Growth patterns in relation to drought-induced mortality at two Scots pine (Pinus sylvestris L.) sites in NE Iberian Peninsula

Drought-related tree mortality has become a widespread phenomenon. Scots pine (Pinus sylvestris L.) is a boreal species with high ecological amplitude that reaches its southwestern limit in the Iberian Peninsula. Thus, Iberian Scots pine populations are particularly good models to study the effects of the increase in aridity predicted by climate change models. A total of 78 living and 39 dead Scots pines trees were sampled at two sites located in the NE of the Iberian Peninsula, where recent mortality events have been recorded. Annual tree rings were used to (1) date dead trees; (2) investigate if there was an association between the occurrence of tree death and severe drought periods characterized by exceptionally low ratios of summer precipitation to potential evapotranspiration (P/PET); and (3) to compare the growth patterns of trees that died with those of surviving ones. Mixed models were used to describe the relationships between tree growth (in terms of basal area increment, BAI, and the percentage of latewood, LW%) and climate variables. Our results showed a direct association between Scots pine mortality and severe drought periods characterized by low summer water availability. At the two sites, the growth patterns of dead trees were clearly distinguishable from those of the trees that survived. In particular, the BAI of dead trees was more sensitive to climate dryness (low P/PET_summer, high temperatures) and started to decline below the values of surviving neighbors 15–40 years before the time of death, implying a slow process of growth decline preceding mortality.     

Temperature sensitivity of blue intensity,maximum latewood density,and ring width data of living black spruce trees in the eastern Canadian taiga

The blue intensity (BI) technique provides opportunities to obtain surrogates to tree-ring density for reconstructing summer temperatures in high-latitude regions. In this study, we compare latewood BI (LBI) and delta BI (DBI), with the conventional X-ray maximum latewood density (MXD) and tree-ring width (TRW) data using 178 living trees of black spruce (Picea mariana (Mill.) B.S.P.), one of the most dominant species of conifers in the Northern Hemisphere, from 17 sites across the eastern Canadian taiga. The regional LBI and DBI chronologies are highly correlated to that of MXD (Pearson’s r = 0.97 and 0.92, respectively), while DBI is also similar to TRW (Pearson’s r = 0.67). Both LBI and DBI exhibit stronger responses to the May–August temperatures than TRW over larger time and spatial scales. However, only DBI is comparable to MXD data from inter-annual to decadal timescales. Low-frequency components of LBI data are likely distorted by color biases even if no obvious discoloration is present, as well as by the potentially low measurement resolution, leading to an overall weaker temperature sensitivity compared to the MXD data. Resampling experiments suggest that a minimum replication of 10 trees is needed to retain ≥90 % of the optimal temperature signal for MXD, LBI, and DBI data, and a minimum of 20 trees is required for TRW data.     

Dendroecological analysis of relict pine forests in the centre of the Iberian Peninsula

Pinus nigra subsp. salzmannii is found in the east and centre of the Iberian Peninsula, in the south of France and in North Africa. This subspecies occupies the westernmost position of the species’ general range. The persistence on the Iberian Peninsula of very long-lived specimens of Pinus nigra subsp. salzmannii, along with their sensitivity to climate, has drawn the attention of many researchers, but to date the importance of dendroecological studies relating to conservation of biodiversity or the genetic resources of this taxon had not been stressed. In the present paper we use dendroecological methods to analyse the relict pine forest in Navalacruz, an interesting and endangered genetic forestry resource on the northern slopes of the Gredos mountains (in Spain’s Central System Range) at the subspecies’ south-western global limit. This forest provides a prime example for demonstrating the potential application of dendroecology for studying the origin, dynamics, local variability, relationships with climate and anthropogenic disturbances of relict forest populations. We dated 93 growth sequences from 47 trees ranging from 1809 to 2006 and we have determined that interspecific competition is the most relevant factor as regards differences in the diameter growth of these trees. Moreover, we detected great variability and numerous common growth disturbances unrelated to climatic oscillations. These quasi-periodic disturbances alternate between suppression and release suggesting continuous management cycles of different intensities. Despite its high level of disturbance, the pine forest presents a certain degree of climatic sensitivity. Comparing with others Pinus nigra subsp. salzmannii populations, we denoted a temporal grading of the growth response to precipitation that is indicative of differences in the start and length of the vegetative period. Furthermore, we compiled different dendroecological and palaeobiogeographical data to demonstrate that this dense, homogeneous and relatively younger P. nigra population is of an indigenous nature. This study aims to provide data for improved management and conservation of this exceptional and highly endangered bastion of biodiversity.     

