Resistance of European tree species to drought stress in mixed versus pure forests: evidence of stress release by inter‐specific facilitation

While previous studies focused on tree growth in pure stands, we reveal that tree resistance and resilience to drought stress can be modified distinctly through species mixing. Our study is based on tree ring measurement on cores from increment boring of 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in South Germany, with half sampled in pure, respectively, mixed stands. Indices for resistance, recovery and resilience were applied for quantifying the tree growth reaction on the episodic drought stress in 1976 and 2003. The following general reaction patterns were found. (i) In pure stands, spruce has the lowest resistance, but the quickest recovery; oak and beech were more resistant, but recover was much slower and they are less resilient. (ii) In mixture, spruce and oak perform as in pure stands, but beech was significantly more resistant and resilient than in monoculture. (iii) Especially when mixed with oak, beech is facilitated. We hypothesise that the revealed water stress release of beech emerges in mixture because of the asynchronous stress reaction pattern of beech and oak and a facilitation of beech by hydraulic lift of water by oak. This facilitation of beech in mixture with oak means a contribution to the frequently reported overyield of beech in mixed versus pure stands. We discuss the far‐reaching implications that these differences in stress response under intra‐ and inter‐specific environments have for forest ecosystem dynamics and management under climate change.     

Site‐adapted admixed tree species reduce drought susceptibility of mature European beech

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.     

Contrasting resistance and resilience to extreme drought and late spring frost in five major European tree species

Extreme climate events (ECEs) such as severe droughts, heat waves, and late spring frosts are rare but exert a paramount role in shaping tree species distributions. The frequency of such ECEs is expected to increase with climate warming, threatening the sustainability of temperate forests. Here, we analyzed 2,844 tree‐ring width series of five dominant European tree species from 104 Swiss sites ranging from 400 to 2,200 m a.s.l. for the period 1930–2016. We found that (a) the broadleaved oak and beech are sensitive to late frosts that strongly reduce current year growth; however, tree growth is highly resilient and fully recovers within 2 years; (b) radial growth of the conifers larch and spruce is strongly and enduringly reduced by spring droughts—these species are the least resistant and resilient to droughts; (c) oak, silver fir, and to a lower extent beech, show higher resistance and resilience to spring droughts and seem therefore better adapted to the future climate. Our results allow a robust comparison of the tree growth responses to drought and spring frost across large climatic gradients and provide striking evidence that the growth of some of the most abundant and economically important European tree species will be increasingly limited by climate warming. These results could serve for supporting species selection to maintain the sustainability of forest ecosystem services under the expected increase in ECEs.     

Phenology and growth of Fagus sylvatica and Quercus robur seedlings in response to temperature variation in the parental versus offspring generation

• Plants are known to respond to warming temperatures. Few studies, however, have included the temperature experienced by the parent plant in the experimental design, in spite of the importance of this factor for population dynamics.
• We investigated the phenological and growth responses of seedlings of two key temperate tree species (Fagus sylvatica and Quercus robur) to spatiotemporal temperature variation during the reproductive period (parental generation) and experimental warming of the offspring. To this end, we sampled oak and beech seedlings of different ages (1–5 years) from isolated mother trees and planted the seedlings in a common garden.
• Warming of the seedlings advanced bud burst in both species. In oak seedlings, higher temperatures experienced by mother trees during the reproductive period delayed bud burst in control conditions, but advanced bud burst in heated seedlings. In beech seedlings, bud burst timing advanced both with increasing temperatures during the reproductive period of the parents and with experimental warming of the seedlings. Relative diameter growth was enhanced in control oak seedlings but decreased with warming when the mother plant experienced higher temperatures during the reproductive period.
• Overall, oak displayed more plastic responses to temperatures than beech. Our results emphasise that temperature during the reproductive period can be a potential determinant of tree responses to climate change.

     

Role of six European tree species and land‐use legacy for nitrogen and water budgets in forests

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 Vallø 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 Vallø. 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.     

Biomass partitioning and growth efficiency in four naturally regenerated forest tree species

Current forest growth models and yield tables are almost exclusively based on data from mature trees, reducing their applicability to young and developing stands. To address this gap, young European beech, sessile oak, Scots pine and Norway spruce trees approximately 0–10 yr old were destructively sampled in a range of naturally regenerated forest stands in Central Europe. Diameter at base and height was first measured in situ for up to 175 individuals per species. Subsequently, the trees were excavated and dry biomass of foliage, branches, stems and roots was measured. Allometric relations were then used to calculate biomass allocation coefficients (BAC) and growth efficiency (GE) patterns in young trees. We found large differences in BAC and GE between broadleaves and conifers, and also between species within these categories. Both BAC and GE are strongly age-specific in young trees, their rapidly changing values reflecting different growth strategies in the earliest stages of growth. We show that linear relationships describing biomass allocation in older trees are not applicable in young trees. To accurately predict forest biomass and carbon stocks, forest growth models need to include species and age-specific parameters of biomass allocation patterns.     

