Drought matters - Declining precipitation influences growth of Fagus sylvatica L. and Quercus robur L. in north-eastern Germany

For north-eastern Germany regional climate models project rising temperatures in combination with decreasing summer and increasing winter precipitation. The resulting overall drier conditions during the growing season will considerably impact forest growth there. We evaluate the consequences of increasing drought on the growth of the two locally most important broadleaf tree species common beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.). Three mixed forests of beech and oak were sampled along a west-east gradient of declining precipitation. In total we used 257 ring-width samples from 133 trees to build six species and site specific chronologies. Additionally, we modelled the soil water budget for each site. We performed continuous and discontinuous (pointer year) analysis of climate-tree-growth relationships with particular emphasis on inter-annual-variations and their dependence upon climatic factors (temperature, precipitation, soil moisture) and on the stability of the obtained relationships. Results of climate-growth correlations together with pointer year analysis indicate a strong dependency of growth of both species from water availability, especially during early summer (June and July). General correlation pattern between growth and climate are similar for both species, but climate sensitivity of beech is generally higher. We identified drought as the main driver of negative growth depressions in both species. Increasing drought stress along the gradient is expressed in higher correlations to climatic variables, higher sensitivity (variance) of growth, and a higher number of negative pointer years for both species. For beech we also found a significant trend of decreasing average growth rates along the gradient. Growth superiority of beech compared to oak declines with decreasing precipitation. The relationships were generally stable throughout the 20th century. A rise of sensitivity together with a higher frequency of negative pointer years during the last decades suggests that increasing climatic variability together with rising temperatures might be influencing growth of Fagus at the more humid sites. If we substitute space by time it seems that already small changes in precipitation regime can have considerable impact, especially on the growth of beech. Other, more drought tolerant species like oak might gain competitive advantages under the projected climatic changes.     

The north-eastern distribution range of European beech a review

Today, European beech (Fagus sylvatica L.) seems to be a markedlysuccessful tree species in the north-east of its distributionrange. The distribution area may be larger than originally assumed;past forest management is probably the main cause of the contractionin the postglacial European beech range. Numerous attempts consistentlyhave failed to locate a distinct distribution edge for Europeanbeech. Therefore, we define northern and eastern Poland andthe southern regions of the Baltic States as margins of Europeanbeech distribution. Ecophysiological approaches have identifiedthe drought constraints for European beech in terms of (1) thecritical limit for xylem cavitation and loss of hydraulic conductivity,reached at a shoot water potential of –1.9 MPa, and (2)a reduction in gross primary production and total ecosystemrespiration when relative extractable soil water reaches 40and 20 per cent, respectively. However, it is difficult to correlateEuropean beech distribution margins with single macro-climaticfactors. Moreover, the adaptation of European beech populationsand provenances to drought and frost varies. The phenotypicplasticity and evolutionary adaptability of European beech appearto be underestimated. These characteristics may counteract afurther contraction of the European beech range arising fromclimate change in the future.     

Seasonal and interannual ecophysiological responses of beech (Fagus sylvatica) at its south-eastern distribution limit in Europe

Due to its wide European distribution and its drought-susceptibility, beech (Fagus sylvatica L.) received intensive attention recently in the light of global warming. Contrary to central European beech ecosystems, little is known about the ecophysiology of beech at its south-eastern European distribution limit. Here we tested whether climatic fluctuations during a three-year period affected the ecophysiology of a beech site in Greece. Attention was paid at comparing our findings to the intense effects the 2003 extreme drought had on beech forests in central Europe.     

Growth–climate relationship of European beech at its northern distribution limit

In general, trees growing at or near their limit of distribution are more sensitive to climate than those growing at their ecological core. Here we examined the growth–climate relationship of European beech (Fagus sylvatica L.) close to its northern distribution limit in southern Sweden. Tree-ring width chronologies were developed from four well-separated sites where the species was dominant (Djupeåsen, DJ; Baldringe, BLD; Komperskulla, KSK and Ryssberget Nature Reserve, RYSS). The chronologies extended from 52 years (BLD) to 150 years (RYSS). Significant negative relationships were found between tree growth and previous summer (July and August) temperatures at three sites. July temperature of the year of growth had a negative relationship with beech growth at BLD and DJ. In contrast, current summer (July and August) precipitation was positively correlated with beech growth at DJ and KSK. This sensitivity of European beech to drought at its northern limit is in line with the previous research. However, following the exceptionally dry summer in southern Sweden in 1970, a marked growth decrease was noted as well as a shift in the relationship between beech growth and current growing-season temperature. Our results show that that the radial growth of European beech has become more sensitive to drought and precipitation than temperature at its northern distribution limit in the last several decades.     

