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The role of future forests in global biogeochemical cycles will depend on how different tree species respond to climate. Interpreting the response of forest growth to climate change requires an understanding of the temporal and spatial patterns of seasonal climatic influences on the growth of common tree species. We constructed a new network of 310 tree-ring width chronologies from three common tree species (Quercus robur, Pinus sylvestris and Fagus sylvatica) collected for different ecological, management and climate purposes in the south Baltic Sea region at the border of three bioclimatic zones (temperate continental, oceanic, southern boreal). The major climate factors (temperature, precipitation, drought) affecting tree growth at monthly and seasonal scales were identified. Our analysis documents that 20th century Scots pine and deciduous species growth is generally controlled by different climate param-eters, and that summer moisture availability is increasingly important for the growth of deciduous species examined. We report changes in the influence of winter climate variables over the last decades, where a decreasing influence of late winter tempera-ture on deciduous tree growth and an increasing influence of winter temperature on Scots pine growth was found. By comparing climate–growth responses for the 1943–1972 and 1973–2002 periods and characterizing site-level growth response stability, a descriptive application of spatial segregation nalysis distinguished sites with stable responses to dominant climate parameters (northeast of the study region), and sites that collectively showed unstable responses to winter climate (southeast of the study region). The findings presented here highlight the temporally unstable and nonuniform responses of tree growth to climate variability, and that there are geo-graphical coherent regions where these changes are similar. Considering continued climate change in the future, our results provide important regional perspectives on recent broad-scale climate–growth relationships for trees across the temperate to boreal forest transition around the south Baltic Sea.

Autumnal leaf senescence signals the end of photosynthetic activities in temperate deciduous trees and consequently exerts a strong control on various ecological processes. Predicting leaf senescence dates (LSD) with high accuracy is thus a prerequisite for better understanding the climate-ecosystem interactions. However, modeling LSD at large spatial and temporal scales is challenging. In this study, first, we used 19972 site-year records (848 sites and four deciduous tree species) from the PAN European Phenology network to calibrate and evaluate six leaf senescence models during the period 1980-2013. Second, we extended the spatial analysis by repeating the procedure across Europe using satellite-derived end of growing season and a forest map. Overall, we found that models that considered photoperiod and temperature interactions outperformed models using simple temperature or photoperiod thresholds for Betula pendula, Fagus sylvatica and Quercus robur. On the contrary, no model displayed reasonable predictions for Aesculus hippocastanum. This inter-model comparison indicates that, contrary to expectation, photoperiod does not significantly modulate the accumulation of cooling degree days (CDD). On the other hand, considering the carryover effect of leaf unfolding date could promote the models' predictability. The CDD models generally matched the observed LSD at species level and its interannual variation, but were limited in explaining the inter-site variations, indicating that other environmental cues need to be considered in future model development. The discrepancies remaining between model simulations and observations highlight the need of manipulation studies to elucidate the mechanisms behind the leaf senescence process and to make current models more realistic.

Leaf phenology is one of the most reliable bioindicators of ongoing global warming in temperate and boreal zones because it is highly sensitive to temperature variation. A large number of studies have reported advanced spring leaf-out due to global warming, yet the temperature sensitivity of leaf-out has significantly decreased in temperate deciduous tree species over the past three decades. One of the possible mechanisms is that photoperiod is limiting further advance to protect the leaves against potential damaging frosts. However, the "photoperiod limitation" hypothesis remains poorly investigated and experimentally tested. Here, we conducted a photoperiod- and temperature-manipulation experiment in climate chambers on two common deciduous species in Europe: Fagus sylvatica (European beech, a typically late flushing species) and Aesculus hippocastanum (horse chestnut, a typically early flushing species). In agreement with previous studies, we found that the warming significantly advanced the leaf-out dates by 4.3 and 3.7 days/degrees C for beech and horse chestnut saplings, respectively. However, shorter photoperiod significantly reduced the temperature sensitivity of beech only (3.0 days/degrees C) by substantially increasing the heat requirement to avoid leafing-out too early. Interestingly, the photoperiod limitation only occurs below a certain daylength (photoperiod threshold) when the warming increased above 4 degrees C for beech trees. In contrast, for chestnut, no photoperiod threshold was found even when the ambient air temperature was warmed by 5 degrees C. Given the species-specific photoperiod effect on leaf phenology, the sequence of the leaf-out timing among forest tree species may change under future climate warming conditions. Nonphotoperiodic species may benefit from warmer springs by starting the growing season earlier than photoperiodic sensitive species, modifying forest ecosystem structure and functions, but this photoperiod limitation needs to be further investigated experimentally in numerous species.