Dendroclimatic signals deduced from riparian versus upland forest interior pines in North Karelia, Finland

Radial growth of boreal tree species is only rarely studied in riparian habitats. Here we investigated chronologies of earlywood, latewood, and annual ring widths and blue intensity (BI; a surrogate to latewood density) from riparian lake shore and upland forest interior pines (Pinus sylvestris L.) growing in boreal forest in eastern Finland. Riparian and upland chronologies were compared to examine differences in the pine growth variability and growth response to climatic variation in the two habitats. It was found that the climatic variables showing statistically significant correlations with the tree-ring chronologies were related to snow conditions at the start of the growing season. Deeper snowpack led to reduced upland pine growth, possibly due to delayed snowmelt and thus postponed onset of the growing season. Warm late winters were followed by increased riparian pine growth because of earlier start of the snow-melt season and thus a lower maximum early summer lake level. Moreover, riparian pines reacted negatively to increased rainfall in June, whereas the upland pines showed a positive response. Latewood growth reacted significantly to summer temperatures. The BI chronology showed a strong correlation with warm-season temperatures, indicating an encouraging possibility of summer temperature reconstruction using middle/south boreal pine tree-ring archives.     

Summer rainfall variability in European Mediterranean mountains from the sixteenth to the twentieth century reconstructed from tree rings

Since the end of the last glacial period, European Mediterranean mountains have provided shelter for numerous species of Eurosiberian and Boreal origin. Many of these species, surviving at the southern limit of their range in Europe and surrounded by Mediterranean ones, are relatively intolerant to summer drought and are in grave danger of loss, as a result of increasingly long and frequent droughts in this region. This is the case of the Scots pine (Pinus sylvestris) and the Austrian pine (Pinus nigra ssp. salzmannii) which are found on Central Iberian Peninsula at the edge of their natural range. We used a tree ring network of these two species to reconstruct past variations in summer rainfall. The reconstruction, based upon a tree ring composite chronology of the species, dates back to 1570 (adjusted R ²?=?0.49, P?<?0.000001) and captures interannual to decadal scale variability in summer precipitation. We studied the spatial representativeness of the rainfall patterns and described the occurrence rate of extremes of this precipitation. To identify associations between macroclimatic factors and tree radial growth, we employed a principal component analysis to calculate the resultant of the relationship between the growth data of both species, using this resultant as a dependent variable of a multiple regression whose independent variables are monthly mean temperature and precipitation from the average records. Spatial correlation patterns between instrumental precipitation datasets for southern Europe and reconstructed values for the 1950–1992 period indicate that the reconstruction captures the regional signal of drought variability in the study region (the origin of this precipitation is convective: thermal low pressure zones induced in the inland northeastern areas of the Iberian Peninsula). There is a clear increase in the recurrence of extreme dry events as from the beginning of twentieth century and an abrupt change to drier conditions. There appears to be a tendency toward recurrent exceptionally dry summers, which could involve a significant change for the Eurosiberian refugee species.     

