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Wood density (WD), a key trait in the trait‐based approach of plant ecology, represents a carbon investment trait that varies across species and reflects a trade‐off between metabolism and longevity. Across species, WD has been found to vary with phylogeny, moisture, temperature and xylem anatomy (e.g. vessel diameter). However, we know little about WD variation at the intraspecific level. Here, we examined how ecologically important functional traits vary in relation to WD in three generalist, exceptionally wide‐niche breadth tree species from southern Chile, making use of broad precipitation and temperature gradients.We collected branches from Embothrium coccineum and Nothofagus antarctica across a wide W–E precipitation gradient (2,500–600 mm of annual precipitation) and from N. pumilio across an elevational gradient (from low to treeline elevation). For each individual, we determined WD, several xylem anatomical features, for example, mean vessel diameter (MVD), hydraulic diameter (Dh), hydraulic conductivity (Kt), mean vessel area (MVA), vessel size distribution (MVA/VD) and other traits including concentrations of non‐structural carbohydrates (NSCs), secondary growth, leaf traits, water‐use efficiency (iWUE) and growth rate (AI). We quantified the correlation between WD and other traits and evaluated whether WD‐trait relationships differed across species using linear mixed‐effects models.We found consistent and significant negative relationships between WD and several xylem anatomy traits, including Dh, Kt, MVA and MVD. Contrary to our expectations, WD was not related to leaf traits, NSCs, iWUE and AI. Wood density had a significantly negative relationship to MVA/VD (and to a lesser extent, MVD) for both E. coccineum and N. antarctica (the precipitation gradient), while these relationships were positive for N. pumilio (the temperature gradient).Few of the WD and other trait relationships examined at the intraspecific level in this study paralleled those found across species. At the intraspecific level, WD proved to be much more related to xylem anatomical traits than to other traits. Importantly, in some cases, WD‐xylem anatomical trait relationships showed opposite trends depending on the environmental gradient. We must be cautious when using interspecific studies as the basis for inferring trait‐based responses to environmental change across other organisation levels. Read the free Plain Language Summary for this article on the Journal blog. [ABSTRACT FROM AUTHOR]

Aims: Rising temperature and declining summer precipitation due to the 1970s‐climate shift in southern South America have reduced forest productivity at dry sites. Here, we worked with the most widespread Southern Hemisphere tree line species, Nothofagus pumilio, across contrasting climatic conditions and determined whether rising atmospheric CO2 concentrations as well as warmer and drier climatic conditions provoked by the 70s‐climatic shift have been causing systematic changes in tree line growth rates and intrinsic water‐use efficiency (iWUE). Location: 36–54°S, southern Andes. Time period: 1950–2010. Major taxa studied: Nothofagus pumilio. Methods: We worked at five disparate climatic tree line locations, spanning 18 degrees of latitude; at each location, we sampled trees at four different elevations, including tree line elevation. We quantified the variation in annual tree‐ring width (TRW) as a function of climate, elevation, tree age, size, annual CO2 concentrations and location, using linear mixed‐effects models and interpreted TRW trends in relation to iWUE and isotope (δ13C and δ18O) signalling. Results: Across locations, the patterns of tree line growth occurring in the 1980–2010 period exhibited a clear and significant negative trend, in contrast to the previous 1950–1980 period. We found an increase in iWUE and δ18O across time and locations. Given that an increase in δ18O indicates a decrease in stomatal conductance, we assert that drought‐induced stomatal closure appears to be causing the reduction in growth. Main conclusions: We show unequivocal evidence that warmer and drier summer conditions translated into a decrease in growth rates along the elevational tree line of the southern Andes, reverting previous growth improvements linked to climate warming. An improvement in iWUE at all locations is most likely explained by decreased stomatal conductance given the rising δ18O signal. An iWUE–growth decoupling may act as an ecological strategy to respond to drought. [ABSTRACT FROM AUTHOR]