A comparison of litterfall and element fluxes in even aged Norway spruce,sitka spruce and beech stands in Denmark

Litterfall was investigated in three even-aged Norway spruce (Picea abies), sitka spruce (Picea sitchensis) and beech (Fagus sylvatica) stands on a nutrient poor-soil in Southern Denmark. Dry weights and N, P, K, S, Mg, Ca, Na, Al, and Fe concentrations and fluxes were examined in litterfall fractions. Foliage litter amounted to 90% of total litterfall. The tree stands showed a similar mean annual litterfall. In the spruce stands, annual litterfall was correlated negatively with the current year increment and positively with the previous year increment. Annual litterfall in beech was constant during the 6 study years whereas Norway spruce and sitka spruce showed large fluctuations between years caused by drought, spruce aphid infestations and probably sea salt stress. Norway spruce responded with a long lasting elevated needle loss. Sitka spruce responded to infestations with premature needle loss during short periods. The presence of a large syrphid (Coccinellidae) population was important in regulating aphid (Elatobium abietinum) population density. The between-year variation in element concentrations of litterfall was small whereas variations during the year were large. Interspecific levels were recognized: Norway spruce>beech>sitka spruce. High concentrations in Norway spruce were ascribed to a combination of drought, sea salt stress and elevated transpiration. In sitka spruce, aphid infestations reduced the litterfall N content. Sitka spruce showed the smallest amount of base cation fluxes with litterfall. In contrast, spruce and beech exhibited even litterfall element fluxes. Litterfall studies revealed reduced vitality in the non-native spruce stands and underlined the perception of a healthy stand of native beech.     

Compared effects of competition by grasses (Graminoids) and broom (Cytisus scoparius) on growth and functional traits of beech saplings (Fagus sylvatica)

? The effect of two weeds (grasses and broom, Cytisus scoparius) competition on the growth and functional traits of European beech saplings (Fagus sylvatica) was investigated in an experimental plantation in the French Massif central.

? We hypothesized that grasses would have a much more harmful effect than broom on beech growth through strong competition for soil water and nitrogen.

? A randomized block design was used with three separate blocks, each possessing three types of vegetation; grasses (mainly Agrostis capillaris, Holcus mollis and H. lanatus), broom, and bare soil as a control.

? Two years after plantation beech sapling growth was significantly greater in bare soil than with competition from grasses, and was intermediate on broom. Growth was positively correlated with beech leaf nitrogen content (N_m) and xylem water potential (Ψ_x). N and Ψ_x were dependent, with nitrogen uptake probably decreasing with increasing drought. There was no clear correlation between growth and light transmitted 10 cm below the apex of the saplings.

? Beech leaf mass on an area basis (LMA) was correlated with the three resource variables (water, nitrogen and light) and with growth, confirming that LMA is a sensitive functional trait integrating the degree of stress experienced by the saplings.

? The results suggest that broom is less competitive than grasses and, if properly managed on a sufficiently well-watered site, can generate satisfactory conditions for beech establishment and growth.

     

Differential response to soil drought among co-occurring broad-leaved tree species growing in a 15- to 25-year-old mixed stand

? Introduction

In the context of global environmental changes, better understanding of tree response to soil drought in young mixed species stands is needed to anticipate forest adaptation and management practices for the future.

? Materials and methods

We investigated the functional response of five co-occurring broad-leaved tree species growing in a 15- to 25-year-old mixed stand in northeastern France during the 2006 summer drought. We measured functional traits related to water acquisition (phenology, rooting pattern and vulnerability of xylem to cavitation) and the ecophysiological response (sap flux density, leaf water potential) of these species to soil water shortage.