First measurements of Blue intensity from Pinus peuce and Pinus heldreichii tree rings and potential for climate reconstructions

Climate changes, their regional patterns, origin, and prediction are currently one of the most important scientific challenges. Tree-rings are among the most widely used proxies for past climate variation. However tree-ring width (TRW) from certain tree species and regions often do not contain robust climate signal. Other parameters such as Maximum latewood density (MXD) of conifer tree rings are more sensitive to summer temperatures, but the measurements have high costs. A potential surrogate for MXD is blue intensity (BI), which is based on the measurements of high-resolution images of the wood. The method has been tested and applied for several species. However it has not been tested up to now for Pinus heldreichii Christ (PIHE) and Pinus peuce Griseb. (PIPE). Those species are with limited distribution on the Balkan Peninsula and due to their longevity (frequently more than 500 years with potential to more than 1000 years) may serve as one of the best proxies for past climate variation in SE Europe.We composed BI chronologies following standard procedures from PIHE and PIPE trees from subalpine locations in the Pirin Mts, Bulgaria. The correlation analysis with climate parameters revealed strong and significant positive correlations of PIHE BI (ΔBI and Latewood BI (LwBI) series) and summer temperatures. The highest were 0.74 (p < 0.05) with July-August average temperatures for the period 1933–1983. The correlation values of ΔBI remained higher than 0.6 for the whole period (1933–2014) and sub-periods both for average monthly and average maximum temperatures. LwBI values of the PIHE chronology had correlation coefficients above 0.52 (highest 0.64) with August and August-July temperatures. The PIPE BI chronologies were also positively correlated with summer temperatures, but showed lower values than PIHE with lower temporal stability. The most stable were the correlations with LwBI average August temperatures, which were above 0.52. Delta BI series displayed high correlations for the 1933–1983 period, but then decreased and this caused overall lower correlations with August temperatures. Our data shows that there is potential to develop long BI chronologies and proxy climate reconstructions from the studied species and in this way complement the knowledge of the past climate of SE Europe.     

Different climate response of three tree ring proxies of Pinus sylvestris from the Eastern Carpathians,Romania

The aim of this study was to compare the climatic responses of three tree rings proxies: tree ring width (TRW), maximum latewood density (MXD), and blue intensity (BI). For this study, 20 cores of Pinus sylvestris covering the period 1886–2015 were extracted from living non-damaged trees from the Eastern Carpathian Mountains (Romania). Each chronology was compared to monthly and daily climate data. All tree ring proxies had a stronger correlation with the daily climate data compared to monthly data. The highest correlation coefficient was obtained between the MXD chronology and daily maximum temperature over the period beginning with the end of July and ending in the middle of September (r = 0.64). The optimal intervals for the temperature signature were 01 Aug – 24 Sept for the MXD chronology, 05 Aug – 25 Aug for the BI chronology, and both 16 Nov of the previous year – 16 March of the current year and 15 Apr – 05 May for the TRW chronology. The results from our study indicate that MXD can be used as a proxy indicator for summer maximum temperature, while TRW can be used as a proxy indicator for just March maximum temperature. The weak and unstable relationship between BI and maximum temperature indicates that BI is not a good proxy indicator for climate reconstructions over the analysed region.     

Comparing Scots pine tree-ring proxies and detrending methods among sites in Jämtland,west-central Scandinavia

Scots pine tree-ring width (TRW) data from Jämtland in the Central Scandinavian Mountains has been used to reconstruct summer temperatures back to 1630 BC. However, it was recently shown that this reconstruction was of limited spatial importance. In this paper, we aim to explain this limitation in the TRW data as a temperature proxy, as well as assess the temperature information from new maximum latewood density (MXD) data. Furthermore, the effect of two standardization methods is evaluated: regional curve standardization (RCS) and a more traditional standardization, termed “non-RCS” standardization. Three TRW and two MXD sites were analyzed. Our results showed that despite the proximity to the Norwegian Sea, the MXD data is a powerful temperature proxy. Difference among sites in TRW data, especially on decadal timescales, together with a lower temperature association, suggests that other factors, such as changes in the local climate regimes, weakens the temperature signal. In general the RCS method overestimates pine growth trends in the latter half of the twentieth century, a feature not seen when using “non-RCS” standardization. This is likely due to an age-bias of older trees in most recent parts of the tree-ring chronologies. This effect will have consequences when reconstructing climate with tree-ring data. To overcome this problem, all age-classes should be represented throughout a chronology. If this is not possible, the use of “non-RCS” standardization is recommended, although this method results in a loss of low-frequency variability.     