Although the principal mechanism determining tree line formation appears to be carbon (C)-sink limitations due to low temperatures, few studies have assessed the complementary role of reduced soil nutrient availability with elevation. We tested the hypothesis that nutrient (especially nitrogen, N) limitations at tree line may directly (via C-source) or indirectly (via C-sink) reduce the growth of a winter deciduous tree line species., If a shortage of soil N with elevation is involved in tree line formation, it should occur in two alternative ways: (i) through sink limitations because N is required for tissue formation, which would indirectly limit C investments (N decreases and C reserves increase with elevation), and (ii) through C limitations because this would lead directly to a reduction of photoassimilates (N and C reserves decrease with elevation)., In testing our hypothesis, we analysed tree growth rates (basal area increment), twig non-structural carbohydrate ( NSC) and N concentrations, leaf N, phosphorus (P), N:P ratio concentrations, and soil nutrient levels ( NO3−, NH4+, Olsen-P) in four disparate climate and soil Nothofagus pumilio tree lines spanning 18 degrees of latitude in the southern Andes of Chile., We found a significant decrease in tree growth with elevation. Twig NSC concentrations pooled across locations also decreased significantly with elevation (starch constituted most of the NSC and was highly responsible for the negative trend), although this trend was mostly driven by the northernmost locations. Contrary to soil N availability, leaf N and P concentrations increased significantly with elevation. Twig N concentrations, soil P and leaf N:P ratios did not change with elevation., Synthesis. The elevational decrease in NSC concentrations supports C-source limitation in N. pumilio trees at tree line elevation. In the light of this, we assert that the current global explanation for tree line formation (C-sink-limitation driven by low temperatures) must be revisited. Given that leaf N and P concentrations increased and twig N concentrations did not change with elevation, nutrient limitation is not likely to be involved in the C-limitations and could not therefore be an explanation for tree line formation. [ABSTRACT FROM AUTHOR]

The transition from seedlings into trees at alpine tree lines is a temperature-limited process that ultimately sets the tree line elevation at a global scale. As such, tree lines may be key bioassays of global warming effects on species distributions. For global warming to promote upward tree line migration, as predicted, seedlings must be available. We examined, for the first time at a global scale, elevational patterns and drivers of seedling availability at tree lines., Working at 10 sites across five mountain regions (dry Andes, humid Andes, Patagonian Andes, Swiss Alps and US Rocky Mountains) with different tree line forms (abrupt and diffuse) and dominated by different tree species (broadleaves and conifers), we answered the following question: How is seedling abundance affected by elevation (as a coarse grain surrogate of temperature), light exposure (openness immediately above plots) or wind exposure (an index for openness in the horizontal direction), or combinations thereof and what is the relative importance of each factor?, We tested five biological hypotheses to determine the relative strength of these tree line drivers on variable-size sampling plots of seedling abundance ( S) ( n = 1056). Specifically, we tested likely combinations of temperature limitation ( T), light as a resource (light, L) and as a radiation stress (via high light at low temperature, R), wind exposure as a tree line stressor ( W) and tree line form (a coarse scale test: abrupt vs. diffuse, D)., We found strong, moderate and weak negative effects of our estimates of wind exposure, radiation stress and elevation-related temperature on seedling abundance, respectively. We also found a positive effect, at tree line, for site-level tree line diffuseness. Two distinct facilitation mechanisms likely improved seedling abundance at tree line elevation: wind blockage by neighbourhood trees (the sheltering effect) and partial shading by overhead trees., Synthesis. Seedling abundance at alpine tree lines is limited by multiple simultaneous factors with the temperature decrease with elevation playing a relatively minor role. We therefore note that if the temperature threshold limiting the conversion from seedlings to adult trees is relaxed because of global warming, upward tree line migration will depend on the availability of shelter sites for seedlings. [ABSTRACT FROM AUTHOR]

PREMISE OF THE STUDY: The study of scaling examines the relative dimensions of diverse organismal traits. Understanding whether global scaling patterns are paralleled within species is key to identify causal factors of universal scaling. I examined whether the foliage-stem (Corner's rules), the leaf size-number, and the leaf mass-leaf area scaling relationships remained invariant and isometric with elevation in a wide-distributed treeline species in the southern Chilean Andes. METHODS: Mean leaf area, leaf mass, leafing intensity, and twig cross-sectional area were determined for 1-2 twigs of 8-15 Nothofagus pumilio individuals across four elevations (including treeline elevation) and four locations (from central Chile at 36°S to Tierra del Fuego at 54°S). Mixed effects models were fitted to test whether the interaction term between traits and elevation was nonsignificant (invariant). KEY RESULTS: The leaf-twig cross-sectional area and the leaf mass-leaf area scaling relationships were isometric (slope = 1) and remained invariant with elevation, whereas the leaf size-number (i.e., leafing intensity) scaling was allometric (slope ≠ -1) and showed no variation with elevation. Leaf area and leaf number were consistently negatively correlated across elevation. CONCLUSIONS: The scaling relationships examined in the current study parallel those seen across species. It is plausible that the explanation of intraspecific scaling relationships, as trait combinations favored by natural selection, is the same as those invoked to explain across species patterns. Thus, it is very likely that the global interspecific Corner's rules and other leaf-leaf scaling relationships emerge as the aggregate of largely parallel intraspecific patterns. [ABSTRACT FROM AUTHOR]

