Worldwide, large wildfires are becoming increasingly common, leading to economic damages and threatening ecosystems and human health. Under future climate change, more frequent fire disturbance may ...push ecosystems into non‐forested alternative stable states. Fire‐prone ecosystems such as those in the Mediterranean Basin are expected to be particularly vulnerable, but the position of tipping points is unclear.
We compare long‐term palaeoecological data from Sardinia with output from a process‐based dynamic vegetation model to investigate the mechanisms controlling the complex interactions between fire, climate, and vegetation in the past and the future.
Our results show that past vegetation changes from Erica‐shrublands to mixed evergreen‐broadleaved Quercus ilex‐dominated forests were driven by a climate‐induced fire regime shift. By simulating vegetation dynamics under varying fire regimes, we could reproduce Holocene vegetation trajectories and mechanistically identify tipping points.
Without an immediate reduction of greenhouse gas emissions, we simulate future expansion of fire‐prone Mediterranean maquis and increasing fire occurrence. Similarly, high anthropogenic ignition frequencies and plantations of non‐native, highly flammable trees could induce a shift to fire‐adapted Erica shrublands. However, our simulations indicate that if global warming can be kept below 2°C, Quercus ilex forests will be able to persist and effectively reduce fire occurrences and impacts, making them a valuable restoration target in Mediterranean ecosystems.
Synthesis. By combining long‐term records of ecosystem change with a dynamic vegetation model, we show that past climate‐driven fire regime shifts were the main driver of vegetation change, creating alternative stable states that persisted over centuries. Projected future climate change exceeding Holocene variability leads to pronounced vegetation changes and increased fire risks in our simulations, requiring new fire management strategies to maintain current ecosystem services.
Wildfires are increasing in Mediterranean ecosystems. By combining long‐term records of past vegetation and fire with a dynamic vegetation model, the authors show that past fire regime shifts resulted in marked vegetation changes. Future climate change exceeding Holocene variability will push ecosystems into new stable states, increasing fire risks and threatening ecosystem services.
Editor's Choice
Aim
Knowing a species' response to historical climate shifts helps understanding its perspectives under global warming. We infer the hitherto unresolved postglacial history of Pinus cembra. Using ...independent evidence from genetic structure and demographic inference of extant populations, and from palaeoecological findings, we derive putative refugia and re‐colonisation routes.
Location
European Alps and Carpathians.
Taxa
Pinus cembra.
Methods
We genotyped nuclear and chloroplast microsatellite markers in nearly 3000 individuals from 147 locations across the entire natural range of P. cembra. Spatial genetic structure (Bayesian modelling) and demographic history (approximate Bayesian computation) were combined with palaeobotanical records (pollen, macrofossils) to infer putative refugial areas during the Last Glacial Maximum (LGM) and re‐colonisation of the current range.
Results
We found distinct spatial genetic structure, despite low genetic differentiation even between the two disjunct mountain ranges. Nuclear markers revealed five genetic clusters aligned East–West across the range, while chloroplast haplotype distribution suggested nine clusters. Spatially congruent separation at both marker types highlighted two main genetic lineages in the East and West of the range. Demographic inference supported early separation of these lineages dating back to a previous interstadial or interglacial c. 210,000 years ago. Differentiation into five biologically meaningful genetic clusters likely established during postglacial re‐colonisation.
Main Conclusions
Combining genetic and palaeoecological evidence suggests that P. cembra primarily survived the LGM in ‘cold period’ refugia south of the Central European Alps and near the Carpathians, from where it expanded during the Late Glacial into its current Holocene ‘warm period’ refugia. This colonisation history has led to the distinct East–West structure of five genetic clusters. The two main genetic lineages likely derived from ancient divergence during an interglacial or interstadial. The respective contact zone (Brenner line) matches a main biogeographical break in the European Alps also found in herbaceous alpine plant species.
Aim
Pinewood decline and scrubland expansion are major features of Late Holocene vegetation history in the Cantabrian Range. However, the drivers of this remarkable vegetation shift remain to be ...investigated. Here, we aim at disentangling the role of past land use and climate in shaping the high‐elevation Cantabrian landscape during the past two millennia.
Location
Cantabrian Range (northern Iberia).
Taxa
Pinus sylvestris, Betula, Ericaceae, Juniperus, Poaceae.
