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Worldwide, the implementation of forest management guidelines has tried to mimic natural forest dynamics. The Oliver forest stand dynamics model has been successful in bridging natural forest dynamics associated with the disturbances of various spatial scales and silviculture also related to various spatial scales of management. However, the application of this model (and others) is restricted to tall forests. We claim here that the general research and commercial interest bias towards tall tree species has permeated silviculture and forest management, and that both the stand dynamics of short‐stature tree species and their treatment have remained largely unexplored and unknown. Using one short‐stature tree species, Nothofagus antarctica, as a model system, we show that this species indeed follows a stand dynamic different from other tall Nothofagus‐dominated forests in southern South America. This species forms short, mature, even‐aged stands without large canopy gaps that promote tree regeneration. With this in mind, we propose a conceptual ecology‐based silvicultural guideline focusing on the creation of strip cuts or gaps along with topsoil scarification to ensure regeneration and forest reorganization after disturbances. Policy implications. The acknowledgement that short‐stature tree species follow a stand dynamic different from tall tree species necessarily implies that they must be managed differently. This recognition, along with the implementation of ecology‐based silvicultural recommendations, represents strong arguments to make modifications to forest legislation that, together, must be aimed at conservation and sustainable forest management. [ABSTRACT FROM AUTHOR]

Functional traits have been proposed to define key dimensions of plant ecological strategies, but we lack consensus on whether traits can accurately predict plant demography. Despite theoretical expectations, it has been challenging to find consistent relationships between functional traits and growth.In this study, we quantified inter‐ and intraspecific trait variation (ITV) and individual growth rates of woody plants across a steep moisture gradient that varies 10‐fold in annual precipitation (350–3700 mm) in southern Chile and used a hierarchical Bayesian model to predict growth as a function of trait values.We show that large‐leaved species with lower stem tissue density exhibited the fastest growth rates, and these two traits exhibited the highest proportion of interspecific variation. Predictions of growth improved considerably (R2 of the best model increased from 0.28 to 0.49) when species‐level multiple traits and their interactions were considered. The inclusion of ITV, however, did not improve models of growth rate.We found that trait–growth rate relationships were not always consistent across levels of biological organization; relationships observed at the interspecific level did not necessarily hold at the intraspecific level. We found that the relationships between wood density or leaf size and growth were consistent in direction across the precipitation gradient, and the relationships between leaf economics traits and growth were weak and site‐specific.Synthesis. Although using more than one functional trait considerably improved growth predictions, wood density and leaf size successfully predicted growth rates across (not within) species, which is consistent with a whole‐plant carbon economy. We assert that these two traits are intimately linked and ultimately describe a continuum of plant architecture and carbon economy that covers multiple trait syndromes. [ABSTRACT FROM AUTHOR]

Summary Statement: We propose to add some aspects that are of great complementary importance to broaden the kin recognition, discrimination and selection framework in plants and that need to be considered in future studies. [ABSTRACT FROM AUTHOR]

Background and Aims The vulnerability and responsiveness of forests to drought are immensely variable across biomes. Intraspecific tree responses to drought in species with wide niche breadths that grow across contrasting climatically environments might provide key information regarding forest resistance and changes in species distribution under climate change. Using a species with an exceptionally wide niche breath, we tested the hypothesis that tree populations thriving in dry environments are more resistant to drought than those growing in moist locations. Methods We determined temporal trends in tree radial growth of 12 tree populations of Nothofagus antarctica (Nothofagaceae) located across a sharp precipitation gradient (annual precipitation of 500–2000 mm) in Chile and Argentina. Using dendrochronological methods, we fitted generalized additive mixed-effect models to predict the annual basal area increment as a function of year and dryness (De Martonne aridity index). We also measured carbon and oxygen isotope signals (and estimated intrinsic water-use efficiency) to provide potential physiological causes for tree growth responses to drought. Key Results We found unexpected improvements in growth during 1980–1998 in moist sites, while growth responses in dry sites were mixed. All populations, independent of site moisture, showed an increase in their intrinsic water-use efficiency in recent decades, a tendency that seemed to be explained by an increase in the photosynthetic rate instead of drought-induced stomatal closure, given that δ18O did not change with time. Conclusions The absence of drought-induced negative effects on tree growth in a tree species with a wide niche breadth is promising because it might relate to the causal mechanisms tree species possess to face ongoing drought events. We suggest that the drought resistance of N. antarctica might be attributable to its low stature and relatively low growth rate. [ABSTRACT FROM AUTHOR]

