Historical ecology identifies long-term rewilding strategy for conserving Mediterranean mountain forests in south Italy.

In the context of global decline in old-growth forest, historical ecology is a valuable tool to derive insights into vegetation legacies and dynamics and develop new conservation and restoration strategies. In this cross-disciplinary study, we integrate palynology (Lago del Pesce record), history, dendrochronology, and historical and contemporary land cover maps to assess drivers of vegetation change over the last millennium in a Mediterranean mountain forest (Pollino National Park, southern Italy) and discuss implications in conservation ecology. The study site hosts a remnant beech-fir (Fagus sylvatica-Abies alba) mixed forest, a priority habitat for biodiversity conservation in Europe. In the 10th century, the pollen record showed an open environment that was quickly colonized by silver fir when sociopolitical instabilities reduced anthropogenic pressures in mountain forests. The highest forest cover and biomass was reached between the 14th and the 17th centuries following land abandonment due to recurring plague pandemics. This rewilding process is also reflected in the recruitment history of Bosnian pine (Pinus heldreichii) in the subalpine elevation belt. Our results show that human impacts have been one of the main drivers of silver fir population contraction in the last centuries in the Mediterranean, and that the removal of direct human pressure led to ecosystem renovation. Since 1910, the Rubbio State Forest has locally protected and restored the mixed beech-fir forest. The institutions in 1972 for the Rubbio Natural Reserve and in 1993 for Pollino National Park have guaranteed the survival of the silver fir population, demonstrating the effectiveness of targeted conservation and restoration policies despite a warming climate. Monitoring silver fir populations can measure the effectiveness of conservation measures. In the last decades, the abandonment of rural environments (rewilding) along the mountains of southern Italy has reduced the pressure on ecosystems, thus boosting forest expansion. However, after four decades of natural regeneration and increasing biomass, pollen influx and forest composition are still far from the natural attributes of the medieval forest ecosystem. We conclude that long-term forest planning encouraging limited direct human disturbance will lead toward rewilding and renovation of carbon-rich and highly biodiverse Mediterranean old-growth forests, which will be more resistant and resilient to future climate change.

Palli J ，Mensing SA ，Schoolman EM ，Solano F ，Piovesan G ... -  《ECOLOGICAL APPLICATIONS》

The return of tall forests: Reconstructing the canopy resilience of an extensively harvested primary forest in Mediterranean mountains.

Understanding recovery times and mechanisms of ecosystem dynamics towards the old-growth stage is crucial for forest restoration, but still poorly delineated in Mediterranean. Through tree-ring methods, we reconstructed the return of a tall canopy after severe human disturbance in a mixed beech (Fagus sylvatica) and silver fir (Abies alba) forest, located at a mountain site in the southern edge of both species' range (Gariglione, south Italy). The primary forest was extensively harvested between 1930 and 1950, removing up to 91 % of the biomass. Growth histories, climate-growth relationships and time-series of growth dominance in Gariglione were compared with a network of protected mature and old-growth beech forests distributed along a wide elevational gradient in the same region. We found that the renewed tall canopy of Gariglione is mainly composed of remnant trees, which include uncut trees and saplings, and the post-harvesting regeneration mostly represented by fir. Canopy beech trees reached maximum basal area increment (BAI) in the 1970s, 40-50 years after cutting. Then, beech BAI shifted towards negative trends in phase with drying climate (PDSI), while fir maintained a sustained growth until 2000. This growth asynchrony between the two species conferred community stability over the last decades. The network comparison highlighted the common negative impact of summer drought on high-frequency growth signals of beech in south Italy. However, analysis of long-term mean growth trends indicates decreasing BAI limited to Gariglione beech, revealing relevant differences due to site ecology and its interactions with legacy effects of past management in driving growth responses to climate change. Indeed, lowland mature beech forests showed increasing BAI in the last decades, while primary high-mountain forests displayed a remarkably stable low oscillating growth. In all the Mediterranean forests we studied, large and old trees showed a marked growth acclimation despite ongoing climate warming, demonstrating the effectiveness of landscape rewilding.

Baliva M ，Palli J ，Perri F ，Iovino F ，Luzzi G ，Piovesan G ... -  《-》

Long-Term Responses of Mediterranean Mountain Forests to Climate Change, Fire and Human Activities in the Northern Apennines (Italy).