Methods
We conducted high‐resolution multiproxy palaeoecological analyses (pollen, plant macrofossils, microscopic charcoal and dung fungi) on lake sediments from Lago del Ausente to reconstruct vegetation, fire occurrence and grazing through time. The chronology is based on 14C (terrestrial plant macrofossils) and 210Pb dating, and Bayesian age‐depth modelling (‘rbacon’). We carried out cross‐correlation analysis to quantify vegetation responses to fire.
Results
Between 250 and 900 CE, the vegetation above 1700 m a.s.l. consisted of subalpine scrubland and scattered P. sylvestris trees/stands. Pinewoods with Betula were widespread at slightly lower elevation. This vegetation was resilient to moderate fire disturbance associated with limited pastoral activities. In contrast, enhanced fire occurrence alongside heavier pastoralism led to the demise of pinewoods and their replacement with Betula stands, subalpine scrublands, and meadows between 900 and 1100 CE. Later, the subalpine scrubland‐birch tree line did not respond to Little Ice Age cooling. However, further intensification of transhumant herding between 1300 and 1860 CE (‘La Mesta’) triggered birch decline and the establishment of the modern treeless landscape.
Main conclusions
The extant high‐elevation vegetation of the Cantabrian Range is largely the legacy of intensive land use starting more than one millennium ago. Recurrent and severe fires to promote pasturelands led to the regional extirpation of the previously widespread Pinus sylvestris. Future management should aim at preserving the valuable cultural open landscape of mountain scrubland and meadows and also at restoring patches of ancient pine‐birch woodlands.
Mediterranean rear-edge populations of Betula, located at the southwestern Eurasian margin of the distribution range, represent unique reservoirs of genetic diversity. However, increasing densities ...of wild ungulates, enhanced dryness, and wildfires threaten their future persistence. A historical perspective on the past responses of these relict populations to changing herbivory, fire occurrence and climatic conditions may contribute to assessing their future responses under comparable scenarios. We have reconstructed vegetation and disturbance (grazing, fire) history in the Cabañeros National Park (central-southern Spain) using the paleoecological records of two small mires. We particularly focused on the historical range of variation in disturbance regimes, and the dynamics of rear-edge Betula populations and herbivore densities. Changes in water availability, probably related to the North Atlantic Oscillation (NAO) index, and land-use history have played a crucial role in vegetation shifts. Our data suggest that heathlands (mainly Erica arborea and E. scoparia) and Quercus woodlands dominated during dry phases while Sphagnum bogs and Betula stands expanded during wet periods. Betula populations survived past moderately dry periods but were unable to cope with enhanced land use, particularly increasing livestock raising since ~1,100–900 cal. yr BP (850–1,050 CE), and eventually underwent local extinction. High herbivore densities not only contributed to the Betula demise but also caused the retreat of Sphagnum bogs. Ungulate densities further rose at ~200–100 cal. yr BP (1750–1850 CE) associated with the historically documented intensification of land use around the Ecclesiastical Confiscation. However, herbivory reached truly unprecedented values only during the last decades, following rural depopulation and subsequent promotion of big game hunting. For the first time in temperate and Mediterranean Europe, we have used the abundances of fossil dung fungal spores to assess quantitatively that current high herbivore densities exceed the historical range of variation. In contrast, present fire activity lies within the range of variation of the last millennia, with fires (mainly human-set) mostly occurring during dry periods. Our paleodata highlight the need of controlling the densities of wild ungulates to preserve ecosystem composition and functioning. We also urge to restore Betula populations in suitable habitats where they mostly disappeared because of excessive human activities.
Tree lines are supposed to react sensitively to the current global change. However, the lack of a long‐term (millennial) perspective on tree line shifts in the Pyrenees prevents understanding the ...underlying ecosystem dynamics and processes.
We combine multiproxy palaeoecological analyses (fossil pollen, spores, conifer stomata, plant macrofossils, and ordination) from an outstanding ice cave deposit located in the alpine belt c. 200 m above current tree line (Armeña‐A294 Ice Cave, 2,238 m a.s.l.), to assess for the first time in the Pyrenees, tree line dynamics, and ecosystem resilience to climate changes 5,700–2,200 (cal.) years ago.