? Results

Our study highlights contrasted drought response strategies among these species and a trade-off between leaf phenology, resistance of xylem to cavitation and root system depth.

? Conclusion

At this site, a deep root system seemed to be a key functional trait for the species to cope with drought. Quercus robur and Salix capreae can be characterised as drought-avoidance species as they possess a deep root system and therefore did not strongly experience soil drought. Despite deep rooting capacity, Betula pendula did not really avoid soil drought and strongly regulated transpiration during dry periods. Nevertheless, the earliness of budburst of this species contributes to high annual growth rate. In contrast, Carpinus betulus and Fagus sylvatica both displayed typical characteristics of drought-sensitive species.     

Thinning increases drought tolerance of European beech: a case study on two forested slopes on opposite sides of a valley

European beech (Fagus sylvatica L.) is one of the economically most important broadleaved tree species in Central Europe. However, beech shows high drought sensitivity and calls for profound research to test its ability to cope with limited water resources. Here, we investigated the drought tolerance of beech to the 2003 drought as influenced by Kraft class, aspect and thinning intensity. Annual basal area increment data of 126 sample trees from southwest Germany were used to assess the variability of drought tolerance indices, by comparing three social classes (predominant, dominant and co-dominant), two contrasting sites [a dry southwest (SW) aspect and a moist northeast (NE) aspect], and three treatments [control, strong thinning (stand basal area 15 m² ha^?1) and very strong thinning (stand basal area 10 m² ha^?1)] in mature beech stands. Our results show that the co-dominant and dominant trees had lower growth recovery and lower growth resilience after the drought, compared to the predominant trees. The differences between aspects pointed to a growth–drought tolerance trade-off, in which trees on the SW aspect displayed lower growth rates but higher resilience indices than trees on the moist NE aspect. Furthermore, our results suggest that the resistance to and resilience after the 2003 drought significantly increased for the thinned trees. Our results provide novel insights into the linkage between the forest stand management and drought tolerance of beech under contrasting sites. We conclude that thinning can partially alleviate effects of severe drought on European beech forests in southwest Germany and can be applied as an adaptive measure to increase the mitigation potential of beech stands.     

Genetic structure and variability of phenological forms in the European beech (Fagus sylvatica L.)

? Microsatellite markers were used to describe the genetic structure and variability of early, intermediate and late phenological forms of European beech (Fagus sylvatica L.). Two hundred and seventy individuals from three populations located in southern Poland were divided into three forms according to the phenological criterion — bud burst, and analyzed for allelic variation at five highly polymorphic microsatellite loci.

? Population differentiation was moderate and differed significantly among phenological forms. Average values of F _ST and R _ST decreased across phenological forms and amounted to F _ST values of 0.135, 0.110 and 0.108 and R _ST values of 0.365, 0.231 and 0.098 for early, intermediate and late forms of beech, respectively.

? Analysis of Molecular Variance (AMOVA) revealed different genetic structures characteristic of respective phenological forms of beech. The amount of within-population variability increased with the delay of the beginning of vegetation and amounted to 64%, 77% and 90% of total variability, depending on phenological form. A similar trend was found in average pairwise genetic distance between individuals belonging to a given phenological form (11.78, 11.85 and 12.22, from early to late forms).

? Our results demonstrate the importance of late spring frosts as a factor influencing the genetic structure of beech, and as a cause of the decrease in genetic variability as well as the increase in population differentiation proportional to the degree of phenological earliness.

     

Effects of drought and rewatering on growth and transpiration in European beech seedlings late in the growing season

Drought stress is one of the most important environmental factors affecting plant growth and survival. To date, most studies aim at understanding of post-stress physiological and anatomical adaptation to drought stress; however only few studies focus on plant recovery. In the present study, transpiration, shoot water potential, and anatomical and morphological measurements were performed on 4-year-old European beech seedlings with fully developed leaves. The seedlings were exposed to three levels of soil water potential (well-watered, moderate drought stress and severe drought stress) and followed by rewatering under greenhouse conditions. Reduced transpiration rates were observed in the stressed seedlings as a response to drought stress, whereas anatomical and morphological variables remained unchanged. Three days after rewatering, transpiration rates in both moderately and severely stressed seedlings recovered to the levels of those of well-watered seedlings. Drought stress promoted leaf budding, resulting in higher shoot dry mass of stressed seedlings. Our findings indicate that anatomical and morphological adaptations of European beech seedlings to drought stress are visibly limited during late-season growth stages. These results will help us to further understand factors involved in drought adaptation potential of European beech seedlings faced with expected climate-related environmental changes. To complete our findings, further experiments on plant recovery from drought stress should be focused on different periods of growing season.     