Impact of waterlogging on the N-metabolism of flood tolerant and non-tolerant tree species

The present study was conducted to characterize the N‐metabolism of important European tree species with different degrees of flooding tolerance. The roots of Fagus sylvatica (sensitive to flooding), Quercus robur (moderately flood tolerant) and Populus tremula × P. alba (flood tolerant) saplings were exposed to different flooding regimes and N uptake, amino acid, protein and chlorophyll concentrations as well as gas exchange were measured. The effects of these treatments on the tree species varied distinctly. In general, the N metabolism of beech was severely affected whereas less impacts were observed on oaks and almost no effects on poplars. The concentrations of amino compounds, particularly of Asp, Asn, Glu and Gln, were lower in the roots of flooded trees than in controls. By contrast, γ‐amino butyric acid concentrations increased. Root protein concentrations remained unaffected in oak and poplar but decreased in beech in response to flooding. The concentrations of pigments remained unaffected by flooding in all tree species investigated. However, photosynthesis and transpiration were severely affected in beech but much less in oak and poplar. The data obtained show a clear correlation between the different flooding tolerances of the trees investigated and the impacts of flooding on N uptake and N metabolism.     

Using X-ray CT based tree-ring width data for tree growth trend analysis

Changes in the environment influence the growth of tree species in Europe. Understanding the drivers of these growth changes is important to predict further growth and adapt forest management. To disentangle the different drivers of growth changes, it is common practice to apply mixed modeling techniques to tree-ring width series. Mixed modeling requires precise, replicated and well cross-dated tree-ring width series. The goal of this study was to compare a recently developed ring width measuring method based on X-ray Computed Tomography images (CT scan) with the standard LINTAB measuring method and to examine whether the same growth trends are detected with both methods using common beech (Fagus sylvatica) and sessile oak trees (Quercus petraea) as a case study. Although the CT scan method has a lower resolution than LINTAB measurements, it is of interest since it measures wood density in addition to ring width and it is less laborious in comparison to standard ring width measuring methods. No significant differences in ring width were found between the two measuring methods. The small non-significant difference between the two methods could largely be explained by the drying of cores needed for CT scanning. The same growth trends were detected with both methods: for common beech and sessile oak in Southern Belgium. These findings suggest that ring widths measured on CT scan images can be used as input for long-term modeling of tree growth changes for the targeted tree species.     

Comparative analysis of growth and movements of the aboveground parts in some deciduous woody species

Vertical projections of nutation movements in the epicotyls of sessile oak, Turkey oak, horse chestnut and hypocotyls of European beech were recorded by means of photography and time-lapse cinematography at the early stages of their ontogeny. Attention was paid to the kinetics of growth rate, diameter of the lower part of the elongating zone, and the form, amplitude and frequency of nutation turns. At the beginning of ontogeny the vertical projection of the movements of all woody species under study was represented by irregular curves, followed by elliptical trajectories, particularly when reaching the maximum growth rate. The highest average growth rate was recorded in horse chestnut, followed by oak, and the lowest values were exhibited by beech. As to the size of amplitudes opposite results were obtained. The frequency of turns proved to be a less sensitive parameter characterizing the movement. It is suggested that the geocontrol system of horse chestnut is more effective than that of oak and, especially, of beech.     

Physiological responses of beech and sessile oak in a natural mixed stand during a dry summer

Responses of CO2 assimilation and stomatal conductance to decreasing leaf water potential, and to environmental factors, were analysed in a mixed natural stand of sessile oak (Quercus petraea ssp. medwediewii) and beech (Fagus svlvatica L.) in Greece during the exceptionally dry summer of 1998. Seasonal courses of leaf water potential were similar for both species, whereas mean net photosynthesis and stomatal conductance were always higher in sessile oak than in beech. The relationship between net photosynthesis and stomatal conductance was strong for both species. Sessile oak had high rates of photosynthesis even under very low leaf water potentials and high air temperatures, whereas the photosynthetic rate of beech decreased at low water potentials. Diurnal patterns were similar in both species but sessile oak had higher rates of CO2 assimilation than beech. Our results indicate that sessile oak is more tolerant of drought than beech, due, in part, to its maintenance of photosynthesis at low water potential.     