Climatic impact on tree-ring widths in Abies borisii-regis forests from South-East Albania

The long-term radial growth responses to drought and climatic variability of less-studied species such as Abies borisii-regis (Mattf.) remain poorly understood.We tested the hypothesis that severe short-term drought conditions during summer months will impact the radial growth of A. borisii-regis (Mattf.) trees and such impact will have a more pronounced effect on latewood (LW) than earlywood (EW) width.Correlation analysis was employed to investigate the impact of climatic drivers (temperature, precipitation) and drought, using the Standardized Precipitation Evapotranspiration Index (SPEI) calculated at cumulative time scales (1–12 months), on EW, LW and tree-ring width (TRW) in A. borisii-regis (Mattf.) trees from South-Eastern Albania.We found that EW width was positively correlated with precipitation in July and previous September, while the LW width and TRW was enhanced by the current June–July precipitation. Previous autumn and current summer high temperatures constrained the radial growth in A. borisii-regis (Mattf.) trees, particularly the LW and TRW. All the tree-ring widths components showed the highest significant response to drought at short cumulative time scales (<4 months) mainly during July, August and September. The highest impact of drought was observed for the LW width.Under a future reduction of summer precipitation and temperature increase, the A. borisii-regis (Mattf.) may show a decrease in EW formation, causing a decline of radial growth, leading to a reduction in hydraulic conductivity and carbon uptake in these forests.     

High-resolution climatic analysis of wood anatomical features in Corsican pine from Corsica (France) using latewood tracheid profiles

Key message

We propose a new methodology to identify intra-annual density fluctuations in latewood using cell features and relative radial position within the latewood of pine trees growing on Corsica, France. Climatic forcing of latewood wood anatomical anomalies was analyzed.

Abstract

We analyzed latewood anatomical features from Corsican pine (Pinus nigra ssp. laricio) of high-elevation sites in Corsica (France) derived from digital images of the wood surface. Latewood of each ring during the period 1950–2008 was partitioned into ten equal parts P1–P10. Mean values of the cell parameters cell lumen area (CLA), radial cell width (RCW), radial cell wall thickness (CWT), and modeled latewood density (MLD) were calculated for P1–P10. The cellular profiles for each cell parameter were subjected to principal component analyses. It was possible to quantify macroscopically visible variations of wood anatomy like intra-annual density fluctuations (IADFs) by latewood profiles of different cell parameters. A combination of cell parameter characteristics including their relative radial position within latewood provides a quantification of the cell anatomical variations in an IADF. Individual cell parameter chronologies and principal components of cell parameter profiles were correlated with climate data to determine the climatic forcing on latewood formation. Average cell parameter profiles and deviations from the long-term means are able to describe “normal” and “anomalous” environmental conditions during latewood formation. Cell feature anomalies throughout the latewood during individual years allow the reconstruction of past weather conditions with a high temporal resolution.     

Structural attributes,tree-ring growth and climate sensitivity of Pinus nigra Arn. at high altitude: common patterns of a possible treeline shift in the central Apennines (Italy)

European black pine (Pinus nigra ssp. nigra Arnold) encroachment at increasing elevation has been analyzed at four treeline ecotones of the central Apennines (Italy). The study sites are located along a North-South gradient of 170 km across Marche and Abruzzo regions in Central Italy. The aims of this study were: (i) to detect possible common patterns of structural attributes of black pine regeneration at the treeline ecotone; (ii) to date the seedlings germination and (iii) to assess the climate influence on the pine upward encroachment process also using intra-annual density fluctuations (IADFs) in tree-rings. We sampled 658 encroached black pine trees above the current treeline to the mountain top. All individuals were mapped and their basal stem diameter, total height, annual height increments and other structural attributes measured. One increment core was extracted from stem base of most samples for cambial age determination and detection of intra-annual density fluctuations (IADF). At two sites we also extracted cores at DBH from forest trees to assess climate–growth relationships of black pine. We used multivariate analysis (PCA) to explore the correlation structure of the main tree attributes, regression analysis to relate radial and height increment and dendroclimatic analysis to assess the influence of climate on tree growth and IADF formation.Most black pine trees were located at high altitude and their structural attributes were similar at the four sites where the pine encroachment process started between 30 and 40 years ago featuring similar germination peaks and growth patterns. Black pine is particularly sensitive to maximum temperatures and IADF occurred in mid-late summer with highest frequency peaks between 2003 and 2004. The pine encroachment process, besides the differences of environmental features and land use histories of the four study sites, appears synchronic and spatially diffused. Consistent tree-growth dynamics and the species adaptation to a warming climate are signals envisaging a possible treeline upward shift.     