Premise: Climate change may lead to C stress (negative C balance) in trees. Because nonstructural carbohydrates (NSC) are required during metabolic reactivation in the spring, C stress might delay budbreak timing. This effect is expected to be greater in shade‐intolerant than in shade‐tolerant species, owing to the faster C economy in the shade‐intolerant. Methods: We experimentally induced C stress in saplings of six temperate tree species that differed in their light requirements by exposing them to either full light or shade from summer to spring, then recorded the date of first budbreak for the individuals. Because the levels of C reserves that represent effective C stress may differ among species, we estimated the degree of C stress by recording survival during the experiment and measuring whole‐sapling NSC concentrations after budbreak. Results: Shade reduced NSC concentrations and increased the sugar fraction in the NSC in all species. In the shade, shade‐intolerant species had higher mortality and generally lower NSC concentrations than the shade‐tolerant species, indicating a trend for more severe C stress in species with faster C economy. In shade‐intolerant species, budbreak started earlier and proceeded faster in full light than in shade, but in shade‐tolerant species budbreak was delayed in full light. The effects of the light environments on budbreak were not greater in shade‐intolerant than in shade‐tolerant species. Conclusions: Our study reveals a correspondence between budbreak responses to light and the light requirements of the species. This finding confirms that C metabolism has a significant role in triggering budbreak and demonstrates that whether C stress accelerates or delays budbreak depends on the species' light requirements. [ABSTRACT FROM AUTHOR]

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]

The effective use of functional traits to explain species coexistence has been hindered by the scarce use of demographic variables, such as growth, with the association between functional traits and growth often assumed rather than tested. In this study, using spatial point‐pattern analyses, we spatially related three functional traits linked to plant competitive ability (maximum tree height [HMAX], leaf mass per area [LMA] and wood density [WD]) to tree growth rates (basal area increment [BAI10] and growth efficiency [GE]) in a stem‐mapped plot of a temperate rainforest in southern Chile. With this, we tested the competition‐trait similarity and the competition‐trait hierarchy hypotheses, and further assessed the relationship between functional trait values' spatial distribution and growth rate. We used spatial point‐pattern analyses, in particular, pair‐wise spatial association patterns, the Schlather's I index and mark variograms associated to specific null models to infer spatial correlations among individuals of several species with different traits and growth values. First, we found spatial similarity in trees' functional traits on a fine‐scale, a result supporting the competition‐trait hierarchy hypothesis. Second, while the HMAX and LMA specific values of the focal tree were spatially positively correlated with their neighbours' growth rates, the WD of the focal tree had a negative spatial correlation with its neighbours' growth rates. We also found a positive spatial autocorrelation among HMAX, LMA and WD at a fine‐scale; neighbouring trees tended to have similar values. In agreement with our a priori statements, we infer that HMAX and LMA provide a competitive tolerance, while WD of the focal tree has a competitive effect on its immediate neighbours. As such, we have demonstrated that functional traits mediate species interactions, which is a step forward in predictive community ecology, thus we can further our understanding regarding how traits mediate community assembly and species coexistence. [ABSTRACT FROM AUTHOR]

Summary: The success of invasive plants is influenced by many interacting factors, but evaluating multiple possible mechanisms of invasion success and elucidating the relative importance of abiotic and biotic drivers is challenging, and therefore rarely achieved.We used live, sterile or inoculated soil from different soil origins (native range and introduced range plantation; and invaded plots spanning three different countries) in a fully factorial design to simultaneously examine the influence of soil origin and soil abiotic and biotic factors on the growth of invasive Pinus contorta.Our results displayed significant context dependency in that certain soil abiotic conditions in the introduced ranges (soil nitrogen, phosphorus or carbon content) influenced responses to inoculation treatments.Our findings do not support the enemy release hypothesis or the enhanced mutualism hypothesis, as biota from native and plantation ranges promoted growth similarly. Instead, our results support the missed mutualism hypothesis, as biota from invasive ranges were the least beneficial for seedling growth. Our study provides a novel perspective on how variation in soil abiotic factors can influence plant–soil feedbacks for an invasive tree across broad biogeographical contexts. [ABSTRACT FROM AUTHOR]

