Identifying ontogenetic, environmental and individual components of forest tree growth

Background and Aims

This study aimed to identify and characterize the ontogenetic, environmental and individual components of forest tree growth. In the proposed approach, the tree growth data typically correspond to the retrospective measurement of annual shoot characteristics (e.g. length) along the trunk.

Methods

Dedicated statistical models (semi-Markov switching linear mixed models) were applied to data sets of Corsican pine and sessile oak. In the semi-Markov switching linear mixed models estimated from these data sets, the underlying semi-Markov chain represents both the succession of growth phases and their lengths, while the linear mixed models represent both the influence of climatic factors and the inter-individual heterogeneity within each growth phase.

Key Results

On the basis of these integrative statistical models, it is shown that growth phases are not only defined by average growth level but also by growth fluctuation amplitudes in response to climatic factors and inter-individual heterogeneity and that the individual tree status within the population may change between phases. Species plasticity affected the response to climatic factors while tree origin, sampling strategy and silvicultural interventions impacted inter-individual heterogeneity.

Conclusions

The transposition of the proposed integrative statistical modelling approach to cambial growth in relation to climatic factors and the study of the relationship between apical growth and cambial growth constitute the next steps in this research.     

Markov and Semi‐Markov Switching Linear Mixed Models Used to Identify Forest Tree Growth Components

Summary Tree growth is assumed to be mainly the result of three components: (i) an endogenous component assumed to be structured as a succession of roughly stationary phases separated by marked change points that are asynchronous among individuals, (ii) a time‐varying environmental component assumed to take the form of synchronous fluctuations among individuals, and (iii) an individual component corresponding mainly to the local environment of each tree. To identify and characterize these three components, we propose to use semi‐Markov switching linear mixed models, i.e., models that combine linear mixed models in a semi‐Markovian manner. The underlying semi‐Markov chain represents the succession of growth phases and their lengths (endogenous component) whereas the linear mixed models attached to each state of the underlying semi‐Markov chain represent—in the corresponding growth phase—both the influence of time‐varying climatic covariates (environmental component) as fixed effects, and interindividual heterogeneity (individual component) as random effects. In this article, we address the estimation of Markov and semi‐Markov switching linear mixed models in a general framework. We propose a Monte Carlo expectation–maximization like algorithm whose iterations decompose into three steps: (i) sampling of state sequences given random effects, (ii) prediction of random effects given state sequences, and (iii) maximization. The proposed statistical modeling approach is illustrated by the analysis of successive annual shoots along Corsican pine trunks influenced by climatic covariates.     

Dissecting apple tree architecture into genetic, ontogenetic and environmental effects: QTL mapping

The present study aimed to dissect tree architectural plasticity into genetic, ontogenetic and environmental effects over the first 4 years of growth of an apple F1 progeny by means of quantitative traits loci (QTL) mapping. Both growth and branching processes were phenotyped on the consecutive annual shoots of different axes within a tree. For each studied trait, predicted values (best linear unbiased predictors, BLUPs) of the genotypic (G) effect or its interaction with tree age (G×A) and climatic year (G×Y) were extracted from mixed linear models of repeated data. These BLUPs, which are independent from autocorrelations between repeated measurements, were used for QTL mapping. QTL detection power was improved by this two-step approach. For each architectural process, numerous QTLs were detected and some particularly interesting co-localised in common genomic regions, for internode lengthening, top diameter, and number and percentage of axillary shoots. When several QTLs were detected for a given trait, global models were estimated, which explained a maximum of 40% of the total variance for both internode length and top diameter and 28% for branching. QTLs detected for BLUPs of G×Y effects were interpreted as resulting from the interaction between genetic maximal potential of growth and climatic factors, while those for G×A effects were interpreted in relation to tree ontogeny. Most of the latter ones were found to be concomitant with key development stages during which the trait average started to decrease, but with different magnitudes depending on genotype. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Environmental Correlates of Tree and Seedling–Sapling Distributions in a Mexican Tropical Dry Forest