Size-dependent responses to summer drought in Scots pine, Norway spruce and common oak

In this study, we provide a detailed analysis of tree growth and water status in relation to climate of three major species of forest trees in lower regions of Bavaria, Southern Germany: Scots pine (Pinus sylvestris), Norway spruce (Picea abies) and common oak (Quercus robur). Tree-ring chronologies and latewood δ¹³C were used to derive measures for drought reaction across trees of different dimensions: growth reduction associated with drought years, long-term growth/climate relations and stomatal control on photosynthesis. For Scots pine, growth/climate relations indicated a stronger limitation of radial growth by high summer temperatures and low summer precipitation in smaller trees in contrast to larger trees. This is corroborated by a stronger stomatal control on photosynthesis for smaller pine trees under average conditions. In dry years, however, larger pine trees exhibited stronger growth reductions. For Norway spruce, a significantly stronger correlation of tree-ring width with summer temperatures and summer precipitation was found for larger trees. Additionally, for Norway spruce there is evidence for a change in competition mode from size-asymmetric competition under conditions with sufficient soil water supply to a more size-symmetric competition under dry conditions. Smaller oak trees showed a weaker stomatal control on photosynthesis under both dry and average conditions, which is also reflected by a significantly faster recovery of tree-ring growth after extreme drought events in smaller oak trees. The observed patterns are discussed in the context of the limitation-caused matter partitioning hypothesis and possible species-specific ontogenetic modifications.     

Woody macroremains from the Acheulian site of Torralba: Occurrence and palaeoecology of Pinus cf. sylvestris in the Middle Pleistocene of the Iberian Peninsula

The pieces of wood extracted by the Marquis of Cerralbo at the Acheulian site of Torralba constitute one of the few materials from this famous locality that have not been subjected to detailed study by researchers. Over a century after their extraction, the present paper provides the first anatomical results of these 34 woody remains. The results highlight the presence of the taxon Pinus sylvestris–nigra group (Pinus cf. sylvestris) at the site, thus constituting one of the few and oldest references attributable to the aforementioned taxon for the Iberian Peninsula. The anatomical diagnosis confirms the existence of Middle Pleistocene landscapes with pine woods at elevations at least 200 m lower than at present in the site area. From a taphonomic point of view, the wood samples exhibit the typical morphologies of woody macroremains preserved in continental sedimentary environments, and there was no evidence at all that they had previously been manipulated by man.     

Assessing the Conservation Status of an Iberian Moth Using Pseudo-Absences

ABSTRACT Knowing the distribution of endangered species is of substantial importance for conservation. We considered a useful approach for modeling species distribution when managing information from atlases and museums but when absence data is not available. By modeling the distribution for Graellsia isabelae, a threatened moth species, we assessed its current conservation status and identified its most relevant distribution explanatory variables using Geographic Information System and Generalized Linear Models. The distribution model was built from 136 occurrence records and 25 digitized explanatory variables at a 10 × 10 km resolution. Model predictions from logistic-regressed pseudo-absences, obtained from a presence-only method (Ecological-Niche Factor Analysis), explained 96.23% of the total deviance. We found that the best predictor variables were summer precipitation, aridity, and mean elevation. With respect to host plants, the presence of G. isabelae associated mainly with Scots pine (Pinus sylvestris) and Austrian pine (P. nigra). The finding of 8 areas, exclusively in the eastern Iberian territory, and a larger unoccupied habitat in the western Iberian Peninsula indicates that this species is probably not in equilibrium with its environment by historical factors. Sites of Community Importance under protection do not seem sufficient to maintain current populations, necessitating the protection of suitable neighboring habitats. Our methodology is useful to manage the conservation status of species for which reliable absence data is not available. It is possible to determine those variables that most affect the distribution of species as well as the potential suitable areas with the purpose of evaluating protected areas, connectivity among populations, and possible reintroductions.     