Worldwide drought events have been reported to cause tree growth decline and mortality, thus altering the carbon (C) balance of forest ecosystems. While most of the attention has been focused on the physiological mechanisms associated with drought‐induced tree responses of a few species at specific locations, the ecological attributes of these species, like their niche breadth, may be also important in determining species' sensitivity or resilience to drought. We postulated that wide‐niche breadth tree species should be more drought‐resilient than narrow‐niche breadth species.Using the most severe 2015–2016 El Niño drought event in the last 70 years in Patagonia, we determined pre‐ and post‐drought growth (BAI, basal area increment), C reserves in the form of non‐structural carbohydrate concentrations (NSCs = starch and soluble sugars), wood isotope (δ13C, iWUE and δ18O) signalling and xylem anatomy (mean vessel diameter, mvd) in eight angiosperm tree species of contrasting niche breadth across a sharp precipitation gradient in southern Chile.All species responded in unison after the drought with a non‐water‐conservative response, maintaining BAI and NSCs, decreasing δ13C, and increasing both mvd and the soluble sugars:NSCs ratio relative to pre‐drought time. Contrary to previous results reporting species‐specific drought responses, our results show unequivocally a functional coordination of organisms' vital traits associated with a non‐water‐conservative strategy, and a drought‐induced acclimation based on starch conversion into soluble sugars in all of the tree species we examined, regardless of their niche breadth and habitat preference.We state that abiotic drivers such as drought may have selected similar interspecific responses provided that they operate at the community level rather than at the species level. These findings mark the need to revise current views about the ultimate interspecific functional coordination of organisms' vital traits when facing more frequent and intensive drought events. A free Plain Language Summary can be found within the Supporting Information of this article. [ABSTRACT FROM AUTHOR]

Background and Aims Twig cross-sectional area and the surface area of leaves borne on it are expected to be isometrically correlated across species (Corner's rules). However, how stable this relationship remains in time is not known. We studied inter- and intraspecific twig leaf area–cross-sectional area (l a– c s) and other scaling relationships, including the leaf–shoot mass (l m– s m) scaling relationship, across a complete growing season. We also examined the influence of plant height, deciduousness and the inclusion of reproductive buds on the stability of the scaling relationships, and we discuss results from a functional perspective. Methods We collected weekly current-year twigs of six Patagonian woody species that differed in growth form and foliar habit. We also used prominent inflorescences from Embothrium coccineum (Proteaceae) to assess whether reproductive buds alter the l a– c s isometric relationship. Mixed effects models were fitted to obtain parameter estimates and to test whether interaction terms were non-significant (invariant) for the scaling relationships. Key Results The slope of the l a– c s scaling relationship remained invariant across the growing season. Two species showed contrasting and disproportional (allometric) l a– c s scaling relationships (slope ≠ 1). Scaling relationships varied significantly across growth form and foliar habit. The l m– s m scaling relationship differed between reproductive- and vegetative-origin twigs in E. coccineum , which was explained by a significantly lower leaf mass per area in the former. Conclusions Although phenology during the growing season appeared not to change leaf–shoot scaling relationships across species, we show that scaling relationships departed from the general trend of isometry as a result of within-species variation, growth form, foliar habit and the type of twig. The identification of these functional factors helps to understand variation in the general trend of Corner's rules. [ABSTRACT FROM AUTHOR]

