Climate sensitivity of Mediterranean pine growth reveals distinct east-west dipole

The European Mediterranean region is governed by a characteristic climate of summer drought that is likely to increase in duration and intensity under predicted climate change. However, large-scale network analyses investigating spatial aspects of pre-instrumental drought variability for this biogeographic zone are still scarce. In this study we introduce 54 mid- to high-elevation tree-ring width (TRW) chronologies comprising 2186 individual series from pine trees (Pinus spp.). This compilation spans a 4000-km east–west transect from Spain to Turkey, and was subjected to quality control and standardization prior to the development of site chronologies. A principal component analysis (PCA) was applied to identify spatial growth patterns during the network's common period 1862–1976, and new composite TRW chronologies were developed and investigated. The PCA reveals a common variance of 19.7% over the 54 Mediterranean pine chronologies. More interestingly, a dipole pattern in growth variability is found between the western (15% explained variance) and eastern (9.6%) sites, persisting back to 1330 AD. Pine growth on the Iberian Peninsula and Italy favours warm early growing seasons, but summer drought is most critical for ring width formation in the eastern Mediterranean region. Synoptic climate dynamics that have been in operation for the last seven centuries have been identified as the driving mechanism of a distinct east–west dipole in the growth variability of Mediterranean pines.

Contrasting vulnerability and resilience to drought-induced decline of densely planted vs. natural rear-edge Pinus nigra forests

The southernmost European natural and planted pine forests are among the most vulnerable areas to warming-induced drought decline. Both drought stress and management factors (e.g., stand origin or reduced thinning) may induce decline by reducing the water available to trees but their relative importances have not been properly assessed. The role of stand origin - densely planted vs. naturally regenerated stands - as a decline driver can be assessed by comparing the growth and vigor responses to drought of similar natural vs. planted stands. Here, we compare these responses in natural and planted Black pine (Pinus nigra) stands located in southern Spain. We analyze how environmental factors - climatic (temperature and precipitation anomalies) and site conditions - and biotic factors - stand structure (age, tree size, density) and defoliation by the pine processionary moth - drive radial growth and crown condition at stand and tree levels. We also assess the climatic trends in the study area over the last 60 years. We use dendrochronology, linear mixed-effects models of basal area increment and structural equation models to determine how natural and planted stands respond to drought and current competition intensity. We observed that a temperature rise and a decrease in precipitation during the growing period led to increasing drought stress during the late 20th century. Trees from planted stands experienced stronger growth reductions and displayed more severe crown defoliation after severe droughts than those from natural stands. High stand density negatively drove growth and enhanced crown dieback, particularly in planted stands. Also pine processionary moth defoliation was more severe in the growth of natural than in planted stands but affected tree crown condition similarly in both stand types. In response to drought, sharp growth reduction and widespread defoliation of planted Mediterranean pine stands indicate that they are more vulnerable and less resilient to drought stress than natural stands. To mitigate forest decline of planted stands in xeric areas such as the Mediterranean Basin, less dense and more diverse stands should be created through selective thinning or by selecting species or provenances that are more drought tolerant. (C) 2013 Elsevier B.V. All rights reserved.

Pinus nigra (European black pine) as the dominant species of the last glacial pinewoods in south-western to central Iberia: a morphological study of modern and fossil pollen

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.

Are ectomycorrhizal fungi alleviating or aggravating nitrogen limitation of tree growth in boreal forests?

•Symbioses between plant roots and mycorrhizal fungi are thought to enhance plant uptake of nutrients through a favourable exchange for photosynthates. Ectomycorrhizal fungi are considered to play this vital role for trees in nitrogen (N)-limited boreal forests. •We followed symbiotic carbon (C)–N exchange in a large-scale boreal pine forest experiment by tracing 13CO2 absorbed through tree photosynthesis and 15N injected into a soil layer in which ectomycorrhizal fungi dominate the microbial community. •We detected little 15N in tree canopies, but high levels in soil microbes and in mycorrhizal root tips, illustrating effective soil N immobilization, especially in late summer, when tree belowground C allocation was high. Additions of N fertilizer to the soil before labelling shifted the incorporation of 15N from soil microbes and root tips to tree foliage. •These results were tested in a model for C–N exchange between trees and mycorrhizal fungi, suggesting that ectomycorrhizal fungi transfer small fractions of absorbed N to trees under N-limited conditions, but larger fractions if more N is available. We suggest that greater allocation of C from trees to ectomycorrhizal fungi increases N retention in soil mycelium, driving boreal forests towards more severe N limitation at low N supply.