Bray and Curtis ordination was used to explore which environmental variables explained importance values and the presence–absence of tropical tree seedlings, saplings and adults in La Escondida-La Cabaña, Sierra de Manantlán, Jalisco, Mexico. The diameters of trees ≥2.5 cm DBH and the presence and height of seedlings and saplings were measured in nine 0.1 ha sites. Four matrices including presence–absence data and importance value indices for trees and seedlings and saplings were analyzed through Bray and Curtis ordination. The matrices were based on density, frequency, and dominance of adult trees as well as seedlings and saplings. The environmental matrix consisted of 18 variables, including elevation, slope, canopy gaps, disturbance, and soil variables. We recorded 63 tree species and 38 seedling and sapling species in the nine sites. The ordination explained 70.9% of the variation in importance value data for trees and 62.6% for seedlings and saplings. The variation explained in presence–absence data for trees was 67.1 and 77.4% for seedlings and saplings. The variance in the ordination axes of seedlings and sapling presence–absence data was poorly explained by the number of gaps in the tree, shrub, or herb layer, suggesting little light specialization by seedlings and saplings. Habitat specialization for soil nutrients appears to be important in explaining the presence–absence of seedlings and saplings. Seedling and sapling specialization along different soil microsites could promote species coexistence in this forest, while heterogeneity in light conditions may instead determine differences in growth and, thus, importance value of trees. We hypothesize that in tropical dry forest in Jalisco, Mexico, a habitat specialization for soil resources is likely more importance at early stages in tree life histories than in later life history.     

Ecological and methodological drivers of non-stationarity in tree growth response to climate

Radial tree growth is sensitive to environmental conditions, making observed growth increments an important indicator of climate change effects on forest growth. However, unprecedented climate variability could lead to non-stationarity, that is, a decoupling of tree growth responses from climate over time, potentially inducing biases in climate reconstructions and forest growth projections. Little is known about whether and to what extent environmental conditions, species, and model type and resolution affect the occurrence and magnitude of non-stationarity. To systematically assess potential drivers of non-stationarity, we compiled tree-ring width chronologies of two conifer species, Picea abies and Pinus sylvestris, distributed across cold, dry, and mixed climates. We analyzed 147 sites across the Europe including the distribution margins of these species as well as moderate sites. We calibrated four numerical models (linear vs. non-linear, daily vs. monthly resolution) to simulate growth chronologies based on temperature and soil moisture data. Climate–growth models were tested in independent verification periods to quantify their non-stationarity, which was assessed based on bootstrapped transfer function stability tests. The degree of non-stationarity varied between species, site climatic conditions, and models. Chronologies of P. sylvestris showed stronger non-stationarity compared with Picea abies stands with a high degree of stationarity. Sites with mixed climatic signals were most affected by non-stationarity compared with sites sampled at cold and dry species distribution margins. Moreover, linear models with daily resolution exhibited greater non-stationarity compared with monthly-resolved non-linear models. We conclude that non-stationarity in climate–growth responses is a multifactorial phenomenon driven by the interaction of site climatic conditions, tree species, and methodological features of the modeling approach. Given the existence of multiple drivers and the frequent occurrence of non-stationarity, we recommend that temporal non-stationarity rather than stationarity should be considered as the baseline model of climate–growth response for temperate forests.     

Dry season watering alters the significance of climate factors influencing phenology and growth of saplings of savanna woody species in central Zambia,southern Africa

Although tree growth in southern African savannas is correlated with rainfall in the wet season, some studies have shown that tree growth is controlled more by rainfall in the dry season. If more rainfall occurred in the dry season in future climates, it would affect the growth of savanna trees, especially saplings that have shallower roots which limit access to subsoil water during the dry season when leaf flush and shoot extension occur. Recent paleobotanical evidence has revealed that there was relatively more precipitation in the dry season in eastern Africa in the Eocene than under the current climate. Saplings therefore can be expected to respond more to water addition during the dry season than mature trees that have more stored water and deeper roots that access subsoil water. Accordingly, I hypothesized that irrigation in the dry season should (i) advance the onset of the growing season, (ii) increase growth rates and (iii) alter the growth responses of saplings to climate factors. To test these hypotheses saplings of five savanna woody species were irrigated during the hot‐dry season at a site in central Zambia and their monthly and annual growth rates compared to those of conspecifics growing under control conditions. Although the responses among the species were variable, all irrigated saplings had significantly higher monthly and annual growth rates than control plants. In addition, dry season watering significantly altered the climatic determinants of sapling growth by either strengthening the role of the same climatic factors that were important under control conditions or displacing them altogether. In conclusion, more precipitation during the hot‐dry season is likely to have significant positive effects on sapling growth and consequently reduce the sapling‐tree transition periods and promote future tree population recruitment in some southern African savanna tree species.     