Background and Aims Plants have the potential to adjust the configuration of their hydraulic system to maintain its function across spatial and temporal gradients. Species with wide environmental niches provide an ideal framework to assess intraspecific xylem adjustments to contrasting climates. We aimed to assess how xylem structure in the widespread species Nothofagus pumilio varies across combined gradients of temperature and moisture, and to what extent within-individual variation contributes to population responses across environmental gradients. Methods We characterized xylem configuration in branches of N. pumilio trees at five sites across an 18° latitudinal gradient in the Chilean Andes, sampling at four elevations per site. We measured vessel area, vessel density and the degree of vessel grouping. We also obtained vessel diameter distributions and estimated the xylem-specific hydraulic conductivity. Xylem traits were studied in the last five growth rings to account for within-individual variation. Key Results Xylem traits responded to changes in temperature and moisture, but also to their combination. Reductions in vessel diameter and increases in vessel density suggested increased safety levels with lower temperatures at higher elevation. Vessel grouping also increased under cold and dry conditions, but changes in vessel diameter distributions across the elevational gradient were site-specific. Interestingly, the estimated xylem-specific hydraulic conductivity remained constant across elevation and latitude, and an overwhelming proportion of the variance of xylem traits was due to within-individual responses to year-to-year climatic fluctuations, rather than to site conditions. Conclusions Despite conspicuous adjustments, xylem traits were coordinated to maintain a constant hydraulic function under a wide range of conditions. This, combined with the within-individual capacity for responding to year-to-year climatic variations, may have the potential to increase forest resilience against future environmental changes. [ABSTRACT FROM AUTHOR]

Summary: Variation in xylem conduit diameter traditionally has been explained by climate, whereas other evidence suggests that tree height is the main driver of conduit diameter. The effect of climate versus stem length on vessel diameter was tested in two tree species (Embothrium coccineum, Nothofagus antarctica) that both span an exceptionally wide precipitation gradient (2300–500 mm).To see whether, when taking stem length into account, plants in wetter areas had wider vessels, not only the scaling of vessel diameter at the stem base across individuals of different heights, but also the tip‐to‐base scaling along individuals of similar heights across sites were examined.Within each species, plants of similar heights had similar mean vessel diameters and similar tip‐to‐base widening of vessel diameter, regardless of climate, with the slopes and intercepts of the vessel diameter–stem length relationship remaining invariant within species across climates.This study focusing on within‐species variation––thus, avoiding noise associated with the great morphological variation across species––showed unequivocally that plant size, not climate, is the main driver of variation in vessel diameter. Therefore, to the extent that climate selects for differing vessel diameters, it will inevitably also affect plant height. [ABSTRACT FROM AUTHOR]

Background and aims Cluster roots (CRs) constitute a special root adaptation that enables plants to take up nutrients, especially phosphorus (P), from soils with low nutrient availability, including recent volcanic deposits. It is unclear, however, how CR species interact with non-cluster root-bearing (NCR) species, and how substrates' fertility modulates potential interactions. Methods We experimentally assessed the net interaction between CR and NCR species using two substrates of contrasting fertility: nutrient-rich nursery mix and tephra (low P availability). We planted seedlings of two southern South American (SSA) Proteaceae, CR species and two NCR Nothofagus species in pairs (conspecifics and heterospecifics) and as singles. We analysed the effect of seedling neighbours on survival, growth performance (e.g. total biomass and leaf area) and leaf and substrate nutrient concentrations (including manganese, a proxy for P-acquisition efficiency through CR activity) using the relative interaction index. Key Results After three growing seasons, we found that (1) Proteaceae species had fewer CRs and lower CR biomass and grew less in the tephra than in the nursery substrate; (2) Nothofagus species did not improve their survival and growth in the presence of Proteaceae species in any substrate; (3) contrary to Nothofagus , Proteaceae species improved their growth more when planted with any neighbour (including conspecifics) than when planted alone, which was accompanied by a significant accretion of leaf P; and (4) the presence of a neighbour increased the final nitrogen and P concentrations in the nursery substrate, regardless of species identity. Conclusions CRs provide Proteaceae a competitive advantage over NCR species at the seedling stage, which may have important consequences for species coexistence and community structuring. The investigated SSA Proteaceae, which have not evolved in nutrient-impoverished soils, as have their relatives in south-western Australia and South Africa, improve their growth when cultivated in pairs, especially in nutrient-rich substrates. [ABSTRACT FROM AUTHOR]

