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.

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Paleoecology can provide valuable insights into the ecology of species that complement observation and experiment‐based assessments of climate impact dynamics. New paleoecological records (e.g., pollen, macrofossils) from the Italian Peninsula suggest a much wider climatic niche of the important European tree species Abies alba (silver fir) than observed in its present spatial range. To explore this discrepancy between current and past distribution of the species, we analyzed climatic data (temperature, precipitation, frost, humidity, sunshine) and vegetation‐independent paleoclimatic reconstructions (e.g., lake levels, chironomids) and use global coupled carbon‐cycle climate (NCAR CSM1.4) and dynamic vegetation (LandClim) modeling. The combined evidence suggests that during the mid‐Holocene (∼6000 years ago), prior to humanization of vegetation, A. alba formed forests under conditions that exceeded the modern (1961–1990) upper temperature limit of the species by ∼5–7°C (July means). Annual precipitation during this natural period was comparable to today (>700–800 mm), with drier summers and wetter winters. In the meso‐Mediterranean to sub‐Mediterranean forests A. alba co‐occurred with thermophilous taxa such as Quercus ilex, Q. pubescens, Olea europaea, Phillyrea, Arbutus, Cistus, Tilia, Ulmus, Acer, Hedera helix, Ilex aquifolium, Taxus, and Vitis. Results from the last interglacial (ca. 130 000–115 000 BP), when human impact was negligible, corroborate the Holocene evidence. Thermophilous Mediterranean A. alba stands became extinct during the last 5000 years when land‐use pressure and specifically excessive anthropogenic fire and browsing disturbance increased. Our results imply that the ecology of this key European tree species is not yet well understood. On the basis of the reconstructed realized climatic niche of the species, we anticipate that the future geographic range of A. alba may not contract regardless of migration success, even if climate should become significantly warmer than today with summer temperatures increasing by up to 5–7°C, as long as precipitation does not fall below 700–800 mm/yr, and anthropogenic disturbance (e.g., fire, browsing) does not become excessive. Our finding contradicts recent studies that projected range contractions under global‐warming scenarios, but did not factor how millennia of human impacts reduced the realized climatic niche of A. alba.

Few examples of natural forest remain near the Mediterranean coast. Therefore, it is difficult to study how coastal forests respond to climatic change or their resilience to human impact. We developed new sedimentary record of Holocene vegetation and fire history at Lago Preola, a coastal lake in southwestern Sicily (Italy). In order to verify the existence of forest at large scale on the coast, we compare pollen from Lago Preola, a medium-sized lake (33 ha), to Gorgo Basso, a small lake (3 ha) located nearby with the aim of separating local from extra-local vegetation dynamics through time using pollen percentages and influx. We then compare Lago Preola pollen to the record from Biviere di Gela, a large lagoon (120 ha) situated 160 km to the east in southern Sicily, to examine differences in vegetation dynamics between the two coastal areas during the Holocene. Lake-level reconstructions and ostracode analyses from Lago Preola provide vegetation-independent evidence of climate change, and help to disentangle human and climatic impacts on vegetation. Pollen data indicate Pistacia-dominated shrublands replaced open grasslands in the region surrounding Lago Preola by 9500 cal yr BP. This change coincided with rising lake levels and the development of an ostracode fauna typical of fresh waters. Evergreen forest dominated by Quercus ilex and Olea europaea started to expand by 7000 cal BP and consolidated at 6500 cal yr BP, when lake levels were near their Holocene high. Similarities between pollen from Lago Preola and Gorgo Basso demonstrate that forest was the dominant vegetation type in coastal Sicily during the middle Holocene at both regional and local scales, and even developed in the drier climatic setting around Biviere di Gela. Lake levels fell at Lago Preola after 7000 cal yr BP, with a strong decline accompanied by increasing salinity after 4500 cal yr BP. However, no transition in vegetation matched these inferred hydrological changes. Instead, forests persisted in the surrounding region until 2200 cal BP when human disturbance intensified. We propose that different climatic factors control lake levels and vegetation in coastal Mediterranean ecosystems. Whereas lake levels are most sensitive to the abundance of winter precipitation, coastal forests depend on spring precipitation and are limited by the length of summer drought. Moisture availability remained suitable for evergreen forests in coastal Sicily during the late Holocene, and humans, not a drier climate drove the regional forest decline.