The growth limitation hypothesis ( GLH) is the most accepted explanation for treeline formation, but it has been scarcely examined in Mediterranean regions, where treelines are located at lower elevations than in temperate regions. The GLH states that low temperature is the ultimate environmental driver for treeline formation, constraining C-sinks (i.e. tissue formation) more than C-sources. The GLH predicts similar or increasing (but not decreasing) non-structural carbohydrate ( NSC) concentrations with elevation throughout the course of the growing season. We hypothesized that elevational trends in growth and NSC in Mediterranean regions are not determined by low temperature alone., We tested the GLH by examining the mean annual basal area and NSC concentrations in developing (new twigs) and ripened tissues (branch, stem) of trees growing at three elevations in three Mediterranean and three temperate treelines in the Chilean southern Andes (33° S, Kageneckia angustifolia; 36, 40 and 46° S, Nothofagus pumilio) and in Spain (36° N, Pinus sylvestris; 42° N, Pinus uncinata). Samples for NSC were taken at the onset of summer and autumn, which represent periods of contrasting drought intensities in Mediterranean regions., Tree growth decreased significantly with elevation in temperate treelines but not in Mediterranean treelines. In Mediterranean treelines, new twig NSC concentrations increased significantly with elevation in the early summer but not in the early autumn. In temperate treelines, in contrast, no elevational or seasonal variation (or interaction between them) was observed in new twig NSC concentrations. The NSC concentrations of the branches and stems from both climates showed no elevational trends. The soluble sugars' NSC fraction increased over the season in Mediterranean treelines and decreased in temperate treelines., Synthesis. Although we found support for the growth limitation hypothesis ( GLH) in temperate and Mediterranean treelines, our study shows that the tree growth and C balance in Mediterranean treelines are not controlled by low temperature alone. We suggest that environmental factors other than temperature explain the lower global elevation of Mediterranean treelines when compared to temperate treelines. [ABSTRACT FROM AUTHOR]

Aims: While treeline positions are globally correlated to growing season temperatures, seedling establishment, an important process of alpine treeline dynamics, is additionally controlled by regional-scale factors such as snow cover duration, desiccating winds and biotic interactions. Knowing that alpine treelines have shown contrasting responses to climate change, we determined the relative importance of key abiotic and biotic factors involved in seedling survival and growth. Location: McGerrigle Mountains, Parc National de la Gaspésie, Appalachian Range, eastern Quebec, Canada. Methods: In two white spruce (Picea glauca) treeline sites, we used the microclimate in the vicinity of tree islands, densely packed clusters of trees isolated from each other by alpine tundra vegetation, to assess the effects of abiotic variables (sum of degree days [DD], snowpack duration and a wind exposure index) as well as the effects of biotic interactions with neighbouring vegetation on the survival and growth of transplanted white spruce seedlings. For 3 yr, we surveyed seedling survival twice a year to discriminate between winter and summer survival, and measured seedling growth at the end of each growing season. We used Bayesian hierarchical models to estimate the relative effects of covariates on survival and growth. Results: Survival probability decreased in microsites wherewinter DD was high, and increased in microsites with longer snowpack duration. In wind-exposed microsites, seedling survival increased when neighbouring vegetation was present, indicating facilitative mechanisms. Seedling growth was positively affected by the duration of snow cover and tended to increase with higher DD during the previous year. In wind-sheltered microsites, seedling growth tended to be negatively affected by neighbouring vegetation, indicating competitive mechanisms. Conclusions: Our study demonstrates that seedling establishment is more sensitive to winter conditions, notably to the length of snow cover (which protects seedlings from frost and desiccation), than to summer temperature. Biotic interactions increased seedling establishment when environmental stresses were higher. We suggest that regional-scale factors such as winter climate and biotic interactions should be included inmodelling exercises to improve future treeline location forecasts. [ABSTRACT FROM AUTHOR]

If isotherms shift upslope or poleward due to climate warming, treeline trees will be exposed to more or less thermal constraints of growth depending on treeline shift rates. Climate warming is expected to enhance tree growth at alpine treelines. Isotherms may move upslope, but treeline trees may not; our study focused on trees growing at the altitudinal or latitudinal limits of tree existence today regardless of where the treeline will be in the future. [Extracted from the article]