Radial growth response of four dominant boreal tree species to climate along a latitudinal gradient in the eastern Canadian boreal forest

To address the central question of how climate change influences tree growth within the context of global warming, we used dendroclimatological analysis to understand the reactions of four major boreal tree species –Populus tremuloides, Betula papyrifera, Picea mariana, and Pinus banksiana– to climatic variations along a broad latitudinal gradient from 46 to 54°N in the eastern Canadian boreal forest. Tree‐ring chronologies from 34 forested stands distributed at a 1° interval were built, transformed into principal components (PCs), and analyzed through bootstrapped correlation analysis over the period 1950–2003 to identify climate factors limiting the radial growth and the detailed radial growth–climate association along the gradient. All species taken together, previous summer temperature (negative influences), and current January and March–April temperatures (positive influences) showed the most consistent relationships with radial growth across the gradient. Combined with the identified species/site‐specific climate factors, our study suggested that moisture conditions during the year before radial growth played a dominant role in positively regulating P. tremuloides growth, whereas January temperature and growing season moisture conditions positively impacted growth of B. papyrifera. Both P. mariana and P. banksiana were positively affected by the current‐year winter and spring or whole growing season temperatures over the entire range of our corridor. Owing to the impacts of different climate factors on growth, these boreal species showed inconsistent responsiveness to recent warming at the transition zone, where B. papyrifera, P. mariana, and P. banksiana would be the most responsive species, whereas P. tremuloides might be the least. Under continued warming, B. papyrifera stands located north of 49°N, P. tremuloides at northern latitudes, and P. mariana and P. banksiana stands located north of 47°N might benefit from warming winter and spring temperatures to enhance their radial growth in the coming decades, whereas other southern stands might be decreasing in radial growth.     

Climate–growth relationship of tree species from a mixed stand of Apuseni Mts., Romania

We present the first results of a dendroecological investigation on three prevailing species growing in the Apuseni Mountains. In this study we succeeded in building up about 200-year long local master chronologies for spruce, fir and beech. These chronologies will serve as a reference for crossdating of (sub-fossil) logs that were preserved in ice caves at the Apuseni Mts.

The analysis of climate–growth relationship provided interesting and different results for the three studied species: we found significant correlations between the beech chronology and the mean temperature of the prior July, autumn and January of the actual year as well as the weather conditions at the beginning of the vegetation period. Conifers yielded coherent results for the mean temperature of prior July and sum of precipitation in April. In addition, fir seems to be favouring above-average temperatures in January and February. The coherent and significant correlations with the April precipitation suggest a common coniferous phenomenon. Finally, beech index showed negative correlation with April temperature.     

Dating of non-oak species in the United Kingdom historical buildings archive using stable oxygen isotopes

Stable oxygen isotope dendrochronology is an effective precision-dating method for fast grown, invariant (complacent) tree-rings and for trees growing in moist, temperate climatic regions where growth may not be strongly controlled by climate. The method works because trees preserve a strong common isotopic signal, from summer precipitation, and therefore do not need to be physiologically stressed to record a dating signal. This study explores the working hypothesis that whilst tree species may differ in their eco-physiology, leaf morphology and wood anatomy they will record an isotopic signal in their growth rings that is sufficiently similar to enable their precise dating against isotopic reference chronologies developed using dated oak tree rings from the same region. Modern and historical samples from six species (sweet chestnut, English elm, ash, alder, European beech and black poplar) were analysed and their oxygen isotopic variability was compared against an oak master chronology previously developed for central southern England. Whilst differences in the relative strength of the agreement between the different species and the master chronology are apparent, the potential for interspecies dating is demonstrated convincingly. The ability to date non-oak species using stable oxygen isotopes opens-up new opportunities for science-based archaeology and will improve understanding of a largely-unexplored, but significant part of the European historical buildings archive.     

