Patterns of post-glacial spread and the extent of glacial refugia of European beech (Fagus sylvatica)

Aim  Establishing possible relationships between the magnitudes of the glacial distribution of the European beech, Fagus sylvatica L., and its post-glacial spread.
Location  Europe.
Methods  A database of over 400 pollen records has been used to locate Fagus populations at the end of the last glacial and during the post-glacial in Europe and to assess the areal extent of their past distribution.
Results  The rate of late-glacial and post-glacial increase in the number of pollen sites where Fagus was locally present conforms well to a logistic model of population growth. This suggests that the area occupied by beech populations expanded exponentially from the glacial refugia for a duration of over 10,000 years, until about 3500 yr bp . In the past three millennia beech populations increased at a slower rate, tending towards an equilibrium value.
Main conclusions  The conformity of the increase in beech distribution to the classical logistic model of population growth indicates that: (1) a multiplicative biological process was the main factor shaping the pattern of the post-glacial expansion of F. sylvatica in Europe, (2) climate conditions, human activity and competition may have influenced its rate of spread, and (3) beech populations did not expand with a moving closed front, but with a diffuse spread from scattered nuclei. The distribution of Fagus in Europe at the end of the last glacial appears to have been of two orders of magnitude less extensive than at present. Pleistocene refugia were likely to have been a mosaic of sparse stands of small populations scattered in multiple regions. Fagus populations appear to have increased very slowly and to a moderate extent in southern Europe, where they are now declining slightly. The central European populations increased quickly and extensively, reaching northern Europe, and are now approaching their carrying capacity.     

欧洲水青冈(Fagus sylvatical L.)构筑型与形态多样性研究

通过对植物构件和构筑型的分析,可以了解植物的整体结构与系统演化关系。高等植物构筑型的研究以树木最为深入,植物学家对全球的树木进行了构筑模式的分类,并确定了２３个基本的构筑类型。植物的构筑型和其形态的多样性是密不可分的。构筑奕型确定了植物所属的整体形态类型,而对每种植物具体的形态多样性分析可以深入了解每种构筑型的数量特征及其形成过程。在概述了树木树筑型研究的基础上,系统说明了欧洲水青冈构筑才形态多样性分析的步骤和步骤和分析的意义。水青冈属（Fagus)为Ｔroll型构筑模式,处于不同生态条件下的同一种的不同种群之间,在形态、解剖、生理和遗传多样性方面都存在着差异。树木的形态多样性虽然主要受遗传因素的控制,但生态条件对其形态和遗传多样性也有非常显著的影响。在对树木进行构筑型分析时,道德要根据构筑要素（主要是形态与生长特性）确定它所属的构筑类型,其次是分析它的形态特征与环境条件的关系,进行进一步分析其遗传多样性。     

Mechanism of freeze-induced embolism in Fagus sylvatica L.

 The mechanism of freeze stress-induced embolism in Fagus sylvatica L. branches was analyzed under controlled conditions. Excised branches were exposed to successive freeze-thaw cycles in temperature controlled chambers. Thermocouples were placed on the bark to detect sap freezing exotherms. The degree of xylem embolism was estimated after each cycle by the loss of hydraulic conductivity. After one freeze-thaw cycle the degree of embolism was found to decrease with xylem specific hydraulic conductivity, small apical shoots being more susceptible to embolism. Exotherms revealed that distal shoots were freezing first and exuded sap as a result of water expansion. The lower water content in apical shoots upon freezing probably induced higher sap tensions which promoted air bubble expansion and vessel cavitation preferentially near the apices. When the decrease in water content was experimentally prevented, embolism developed to a lesser extent. The higher vulnerability of shoot apices may protect the rest of the branch from winter damage. Received: 29 May 1998 / Accepted: 15 August 1998     

Is the spatial distribution of European beech (Fagus sylvatica L.) limited by its potential height growth?