Unusual Fine Root Distributions of Two Deciduous Tree Species in Southern France: What Consequences for Modelling of Tree Root Dynamics?

The spatial distribution of fine roots of two deciduous tree species was investigated in contrasting growing conditions in southern France. Hybrid walnut trees (Juglans regia×nigra cv. NG23) and hybrid poplars (Populus euramericana cv. I214) were both cultivated with or without annual winter intercrops for 10 years on deep alluvial soils. Soil samples for measuring the fine root distribution of both trees and crops were obtained by soil coring down to 3-m depth at several distances and orientations from the tree trunk. The distribution of live fine roots from walnut and poplar trees was patchy and sometimes unexpected. In the tree-only stands, fine root profiles followed the expected pattern, as fine root density decreased with increasing depth and distance from the tree trunk. However, many fine root profiles under intercropped trees were uniform with depth, and some inverse profiles were observed. These distributions may result from a high degree of plasticity of tree root systems to sense and adapt to fluctuating and heterogeneous soil conditions. The distortion of the tree root system was more pronounced for the walnut trees that only partially explored the soil volume: in the tree-only stand, the walnut rooting pattern was very superficial, but in the intercropped stand walnut trees developed a deep and dense fine root network below the crop rooting zone. The larger poplars explored the whole available soil volume, but the intercrop significantly displaced the root density from the topsoil to layers below 1 m depth. Most tree root growth models assume a decreasing fine root density with depth and distance from the tree stem. These models would not predict correctly tree–tree and tree–understorey competition for water and nutrients in 3D heterogeneous soil conditions that prevail under low-density tree stands. To account for the integrated response of tree root systems to such transient gradients in soils, we need a dynamic model that would allow for both genotypic plasticity and transient environmental local soil conditions.     

Dissecting apple tree architecture into genetic, ontogenetic and environmental effects: mixed linear modelling of repeated spatial and temporal measures

The present study aimed to dissect tree architectural plasticity into genetic, ontogenetic and environmental effects over the first 4 yr of growth of an apple (Malus x domestica) F1 progeny by means of mixed linear modelling of repeated data. Traits related to both growth and branching processes were annually assessed on different axes of the trees planted in a staggered-start design. Both spatial repetitions, (i.e. different axis types) and temporal repetitions (i.e. successive ages of trees) were considered in a mixed linear model of repeated data. A significant genotype effect was found for most studied traits and interactions between genotype and year and/or age were also detected. The analysis of repeated temporal measures highlighted that the magnitude of the decrease in primary growth is mainly determined by the first year of growth, and the decrease in bottom diameter increment is concomitant with the first fruiting occurrence. This approach allowed us to distinguish among the traits that were under genetic control, those for which this control is exerted differentially throughout tree life or depending on climatic conditions or an axis type. Mapping quantitative trait loci (QTL) that are specific to these different effects will constitute the next step in the research.     

Interannual changes in folivory and bird insectivory along a natural productivity gradient in northern Patagonian forests

Trophic regulation models suggest that the magnitude of herbivory and predation (top-down forces) should vary predictably with habitat productivity. Theory also indicates that temporal abiotic variation and within-trophic level heterogeneity both affect trophic dynamics, but few studies addressed how these factors interact over broad-scale environmental gradients. Here we document herbivory from leaf-feeding insects along a natural rainfall/productivity gradient in Nothofagus pumilio forests of northern Patagonia, Argentina, and evaluate the impact of insectivorous birds on foliar damage experienced by tree saplings at each end of the gradient. The study ran over three years (1997–2000) comprising a severe drought (1998–1999), which allowed us to test how climatic events alter top-down forces. Foliar damage tended to increase towards the xeric, least productive forests. However, we found a predictable change of insect guild prevalence across the forest gradient. Leaf miners accounted for the greater damage recorded in xeric sites, whereas leaf chewers dominated in the more humid and productive forests. Interannual folivory patterns depended strongly on the feeding guild and forest site. Whereas leaf-miner damage decreased during the drought in xeric sites, chewer damage increased after the drought in the wettest site. Excluding birds did not affect leaf damage from miners, but generally increased chewer herbivory on hydric and xeric forest saplings. Indirect effects elicited by bird exclusion became most significant after the drought, when total folivory levels were higher. Thus, interannual abiotic heterogeneity markedly influenced the amount of folivory and strength of top-down control observed across the forest gradient. Moreover, our results suggest that spatial turnovers between major feeding guilds may need be considered to predict the dynamics of insect herbivory along environmental gradients.     