Different responses of Korean pine (Pinus koraiensis) and Mongolia oak (Quercus mongolica) growth to recent climate warming in northeast China

Different tree species growing in the same area may have different, or even contrasting growth responses to climate change. Korean pine (Pinus koraiensis) and Mongolia oak (Quercus mongolica) are two crucial tree species in temperate forest ecosystems. Six tree-ring chronologies for Korean pine and Mongolia oak were developed by using the zero-signal method to explore their growth response to the recent climate warming in northeast China. Results showed that Mongolia oak radial growth was mainly limited by precipitation in the growing season, while Korean pine growth depended on temperature condition, especially monthly minimum temperature. With the latitude decrease, the relationships between Korean pine growth and monthly precipitation changed from negative to positive correlation, while the positive correlation with monthly temperature gradually weakened. In the contrary, Mongolia oak growth at the three sampling sites was significantly and positively correlated with precipitation in the growing season, while it was negatively correlated with temperature and this relationship decreased with the latitude decrease. The radial growth of Korean pine at different sites showed a clearly discrepant responses to the recent warming since 1980. Korean pine growth in the north site increased with the temperature increase, decreased in the midwest site, and almost unchanged in the southeast site. Conversely, Mongolia oak growth was less affected by the recent climate warming. Our finding suggested that tree species trait and sites are both key factors that affect the response of tree growth to climate change. In addition, the suitable distribution area of Korean pine may be moved northward with the continued global warming in the future, but Mongolia oak may not shift in the same way.     

Mesophication in temperate Europe: A dendrochronological reconstruction of tree succession and fires in a mixed deciduous stand in Białowieża Forest

1. The shift from shade‐intolerant species to shade‐tolerant mesophytic species in deciduous and mixed forests of the temperate zone is well described in studies from North America. This process has been termed mesophication and it has been linked to changes in fire regime. Fire suppression results in the cessation of establishment of heliophytic, fire‐dependent tree species such as oak (Quercus) and pine (Pinus). Due to the scarcity of old‐growth forests in Europe, data on long‐term compositional changes in mixed forests are very limited, as is the number of studies exploring whether fire played a role in shaping the dynamics.
2. The aim of this study was to reconstruct tree succession in a 43‐ha natural mixed deciduous forest stand in Bia?owie?a Forest (BF), Poland using dendrochronological methods. In addition, the presence of aboveground fire legacies (charred and fire‐scarred deadwood) enabled the fire history reconstruction.
3. Dendrochronological data revealed tree establishment (Quercus) back to the end of the 1500s and fires back to 1659. Under a regime of frequent fires until the end of the 18th century, only oak and pine regenerated, sporadically. A shift in the fire regime in the first half of the 19th century triggered oak and pine cohort regeneration, then gradually spruce (Picea) encroached. Under an increasingly dense canopy and less flammable conditions, regeneration of shade‐tolerant Carpinus, Tilia, and Acer began simultaneously with the cessation of oak and pine recruitment.
4. Synthesis. The study reports the first evidence of mesophication in temperate Europe and proves that fire was involved in shaping the long‐term dynamics of mixed deciduous forest ecosystems. Our data suggest that fire exclusion promoted a gradual recruitment of fire‐sensitive, shade‐tolerant species that inhibited the regeneration of oak and pine in BF.

     

Temporal variation of competition and facilitation in mixed species forests in Central Europe

Facilitation, reduced competition or increased competition can arise in mixed stands and become essential to the performance of these stands when compared to pure stands. Facilitation and over‐yielding are widely held to prevail on poor sites, whereas neutral interactions or competition, leading to under‐yielding of mixed versus pure stands, can occur on fertile sites. While previous studies have focused on the spatial variation of mixing effects, we examine the temporal variation of facilitation and competition and its effect on growth. The study is based on tree ring measurement on cores from increment borings from 559 trees of Norway spruce (Picea abies [L.] Karst.), European beech (Fagus sylvatica [L.]) and sessile oak (Quercus petraea (Matt.) Liebl.) in southern Germany, half of which were in pure stands and half in adjacent mixed stands. Mean basal area growth indices were calculated from tree ring measurements for pure and mixed stands for every species and site. The temporal variation, with positive correlations between species‐specific growth indices during periods of low growth and neutral or negative correlations during periods of high growth, is more distinct in mixed than in neighbouring pure stands. We provide evidence that years with low growth trigger over‐yielding of trees in mixed as opposed to pure stands, while years with high growth lead to under‐yielding. We discuss the relevance of the results in terms of advancing our understanding and modelling of mixed stands, extension of the stress gradient hypothesis, and the performance of mixed versus pure stands in the face of climate change.     

Osservazioni Preliminari Sull'ecologia di Semenzali di Rovere (Quercus Petraea (Matt.) Liebl.)