Aim To improve our understanding of species range limits by studying how height growth, a trait related to plant survival, varies throughout the geographic range of Fagus sylvatica L. in France. Location The geographic range of beech in France, representing the western area of its European distribution, within which this species exhibits range distribution limits in both plains and mountainous areas. Methods A generalized linear regression model was used to link beech growth performance to environmental variables using data from 819 plots of the French National Forest Inventory (IFN) database. This model was applied to predict potential growth on 97,281 IFN plots covering the geographic range of beech in France. A kriging technique was used to interpolate estimated growth potential. Finally, the performance of plot‐based predictions of potential growth from the map (i.e. map quality) was evaluated against an independent data set. Results The beech growth performance model highlighted the major impact of climate on potential tree growth at a broad spatial scale. The relevant climatic factors were related mainly to spring cold, summer heat, and winter temperatures and rainfall. The study also revealed the predictive power of soil parameters, which explained a large proportion of the variation in potential beech growth (c. 30%). Analyses of height growth patterns near the boundary of the species range in France showed that the limit only partly coincides with the growth decline caused by climatic and soil factors. Along parts of the range limit, the predicted potential for growth was high, suggesting that in these areas the limit of the range could be explained by other factors, such as competition or constraints on reproduction. Main conclusions The spatial variation in the potential height growth of Fagus sylvatica can be explained by environmental factors and is partly correlated with its regional range limits. By identifying areas where growth potential constrains the geographic range of species, environmental growth models can help to improve our knowledge of the spatial drivers of species geographic range limits and shed light on their response to future environmental changes.     

Variazioni stagionali di nutrienti minerali ed elementi in traccia nelle foglie di Fagus sylvatica L. a Vallombrosa (FI)

Abstract

Seasonal variations of mineral nutrients and trace elements in Fagus sylvatica L. leaves at Vallombrosa (FI). - During the vegetative periodo 1987 monthly samples of beech leaves were collected in Vallombrosa forest area to determine the nutritive situation of beech trees by leaves analysis. Among nutrients, potassium appears the worst supplied one. Even copper is present in very low values, under the critical level. Heavy metals as manganese, aluminium, lead, cadmium and also iron show a clear increase during the vegetative season.     

Water relations and gas exchange in Fagus sylvatica L. and Quercus petraea (Mattuschka) Liebl. in a mixed stand at their southern limit of distribution in Europe

Water status and gas exchange of beech (Fagus sylvatica L.) and sessile oak [Quercus petraea (Mattuschka) Liebl.] were studied in a mixed stand in the Montejo de la Sierra forest (central Iberian Peninsula), one of the southernmost locations of both species in Europe. Gas exchange and water potential were measured in leaves at different canopy levels over several days in two growing seasons. The daily variation pattern was established with the measurements of three selected dates per year, representative of the soil moisture content situations in early, mid- and late summer. A similar daily time course of leaf water potential was found for the two species. Nevertheless, beech showed a most noticeable decrease of water potential at midmorning and maintained lower leaf water potential than oak in the early afternoon. In 1994 the sessile oak saplings showed higher values of predawn water potential (Ψ_pd) than beech at the end of summer, when soil moisture content was lowest (20 cm depth). Beech showed a significantly lower net assimilation rate (A) than sessile oak for leaves under the same PPFD. Maximum net photosynthesis values (A _max) for beech and sessile oak on sunny leaves were 10.1±0.4 μmol m^–2 s^–1 and 17.8±1.7 μmol m^–2 s^–1 respectively, and those for water vapour stomatal conductance (g _wv) were 265±31 mmol m^–2 s^–1 and 438±74 mmol m^–2 s^–1. Differences in A and g _wv between the two species were maintained throughout the day on all measurement dates. No clear relationship was found between water status of saplings and stomata performance; there was only a negative correlation between Ψ_pd and g _wvmid in beech. Nevertheless, a significant response to the air vapour pressure gradient between leaf and air was translated into stomata closure on an hourly basis, more intensively in beech. Received: 4 March 1999 / Accepted: 21 December 1999     

Gene Flow in European Beech (Fagus sylvatica L.)