Temperature versus species-specific influences on the stable oxygen isotope ratio of tree rings

Stable isotopic ratios integrate ecosystem variability while reflecting change in both environmental and biological processes. At sites, where climate does not strongly limit tree growth, co-occurring trees may display large discrepancies in stable oxygen isotopic ratios (δ¹⁸O) due to the interplay between biological processes (competition for light and nutrients, individual tree physiology, etc.) and climate. For a better quantification of the isotope variability within and among trees, the climatic and/or individual tree effects on seasonal δ¹⁸O variations in precipitation, soil water, leaf water and leaf organic material (whole leaf, cellulose and starch) and annual δ¹⁸O variations in tree-ring cellulose for Fagus sylvatica (Fs), Quercus robur (Qr), Carpinus betulus (Cb) and Pinus sylvestris (Ps) were studied in a mature temperate forest in Switzerland, using a mixed linear regression model technique. Furthermore, the influence of environmental factors on δ¹⁸O was assessed by means of three common isotope fractionation models. Our statistical analysis showed that except for Ps, a greater portion of δ¹⁸O variance in leaf compounds can be explained by individual tree effects, compared to temperature. Concerning tree-ring cellulose, only Fs and Ps show a significant temperature signal (maximum 12% of the variance explained), while the individual tree effect significantly explains δ¹⁸O for all species for a period of 38 years. Large species differences resulted in a limited ability of the isotope fractionation models to predict measured values. Overall, we conclude that in a diverse mixed forest stand, individual tree responses reduce the potential extraction of a temperature signal from δ¹⁸O.     

Environmental heterogeneity and neighbourhood interference modulate the individual response of Juniperus thurifera tree-ring growth to climate

Individual variation of tree-ring growth response to climate and heterogeneity of the local environment are usually neglected in dendrochronological research. Even if there is evidence showing that individual responsiveness to climate may depend on intrinsic traits such as tree age, size or sex, its modulation by the local heterogeneity of extrinsic factors has been less studied. Using an extensive, strictly regular sampling scheme across a 3300 ha woodland, we assessed the individual variation of tree-ring growth responses to climate in 100 Juniperus thurifera L. trees. The climatic response was evaluated by bootstrapped correlations of both population- and individual-based tree-ring chronologies with monthly records of precipitation, cloudiness, minimum and maximum temperatures. We studied also the influence of extrinsic abiotic (elevation, slope, heat load, tree location) and biotic (competition from neighbouring trees) factors on the individual growth variation and its climatic response. At a population level, growth was controlled by February–March precipitation, April minimum temperature, and June water stress. A significant proportion of individuals did not respond to those variables, but were sensitive to others not relevant at the population level. Inter-annual growth variation was strongly modulated by competition, whereas trees under lower competition levels, in eastern and warmer areas, were the most responsive to climate. The individual climatic response was, at least partially, modulated by the local heterogeneity of extrinsic factors. By considering environmental heterogeneity and neighbourhood interference we can identify the spectrum of site-dependent climatic responses in a population, which in turn will enable more realistic predictions of tree responses to ongoing climate change.     

Deciphering the ontogeny of a sympodial tree

This paper addresses the identification and characterization of developmental patterns in the whole structure of a sympodial species, the apple tree. Dedicated stochastic models (hidden variable-order Markov chains) were used to (i) categorise growth units (GUs) on the basis of their morphological characteristics (number of nodes and presence/absence of flowering) and position along axes, (ii) analyse dependencies between successive GUs and (iii) identify repeated patterns in GU sequences. Two successive phases, referred to as “adolescent” and “adult”, were identified in two apple tree cultivars. In the adolescent phase, “very” long monocyclic GUs were followed by long polycyclic GUs, whereas in the adult phase medium GUs were preferentially followed by short GUs. Flowering GUs constituted a preferential pathway between vegetative GUs of decreasing vigour (long, medium and short) and generated patterns that were interpreted with respect to fruiting regularity. The proposed modelling gave a global and quantitative picture of the two-scale structuring of apple tree ontogeny: a coarse scale corresponding to the succession of the previously mentioned phases and a fine scale corresponding to the alternation between flowering and vegetative GUs. This led us to propose a synthetic scheme of apple tree ontogeny that combines growth phases, polycyclism and flowering, and which could be transposed to other sympodial trees.     