Some forest-related studies on possible effects of climate change conclude that growth potential of European beech (Fagus sylvatica L.) might be impaired by the predicted increase in future serious drought events during the growing season. Other recent research suggests that not only multiyear increment rates but also growth resistance and recovery of beech during, respectively, after dry years may differ between pure and mixed stands. Thus, we combined dendrochronological investigations and wood stable isotope measurements to further investigate the impact of neighborhood diversity on long-term performance, short-term drought response and soil water availability of European beech in three major geographic regions of Germany. During the last four decades, target trees whose competitive neighborhood consisted of co-occurring species exhibited a superior growth performance compared to beeches in pure stands of the same investigation area. This general pattern was also found in exceptional dry years. Although the summer droughts of 1976 and 2003 predominantly caused stronger relative growth declines if target trees were exposed to interspecific competition, with few exceptions they still formed wider annual rings than beeches growing in close-by monocultures. Within the same study region, recovery of standardized beech target tree radial growth was consistently slower in monospecific stands than in the neighborhood of other competitor species. These findings suggest an improved water availability of beech in mixtures what is in line with the results of the stable isotope analysis. Apparently, the magnitude of competitive complementarity determines the growth response of target beech trees in mixtures. Our investigation strongly suggest that the sensitivity of European beech to environmental constrains depends on neighborhood identity. Therefore, the systematic formation of mixed stands tends to be an appropriate silvicultural measure to mitigate the effects of global warming and droughts on growth patterns of Fagus sylvatica.

Forest ecosystems in central Europe are predicted to face an increasing frequency and severity of summer droughts because of global climate change. European beech and Norway spruce often coexist in these forests with mostly positive effects on their growth. However, their different below-ground responses to drought may lead to differences in ectomycorrhizal (ECM) fungal community composition and functions which we examined at the individual root and ecosystem levels. We installed retractable roofs over plots in Kranzberg Forest (11 degrees 39'42 '' E, 48 degrees 25'12 '' N; 490 m a.s.l.) to impose repeated summer drought conditions and assigned zones within each plot where trees neighboured the same or different species to study mixed species effects. We found that ECM fungal community composition changed and the numbers of vital mycorrhizae decreased for both tree species over 3 drought years (2014-2016), with the ECM fungal community diversity of beech exhibiting a faster and of spruce a stronger decline. Mixed stands had a positive effect on the ECM fungal community diversity of both tree species after the third drought year. Ectomycorrhizae with long rhizomorphs increased in both species under drought, indicating long-distance water transport. However, there was a progressive decline in the number of vital fine roots during the experiment, resulting in a strong reduction in enzyme activity per unit volume of soil. Hydrolytic enzyme activities of the surviving ectomycorrhizae were stable or stimulated upon drought, but there was a large decline in ECM fungal species with laccase activity, indicating a decreased potential to exploit nutrients bound to phenolic compounds. Thus, the ectomycorrhizae responded to repeated drought by maintaining or increasing their functionality at the individual root level, but were unable to compensate for quantitative losses at the ecosystem level. These findings demonstrate a strong below-ground impact of recurrent drought events in forests.

International audience; Tree populations at the low-altitudinal or -latitudinal limits of species' distributional ranges are predicted to retreat toward higher altitudes and latitudes to track the ongoing changes in climate. Studies have focused on the climatic sensitivity of the retreating species, whereas little is known about the potential replacements. Competition between tree species in forest ecotones will likely be strongly influenced by the ecophysiological responses to heat and drought. We used tree-ring widths and δ13 C and δ18 O chronologies to compare the growth rates and long-term ecophysiological responses to climate in the temperate-Mediterranean ecotone formed by the deciduous Fagus sylvatica and the evergreen Quercus ilex at the low altitudinal and southern latitudinal limit of F. sylvatica (NE Iberian Peninsula). F. sylvatica growth rates were similar to those of other southern populations and were surprisingly not higher than those of Q. ilex, which were an order of magnitude higher than those in nearby drier sites. Higher Q. ilex growth rates were associated with high temperatures, which have increased carbon discrimination rates in the last 25 years. In contrast, stomatal regulation in F. sylvatica was proportional to the increase in atmospheric CO2 . Tree-ring δ18 O for both species were mostly correlated with δ18 O in the source water. In contrast to many previous studies, relative humidity was not negatively correlated with tree-ring δ18 O but had a positive effect on Q. ilex tree-ring δ18 O. Furthermore, tree-ring δ18 O decreased in Q. ilex over time. The sensitivity of Q. ilex to climate likely reflects the uptake of deep water that allowed it to benefit from the effect of CO2 fertilization, in contrast to the water-limited F. sylvatica. Consequently, Q. ilex is a strong competitor at sites currently dominated by F. sylvatica and could be favored by increasingly warmer conditions.