Several plant species have the potential to facilitate the presence of other plant species in the community, and yet most of our knowledge on this phenomenon comes from studies considering only one facilitator species. What happens when there are two facilitator species in the community? Are the facilitation effects by one species being altered by the presence of a potentially second facilitator species and vice‐versa (i.e. bidirectional facilitation)?Knowing that cushion plants ameliorate stress and improve soil fertility and that cluster root‐bearing species mobilize soil phosphorus (P), we investigated the potential occurrence of bidirectional facilitation via stress amelioration and nutrient exchange in the cushion plant Acaena integerrima (Rosaceae) and the cluster root‐bearing tree species Embothrium coccineum (Proteaceae).In southern Chile, where both species naturally occur, we compared aspects of the vegetative and reproductive performance of each species growing in association or not. We also determined if Embothrium alters the nurse effects of Acaena cushions on other species by evaluating species richness, frequency, and abundance in cushion plants with and without Embothrium. To infer the existence of nutrient exchange, we determined soil and leaf nutrient levels of both species.Species richness, frequency and abundance were higher within than outside Acaena cushions. Although the presence of Embothrium did not enhance the overall positive effects of Acaena on species composition, nine species occurred more frequently when Embothrium was present. Embothrium occurred more frequently and had a higher survival and leaf nitrogen (N) concentration when growing with Acaena than without it, although its growth and leaf P concentration were lower. In contrast, Acaena had a lower number of and mass of fruits per inflorescence and a higher leaf P concentration when growing with versus growing without Embothrium.Synthesis. The cushion plant Acaena integerrima facilitates the occurrence of many species in the community, including the cluster root‐bearing species Embothrium coccineum, which in turn enhances the facilitation effect of Acaena on certain species. When Embothrium and Acaena occur together, the latter improves its nutritional status but reduces its fitness, implying a lack of bidirectional facilitation. [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]

It is assumed that widespread, generalist species have high phenotypic variation, but we know little about how intraspecific trait variation (ITV) relates to species abundance and niche breadth. In the temperate rainforest of southern Chile, we hypothesized that species with wide niche breadth would exhibit 1) high among‐plot ITV, 2) a strong relationship between trait values and the environment, and 3) a close fit between traits and local environment trait optima. We measured leaf functional traits (leaf area, LMA, leaf N and P concentrations) of saplings in woody species, and compared the relative abundance of each species with its niche breadth, measured as the range of light, soil N and P availability. We used the slope of the linear regression of species' trait–environment relationships to assess the strength and direction of these relationships, and measured the degree to which species' trait values track the environmental optimum across plots. In some cases, species having wide niche breadth had high ITV in leaf N and also matched traits (LMA and leaf P) to local optima along the light gradient; they also had high ITV in general and matched leaf P to local optima along the soil P gradient. The relationship between species with wide niche breadth and the strength of intraspecific trait–environment relationships was generally weak and varied depending on the niche dimension and trait in question. Species varied considerably in the strength of trait–environment relationships and total magnitude of ITV, and this variation was not generally strongly related to species abundances or niche breadth patterns. In conclusion, trait variation at the community level is not driven by a few abundant, widely distributed species, but depends on the aggregate trait responses of both abundant and rare species. This makes it difficult to scale individual species trait responses up to the community level. [ABSTRACT FROM AUTHOR]

With their imposing grandeur, the small number of very tall tree species attract a disproportionate amount of scientific study. We right this bias by focusing here on the shorter trees, which often grow in the shade of the giants and many other places besides. That tall trees are so restricted in distribution indicates that there are far more habitats available for small trees. We discuss some leading candidates for the mechanisms that limit maximum plant height in any given habitat, as well as why every habitat has a range of plant sizes. At least two attributes – greater adaptation capacity and higher drought resistance – suggest that the forests of the future belong to short trees. Highlights Global warming threatens many tree species, particularly through drought. We give reasons to think that short-statured tree species are best equipped to face drought and global warming in general because they have a high adaptation capacity and a high resistance to drought-induced cavitation. Short trees are able to persist in a higher number of niches than tall species, which are much restricted to a narrow climatic belt. [ABSTRACT FROM AUTHOR]