Tree species distribution in temperate forests is more influenced by soil than by climate

Knowledge of the ecological requirements determining tree species distributions is a precondition for sustainable forest management. At present, the abiotic requirements and the relative importance of the different abiotic factors are still unclear for many temperate tree species. We therefore investigated the relative importance of climatic and edaphic factors for the abundance of 12 temperate tree species along environmental gradients. Our investigations are based on data from 1,075 forest stands across Switzerland including the cold‐induced tree line of all studied species and the drought‐induced range boundaries of several species. Four climatic and four edaphic predictors represented the important growth factors temperature, water supply, nutrient availability, and soil aeration. The climatic predictors were derived from the meteorological network of MeteoSwiss, and the edaphic predictors were available from soil profiles. Species cover abundances were recorded in field surveys. The explanatory power of the predictors was assessed by variation partitioning analyses with generalized linear models. For six of the 12 species, edaphic predictors were more important than climatic predictors in shaping species distribution. Over all species, abundances depended mainly on nutrient availability, followed by temperature, water supply, and soil aeration. The often co‐occurring species responded similar to these growth factors. Drought turned out to be a determinant of the lower range boundary for some species. We conclude that over all 12 studied tree species, soil properties were more important than climate variables in shaping tree species distribution. The inclusion of appropriate soil variables in species distribution models allowed to better explain species' ecological niches. Moreover, our study revealed that the ecological requirements of tree species assessed in local field studies and in experiments are valid at larger scales across Switzerland.     

江西九连山亚热带常绿阔叶林优势种空间分布格局

于江西九连山国家自然保护区设置4 hm2亚热带常绿阔叶林固定监测样地,用样方法对亚热带常绿阔叶林进行调查分析。应用点格局方法分析生境异质性是否影响树木分布并分析优势种红钩栲(Castanopsis lamontii)、米槠(Castanopsis carlesii)、罗浮柿(Diospyros morrisiana)和细枝柃(Eurya loquaiana)分布格局,对比存在生境异质性和排除生境异质性两种情况下优势树种不同生长阶段(幼树、小树、成年树)的空间分布格局以及不同生长阶段之间的空间关联性,探讨生境异质性之外的其它种群分布影响因子。结果表明:1)生境异质性效应显著影响九连山树木分布,4个优势树种在大尺度上存在明显的生境偏好;2)用完全随机零模型不排除生境异质性时,4个优势种总体及其不同生长阶段在0—30 m所有尺度上主要呈现聚集分布;3)用异质性随机零模型排除生境异质性后,4个优势种及其不同生长阶段的聚集程度显著下降,只在小尺度上(0—5 m)呈现聚集分布;4)4个优势种的幼树与小树均表现显著的正相关。红钩栲的成年树与幼树及成年树与小树总体表现不相关。米槠的成年树与幼树总体以无相关为主;成年树与小树之间总体呈正相关。小乔木罗浮柿的成年树与幼树之间在小尺度上(0—5 m)表现负相关或无相关,在较大尺度范围上总体表现正相关关系;小树与成年树在研究尺度上表现出正相关。灌木细枝柃的成年树与幼树,成年树与小树及小树与幼树之间在研究尺度范围内均呈现正相关关系;5)研究发现九连山优势乔木物种通过密度制约和Janzen-Connell效应释放空间,为其他物种共存提供条件,而优势灌木物种细枝柃没有表现出这两个效应。     

Relative effects of different leaf attributes on sapling growth in tropical dry forest