Abstract

Preliminary observations on the ecology of sessile oak (Quercus petraea (Matt.) Liebl.) seedlings.—The responses of sessile oak seedlings to four different light intensities and to increasing drought (three soil moisture contents) have been investigated in a nursery experiment in order to improve the knowledge of the ecology of this species in the Mediterranean area. Data have been compared with observations on natural regeneration established in different shade and soil moisture conditions in a sessile oak stand. The results of the two experiments showed that low light intensities (transmittance <5%) did not limit the seedlings establishment but induced lowest growth rates, while in full daylight seedlings mortality was very high (61%). In accordance with the observations in natural conditions, seedlings showed the best performances at intermediate light levels (50–80% of full daylight) and with increasing soil moisture: mortality was very low and average height increment, dry weight, relative growth rate (RGR) and net assimilation rate (NAR) showed the highest values. The differences in specific leaf area (SLA), leaf area ratio (LAR), root/stem ratio enabled to discuss the sessile oak seedlings ecology and adaptations to different light intensities and soil moisture content.     

Climate warming and precipitation redistribution modify tree–grass interactions and tree species establishment in a warm‐temperate savanna

Savanna tree–grass interactions may be particularly sensitive to climate change. Establishment of two tree canopy dominants, post oak (Quercus stellata) and eastern redcedar (Juniperus virginiana), grown with the dominant C₄ perennial grass (Schizachyrium scoparium) in southern oak savanna of the United States were evaluated under four climatic scenarios for 6 years. Tree–grass interactions were examined with and without warming (+1.5 °C) in combination with a long‐term mean rainfall treatment and a modified rainfall regime that redistributed 40% of summer rainfall to spring and fall, intensifying summer drought. The aim was to determine: (1) the relative growth response of these species, (2) potential shifts in the balance of tree–grass interactions, and (3) the trajectory of juniper encroachment into savannas, under these anticipated climatic conditions. Precipitation redistribution reduced relative growth rate (RGR) of trees grown with grass. Warming increased growth of J. virginiana and strongly reduced Q. stellata survival. Tiller numbers of S. scoparium plants were unaffected by warming, but the number of reproductive tillers was increasingly suppressed by intensified drought each year. Growth rates of J. virginiana and Q. stellata were suppressed by grass presence early, but in subsequent years were higher when grown with grass. Quercus stellata had overall reduced RGR, but enhanced survival when grown with grass, while survival of J. virginiana remained near 100% in all treatments. Once trees surpassed a threshold height of 1.1 m, both tiller number and survival of S. scoparium plants were drastically reduced by the presence of J. virginiana, but not Q. stellata. Juniperus virginiana was the only savanna dominant in which neither survival nor final aboveground mass were adversely affected by the climate scenario of warming and intensified summer drought. These responses indicate that climate warming and altered precipitation patterns will further accelerate juniper encroachment and woody thickening in a warm‐temperate oak savanna.     

The influence of winter temperatures on the annual radial growth of six northern range margin tree species

This study explores the influence of temperature on the growth of six northern range margin (NRM) tree species in the Hudson River Valley (HRV). The HRV has excellent geographic and floristic qualities to study the influence of climate change on forested ecosystems. Indices of radial growth for three populations per species are developed and correlated against average minimum and maximum monthly temperatures from 1897 to 1994. Only positive correlations to temperature are considered for this analysis. Principal component analysis (PCA) is performed on chronologies over the entire HRV and at four subregions. PCA reveals a strong common signal among populations at subregional and regional scales. January temperatures most limit growth at the ecosystem level, supporting the hypothesis that winter temperatures may control vegetational ecotones. Surprisingly, growth of the oak–hickory ecosystem is most limited by January temperatures only in the southern half of the study region. Chestnut and white oak are the primary species driving the geographic pattern. As winter xylem embolism is a constant factor for ring-porous species, snow cover and its interaction on fine root mortality may be the leading factors of the pattern of temperature sensitivity. Species-specific differences in temperature sensitivity are apparent. Atlantic white-cedar (AWC) and pitch pine are more sensitive to the entire winter season (December–March) while oak and hickory are most sensitive to January temperatures. AWC is most sensitive species to temperature. Chestnut and white oak in the HRV are more sensitive to winter temperature than red oak. Pignut hickory has the most unique response with significant relations to late growing season temperatures. Interestingly, AWC and pitch pine are sensitive to winter temperatures at their NRM while oak and hickory are not. Our results suggest that temperature limitations of growth may be species and phylogenetically specific. They also indicate that the influence of temperature on radial growth at species and ecosystem levels may operate differently at varying geographic scales. If these results apply broadly to other temperate regions, winter temperatures may play an important role in the terrestrial carbon cycle.