Elevated temperature has potential to influence the biological mechanisms regulating ecosystem–atmosphere carbon exchange. The relationship between warming and heterotrophic microbial respiration remains poorly understood, not least in terms of the differential sensitivity of microbial groups to temperature and the complexity of interactions with other biota. Cord-forming basidiomycete fungi are dominant primary decomposers in temperate woodland. Decomposition rates are determined by the composition of the decomposer community, ecophysiological relationships between these fungi and abiotic variables and interactions with other organisms. Amongst the latter, a major determinant is the balance between mycelial growth and removal by soil invertebrate grazers, which can themselves be affected by elevated temperature. We investigated the impact of elevated temperature on fungal foraging and decomposition of beech (Fagus sylvatica) wood in soil microcosms to which the invertebrate grazers, Folsomia candida and Protophorura armata (Collembola), were added in factorial combinations with five basidiomycete fungi. Species-specific impacts on mycelial development and function resulted from differential sensitivity of fungi to warming and grazing. Temperature impacts on collembola abundance were resource-specific, causing increased grazing pressure by both species, but on different fungi. Grazing often counteracted warming-induced stimulation of mycelial growth, but occasionally amplified the temperature effect, with implications for colonization rates of new resources. High grazing pressure did not prevent increased fungal-mediated decomposition of colonized wood, as fungi utilized more resource-derived energy to maintain explorative growth. Impacts of elevated temperature on decomposition are likely to depend on local composition of the fungal and invertebrate decomposer community.

When site factors reduce growth rates, tree lifespan tends to increase. This study investigates processes leading to such inverse relationship in Fagus sylvatica stands distributed along two elevation gradients, with an emphasis on climatic response, suppression periods, and growth trends. Dendrochronological records from old-growth beech populations sampled at different elevations within two different bioclimatic regions (Alps vs. Apennines), were used to investigate factors that control tree lifespan. Differences between old-growth (12) and nearby managed (15) stands were used to assess effects of silvicultural practices on maximum age. Logging reduced tree lifespan not only by removing older trees, but also by reducing the number of years beech individuals spent in the shaded understory. Tree lifespan and growth rates were affected by climate (spring–summer temperature) and were inversely related to one another along elevation gradients. The greatest lifespan was observed in old-growth high-mountain populations, and was related not only to slower growth due to a shorter growing season, but also to multidecadal periods of growth suppression during the initial development stages in the understory (i.e., slower growth rates at the youngest cambial ages). Past unfavorable climatic periods (in this case, the Little Ice Age) also helped increase tree lifespan. Using a linear model, we estimated a reduction in beech lifespan of 23 ± 5 years for each degree of warming. Basal area increment of trees with the maximum observed lifespan showed an increasing trend over time. Because growth of old (>300 years) trees has increased in the Alps, while it has recently declined in the Apennines, different bioclimatic regions can have opposite responses to global climatic change. In the next decades, if warming continues, beech lifespan could be reduced in the Alps by faster growth and in the Apennines by drought-induced mortality.

