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

Although it is generally agreed that the Arctic flora is among the youngest and least diverse on Earth, the processes that shaped it are poorly understood. Here we present 50 thousand years (kyr) of Arctic vegetation history, derived from the first large-scale ancient DNA metabarcoding study of circumpolar plant diversity. For this interval we also explore nematode diversity as a proxy for modelling vegetation cover and soil quality, and diets of herbivorous megafaunal mammals, many of which became extinct around 10 kyr bp (before present). For much of the period investigated, Arctic vegetation consisted of dry steppe-tundra dominated by forbs (non-graminoid herbaceous vascular plants). During the Last Glacial Maximum (25-15 kyr bp), diversity declined markedly, although forbs remained dominant. Much changed after 10 kyr bp, with the appearance of moist tundra dominated by woody plants and graminoids. Our analyses indicate that both graminoids and forbs would have featured in megafaunal diets. As such, our findings question the predominance of a Late Quaternary graminoid-dominated Arctic mammoth steppe.

Knowledge of historical fire activity tends to be focused at local to landscape scales with few attempts to examine how local patterns of fire activity scale to global patterns. Generally, fire activity varied globally and continuously since the last glacial maximum (LGM) in response to long-term changes in global climate and shorter-term regional changes in climate, vegetation, and human land use. We have synthesised sedimentary charcoal records of biomass burning since the LGM and present global maps showing changes in fire activity for time slices during the past 21,000 years (as differences in charcoal accumulation values compared to pre-industrial). There is strong broad-scale coherence in fire activity after the LGM, but spatial heterogeneity in the signals increases thereafter. In eastern and western North America and western Europe and southern South America, charcoal records indicate less-than-present fire activity from 21,000 to ~11,000 cal yr BP. In contrast, the tropical latitudes of South America and Africa show greaterthan-present fire activity from ~19,000 to ~17,000 cal yr BP whereas most sites from Indochina and Australia show greater-than-present fire activity from 16,000 to ~13,000 cal yr BP. Many sites indicate greater-than-present or near-present activity during the Holocene with the exception of eastern North America and eastern Asia from 8000 to ~2000 cal yr BP, Indonesia from 11,000 to 4000 cal yr BP, and southern South America from 6000 to 3000 cal yr BP where fire activity was less than present. Regional coherence in the patterns of change in fire activity was evident throughout the postglacial period. These complex patterns can be explained in terms of large-scale climate controls modulated by local changes in vegetation and fuel load.

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.

It is commonly believed that trees were absent in Scandinavia during the last glaciation and first recolonized the Scandinavian Peninsula with the retreat of its ice sheet some 9000 years ago. Here, we show the presence of a rare mitochondrial DNA haplotype of spruce that appears unique to Scandinavia and with its highest frequency to the west-an area believed to sustain ice-free refugia during most of the last ice age. We further show the survival of DNA from this haplotype in lake sediments and pollen of Trøndelag in central Norway dating back ~10,300 years and chloroplast DNA of pine and spruce in lake sediments adjacent to the ice-free Andøya refugium in northwestern Norway as early as ~22,000 and 17,700 years ago, respectively. Our findings imply that conifer trees survived in ice-free refugia of Scandinavia during the last glaciation, challenging current views on survival and spread of trees as a response to climate changes.

Ecosystems across the globe are threatened by climate change and human activities. New rapid survey approaches for monitoring biodiversity would greatly advance assessment and understanding of these threats. Taking advantage of next-generation DNA sequencing, we tested an approach we call metabarcoding: high-throughput and simultaneous taxa identification based on a very short (usually <100 base pairs) but informative DNA fragment. Short DNA fragments allow the use of degraded DNA from environmental samples. All analyses included amplification using plant-specific versatile primers, sequencing and estimation of taxonomic diversity. We tested in three steps whether degraded DNA from dead material in soil has the potential of efficiently assessing biodiversity in different biomes. First, soil DNA from eight boreal plant communities located in two different vegetation types (meadow and heath) was amplified. Plant diversity detected from boreal soil was highly consistent with plant taxonomic and growth form diversity estimated from conventional above-ground surveys. Second, we assessed DNA persistence using samples from formerly cultivated soils in temperate environments. We found that the number of crop DNA sequences retrieved strongly varied with years since last cultivation, and crop sequences were absent from nearby, uncultivated plots. Third, we assessed the universal applicability of DNA metabarcoding using soil samples from tropical environments: a large proportion of species and families from the study site were efficiently recovered. The results open unprecedented opportunities for large-scale DNA-based biodiversity studies across a range of taxonomic groups using standardized metabarcoding approaches.

We studied the pollination ecology and assemblage structure of 31 species of Stylidium (Stylidiaceae) at 25 sites in Western Australia. The number of species per study site varied between two and size. Stylidium species are pollinated by a variety of nectar—seeking solitary bees and bombyliid flies. Within and among species there is significant variation in nectar—tube length (and therefore in the insects that visit the flowers) and in pollen placement on pollinators. Pollen is placed “explosively” on the insect by a motile column of fused staminate and pistillate tissues; the position and reach of the column varies within and among species, thereby causing variation in site of pollen deposition. When discrete pollination niches were defined for all species, only one niche overlap was observed across the 86 interacting pairs of Stylidium species at the 25 sites. To determine whether this was a nonrandom assemblage structure we compared our observation with the outcome of null models. We developed three null models to cover the most likely structuring processes: that communities are organized by (1) ecological sorting, (2) evolution of plant phenotypes, or (3) both processes. We concluded that it was unlikely (P = .055—.002) that so few overlaps in pollination niches would occur by chance. We developed another null model to test whether chance could have created the apparent pattern of character displacement in pollination niches exhibited by the nine species showing intraspecific variation. The analysis indicated that character displacement has probably occurred (P = .014). This study is one of the clearest demonstrations to date of reproductive interactions generating assemblage structure and character displacement in plants.

It is widely accepted, based on data from the last few decades and on model simulations, that anthropogenic climate change will cause increased fire activity. However, less attention has been paid to the relationship between abrupt climate changes and heightened fire activity in the paleorecord. We use 35 charcoal and pollen records to assess how fire regimes in North America changed during the last glacial-interglacial transition (15 to 10 ka), a time of large and rapid climate changes. We also test the hypothesis that a comet impact initiated continental-scale wildfires at 12.9 ka; the data do not support this idea, nor are continent-wide fires indicated at any time during deglaciation. There are, however, clear links between large climate changes and fire activity. Biomass burning gradually increased from the glacial period to the beginning of the Younger Dryas. Although there are changes in biomass burning during the Younger Dryas, there is no systematic trend. There is a further increase in biomass burning after the Younger Dryas. Intervals of rapid climate change at 13.9, 13.2, and 11.7 ka are marked by large increases in fire activity. The timing of changes in fire is not coincident with changes in human population density or the timing of the extinction of the megafauna. Although these factors could have contributed to fire-regime changes at individual sites or at specific times, the charcoal data indicate an important role for climate, and particularly rapid climate change, in determining broad-scale levels of fire activity.biomass burning ͉ charcoal ͉ comet ͉ Younger Dryas