Three relatively isolated stands were used to study gene flow in European beech (Fagus sylvatica L.) in Northern Germany. Nine allozyme loci (Got-B, Idh-A, Lap-A, Mdh-B, Mdh-C, Mnr-A, 6-pgdh-A, Pgi-B and Pgm-A) were utilized for multilocus-genotyping adult trees and seeds. Expected heterozygosity (He) ranged from 0.325 to 0.351 for the three stands. F(ST) revealed that there was small differentiation among stands (mean F(ST) = 0.013). The indirect estimates of gene flow (Nm) based on the mean F(ST) were high and the average Nm was 19.14. External gene flow by pollen ranged from 0.7 to 1.2% inferred from new alleles in seed samples. Moreover, paternity analysis was used to assess effective pollen dispersal by inferring paternity of offspring. The weighted mean distances of pollen dispersal for these three stands were 36.8 and 37.1 m based on simple exclusion procedure and most-likely method, respectively. Two of the trees in one stand had rare allozyme alleles (Lap-A1 and Idh-A4, respectively), which were used to directly measure pollen movement away from those trees. The frequency of the rare Lap and Idh alleles in seeds declines as the distance from the source tree increases. The weighted mean distance of pollen dispersal with rare allele Lap-A1 or Idh-A4 was 26.3 m.     

Mesophilous woods with Fagus sylvatica L. of northern Latium (Tyrrhenian Central Italy): synecology and syntaxonomy

ABSTRACT

The results of a synecological and syntaxonomic study of mesophilous forest coenoses, mixed or with a prevalence of Beech, in northern Latium (Roman Tuscia), are reported. The main parameters determining the distribution pattern and floristic composition of these coenoses were altitude, exposure, acclivity, asperity and ecotope. The presence of Aquifolio-Fagetum (Doronico-Fagion) at la Tolfa and in the Sabatini Mts. is confirmed, together with the autonomy of Teucrio siculi-Quercion cerridis and Lathyro-Quercion cerridis in the Cimini-Vicani and Sabatini Mts. and on the sandstone substrates of Mt. Rufeno, respectively. Variants of Polysticho-Fagetum aceretosum pseudoplatani (Galio odorati-Fagenion) with Polystichum setiferum and with Veronica hederifolia were recognized in certain parts of the Cimini and Vicani Mts. The presence of Carpinion betuli, especially with Carpino-Coryletum avellanae and the new subassociation phyllitiaetosum scolopendrii of the latter, is recognized in gullies and ravines eroded in tufo of the Vulsino and Cimino complexes.     

The effects of acclimation to sunlight on the xylem vulnerability to embolism in Fagus sylvatica L.

We assessed the effects of irradiance received during growth on the vulnerability of Fagus sylvatica L. xylem vessels to water-stress-induced embolism. The measurements were conducted on (1) potted saplings acclimated for 2 years under 100% and 12% incident global radiation and (2) branches collected from sun-exposed and shaded sides of adult trees. Both experiments yielded similar results. Light-acclimated shoots were less vulnerable to embolism. Xylem water potential levels producing 50% loss of hydraulic conductivity were lower in sun-exposed branches and seedlings than in shade-grown ones (–3·0 versus –2·3 MPa on average). The differences in vulnerability were not correlated with differences in xylem hydraulic conductivity nor vessel diameter. Resistance to cavitation was correlated with transpiration rates, midday xylem and leaf water potentials in adult trees. We concluded that vulnerability to cavitation in Fagus sylvatica may acclimate to contrasting ambient light conditions.     

Potential impacts of climate change on habitat suitability of Fagus sylvatica L. forests in spain

Abstract

Fagus sylvatica forests are considered to be of Community interest according to Directive 92/43/EEC. Climate change predictions for Spain point to a warming scenario, coupled with decreasing rainfall, which may have an impact on their future distribution particularly at the extremes of its distribution area. Species distribution models incorporating bioclimatic, topographic and phytogeographic variables were used as predictors to assess their habitat suitability under current conditions and a climate change projection. Ten single models were generated and an ensemble-forecasting model was subsequently built by computing a consensus of single-model projections. The results revealed that ombrothermic indices are the main factors controlling the distribution of Spanish beech forests. They are highly vulnerable to climate change and could suffer a decline in their habitat suitability if climate trends observed are maintained in the future. The least favoured areas for them will be located close to the limit between the Temperate and Mediterranean climates, where they could suffer a loss of habitat suitability. Conversely, suitable new areas could be found mainly in western areas of the Cantabrian Range and in the Central Pyrenees.     