Rising atmospheric carbon dioxide [CO₂] can accelerate tree growth by stimulating photosynthesis and increasing intrinsic water-use efficiency (iWUE). Little evidence exists, however, for the long-term growth and gas-exchange responses of mature trees in tropical forests to the combined effects of rising [CO₂] and other global changes such as warming. Using tree rings and stable isotopes of carbon and oxygen, we investigated long-term trends in the iWUE and stem growth (basal area increment, BAI) of three canopy tree species in a tropical monsoon forest in western Thailand (Chukrasia tabularis, Melia azedarach, and Toona ciliata). To do this, we modelled the contribution of ontogenetic effects (tree diameter or age) and calendar year to variation in iWUE, oxygen isotopes, and BAI using mixed-effects models. Although iWUE increased significantly with both tree diameter and calendar year in all species, BAI at a given tree diameter was lower in more recent years. For one species, C. tabularis, differences in crown dominance significantly influence stable isotopes and growth. Tree ring Δ¹⁸O increased with calendar year in all species, suggesting that increasing iWUE may have been driven by relatively greater reductions in stomatal conductance - leading to enrichment in Δ¹⁸O - than increases in photosynthetic capacity. Plausible explanations for the observed declines in growth include water stress resulting from rising temperatures and El Nino events, increased respiration, changes in allocation, or more likely, a combination of these factors.

Water and nutrient fluxes for single stands of different tree species have been reported in numerous studies, but comparative studies of nutrient and hydrological budgets of common European deciduous tree species are rare. Annual fluxes of water and inorganic nitrogen (N) were established in a 30-year-old common garden design with stands of common ash (Fraxinus excelsior), European beech (Fagus sylvatica L.), pedunculate oak (Quercus robur), small-leaved lime (Tilia cordata Mill.), sycamore maple (Acer pseudoplatanus) and Norway spruce (Picea abies [L.] Karst.) replicated at two sites in Denmark, Mattrup and Vallo during 2 years. Mean annual percolation below the root zone (mm yr−1±SE, n=4) ranked in the following order: maple (351±38)>lime (284±32), oak (271±25), beech (257±30), ash (307±69)≫ spruce (75±24). There were few significant tree species effects on N fluxes. However, the annual mean N throughfall flux (kg N ha−1 yr−1±SE, n=4) for spruce (28±2) was significantly larger than for maple (12±1), beech (11±1) and oak (9±1) stands but not different from that of lime (15±3). Ash had a low mean annual inorganic N throughfall deposition of 9.1 kg ha−1, but was only present at Mattrup. Annual mean of inorganic N leaching (kg ha−1 yr−1±SE, n=4) did not differ significantly between species despite of contrasting tree species mean values; beech (25±9)>oak (16±10), spruce (15±8), lime (14±8)≫ maple (1.9±1), ash (2.0±1). The two sites had similar throughfall N fluxes, whereas the annual leaching of N was significantly higher at Mattrup than at Vallo. Accordingly, the sites differed in soil properties in relation to rates and dynamics of N cycling. We conclude that tree species affect the N cycle differently but the legacy of land use exerted a dominant control on the N cycle within the short-term perspective (30 years) of these stands.

Productivity of old-growth beech forests in the Mediterranean Basin was measured by average stem basal area increment (BAI) of dominant trees at two mountain sites in the Italian Apennines. Both forests could be ascribed to the old-growth stage, but they differed markedly with regard to elevation (1000 vs. 1725m a.s.l.), soil parent material (volcanic vs. calcareous), mean tree age (less than 200 years vs. 300 years), and stand structure (secondary old-growth vs. primary old-growth forest). Drought at the two sites was quantified by the self-calibrated Palmer Moisture Anomaly Index (Z-index), and by the self-calibrating Palmer Drought Severity Index (PDSI) for summer (June through August) and the growing season (May through September). Dendroclimatological analyses revealed a moisture limitation of beech BAI at interannual (water availability measured by Z-index) and decadal scales (water availability measured by PDSI). Both BAI and water availability increased from 1950 to 1970, and decreased afterwards. Trees were grouped according to their BAI trends in auxological groups (growth-type chronologies), which confirmed that growth of most trees at both sites declined in recent decades, in agreement with increased drought. Because BAI is not expected to decrease without an external forcing, the patterns we uncovered suggest that long-term drought stress has reduced the productivity of beech forests in the central Apennines, in agreement with similar trends identified in other Mediterranean mountains, but opposite to growth trends reported for many forests in central Europe.