Aims Soil moisture content (SMC) influences establishment, survival and development of plant species and is considered as the most important limiting factor in tropical dry forest (TDF). In this study we attempt to establish the relationship between leaf attributes and of tree saplings in TDF and address the following questions: (i) how are the functional attributes of dominant tree saplings of TDF affected by seasonal changes in SMC at different habitats?, (ii) what is the relationship of functional attributes with each other?, (iii) how are the functional attributes and their plasticity affected by habitat conditions? and (iv) can the functional attributes in single or in combination predict the growth rate of tree saplings of TDF? The study was conducted on four sites (Hathinala, Gaighat, Harnakachar and Ranitali, listed in order of decreasing SMC) within the tropical dry deciduous forest in northern India.Methods We analysed eight leaf attributes, specific leaf area (SLA); leaf dry matter content (LDMC); leaf nitrogen concentration (leaf N); leaf phosphorus concentration (leaf P); chlorophyll concentration (Chl); mass-based photosynthetic rate (A mass); mass-based stomatal conductance (Gs mass); intrinsic water use efficiency (WUEi) and three growth attributes, relative diameter increment (RDI); relative height increment (RHI); relative growth rate (RGR) of the 10 dominant tree saplings (viz., Acacia catechu, Anogeissus latifolia, Boswellia serrata, Buchanania lanzan, Diospyros melanoxylon, Hardwickia binata, Lagerstroemia parviflora, Lannea coromandelica, Shorea robusta and Terminalia tomentosa) of a TDF and observed the effects of site, season and species for a period of 2 years. Saplings were selected in gradients of deciduousness. Step-wise multiple regression was performed to predict RDI, RHI and RGR from mean values of SMC and leaf attributes.Important findings All the 11 attributes were interrelated and differed significantly among the 10 saplings. Species response varied across sites and seasons. Across the SMC gradient, the attributes showed variable plasticity that differed across species. Among the 10 saplings, the highly deciduous Boswellia serrata showed the maximum plasticity in seven functional attributes. According to the step-wise multiple regressions, 65% variability in RDI and 67% variability in RGR were due to Gs mass, and for RHI, 61% variability was due to A mass. SMC and the other attributes, viz ., SLA, Chl, WUEi and LDMC in combination could contribute only for ~2–6% of the variability in RDI, RHI and RGR, which indicates that other traits/factors, not accounted in this study are also important in modulating the growth of tree saplings in TDFs. In conclusion, growth of the tree saplings in the tropical dry environment is determined by soil moisture, whereas the response of saplings of different tree species is modulated by alterations in key functional attributes such as SLA, Chl, WUEi and LDMC.     

Variable strength of top‐down effects in Nothofagus forests: bird predation and insect herbivory during an ENSO event

Abstract Predators are thought to play a key role in controlling herbivory, thus having positive indirect effects on plants. However, evidence for terrestrial trophic cascades is still fragmentary, perhaps due to variation in top‐down forces created by environmental heterogeneity. We examined the magnitude of predation effects on foliar damage by chewing insects and mean leaf size, by excluding birds from saplings in ‘dry’ and ‘wet’Nothofagus pumilio forests in the northern Patagonian Andes, Argentina. The experiment lasted 2 years encompassing a severe drought during the La Niña phase of a strong El Niño/Southern Oscillation event, which was followed by unusually high background folivory levels. Insect damage was consistently higher in wet than in dry forest saplings. In the drought year (1999), bird exclusion increased folivory rates in both forests but did not affect tree leaf size. In the ensuing season (2000), leaf damage was generally twice as high as in the drought year. As a result, bird exclusion not only increased the extent of folivory but also significantly decreased sapling leaf size. The latter effect was stronger in the wet forest, suggesting compensation of leaf area loss by dry forest saplings. Overall, the magnitude of predator indirect effects depended on the response variable measured. Insectivorous birds were more effective at reducing folivory than at facilitating leaf area growth. Our results indicate that bird‐initiated trophic cascades protect N. pumilio saplings from insect damage even during years with above‐normal herbivory, and also support the view that large‐scale climatic events influence the strength of trophic cascades.     

Tree demography suggests multiple directions and drivers for species range shifts in mountains of Northeastern United States

Climate change is expected to lead to upslope shifts in tree species distributions, but the evidence is mixed partly due to land‐use effects and individualistic species responses to climate. We examined how individual tree species demography varies along elevational climatic gradients across four states in the northeastern United States to determine whether species elevational distributions and their potential upslope (or downslope) shifts were controlled by climate, land‐use legacies (past logging), or soils. We characterized tree demography, microclimate, land‐use legacies, and soils at 83 sites stratified by elevation (~500 to ~1200 m above sea level) across 12 mountains containing the transition from northern hardwood to spruce‐fir forests. We modeled elevational distributions of tree species saplings and adults using logistic regression to test whether sapling distributions suggest ongoing species range expansion upslope (or contraction downslope) relative to adults, and we used linear mixed models to determine the extent to which climate, land use, and soil variables explain these distributions. Tree demography varied with elevation by species, suggesting a potential upslope shift only for American beech, downslope shifts for red spruce (more so in cool regions) and sugar maple, and no change with elevation for balsam fir. While soils had relatively minor effects, climate was the dominant predictor for most species and more so for saplings than adults of red spruce, sugar maple, yellow birch, cordate birch, and striped maple. On the other hand, logging legacies were positively associated with American beech, sugar maple, and yellow birch, and negatively with red spruce and balsam fir – generally more so for adults than saplings. All species exhibited individualistic rather than synchronous demographic responses to climate and land use, and the return of red spruce to lower elevations where past logging originally benefited northern hardwood species indicates that land use may mask species range shifts caused by changing climate.     