The tree line ecotone was located at the cave altitude from 5,700 to 4,650 cal year bp, when vegetation consisted of open Pinus uncinata Ramond ex DC and Betula spp. Woodlands and timberline were very close to the site. Subsequently, tree line slightly raised and timberline reached the ice cave altitude, exceeding its today's uppermost limit by c. 300–400 m during more than four centuries (4,650 and 4,200 cal year bp) at the end of the Holocene Thermal Maximum. After 4,200 cal year bp, alpine tundra communities dominated by Dryas octopetala L. expanded while tree line descended, most likely as a consequence of the Neoglacial cooling. Prehistoric livestock raising likely reinforced climate cooling impacts at 3,450–3,250 cal year bp. Finally, a tree line ecotone developed around the cave that was on its turn replaced by alpine communities during the past 2,000 years.
Synthesis. The long‐term Pyrenean tree line ecotone sensitivity suggests that rising temperatures will trigger future P. uncinata and Betula expansions to higher elevations, replacing arctic–alpine plant species. Climate change is causing the rapid melting of the cave ice; rescue investigations would be urgently needed to exploit its unique ecological information.
We present the first millennial to centennial scale reconstruction of tree line dynamics for the Pyrenees. Our inferences are based on biological proxies (fossil pollen, plant macrofossils, and stomata) found in a unique palaeoenvironmental archive: an ice cave deposit discovered 200 m above current tree line. One main outcome is that subalpine forests were generally more widely distributed during the last 5,700 years and subalpine–alpine vegetation responded sensitively to both climate changes and human activities.
Editor's Choice
1. Pinus nigra Am. forests dominated over extensive areas of the Northern Iberian Plateau (Spain) during the Holocene, but a strong decline during the historical period (c. 1300–700 cal. BP) led to ...the present fragmented populations. This demise has been generally attributed to land-use changes or climate, but the specific roles of disturbance regimes such as fire variability and grazing on the long-term are not fully understood yet. 2. We combine multi-proxy palaeoecological data (fossil pollen, spores, conifer stomata, microscopic and macroscopic charcoal) together with quantitative analyses (ordination and peak detection) from a high-resolution sedimentary sequence (Tubilla del Lago, 900 m a.s.l.) to assess the causes of pine forests demise. A new microscopic charcoal record from an additional sequence (Espinosa de Cerrato, 885 m a.s.l.) is used to assess burning and pine decline at a more regional (100-km radius) scale. 3. Pinus nigra forests could cope with drought and fire regime variability (FRI = 110–500 years), with forest recovery taking c. 100–200 years after fires. Only at 1300–1200 cal. BP a long-lasting irrecoverable demise of P. nigra forests occurred when human-induced fires together with arable and pastoral farming became widespread in the area. Subsequently, Quercus woodlands expanded in the remnant patchy pinewoods. This vegetation shift was primarily caused by three particularly important fire episodes in less than a century (c. 1300–1200 cal. BP). 4. Synthesis. Pinus nigra forests have shown a millennial resilience to the natural fire regime of the Northern Iberian Plateau that was characterized by relatively frequent small-moderate fires and rare high-intensity fires. However, frequent human-caused crown fires and the onset of intensive farming caused their demise over an extensive area. Ongoing land-use abandonment in the Iberian mountains could promote the occurrence of high-intensity, severe fires due to the rapid build-up of high fuel loads. Forest management could mimic the natural fire regime by periodically reducing fuel loads for a transitional period until natural disturbance variability is fully restored, thus preserving these relict native plant communities.
Aim
Temperate forests are currently facing multiple stresses due to climate change, biological invasions, habitat fragmentation and fire regime change. How these stressors interact with each other ...influences how, when and whether ecosystems recover, or whether they adapt or transition to a different ecological state. Because forest recovery or collapse may take longer than a human lifetime, predicting the outcomes of different stressor combinations remains difficult. A clearer vision of future forest trajectories in a changing world may be gained by examining collapses of forests in the past. Here, we use long‐term ecological data to conduct a post‐mortem examination of the decline of maritime pine forests (Pinus pinaster Ait.) on the SW Iberian Peninsula 7000–6500 years ago.
Location
Portugal and Spain.