Mega‐dams create highly fragmented archipelagos, affecting biodiversity and ecosystem functioning in remnant forest isolates. This study assessed the long‐term impact of dam‐induced fragmentation on insular tropical tree communities, with the aim of generating robust recommendations to mitigate some of the detrimental biodiversity impacts associated with future dam development.We inventoried adult and sapling trees across 89 permanent plots, located on 36 islands and in three mainland continuous forest sites in the Balbina Dam, Brazilian Amazon. We examined differences in recruitment, structure, and composition of sapling and adult tree communities, in relation to plot‐, patch‐ and landscape‐scale attributes including area, isolation, and fire severity.Islands harboured significantly lower sapling (mean ± 95% CI 48.6 ± 3.8) and adult (5 ± 0.2) tree densities per 0.01 ha, than nearby mainland continuous forest (saplings, 65.7 ± 7.5; adults, 5.6 ± 0.3). Insular sapling and adult tree communities were more dissimilar than in mainland sites, and species compositions showed a directional shift away from mainland forests, induced by fire severity, island area, and isolation.Insular sapling recruitment declined with increasing fire severity; tree communities with higher community‐weighted mean wood density showed the greatest recruitment declines. Our results suggest that insular tree communities are unstable, with rare species becoming extinction‐prone due to reduced tree recruitment and density on islands, potentially leading to future losses in biodiversity and ecosystem functioning across Balbina's >3,500 reservoir islands.Policy implications. In Balbina, fire and reduced habitat area and connectivity were drivers of tree community decay after only 28 years of insularization, despite strict protection provided by the ~940,000 ha Uatumã Biological Reserve. Given that many dams are planned for lowland, moderately undulating Amazonia, we recommend that dam development strategy explicitly considers (a) dam location, aiming to minimize creation of small (<10 ha) and isolated islands, (b) maintaining reservoir water levels during droughts to reduce fire risk, and (c) including aggregate island area in environmental impact and offset calculations. Ideally, we recommend that alternatives to hydropower be sought in lowland tropical regions, due to the far‐reaching biodiversity losses and ecosystem disruption caused by river impoundment. [ABSTRACT FROM AUTHOR]

Premise of the Study: It is unclear to what extent the co‐occurrence of angiosperm and gymnosperm species in some marginal ecosystems is explained by reduced growth in angiosperms due to carbon (C) limitation and by high stress tolerance in gymnosperms associated with lack of vessels and resource conservation. Methods: We examined growth patterns and traits associated with C balance in four evergreen angiosperm species (including one vesselless species, Drimys winteri) and three gymnosperm tree species of a cold‐temperate rainforest in southern Chile. We measured the mean basal area increment for the first 50 (BAI50) and the last 10 years (BAI10), wood density, leaf lifespan, and nonstructural carbohydrate (NSC) concentrations in different organs. Key Results: BAI50 was 6‐fold higher in angiosperms than in gymnosperms and ca. 4‐fold higher in Drimys than in the fastest‐growing gymnosperm. BAI10 and aboveground NSC concentrations were significantly higher and leaf lifespan lower in angiosperms than in gymnosperms; these differences, however, were largely driven by the slow growth and low NSC concentrations of the Cupressaceae species (Pilgerodendron uviferum), while the two Podocarpaceae had BAI10 and NSC concentrations similar to angiosperms. In angiosperms, NSC and starch concentrations were generally higher in species with lower BAI10, indicating no severe C limitation. Conclusions: The co‐occurrence of angiosperms and gymnosperms in cold‐temperate rainforests of southern Chile is not explained by growth disadvantages and C limitation in angiosperms. Long leaf longevity, but not lack of vessels, appeared to favor resource conservation and C balance in some gymnosperms (Podocarpaceae). [ABSTRACT FROM AUTHOR]

