen occurrence at ≥1% abundance is likely only within 42 km of its range limit. 4.Tsuga mertensiana first appears in the pollen and macrofossil record at the highestelevation site at c. 4,100 cal year bp, then at the next highest-elevation site at c. 1,600 cal year bp, and last at the mid-elevation site at 800 cal year bp. Tsuga mertensiana pollen occurs continuously at ≥1% at all three sites by c. 300 cal year bp suggesting regional presence. The timing of arrival suggests that T. mertensiana is a recent component of the forests of Idaho, having arrived during the Holocene via long-distance dispersal from coastal populations over 160 km away. Synthesis.Comparison with palaeoclimate reconstructions from the broader region suggests that climate was a greater limiting factor than dispersal in the Holocene establishment in the interior, indicating little difficulty overcoming a large dispersal barrier. However, T. mertensiana remained at low abundances for millennia untilPaper previously published as Standard Paper

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Continuous sediment, charcoal, and pollen records were developed from a ~7-m sediment core from Prater Canyon in Mesa Verde National Park (MEVE), Colorado, USA. Sediment input into the canyon is episodic and is linked to precipitation runoff and vegetation cover. Pollen recovered from the Prater Canyon sediment core reflect the vegetation changes within the MEVE region. During the period recorded, the vegetation of the region surrounding Prater Canyon transitioned from xeric adapted species in an open environment to a more mesic, Pinus edulis–Juniperus osteosperma (piñon–juniper) woodland over the last 1500 years. Two distinct changes in fire frequency occurred. Before 4080 cal. yr BP, fires occurred at a much more frequent rate (2.5–12 fires/200 years) than from 4060 cal. yr BP to present (0–2 fires/200 years). Most importantly, the variations occurring in the charcoal record for the past 2500 years coincide with both shifts in human occupation and climate fluctuations within the region, with burning increasing during Ancestral Puebloan occupation and moist but increasingly dry conditions, and declines in both at the end of the ‘Medieval Climate Anomaly’ (MCA). The record from Prater Canyon demonstrates the importance of the Ancestral Puebloans in landscape modification during their occupation from ad1 to 1300. Charcoal deposition also increased during the 20th- to 21st-century transition with the highest deposition rates of the core recorded then.