Different reactions of central and marginal provenances of Fagus sylvatica to experimental drought

Climate extremes are expected to increase in frequency and magnitude as a consequence of global warming, threatening the functioning, services and goods of forest ecosystems. Across Europe, the ecologically and economically important tree species Fagus sylvatica is expected to suffer particularly under such conditions. The regional introduction of provenances from drier and warmer climates is one option to adapt beech forest ecosystems to these adverse effects of climate change. Marginal populations from the drought-prone southern and north-eastern edges of the species’ distribution come into focus in search of suitable candidates for Central European deciduous forests. Here, we test three marginal provenances (Spain, Bulgaria and Poland) and three provenances from the centre of the distribution range (Germany) for their response to drought in two different soil types (sand, loam) in a full factorial common garden experiment in Landau, Germany. Drought impacted all growth parameters negatively (leaf damage +22 % (percentage points), height ?40 % and diameter increment ?41 %), and the sandy substrate exacerbated this effect. However, provenances differed in their response to drought and soil type. Evidence for a local adaptation to summer drought was detected, especially in terms of mortality rates. The Bulgarian and Spanish provenance showed a stable performance under drought conditions (BG ?27 % in diameter increment; ES ?32 %), compared to the Polish (?48 %) or the most sensitive German provenances (?57 %), yet for Bulgaria on a low level of total increment. This may indicate a trade-off between drought tolerance and growth. Therefore, a sole focus on drought-resistant marginal provenances seems to not be conducive, as they might be less adapted to other climatic factors, e.g. frost, as well. However, intermixed with local Central European provenances, these may act as functional insurance in future drought-prone forest stands.     

A recent growth increase of European beech (Fagus sylvatica L.) at its Mediterranean distribution limit contradicts drought stress

Future changes in tree growth, associated with a warmer and drier climate, are predicted for many species and locations across the European Mediterranean Basin. However, quantification of the intensity and severity of related consequences for forest ecosystem functioning and productivity remains challenging. Species-specific distribution limits that are particularly sensitive to small changes in the ambient climate may provide an ideal test bed to assess the nature of past growth trends and extremes and their responsible controls. Here, we seek to understand how twentieth century climate change affected the growth of European beech (Fagus sylvatica L.) nearby its south-eastern distribution limit in Albania and Macedonia on the Balkan Peninsula. We sampled 93 living trees from undisturbed mixed forest stands at ~1,450 m a.s.l. and 29 timbers from nearby historical buildings. Application of different tree-ring detrending techniques allowed robust composite chronologies with varying degrees of high- to low-frequency variability to be developed back to 1648 ad. Comparison with local meteorological station measurements and continental grid-box climate indices revealed spatiotemporal instability in growth–climate response patterns. Nevertheless, year-to-year and decadal-long fluctuations in radial beech growth were significantly (P < 0.001) negatively correlated at ?0.61 with June–September temperature over the 1951–1995 period. This (inverse) relationship between increased beech growth and decreased summer temperature is somewhat indicative for the importance of plant-available soil moisture, which likely controls ring width formation near the species-specific south-eastern distribution limit. Significant positive correlations between beech growth and drought (scPDSI; r = 0.57) confirm metabolistic drought constraints. However, an unexpected late twentieth century growth increase not only contradicts the previously observed growth dependency to summer soil moisture, but also denies any putative drought-induced forest ecosystem suppression in this part of the Mediterranean Basin.     