Abstract: Understanding patterns of functional trait variation across environmental gradients offers an opportunity to increase inference in the mechanistic causes of plant community assembly. The leaf economics spectrum (LES) predicts global tradeoffs in leaf traits and trait‐environment relationships, but few studies have examined whether these predictions hold across different levels of organization, particularly within species. Here, we asked (1) whether the main assumptions of the LES (expected trait relationships and shifts in trait values across resource gradients) hold at the intraspecific level, and (2) how within‐species trait correlations scale up to interspecific or among‐community levels. We worked with leaf traits of saplings of woody species growing across light and soil N and P availability gradients in temperate rainforests of southern Chile. We found that ITV accounted for a large proportion of community‐level variation in leaf traits (e.g., LMA and leaf P) and played an important role in driving community‐level shifts in leaf traits across environmental gradients. Additionally, intraspecific leaf trait relationships were generally consistent with interspecific and community‐level trait relationships and with LES predictions—e.g., a strong negative intraspecific LMA–leaf N correlation—although, most trait relationships varied significantly among species, suggesting idiosyncrasies in the LES at the intraspecific level. [ABSTRACT FROM AUTHOR]

Since growth is more sensitive to drought than photosynthesis, trees inhabiting dry regions are expected to exhibit higher carbohydrate storage and less growth than their conspecifics from more humid regions. However, the same pattern can be the result of different genotypes inhabiting contrasting humidity conditions. To test if reduced growth and high carbohydrate storage are environmentally driven by drought, we examined the growth and non-structural carbohydrate (NSC) concentrations in singleprovenance stands of mature trees of Pinus contorta Douglas and Pinus ponderosa Douglas ex C. Lawson planted at contrasting humidity conditions (900 versus 300mm of annual precipitation) in Patagonia, Chile. Individual tree growth was measured for each species and at each location as mean basal area increment of the last 10 years (BAI10), annual shoot elongation for the period 2011-14, and needle length for 2013 and 2014 cohorts. Additionally, needle, branch, stem sapwood and roots were collected from each sampled tree to determine soluble sugars, starch and total NSC concentrations. The two species showed lower mean BAI10 and 2013 needle length in the dry site; P. ponderosa also had lower annual shoot extension for 2011 and 2014, and lower 2014 needle length, in the dry than in the mesic site. By contrast, NSC concentrations of all woody tissues for both species were either similar or higher in the dry site when compared with the mesic site. Patterns of starch and sugars were substantially different: starch concentrations were similar between sites except for roots of P. ponderosa, which were higher in the dry site, while sugar concentrations of all woody tissues in both species were higher in the dry site. Overall, our study provides evidence that reduced growth along with carbon (C) accumulation is an environmentally driven response to drought. Furthermore, the significant accumulation of low-molecular weight sugars in the dry site is compatible with a prioritized C allocation for osmoregulation. However, since this accumulation did not come at the expense of reduced starch, it is unlikely that growth was limited by C supply in the dry site. [ABSTRACT FROM AUTHOR]

Climate change and rising atmospheric CO 2 concentrations ( c a ) are expected to affect forests worldwide. The effects of climate change, however, have not been deeply assessed in humid forest biomes from the southern Hemisphere where climate is not warming but drying. This is the case of the temperate rainforest in southern Chile, where the endemic and threatened long-living gymnosperm Fitzroya cupressoides occurs. We assessed how radial growth, intrinsic water-use efficiency (iWUE) and tree-ring δ 18 O responded to increasing c a and decreasing precipitation in F. cupressoides and companion species. We hypothesized that F. cupressoides , a long-lived and probably less plastic species, will show less acclimation to global-change effects than co-occurring Nothofagus species which show broader climatic niche. Thus, F. cupressoides should display iWUE increases different from the c i / c a constant scenario, which represents an active mechanism to increase intercellular CO 2 concentrations ( c i ) as c a rises. Although cool and wet conditions during the growing season enhanced growth of all species, particularly in F. cupressoides , growth of F. cupressoides declined noticeably since the 1980s in response to a decrease in precipitation. Current drier conditions led to increased iWUE in Nothofagus species. According to δ 18 O values, this increased in iWUE should be due to a decrease in stomatal conductance. Fitzroya cupressoides , however, displayed a decrease in iWUE in response to drier conditions, shifting from an active c i / c a scenario to a more passive c i / c a scenario, and maintaining a relatively constant stomatal conductance. Using multiple bodies of evidence, our findings indicate a poor adaptability of the long-lived F. cupessoides to drier conditions despite rising c a . Thus, not all species are having similar and expected responses to increasing c a , which should be a call of attention in the case of long-lived, endangered and narrow-distributed species, like F. cupressoides . [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]