Drought duration and intensity are expected to increase with global climate change. How changes in water availability and temperature affect the combined plant-soil-microorganism response remains uncertain. We excavated soil monoliths from a beech (Fagus sylvatica L.) forest, thus keeping the understory plant-microbe communities intact, imposed an extreme climate event, consisting of drought and/or a single heat-pulse event, and followed microbial community dynamics over a time period of 28 days. During the treatment, we labeled the canopy with (13) CO2 with the goal of (i) determining the strength of plant-microbe carbon linkages under control, drought, heat and heat-drought treatments and (ii) characterizing microbial groups that are tightly linked to the plant-soil carbon continuum based on (13) C-labeled PLFAs. Additionally, we used 16S rRNA sequencing of bacteria from the Ah horizon to determine the short-term changes in the active microbial community. The treatments did not sever within-plant transport over the experiment, and carbon sinks belowground were still active. Based on the relative distribution of labeled carbon to roots and microbial PLFAs, we determined that soil microbes appear to have a stronger carbon sink strength during environmental stress. High-throughput sequencing of the 16S rRNA revealed multiple trajectories in microbial community shifts within the different treatments. Heat in combination with drought had a clear negative effect on microbial diversity and resulted in a distinct shift in the microbial community structure that also corresponded to the lowest level of label found in the PLFAs. Hence, the strongest changes in microbial abundances occurred in the heat-drought treatment where plants were most severely affected. Our study suggests that many of the shifts in the microbial communities that we might expect from extreme environmental stress will result from the plant-soil-microbial dynamics rather than from direct effects of drought and heat on soil microbes alone.

Studies on Fagus sylvatica show that growth in populations toward the southern limit of this species' distribution is limited strongly by drought. Warming temperatures in the Mediterranean region are expected to exacerbate drought where they are not accompanied by increases in precipitation. We studied levels of annual growth in mature F. sylvatica trees over the last half-century in the Montseny Mountains in Catalonia (northeast Spain). Our results show significantly lower growth of mature trees at the lower limit of this species' distribution when compared with trees at higher altitudes. Growth at the lower Fagus limit is characterized by a rapid recent decline starting in approximately 1975. By 2003, growth of mature trees had fallen by 49% when compared with predecline levels. This is not an age-related phenomenon, nor is it seen in comparable populations at higher altitudes. Analysis of climate-growth relationships suggests that the observed decline in growth is a result of warming temperatures and that, as precipitation in the region has not increased, precipitation is now insufficient to ameliorate the negative effects of increased temperatures on tree growth. As the climate-response of the studied forest is comparable with that of F. sylvatica forests in other southern European regions, it is possible that this growth decline is a more widespread phenomenon. Warming temperatures may lead to a rapid decline in the growth of range-edge populations and a consequent retreat of the species distribution in southern Europe. Assessment of long-term growth trends across the southern range edge of F. sylvatica therefore merits further attention.

Are mature forest trees carbon limited at current CO2 concentrations? Will `mid-life`, 35 m tall deciduous trees grow faster in a CO2-enriched atmosphere? To answer these questions we exposed ca. 100-year-old temperate forest trees at the Swiss Canopy Crane site near Basel, Switzerland to a ca. 540 ppm CO2 atmosphere using web-FACE technology. Here, we report growth responses to elevated CO2 for 11 tall trees (compared with 32 controls) of five species during the initial four treatment years. Tested across all trees, there was no CO2 effect on stem basal area (BA) increment (neither when tested per year nor cumulatively for 4 years). In fact, the 4th year means were almost identical for the two groups. Stem growth data were standardized by pretreatment growth (5 years) in order to account for a priori individual differences in vigor. Although this experiment was not designed to test species specific effects, one species, the common European beech, Fagus sylvatica, showed a significant growth enhancement in the first year, which reoccurred during a centennial drought in the third year. None of the other dominant species (Quercus petraea, Carpinus betulus) showed a growth response to CO2 in any of the 4 years or for all years together. The inclusion or exclusion of single individuals of Prunus avium and Tilia platyphyllos did not change the picture. In elevated CO2, lateral branching in terminal shoots was higher in Fagus in 2002, when shoots developed from buds that were formed during the first season of CO2 enrichment (2001), but there was no effect in later years and no change in lateral branching in any of the other species. In Quercus, there was a steady stimulation of leading shoot length in high-CO2 trees. Phenological variables (bud break, leaf fall, leaf duration) were highly species specific and were not affected by elevated CO2 in any consistent way. Our 4-year data set reflects a very dynamic and species-specific response of tree growth to a step change in CO2 supply. Stem growth after 4 years of exposure does not support the notion that mature forest trees will accrete wood biomass at faster rates in a future CO2-enriched atmosphere.