Assessment of genetic relationships and population discrimination among Fagus sylvatica L. by RAPD

 We assessed the genetic relationships between members of the Fagaceae family by RAPDs in order to better ascertain the taxonomic status of a very particular population of Fagus sylvatica, the ‘tortuosa’ variety. Intra- and inter-population Nei and Li’s mean genetic distances were compared, and the genetic relationships between individuals were clarified on dendrograms by the Neighbor joining method. RAPD analysis was first conducted on three species from three genera, Quercus petraea, Castanea sativa, and Fagus sylvatica, in order to develop an efficient RAPD protocol. The variety level was then studied, and a general tendency of the individuals to cluster by variety was observed. Individuals also clustered by geographic locations, but the genetic distances between populations were not correlated to the distances between sites. Finally, we compared the common beech and ‘tortuosa’ varieties from two different locations, Verzy and Süntel. Both populations from one location were closer than the same variety from two sites. This last result is in agreement with those previously obtained with isozymes. Hypotheses concerning the origin of the ‘tortuosa’ variety are discussed. Received: 9 January 1998 / Accepted: 25 February 1998     

The effect of ozone on the emission of carbonyls from leaves of adult Fagus sylvatica

Under the site conditions of a temperate forest, the exchange of short-chained oxygenated carbonyls (aldehydes, ketones) was assessed from leaves of adult European beech trees. The crowns of the trees were either exposed to an elevated O₃ regime as released by a free-air fumigation system (2 × O₃) or to the unchanged O₃ regime at the site (1 × O₃, ‘control’). Daily fluctuations of oxygenated carbonyls were quantified in relation to environmental and physiological factors. In particular, the effect of O₃ on carbonyl exchange was studied. Measurements of leaf gas exchange were performed with a dynamic cuvette system, and carbonyl fluxes were determined using 2,4-dinitrophenylhydrazine (DNPH)-coated silica gel cartridges. Leaves mainly emitted acetaldehyde, formaldehyde and acetone. Acetaldehyde dominated the emissions, amounting up to 100 nmol m⁻² min⁻¹, followed by formaldehyde (approximately 80 nmol m⁻² min⁻¹) and acetone (approximately 60 nmol m⁻² min⁻¹). Carbonyl emissions were highest during midday and significantly lowered at night, irrespective of the O₃ exposure regime. Trees exposed to 2 × O₃ emitted acetaldehyde and acetone at enhanced rates. The findings are of particular significance for future climate change scenarios that assume increased O₃ levels.     

Characterization of microsatellite markers in Fagus sylvatica L. and Fagus orientalis Lipsky

Using an enrichment procedure, we cloned microsatellite repeats from European beech (Fagus sylvatica L.) and developed primers for the amplification of microsatellite markers. Six polymorphic loci were characterized which produced 3–21 alleles in 70 individuals from one Italian population, with an observed heterozygosity between 0.58 and 0.85. All six loci amplified fragments which were polymorphic in the closely related species, Fagus orientalis, also. Owing to their very high degree of variation, these markers should be very useful in gene flow studies of these species.     

Fagus sylvatica L. recruitment across a fragmented Mediterranean Landscape, importance of long distance effective dispersal, abiotic conditions and biotic interactions

Native tree populations have been fragmented by anthropogenic disturbance worldwide, leaving them at risk from extinction. The possibility of sizable recovery of fragmented populations is a function of their dispersal, the abiotic conditions, and the biotic interactions. The relative importance of these three drivers for the recruitment rate of a fragmented population of the late-successional Fagus sylvatica L. was analysed at the landscape scale in Causse du Larzac, southern margins of the Massif Central, in the South of France. We used regression models on observational data to analyse the response of Fagus recruitment rate to the distance to the nearest mature population, to climatic and geological variables, and to variables describing biotic interactions (pine vs. grassland, light, shrub cover). Distance to the nearest F. sylvatica adult population was the most important explanatory variable. Recruitment rate was also influenced by facilitative biotic interactions with shrubs, and by the climatic conditions of the plot. Recruitment occurred at a greater distance from the nearest mature population of Fagus in pine forests than in grassland. Dispersal was the major limitation to recruitment of F. sylvatica in this landscape. The recruitment rate was then modulated by the climate and positive biotic interactions. The activity of the European jay could be of great importance for such fragmented populations, because it can lead to long-distance dispersal events and may result in a preferential dispersal towards pine forests.