In low temperature-adapted plants, including treeline trees, light-saturated photosynthesis is considerably less sensitive to temperature than growth. As a consequence, all plants tested so far show increased nonstructural carbohydrate (NSC) tissue concentrations when exposed to low temperatures. Reduced carbon supply is thus an unlikely cause for low temperature range limits of plants. For altitudinal treeline trees there is, however, a possibility that high NSC genotypes have been selected., Here, we explored this possibility using afforestations with single-provenance conifers along elevational gradients in the Southern Chilean Andes and the Swiss Alps., Tree growth was measured at each of four approximately equidistant elevations at and below the treeline. Additionally, at the same elevations, needle, branch and stem sapwood tissues were collected to determine NSC concentrations., Overall, growth decreased and NSC concentrations increased with elevation. Along with previous empirical and experimental studies, the findings of this study provide no indication of NSC reduction at the treeline; NSC increased in most species (each represented by one common population) towards their upper climatic limit. The disparity between carbon acquisition and structural carbon investment at low temperature (accumulation of NSC) thus does occur even among genotypes not adapted to treeline environments. [ABSTRACT FROM AUTHOR]

1. Altitudinal tree line ecotones (ATE) are among the most sensitive plant formations facing global warming as the altitudinal decrease in temperature is considered the driver controlling the upper elevation limit of tree lines world-wide. In this study, we attempted to answer the following questions: (i) how have the conditions during the last 2-3 centuries affected ATE tree growth (physiology) and recruitment (demography)? and (ii) how strong is synchrony between these two processes at the ATEs? 2. We used spatial sampling grids at different ATEs in two ecosystems on two subcontinents: Nothofagus pumilio in the Andes of Chilean Patagonia (46° SL) and Pinus albicaulis in the Rockies of Western Montana, USA (46° NL). Basal increment cores were extracted from trees to estimate the growth and recruitment date. An annual detrended basal area increment was estimated for each tree and was modelled against elevation and time. 3. Tree growth improved over multiple centuries at all tree lines. Recently ( c. 50 years), however, improvements are disappearing or reversing. The uppermost tree line trees showed moderate declines in Montana and incipient declines in Patagonia. The declines are most dramatic slightly below current tree line ( c. 200 m). Tree recruitment patterns showed that tree lines have been moving uphill in both regions until at least 40-70 years ago. These movements occurred primarily through abrupt pulses upward with infilling occurring concurrently (Patagonia) or at some time thereafter (Montana). 4. Synchrony between growth and recruitment occurred in the 18th and 19th centuries in both regions. This synchrony was negative in Patagonia and positive in Montana, with varying lag periods. During the 20th century, these patterns of synchrony were lost at all sites. This loss of synchrony suggests that we could be entering a global period in which temperature is no longer the dominant driver of key features of tree lines. 5. Synthesis. Our study shows that at two structurally different tree lines, recent and initial declines in growth and losses of long-term synchrony are occurring in the latter part of the 20th century. These findings are opposite to simplistic expectations of global warming effects on tree line dynamics and call for a model reformulation that uncouples drivers of growth and recruitment. [ABSTRACT FROM AUTHOR]

Two alternative hypotheses attempt to explain the upper elevation limit of tree lines world-wide, the carbon-limitation hypothesis (CLH) and the growth-limitation hypothesis (GLH); the altitudinal decrease of temperature is considered the driver constraining either carbon gain or growth. Using a widely distributed tree line species ( Nothofagus pumilio) we tested whether tree line altitude is explained by the CLH or the GLH, distinguishing local from global effects. We elaborated expectations based on most probable trends of carbon charging with altitude according to both hypotheses, considering the alternative effects of drought. Two climatically contrasting tree line ecotones in the southern Andes of Chile: Mediterranean (36°54′ S) and Patagonia (46°04′ S). At both locations, 35-50 trees of different ages were selected at each of four altitudes (including tree line), and stem and root sapwood tissues were collected to determine non-structural carbohydrate (NSC) concentrations. NSC accumulates whenever growth is more limited than photosynthesis. An altitudinal increase in NSCs means support for the GLH, while the opposite trend supports the CLH. We also determined stable carbon isotope ratios (δC) to examine drought constraints on carbon gain. NSC concentrations were positively correlated with altitude for stem tissue at the Mediterranean and root sapwood tissue at the Patagonia site. No depletion of NSC was found at either site in either tissue type. For both tissues, mean NSC concentrations were higher for the Patagonia site than for the Mediterranean site. Mean root sapwood NSC concentration values were five times higher than those of the corresponding stem sapwood at all altitudes. Values for δC were positively correlated with altitude in the Mediterranean site only. We found support for the GLH at the site without drought effects (Patagonia) and no support for the CLH at either site. It is suggested that drought moderated the effects of low temperature by masking the expected trend of the GLH at the Mediterranean site. [ABSTRACT FROM AUTHOR]