Fagus sylvatica (beech) dominates the montane forests of the Apennines and builds old-growth high-conservation value stands. However, recent severe drought-induced diebacks raise concern on the future persistence of these forests and of Southern European mesophilous woodlands overall, growing at their dry edge. To explore the history of Apennine beech-dominated forests, we draw on the multiproxy paleoecological record from Lago Verdarolo, which includes a robust vegetation-independent temperature reconstruction. Numerical techniques are used to investigate the drivers of long-term Mediterranean mountain forest dynamics. Specifically, we focus on disentangling the ecological factors that caused the shift from high-diversity mixed forests to beech-dominated stands and on assessing the occurrence of legacy effects on present-day forests. Abrupt climate change largely drove vegetation dynamics during the Late Glacial and Early Holocene. Species-rich mixed Abies alba (silver fir) forests dominated about 10,500-5500 years ago, under rather dry and warmer-than-today conditions (+ 1-2 °C) and limited fire occurrence. Cooler and moister summers and increasing fire activity caused declines in several fire-sensitive temperate deciduous trees (for example, Ulmus, Tilia, Fraxinus) and favored the establishment of fir-beech forests around 5500 years ago. Further enhancement of fire activity and farming around 2000 years ago led to local Abies alba extinction and forest impoverishment. We conclude that the currently widespread monospecific Apennine beech forests are the result of multi-millennial land-use intensification superimposed on Late Holocene cooling and moistening. Given their higher drought-tolerance compared to beech stands, reviving ancient species-rich mixed fir forests represents a feasible and 'tested' possibility to adapt forests to climate change.

Morales-Molino C ，Steffen M ，Samartin S ，van Leeuwen JFN ，Hürlimann D ，Vescovi E ，Tinner W ... -  《-》

Climate-human interactions contributed to historical forest recruitment dynamics in Mediterranean subalpine ecosystems.

Long-term tree recruitment dynamics of subalpine forests mainly depend on temperature changes, but little is known about the feedbacks between historical land use and climate. Here, we analyze a southern European, millennium-long dataset of tree recruitment from three high-elevation pine forests located in Mediterranean mountains (Pyrenees, northeastern Spain; Pollino, southern Italy; and Mt. Smolikas, northern Greece). We identify synchronized recruitment peaks in the late 15th and early 16th centuries, following prolonged periods of societal and climate instability. Major European population crises in the 14th and 15th centuries associated with recurrent famines, the Black Death pandemic, and political turmoil are likely to have reduced the deforestation of subalpine environments and caused widespread rewilding. We suggest that a distinct cold phase in the Little Ice Age around 1450 ce could also have accelerated the cessation of grazing pressure, particularly in the Pyrenees, where the demographic crisis was less severe. Most pronounced in the Pyrenees, the enhanced pine recruitment from around 1500-1550 ce coincides with temporarily warmer temperatures associated with a positive phase of the North Atlantic Oscillation. We diagnose that a mixture of human and climate factors has influenced past forest recruitment dynamics in Mediterranean subalpine ecosystems. Our results highlight how complex human-climate interactions shaped forest dynamics during pre-industrial times and provide historical analogies to recent rewilding.

Sangüesa-Barreda G ，Esper J ，Büntgen U ，Camarero JJ ，Di Filippo A ，Baliva M ，Piovesan G ... -  《-》

Mediterranean old-growth forests exhibit resistance to climate warming.

Old-growth mountain forests represent an ideal setting for studying long-term impacts of climate change. We studied the few remnants of old-growth forests located within the Pollino massif (southern Italy) to evaluate how the growth of conspecific young and old trees responded to climate change. We investigated two conifer species (Abies alba and Pinus leucodermis) and two hardwood species (Fagus sylvatica and Quercus cerris). We sampled one stand per species along an altitudinal gradient, ranging from a drought-limited low-elevation hardwood forest to a cold-limited subalpine pine forest. We used a dendrochronological approach to characterize the long-term growth dynamics of old (age > 120 years) versus young (age < 120 years) trees. Younger trees grew faster than their older conspecifics during their juvenile stage, regardless of species. Linear mixed effect models were used to quantify recent growth trends (1950-2015) and responses to climate for old and young trees. Climate sensitivity, expressed as radial growth responses to climate during the last three decades, partially differed between species because high spring temperatures enhanced conifer growth, whereas F. sylvatica growth was negatively affected by warmer spring conditions. Furthermore, tree growth was negatively impacted by summer drought in all species. Climate sensitivity differed between young and old trees, with younger trees tending to be more sensitive in P. leucodermis and A. alba, whereas older F. sylvatica trees were more sensitive. In low-elevation Q. cerris stands, limitation of growth due to drought was not related to tree age, suggesting symmetric water competition. We found evidence for a fast-growth trend in young individuals compared with that in their older conspecifics. Notably, old trees tended to have relatively stable growth rates, showing remarkable resistance to climate warming. These responses to climate change should be recognized when forecasting the future dynamics of old-growth forests for their sustainable management.

Colangelo M ，Camarero JJ ，Gazol A ，Piovesan G ，Borghetti M ，Baliva M ，Gentilesca T ，Rita A ，Schettino A ，Ripullone F ... -  《-》