Biscogniauxia nummularia infecting beech (Fagus sylvatica) trees and sympatric plants of the sedge Carex brevicollis

Biscogniauxia nummularia is known for its association with beech (Fagus sylvatica), on which it occurs as a saprotroph and a pathogen causing strip cankers following water stress. This fungus has also been reported as a dominant endophytic species in plants of the sedge Carex brevicollis growing in the understory of beech forests and adjacent grasslands in Sierra de Urbasa (Navarre, Spain). In this area, stromata of B. nummularia were observed in dead and living wood of beech trees at several locations where plants of C. brevicollis also contained the fungus as an endophyte. Pure cultures obtained from stromata of B. nummularia on F. sylvatica trees were compared to endophytic isolates from symptomless C. brevicollis. Culture morphology and micromorphology as well as rDNA sequences of the internal transcribed spacer (ITS) regions were identical, suggesting that B. nummularia from beech can also live endophytically in C. brevicollis. It is unknown whether the endophytic strains of Carex might have a role as an inoculum source for the infection of beech trees, or whether they represent a dead end in the life cycle of the fungus.     

Are marginal beech (<Emphasis Type="Italic">Fagus sylvatica</Emphasis> L.) provenances a source for drought tolerant ecotypes?

Due to the expected increases of number and intensity of summer droughts in Central Europe the identification of drought tolerant ecotypes becomes more important in future forestry. A common garden experiment with seedlings of Fagus sylvatica provenances from the center (Germany) and eastern margin (Poland) of the species’ distribution range was conducted. Responses of morphological, physiological, chemical and growth parameters to three drought treatments were analyzed. Relative growth rates of the marginal provenance were lower as compared to the central provenance. The marginal seedlings showed a tendency to higher total biomasses because of higher seed masses. In both provenances drought decreased biomass production and root/shoot ratio which was lower in the central provenance. A lower specific root area of the marginal provenance indicated a better adaptation to low xylem water potentials. Under moderate drought, lower leaf δ¹³C signatures may indicate lower stomatal limitation (or a reduced rate of CO₂ assimilation) in the marginal provenance. We conclude that marginal beech provenances may exhibit a better drought adaptation.     

Beech in Wessex - a Perspective on Present Health and Silviculture

The condition of beech (Fagus sylvatica L) growing in thirtywoods in the south of England is described. In older woods,up to 20 per cent of trees were killed or seriously damagedby the summer drought of 1976. Most surviving trees have recoveredfully. Beech snap disease, associated with the fungus Nectria coccineais widespread in younger stands, especially those also affectedby lime-induced chlorosis, beech scale (Cryptococcus fagisuga)or severe competition. Nevertheless, sufficient trees surviveto provide a choice of final crop. In the long term, beech doesnot seem likely to die out through drought or disease. Trees of almost all ages and conditions exhibit a tendency tomultiple forking. The principle remedy is to keep trees in closedcanopy until a 6 to 8 m clean bole has been obtained. Beechresponds well to heavy thinning at this stage. The long term future of beech woodlands depends on timber valuesand on support for their ecological and amenity value. Silviculturally,beech is the best adapted species for the shallowest calcareoussoils, especially on the Jurassic limestone.     

Genetic variation of beech (Fagus sylvatica L.) in Rodopi (N.E. Greece)

The recent taxonomic classification of beech in Europe considers existence of one species (Fagus sylvatica L.) with two subspecies: F. sylvatica ssp. sylvatica and F. sylvatica ssp. orientalis. Four beech populations growing on the Greek part of the Rodopi Mountains were studied using morphological traits as well as DNA molecular markers (AFLPs and chloroplast DNA SSR). The aim of the study was to describe the variation patterns of beech in the Rodopi Mountains and to test the hypothesis of possible introgression between the beech subspecies’ sylvatica and orientalis in this area. Both morphological traits and gene markers revealed a possible influence of F. orientalis on the east side of Rodopi and at the low elevations, while characters resembling F. sylvatica were observed mainly on the western part of the mountains and in higher altitudes. There was a clinal increase of genetic diversity from the west to the east, reaching a level firstly reported for beech populations. These results can be explained either by the existence of a main refugial area for beech during the last glaciation or by the occurrence of a recent hybridization among the subspecies, which were spatially isolated during the last glaciation and came into reproductive contact during their postglacial remigration. These two scenarios are not necessarily mutually exclusive.     