Macrofossils as records of plant responses to rapid Late Glacial climatic changes at three sites in the Swiss Alps

Plant macrofossils from the end of the Younger Dryas were analysed at three sites, Gerzensee (603 m asl), Leysin (1230 m asl), and Zeneggen (1510 m asl). For the first two sites an oxygen-isotope record is also available that was used to develop a time scale (Schwander et al., this volume); dates refer therefore to calibrated years according to the GRIP time scale. Around Gerzensee a pine forest with some tree birches grew during the Younger Dryas. With the onset of the isotopic shift initiating the rapid warming (about 11,535 cal. years before 1950), the pine forest became more productive and denser. At Leysin no trees except some juniper scrub grew during the Younger Dryas. Tree birches, pine, and poplar immigrated from lower altitudes and arrived after the end of the isotopic shift (about 11,487 B.P.), i.e., at the beginning of the Preboreal (at about 11,420 B.P.). Zeneggen is situated somewhat higher than Leysin, but single tree birches and pines survived the Younger Dryas at the site. At the beginning of the Preboreal their productivity and population densities increased. Simultaneously shifts from Nitella to Chara and from silt to gyttja are recorded, all indicating rapidly warming conditions and higher nutrient levels of the lake water (and probably of the soils in the catchment). At Gerzensee the beginning of the Younger Dryas was also analysed: the beginning of the isotopic shift correlates within one sample (about 15 years) to rapid decreases of macrofossils of pines and tree birches.

Declining pine growth in Central Spain coincides with increasing diurnal temperature range since the 1970s

Growing evidence suggests environmental change to be most severe across the semi-arid subtropics, with past, present and projected drying of the Mediterranean Basin posing a key multidisciplinary challenge. Consideration of a single climatic factor, however, often fails to explain spatiotemporal growth dynamics of drought-prone ecosystems. Here, we present annually resolved and absolutely dated ring width measurements of 871 Scots pines (Pinus sylvestris) from 18 individual plot sites in the Central Spanish Pinar Grande forest reserve. Although comprising tree ages from 6 to 175 years, this network correlates surprisingly well with the inverse May–July diurnal temperature range (r = 0.84; p < 0.00011956–2011). Ring width extremes were triggered by pressure anomalies of the North Atlantic Oscillation, and the long-term growth decline coincided with Iberian-wide drying since the mid-1970s. Climate model simulations not only confirm this negative trend over the last decades but also project drought to continuously increase over the 21st century. Associated ecological effects and socio-economic consequences should be considered to improve adaptation strategies of agricultural and forest management, as well as biodiversity conservation and ecosystem service.

Reviving extinct Mediterranean forest communities may improve ecosystem potential in a warmer future

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.

Impacts of changing climate and land use on vegetation dynamics in a Mediterranean ecosystem: insights from paleoecology and dynamic modeling

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 pollen-inferred 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 paleo-environmental 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.

Abies – Circum-Mediterranean firs in Europe: distribution, habitat, usage and threats

Most European firs occur predominantly in small to medium-sized populations in the Mediterranean region, sometimes with fragmented and limited distributions, except for silver fir (Abies alba). They all are genetically closely related and can easily hybridise, perhaps as a consequence of late speciation during the late Quaternary. Circum-Mediterranean firs occur principally in mountain areas with medium to high precipitations rates which are mostly concentrated during the winter period. The species are able to tolerate long droughts in summer and tend to form pure stands when in optimal habitats. In the past firs have been extensively logged for construction and fire wood and their stands were replaced by other more disturbance adapted species or converted into rural areas. Nowadays with the exception of silver fir and Caucasian fir (Abies nordmanniana), circum-Mediterranean firs do not have a wide commercial interest. In Turkey they are still exploited for timber wood, while other firs have an ornamental use in gardening. Great importance is given to their preservation, especially to those populations which have very limited areas and specimens, with the creation of protected reserves and conservation programmes. Wild fires, livestock grazing and genetic drift represent actually their main threats.

What drives European beech (Fagus sylvatica L.) mortality after forest fires of varying severity?

Predicting the timing and amount of tree mortality after a forest fire is of paramount importance for post-fire management decisions, such as salvage logging or reforestation. Such knowledge is particularly needed in mountainous regions where forest stands often serve as protection against natural hazards (e.g., snow avalanches, rockfalls, landslides). In this paper, we focus on the drivers and timing of mortality in fire-injured beech trees (Fagus sylvatica L.) in mountain regions. We studied beech forests in the southwestern European Alps, which burned between 1970 and 2012. The results show that beech trees, which lack fire-resistance traits, experience increased mortality within the first two decades post-fire with a timing and amount strongly related to the burn severity. Beech mortality is fast and ubiquitous in high severity sites, whereas small- (DBH <12 cm) and intermediate-diameter (DBH 12–36 cm) trees face a higher risk to die in moderate-severity sites. Large-diameter trees mostly survive, representing a crucial ecological legacy for beech regeneration. Mortality remains low and at a level similar to unburnt beech forests for low burn severity sites. Beech trees diameter, the presence of fungal infestation and elevation are the most significant drivers of mortality. The risk of beech to die increases toward higher elevation and is higher for small-diameter than for large-diameter trees. In case of secondary fungi infestation beech faces generally a higher risk to die. Interestingly, fungi that initiate post-fire tree mortality differ from fungi occurring after mechanical injury. From a management point of view, the insights about the controls of post-fire mortality provided by this study should help in planning post-fire silvicultural measures in montane beech forests.