Inherent trait differences explain wheat cultivar responses to climate factor interactions: New insights for more robust crop modelling

Climate change predictions foresee a combination of rising CO₂, temperature and altered precipitation. Effects of single climatic variables are well defined, but the importance of combined variables and genotypic effects is less known, although pivotal for assessing climate change impacts, for example, with crop growth models. This study provides developmental and physiological data from combined climatic factors for two distinct wheat cultivars (Paragon and Gladius), as a basis to improve predictions for climate change scenarios. The two cultivars were grown in controlled climate chambers in a fully factorial setup of atmospheric CO₂ concentration, growth temperature and watering regime. The cultivars differed considerably in their developmental rate, response pattern and the parameters responsible for most of their variation. The growth of Paragon was linked to climatic effects on photosynthesis and mainly affected by temperature. Paragon was overall more negatively affected by all treatment combinations compared to Gladius. Gladius was mostly affected by watering regime. The cultivars' acclimation strategies to climate factors varied significantly. Thus, considering a single factor is an oversimplification very likely impacting the accuracy of crop growth models. Intraspecific crop variation could help understanding genotype by environment variation. Cultivars with high phenotypic plasticity may have greater resilience against climatic variability.     

The relative importance of above- versus belowground competition for tree growth during early succession of a tropical moist forest

Competition between neighboring plants plays a major role in the population dynamics of tree species in the early phases of humid tropical forest succession. We evaluated the relative importance of above- versus below-ground competition during the first years of old-field succession on soil with low fertility in Southern Mexico, using the premise that competition for light is size-asymmetric, unlike competition for nutrients. Plant growth is thus expected to be disproportionally impeded by larger neighbors. We studied how growth and survival of 3.5–5.5 m tall saplings of Cecropia peltata and Trichospermum mexicanum, two pioneer species that dominate the secondary forests in the study region, varied with the abundance and size of neighboring trees in 1–2 year old secondary vegetation. We found that local neighborhood basal area varied 10-fold (3 to 30 cm² m^-2) and explained most of the variation in diameter and height growth of the target saplings. Most growth variables were strongly affected by the neighbors bigger than the focal trees with no significant additive effect of the smaller neighbors, indicating asymmetric competition. Smaller neighbors did have a small but significant additive effect on the diameter growth of Cecropia saplings and stem slenderness of Trichospermum saplings. We conclude that competition for light was more important than belowground competition in this initial phase of moist tropical forest successional, despite the low soil fertility.     

Vapor–pressure deficit and extreme climatic variables limit tree growth

Assessing the effect of global warming on forest growth requires a better understanding of species‐specific responses to climate change conditions. Norway spruce and European beech are among the dominant tree species in Europe and are largely used by the timber industry. Their sensitivity to changes in climate and extreme climatic events, however, endangers their future sustainability. Identifying the key climatic factors limiting their growth and survival is therefore crucial for assessing the responses of these two species to ongoing climate change. We studied the vulnerability of beech and spruce to warmer and drier conditions by transplanting saplings from the top to the bottom of an elevational gradient in the Jura Mountains in Switzerland. We (1) demonstrated that a longer growing season due to warming could not fully account for the positive growth responses, and the positive effect on sapling productivity was species‐dependent, (2) demonstrated that the contrasting growth responses of beech and spruce were mainly due to different sensitivities to elevated vapor–pressure deficits (VPD), (3) determined the species‐specific limits to VPD above which growth rate began to decline, and (4) demonstrated that models incorporating extreme climatic events could account for the response of growth to warming better than models using only average values. These results support that the sustainability of forest trees in the coming decades will depend on how extreme climatic events will change, irrespective of the overall warming trend.