Distance between south-European and south-west Asiatic refugial areas involved morphological differentiation: Pinus sylvestris case study

The phenotypic differentiation of relic P. sylvestris in southern Europe and southwestern Asia was verified using thirty-two populations sampled from the Iberian Peninsula, Massif Central, Balkan Peninsula, Crimea and Anatolia. Twenty-one morphological and anatomical needle traits and 18 cone morphological characteristics were examined to describe the population diversity and differentiation. The needle characters were not correlated to those of cone. The differences between regions were significant based on 12 needle and 9 cone characteristics, suggesting spatial isolation. The differentiation between the Iberian and Anatolian populations was the highest, which indicates the isolation by distance. The high level of morphological differentiation was also found among Iberian populations, supporting the already known complex history of the species in that region. Populations within other regions were differentiated at lower levels; however, the West Anatolian populations differed morphologically from the eastern ones. The described pattern of morphological differentiation supports the idea of the long-lasting existence of P. sylvestris in the south-European and Anatolian mountain regions. To conserve this variation, seed transfer between regions in the forest economy should be restricted.     

A long-term chronology of Pinus pinea L. from Parco della Versiliana (Pietrasanta,Italy) derived from treefall induced by a windstorm on March 4th-5th, 2015

Pinewoods are distinctive environmental elements in the Mediterranean coastal area and have both natural and historical significance. From the evening of March 4th to the morning of March 5th, 2015, a severe and unusual windstorm occurred in the Tuscany region of central Italy with wind gusts over 120 km/h. The windstorm caused vast damage to the anthropic and natural environment and wounded numerous trees in the renowned pinewoods of Parco della Versiliana in the Tyrrhenian coastal area. The meteorological calamity provided the opportunity to i) date the onset of the artificial plantation of the present Italian stone pine (Pinus pinea L.) forest to the 1820s, ii) build a long-term tree-ring chronology of the Italian stone pines in the area and iii) analyze the climate-growth relationship of the Italian stone pine in the study area. The resulting Versiliana chronology was derived from 60 trees and spanned from 1828 to 2014 (187 years), representing one of the longest living Italian stone pine forests on the Italian Peninsula. Finally, the climate-growth analysis highlighted that at this site the latewood width is positively influenced by summer temperature, a peculiarity worthy of further investigations.     

Canopy status modulates formation of wood rays in scots pine under hemiboreal conditions

Non-structural carbohydrates (NSC) reserves are crucial for trees to cope with weather extremes, thus to ensure their survival and ecological plasticity. The NSC reserves can depend on social status, suggesting uneven plasticity of trees at the stand level. In stemwood of Scots pine (Pinus sylvestris L.), which is a widespread and important species, NSC reserves are stored in parenchyma in wood rays (WR). The quantity of WRs is adjusted intra-annually, allowing retrospective analysis of factors affecting their formation. Accordingly, the differences in WR quantity in stemwood of dominant and intermediate (canopy trees with reduced and narrow crowns) maturing Scots pine were assessed by quantitative wood anatomy. Tangential cuts from the outermost 30 tree-rings were analysed. The relative ray area was intermediate, i.e. covering about 5% of the tangential cut, yet expressed high individuality among the trees. The size and amount of WR mainly differed between the earlywood and latewood; WRs in latewood were higher although narrower in comparison to earlywood, yet their total amount was higher in earlywood. Canopy status had only a slight effect, as quantity and height of WR tended to be higher for the intermediate trees, particularly in earlywood. The size and quantity of WR expressed inter-annual variation, which was mainly related to the meteorological conditions prior to the formation of the tree-ring (previous summer and autumn) indicating legacy effects of climatic factors on NSC and susceptibility of trees to cumulative effects of weather extremes. However, the climatic signals in the inter-annual variation of WR were weaker than observed before, likely due to location of the studied stand in the mid-part of the species range. Nevertheless, the observed differences in mean values and inter-annual variation of WR suggested a within-species diversity of carbon allocation patterns, supporting adaptability of the species.     