Rising atmospheric carbon dioxide (CO2) concentration is expected to change plant tissue quality with important implications for plant–insect interactions. Taking advantage of canopy access by a crane and long-term CO2 enrichment (530l mol mol � 1 ) of a natural old-growth forest (web-free air carbon dioxide enrichment), we studied the responses of a generalist insect herbivore feeding in the canopy of tall trees. We found that relative growth rates (RGR) of gypsy moth (Lymantria dispar) were reduced by 30% in larvae fed on high CO2-exposed Quercus petraea, but increased by 29% when fed on high CO2-grown Carpinus betulus compared with control trees at ambient CO2 (370l mol mol � 1 ). In Fagus sylvatica, there was a nonsignificant trend for reduced RGR under elevated CO2. Tree species-specific changes in starch to nitrogen ratio, water, and the concentrations of proteins, condensed and hydrolyzable tannins in response to elevated CO2 were identified to correlate with altered RGR of gypsy moth larvae. Our data suggest that rising atmospheric CO2 will have strong species-specific effects on leaf chemical composition of canopy trees in natural forests leading to contrasting responses of herbivores such as those reported here. A future change in host tree preference seems likely with far-ranging consequences for forest community dynamics.

Recent studies have revealed large unexplained variation in heat requirement-based phenology models, resulting in large uncertainty when predicting ecosystem carbon and water balance responses to climate variability. Improving our understanding of the heat requirement for spring phenology is thus urgently needed. In this study, we estimated the species-specific heat requirement for leaf flushing of 13 temperate woody species using long-term phenological observations from Europe and North America. The species were defined as early and late flushing species according to the mean date of leaf flushing across all sites. Partial correlation analyses were applied to determine the temporal correlations between heat requirement and chilling accumulation, precipitation and insolation sum during dormancy. We found that the heat requirement for leaf flushing increased by almost 50% over the study period 1980-2012, with an average of 30 heat units per decade. This temporal increase in heat requirement was observed in all species, but was much larger for late than for early flushing species. Consistent with previous studies, we found that the heat requirement negatively correlates with chilling accumulation. Interestingly, after removing the variation induced by chilling accumulation, a predominantly positive partial correlation exists between heat requirement and precipitation sum, and a predominantly negative correlation between heat requirement and insolation sum. This suggests that besides the well-known effect of chilling, the heat requirement for leaf flushing is also influenced by precipitation and insolation sum during dormancy. However, we hypothesize that the observed precipitation and insolation effects might be artefacts attributable to the inappropriate use of air temperature in the heat requirement quantification. Rather than air temperature, meristem temperature is probably the prominent driver of the leaf flushing process, but these data are not available. Further experimental research is thus needed to verify whether insolation and precipitation sums directly affect the heat requirement for leaf flushing.

Seedlings of six major European temperate forest tree species (Fagus sylvatica, Acer pseudoplatanus, Quercus robur, Taxus baccata, Abies alba, Pinus sylvestris)were exposed to 360, 500, and 660 µ LC 0 2 L -1 in the understorey of a 120-y-old forest over two growing seasons. Seedlings rooted in the natural forest soil within 36 open-top chambers (12 0TCs per C02 treatment), each with a different known quantum fux density (QFD)ranging from 0.36 to 2.16 mol m -2 d -1 (= 0.8% to 4.8% of full sun). In contrast to a frequent assumption the natural C02 concentration in the understorey is close to the ambient concentration in the free atmosphere during daytime. The C02effect on seedling growth differed greatly among species and was strongly codetermined by microsite-specifc QFD. Biomass production in the deep-shade tolerant species Fagus and Taxus increased by 73% and 37% under elevated C02 in low QFD microsites but was not signifcantly different among C02-treatments in high QFD microsites. The less shade-tolerant species Acer, Quercus, and Abies showed no signifcant response to elevated C02 in low QFD microsites, but increased their biomass by 39%, 25%, and 55% in high QFD microsites. In the shade-intolerant Pinus, seedling survival was too low for a safe conclusion. 0ur data showed that the largest relative responses to increasing C02 occurred at a comparatively small increase from 360 to 500 µ LL -1 with only small and non-signifcant changes with a further increase to 660 µ LL -1 . Subtle shifts in the availability of light can totally reverse interspecifc differences in the C02 response. Given these different responses, we conclude that increasing atmospheric C02 is likely to induce changes in species composition of temperate forests due to altered chances of recruitment. However, these shifts will depend on light patterns in the understorey, and thus on canopy structure, disturbance patterns and forest management.