Aim Forest communities in the European Central Alps are highly sensitive to climatic change. Palaeobotanical studies have demonstrated that forests rapidly expanded upslope during Holocene warm intervals and contracted when temperatures fell. However, temperature alone cannot account for important changes in tree species abundance. For example, population expansion by Norway spruce (Picea abies), a dominant subalpine species, lagged suitable temperatures by about 3000 years in eastern and by 6000 years in western Switzerland. We hypothesize that spruce expansion was delayed by limited water availability in weakly developed soils and/or by drier-than-present climatic conditions.Location We examine the impact of reduced moisture availability on forest dynamics using a combined dynamic modelling/palaeoecological approach at two high-elevational lakes in the Swiss Central Alps.Methods We simulate Holocene vegetation dynamics with the LandClim model in landscapes surrounding the two lakes and validate the model output by comparison with palaeobotanical reconstructions from the same sites. We evaluate the impact of shallow soils on vegetation dynamics at these sites by varying soil water-holding capacity (i.e. bucket size) and precipitation abundance in model scenarios.Results Simulations with modern soil conditions and precipitation abundance matched reconstructed vegetation dynamics near the tree line, where temperature limits plant growth, but simulated abundant spruce during the entire Holocene. Spruce was absent only in simulations with a maximum bucket size of less than 7 cm, or when precipitation was reduced by at least 60%. In exploratory simulations of future conditions with average temperatures raised by 4°C, the low water-holding capacity of shallow alpine soils, not temperature, determined the upper elevational limit of spruce.Main conclusions Spruce expanded in the Central Alps only after soils developed sufficient water-holding capacity and precipitation neared its modern abundance. Soil development will probably constrain the future response of tree species to warmer conditions (e.g. upslope migrations), as it did in the past.

Forests near the Mediterranean coast have been shaped by millennia of human disturbance. Consequently, ecological studies relying on modern observations or historical records may have difficulty assessing natural vegetation dynamics under current and future climate. We combined a sedimentary pollen record from Lago di Massacciucoli, Tuscany, Italy with simulations from the LANDCLIM dynamic vegetation model to determine what vegetation preceded intense human disturbance, how past changes in vegetation relate to fire and browsing, and the potential of an extinct vegetation type under present climate. We simulated vegetation dynamics near Lago di Massaciucoli for the last 7,000 years using a local chironomid-inferred temperature reconstruction with combinations of three fire regimes (small infrequent, large infrequent, small frequent) and three browsing intensities (no browsing, light browsing, and moderate browsing), and compared model output to pollen data. Simulations with low disturbance support polleninferred evidence for a mixed forest dominated by Quercus ilex (a Mediterranean species) and Abies alba (a montane species). Whereas pollen data record the collapse of A. alba after 6000 cal yr BP, simulated populations expanded with declining summer temperatures during the late Holocene. Simulations with increased fire and browsing are consistent with evidence for expansion by deciduous species after A. alba collapsed. According to our combined paleoenvironmental and modeling evidence, mixed Q. ilex and A. alba forests remain possible with current climate and limited disturbance, and provide a viable management objective for ecosystems near the Mediterranean coast and in regions that are expected to experience a mediterranean-type climate in the future.

Little is known about the vegetation and fire history of Sardinia, and especially the long-term history of the thermo-Mediterranean belt that encompasses its entire coastal lowlands. A new sedimentary record from a coastal lake based on pollen, spores, macrofossils and microscopic charcoal analysis is used to reconstruct the vegetation and fire history in north-eastern Sardinia. During the mid-Holocene (c. 8,100-5,300 cal BP), the vegetation around Stagno di Sa Curcurica was characterised by dense Erica scoparia and E. arborea stands, which were favoured by high fire activity. Fire incidence declined and evergreen broadleaved forests of Quercus ilex expanded at the beginning of the late Holocene. We relate the observed vegetation and fire dynamics to climatic change, specifically moister and cooler summers and drier and milder winters after 5,300 cal BP. Agricultural activities occurred since the Neolithic and intensified after c. 7,000 cal BP. Around 2,750 cal BP, a further decline of fire incidence and Erica communities occurred, while Quercus ilex expanded and open-land communities became more abundant. This vegetation shift coincided with the historically documented beginning of Phoenician period, which was followed by Punic and Roman civilizations in Sardinia. The vegetational change at around 2,750 cal BP was possibly advantaged by a further shift to moister and cooler summers and drier and milder winters. Triggers for climate changes at 5,300 and 2,750 cal BP may have been gradual, orbitallyinduced changes in summer and winter insolation, as well as centennial-scale atmospheric reorganizations. Open evergreen broadleaved forests persisted until the twentieth century, when they were partly substituted by widespread artificial pine plantations. Our results imply that highly flammable Erica vegetation, as reconstructed for the mid-Holocene, could re-emerge as a dominant vegetation type due to increasing drought and fire, as anticipated under global change conditions.

The Mediterranean Basin is the region of Europe most vulnerable to negative climate-change impacts, including forest decline, increased wildfire, and biodiversity loss. Because humans have affected Mediterranean ecosystems for millennia, it is unclear whether the region's native ecosystems were more resilient to climate change than current ecosystems, and whether they would provide sustainable management options if restored. We simulated vegetation with the LANDCLIM model, using present-day climate as well as future climate-change scenarios, in three representative areas that encompass a broad range of Mediterranean conditions and vegetation types. Sedimentary pollen records that document now-extinct forests help to validate the simulations. Forests modeled under present climate closely resemble the extinct forests when human disturbance is limited; under future scenarios, characterized by increased temperatures and decreased precipitation, extinct forests are projected to re-emerge. When combined with modeling, paleoecological evidence reveals the potential of native vegetation to re-establish under current and future climate conditions, and provides a template for novel management strategies to maintain forest productivity and biodiversity in a warmer and drier future