Direct and Indirect Effects of Climate on Demography and Early Growth of Pinus sylvestris at the Rear Edge: Changing Roles of Biotic and Abiotic Factors

Global change triggers shifts in forest composition, with warming and aridification being particularly threatening for the populations located at the rear edge of the species distributions. This is the case of Scots pine (Pinus sylvestris) in the Mediterranean Basin where uncertainties in relation to its dynamics under these changing scenarios are still high. We analysed the relative effect of climate on the recruitment patterns of Scots pine and its interactions with local biotic and abiotic variables at different spatial scales. Number of seedlings and saplings was surveyed, and their annual shoot growth measured in 96 plots located across altitudinal gradients in three different regions in the Iberian Peninsula. We found a significant influence of climate on demography and performance of recruits, with a non-linear effect of temperature on the presence of juveniles, and a positive effect of precipitation on their survival. Abundance of juveniles of P. sylvestris that underwent their first summer drought was skewed towards higher altitudes than the altitudinal mean range of the conspecific adults and the optimum elevation for seedlings'' emergence. At local level, light availability did not influence juveniles'' density, but it enhanced their growth. Biotic interactions were found between juveniles and the herb cover (competition) and between the number of newly emerged seedlings and shrubs (facilitation). Results also highlighted the indirect effect that climate exerts over the local factors, modulating the interactions with the pre-existing vegetation that were more evident at more stressful sites. This multiscale approach improves our understanding of the dynamics of these marginal populations and some management criteria can be inferred to boost their conservation under the current global warming.     

Close and distant: Contrasting the metabolism of two closely related subspecies of Scots pine under the effects of folivory and summer drought

Metabolomes, as chemical phenotypes of organisms, are likely not only shaped by the environment but also by common ancestry. If this is the case, we expect that closely related species of pines will tend to reach similar metabolomic solutions to the same environmental stressors. We examined the metabolomes of two sympatric subspecies of Pinus sylvestris in Sierra Nevada (southern Iberian Peninsula), in summer and winter and exposed to folivory by the pine processionary moth. The overall metabolomes differed between the subspecies but both tended to respond more similarly to folivory. The metabolomes of the subspecies were more dissimilar in summer than in winter, and iberica trees had higher concentrations of metabolites directly related to drought stress. Our results are consistent with the notion that certain plant metabolic responses associated with folivory have been phylogenetically conserved. The larger divergence between subspecies metabolomes in summer is likely due to the warmer and drier conditions that the northern iberica subspecies experience in Sierra Nevada. Our results provide crucial insights into how iberica populations would respond to the predicted conditions of climate change under an increased defoliation in the Mediterranean Basin.     

Genetic variation of isolated and peripheral populations of Pinus sylvestris (L.) from glacial refugia

Analysis of variation at 10 polymorphic isoenzyme loci in 51 European populations of Pinus sylvestris both from its main continuous range and from isolated, marginal populations out of the continuous range, revealed differences in genetic structure. The highest genetic richness, measured as mean number of alleles per locus, was recorded for populations from the Iberian Peninsula, followed by those from Scotland and the Balkans. Marginal, isolated populations were characterized by much greater interpopulation variation than populations from the continuous range of the species. The highest mean observed heterozygosity was recorded for the Iberian populations. The highest gene flow was observed among populations in the continuous range of the species. The populations could be classified into five groups based on genetic similarities. Populations from the continuous range formed one group, those from Spain two groups, pines from Scotland one group and those from the Balkans one group. One population from Scotland was found to be most distinct from the other Scottish pines and was found to be grouped with the Balkan populations. The occurrence and frequencies of rare alleles in the populations showed a characteristic pattern, suggesting that populations from the Iberian Peninsula probably did not participate in the colonization of Europe by Pinus sylvestris after the last glaciation. In contrast, the migration of populations from the Balkan refugium into Central and Western Europe had a significant effect on the contemporary gene pools of populations of Pinus sylvestris in its continuous range.