The phenology of Fagus sylvatica was unaffected by exposure to an atmosphere of elevated CO2 (600 μL L-1) after two years of fumigation. Non-significant changes in nitrogen and phenolic content of the leaves decreased the nutritional status of beech for female larvae in elevated CO2 such that they responded by eating in a compensatory manner; males were unaffected. Rates of development, mortality and adult biomass of Rhynchaenus fagi were no different from those in ambient CO2 concentrations (355 μL L -1). It is possible that, with the changes in leaf chemistry affecting the females, fecundity will be altered, with important consequences for populations of beech weevil.

As a consequence of land-use change and the burning of fossil fuels, atmospheric concentrations of CO2 are increasing and altering the dynamics of the carbon cycle in forest ecosystems. In a number of studies using single tree species, fine root biomass has been shown to be strongly increased by elevated CO2 . However, natural forests are often intimate mixtures of a number of co-occurring species. To investigate the interaction between tree mixture and elevated CO2 , Alnus glutinosa, Betula pendula and Fagus sylvatica were planted in areas of single species and a three species polyculture in a free-air CO2 enrichment study (BangorFACE). The trees were exposed to ambient or elevated CO2 (580 μmol mol(-1) ) for 4 years. Fine and coarse root biomass, together with fine root turnover and fine root morphological characteristics were measured. Fine root biomass and morphology responded differentially to the elevated CO2 at different soil depths in the three species when grown in monocultures. In polyculture, a greater response to elevated CO2 was observed in coarse roots to a depth of 20 cm, and fine root area index to a depth of 30 cm. Total fine root biomass was positively affected by elevated CO2 at the end of the experiment, but not by species diversity. Our data suggest that existing biogeochemical cycling models parameterized with data from species grown in monoculture may be underestimating the belowground response to global change.

Two simplifying hypotheses have been proposed for whole-plant respiration. One links respiration to photosynthesis; the other to biomass. Using a first-principles carbon balance model with a prescribed live woody biomass turnover, applied at a forest research site where multidecadal measurements are available for comparison, we show that if turnover is fast the accumulation of respiring biomass is low and respiration depends primarily on photosynthesis; while if turnover is slow the accumulation of respiring biomass is high and respiration depends primarily on biomass. But the first scenario is inconsistent with evidence for substantial carryover of fixed carbon between years, while the second implies far too great an increase in respiration during stand development – leading to depleted carbohydrate reserves and an unrealistically high mortality risk. These two mutually incompatible hypotheses are thus both incorrect. Respiration isnotlinearly related either to photosynthesis or to biomass, but it is more strongly controlled by recent photosynthates (and reserve availability) than by total biomass.

Masting-temporally variable seed production with high spatial synchrony-is a pervasive strategy in wind-pollinated trees that is hypothesized to be vulnerable to climate change due to its correlation with variability in abiotic conditions. Recent work suggests that aging may also have strong effects on seed production patterns of trees, but this potential confounding factor has not been considered in previous times series analysis of climate change effects. Using a 54 year dataset for seven dominant species in 17 forests across Poland, we used the proportion of seed-producing trees (PST) to contrast the predictions of the climate change and aging hypotheses in Abies alba, Fagus sylvatica, Larix decidua, Picea abies, Pinus sylvestris, Quercus petraea, and Quercus robur. Our results show that in all species, PST increased over time and that this change correlated most strongly with stand age, while the standardized precipitation-evapotranspiration index, a measure of drought, contributed to temporal trends in PST of F. sylvatica and Q. robur. Temporal variability of PST also increased over time in all species except P. sylvestris, while trends in temporal autocorrelation and among-stand synchrony reflect species-specific masting strategies. Our results suggest a pivotal role of plant ontogeny in driving not only the extent but also variability and synchrony of reproduction in temperate forest trees. In a time of increasing forest regrowth in Europe, we therefore call for increased attention to demographic effects such as aging on plant reproductive behavior, particularly in studies examining global change effects using long-term time series data.