Forest transpiration—targeted through xylem sap flux assessment versus hydrological modeling

The assessment of forest transpiration rates is crucial for determining plant-available soil water consumption and drought risk of trees. Xylem sap flux measurements have been used increasingly to quantify stand transpiration in forest ecosystems. Here, we compare this empirical approach with hydrological modeling on the basis of a stand transpiration dataset of adult beech (Fagus sylvatica), which was acquired across Bavaria, Germany, at eight forest sites. Xylem sap flux sensors were installed in five dominant trees each. Two tree to stand upscaling approaches, related to site-specific (1) sapwood area or (2) to leaf area index, were compared. The outcome was examined each in relation to process-based stand hydrological modeling, using LWF-BROOK90. Distinct relationships between tree diameter at breast height (1.30 m) and sapwood area-weighted sap flux along the radial profile became apparent across the study sites, confirming a generic allometric basis for stand-level upscaling of transpiration. The two upscaling approaches did not differ in outcome, representatively covering stand structure for comparison with modeling. Differential analysis yielded high agreement between the empirical and modeling approaches throughout most of the study period, although LWF-BROOK90 tended to overestimate sap flux measurements under low soil moisture. The two empirical approaches proved reliable for even-aged beech stands, as performance under high stand-structural heterogeneity awaits clarification. Findings advance stand-level hydrological modeling regarding coverage of stomatal behavior during temporary limitation in water availability.     

Large-scale genetic structure and drought-induced effects on European Scots pine (Pinus sylvestris L.) seedlings

Seedlings of ten provenances of Scots pine (Pinus sylvestris L.) from the southwestern to the central part of the species distribution range were investigated in a greenhouse experiment under dry conditions compared to a well-watered control. We conducted an isozyme study and recorded phenology as well as growth traits during the first year of growth. Genetic variability and genetic diversity increased from the westernmost provenances to those central to the distribution. Provenances from the Apennines and Spain revealed the strongest genetic differentiation compared to all others, whereas populations from Germany, Poland and Bulgaria were found to belong to the same gene pool. Seedling development and bud set were faster in northern populations. Shoot length was highest for Polish and German provenances within both regimes, populations from France and Spain had the longest roots. Reduced soil moisture slowed later development stages and delayed bud set for all provenances by about 5 days. Shoot growth decreased considerably under the dry regime for all provenances. German provenances were the most sensitive to drought, whereas Spanish provenances showed adaptation to drought conditions indicated by the lowest reduction in shoot growth relative to optimum conditions. The results of this pilot study show that under more pronounced drought conditions with repeated drought events, the so far established superiority of northeastern provenances compared to southwestern ones could diminish in the future.     

Modelling self-pruning and branch attributes for young Quercus robur L. and Fagus sylvatica L. trees

The aim of this study was to develop statistical models for first order branchiness in young planted forest stands of pedunculate oak (Quercus robur L.) and European beech (Fagus sylvatica L.), and to give an ecological and silvicultural interpretation to these models. The reported models focus on the lower most-valuable stem part (i.e. until 6 m height), and cover different tree development classes to capture the development of branchiness over time. For each species 30 study plots were selected spread over two nearby forests in Flanders (northern Belgium), minimising site and genetic variability. Branches were counted on a total of 399 oak and 376 beech trees. On a subsample of 30 trees per species (one tree per plot), detailed non-destructive branch measurements were performed, yielding data for 555 oak and 438 beech branches. For both species, models for tree self-pruning (i.e. total branch number and dead branch portion), branch mortality and branch architecture (i.e. branch diameter and branch insertion angle) were built. A generalised linear mixed modelling approach was adopted. The models for total branch number and dead branch portion may be interpreted in terms of four processes contributing to self-pruning: (1) stand and tree development, (2) tree competitive status, (3) stand density and (4) site humidity. The reported models reveal similar self-pruning rates in oak and beech, but with different driving factors: early branch dying and slow shedding for oak and the other way around for beech. Mortality of individual branches is further determined by branch position and branch dimension. Branch diameter and branch insertion angle of both species are mainly related to branch cord length and relative branch position. All modelled effects are consistent with known ecological and ecophysiological processes. Silvicultural implications for stand establishment and early tree selection are discussed. The reported models can be used to fine-tune operational silvicultural choices for quality timber production. This is a first step towards the integration of branchiness models for oak and beech into forest growth simulators.