Methods
We compared four palaeoecological records—two with pine declines and two without—using a multiproxy approach. Bioclimatic differences between the four sites were explored. Proxies for past vegetation and disturbance (fire and grazing) were compared with independent palaeoclimatic records. We performed functional traits analysis and used phase plots to examine the causes of pine decline.
Results
The pine decline represents a critical transition in SW Iberia, which lies close to maritime pine's bioclimatic limits. Prolonged drought likely killed trees and suppressed the fires that normally stimulate pine germination and pinewood recovery. Increased grazing pressure facilitated the rapid spread of resprouter shrubs. These competed with pine trees and ultimately replaced them. Our data highlight complex interactions between climate, fire, grazing and forest resilience.
Main Conclusions
The pine decline occurred at least a century after post‐fire resprouters overtook obligate seeders in the vegetation, constituting an early‐warning signal of forest loss. Fire suppression, resprouter encroachment and grazing may threaten the persistence of Mediterranean forests as droughts become more frequent and extreme.
Aim: Our aim was to discriminate different species of Pinus via pollen analysis in order to assess the responses of particular pine species to orbital and millennial-scale climate changes, ...particularly during the last glacial period. Location: Modern pollen grains were collected from current pine populations along transects from the Pyrenees to southern Iberia and the Balearic Islands. Fossil pine pollen was recovered from the south-western Iberian margin core MD95-2042. Methods: We measured a set of morphological traits of modern pollen from the Iberian pine species Pinus nigra, P. sylvestris, P. halepensis, P. pinea and P. pinaster and of fossil pine pollen from selected samples of the last glacial period and the early to mid-Holocene. Classification and regression tree (CART) analysis was used to establish a model from the modern dataset that discriminates pollen from the different pine species and allows identification of fossil pine pollen at the species level. Results: The CART model was effective in separating pollen of P. nigra and P. sylvestris from that of the Mediterranean pine group (P. halepensis, P. pinea and P. pinaster). The pollen of Pinus nigra diverged from that of P. sylvestris by having a more flattened corpus. Predictions using this model suggested that fossil pine pollen is mainly from P. nigra in all the samples analysed. Pinus sylvestris was more abundant in samples from Greenland stadials than Heinrich stadials, whereas Mediterranean pines increased in samples from Greenland interstadials and during the early to mid-Holocene. Main conclusions: Morphological parameters can be successfully used to increase the taxonomic resolution of fossil pine pollen at the species level for the highland pines (P. nigra and P. sylvestris) and at the group of species level for the Mediterranean pines. Our study indicates that P. nigra was the dominant component of the last glacial south-western/central Iberian pinewoods, although the species composition of these woodlands varied in response to abrupt climate changes.
The reconstruction of human impact is pivotal in palaeoecological studies, as humans are among the most important drivers of Holocene vegetation and ecosystem change. Nevertheless, separating the ...anthropogenic footprint on vegetation dynamics from the impact of climate and other environmental factors (disturbances such as fire, erosion, floods, landslides, avalanches, volcanic eruptions) is a challenging and still largely open issue. For this purpose, palynologists mostly rely on cultural indicator pollen types and related indices that consist of sums or ratios of these pollen types. However, the high environmental and biogeographical specificity of cultural indicator plants hinders the application of the currently available indices to wide geographical settings. Furthermore, the achievable taxonomic resolution of cultural indicator pollen types may hamper their indicative capacity. In this study, we propose the agricultural land use probability (LUP) index, a novel approach to quantify human impact intensity on European ecosystems based on cultural indicator pollen types. From the ‘classic’ cultural indicators, we construct the LUP index by selecting those with the best indicator capacity based on bioindication criteria. We first train the LUP index using twenty palynological sequences along a broad environmental gradient, spanning from treeless alpine to subtropical mediterranean evergreen plant communities. We then validate the LUP index using independent pollen datasets and archaeological proxies. Finally, we discuss the suitability of the selected pollen types and the potential of the LUP index for quantifying Holocene human impact in Europe, concluding that careful application of the LUP index may significantly contribute to refining pollen-based land-use reconstructions.
•We develop and test a new agricultural land use probability (LUP) index.•Anthropogenic indicator values (AIV) allow weighting pollen percentages in the index.•LUP index has high indicative power for all major biomes of Europe.•LUP index shows good correlation with archaeological proxies for human activities.
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