• Background and Aims Ecologists are increasingly using plant functional traits to predict community assembly, but few studies have linked functional traits to species' responses to fine-scale resource gradients. In this study, it was tested whether saplings of woody species partition fine-scale gradients in light availability based on their leaf mass per area (LMA) in three temperate rain forests and one Mediterranean forest in southern Chile. • Methods LMA was measured under field conditions of all woody species contained in approx. 60 plots of 2 m2 in each site, and light availability, computed as the gap light index (GLI), was determined. For each site, species' pairwise differences in mean LMA (▵ LMA) and abundance-weighted mean GLI (▵ light response) of 2 m2 plots were calculated and it was tested whether they were positively related using Mantel tests, i.e. if species with different LMA values differed in their response to light availability. Additionally linear models were fitted to the relationship between plot-level mean LMA and GLI across plots for each site. • Key Results A positive and significant relationship was found between species' pairwise differences in mean LMA and differences in light response across species for all temperate rain forests, but not for the Mediterranean forest. The results also indicated a significant positive interspecific link between LMA and light availability for all forests. This is in contrast to what is traditionally reported and to expectations from the leaf economics spectrum. • Conclusions In environments subjected to light limitation, interspecific differences in a leaf trait (LMA) can explain the fine-scale partitioning of light availability gradients by woody plant species. This niche partitioning potentially facilitates species coexistence at the within-community level. The high frequency of evergreen shadeintolerant species in these forests may explain the positive correlation between light availability and LMA. [ABSTRACT FROM AUTHOR]

Carbon (C) storage is considered a key component to plant survival under drought and shade, although the combined effects of these factors on survival remain poorly understood. We investigated how drought and shade alter the C dynamics and survival of tree seedlings, and whether drought limits the access to or usage of stored C. We experimentally applied two levels of soil humidity (wellwatered versus drought, the latter induced by dry-down) and light availability (light versus complete darkness) on 1-year-old seedlings of Acer pseudoplatanus L. for 3 months. We quantified the survival, biomass, growth rate and non-structural carbohydrates (NSC) of seedlings at their time of death or at the end of the experiment for those that survived. We found that the soil dried out faster when drought was combined with light than when it was combined with complete darkness. Seedlings subjected to both drought and light showed reduced growth and reached 100%mortality earlier than any other treatment, with the highest NSC concentrations at the time of death. Seedlings exposed to both drought and complete darkness died significantly earlier than seedlings exposed to complete darkness only, but had similar NSC concentrations at time of their death, suggesting that drought accelerated the use of stored C under complete darkness. Complete darkness significantly reduced seedling growth and whole-plant NSC concentrations regardless of soil humidity, while root NSC concentrations were significantly more reduced when complete darkness was combined with drought conditions. Thus, the C dynamics in A. pseudoplatanus seedlings under complete darkness was not hindered by drought, i.e., the access and use of stored C was not limited by drought. The contrasting growth and C storage responses driven by drought under light versus complete darkness are consistent with a key role of the drought progression in the C dynamics of trees. [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]

A basic assumption of the trait-based approach in plant ecology is that differences in functional trait values are greater between species than within species. We questioned this assumption by assessing (1) the relative extent of inter- and intraspecific leaf trait variation throughout a complete growing season (phenological variation) in a group of deciduous and evergreen woody species, and (2) whether species rankings based on leaf traits were maintained across the growing season. We analysed leaf mass per area (LMA) and leaf nutrient concentrations (C, N, P), including the C:N and N:P ratios. Intraspecific trait variation (ITV) due to phenology was significantly greater than interspecific variation for leaf N concentration on a mass basis (N; 68.90 %) and for the leaf C:N ratio (60.60 %), whereas interspecific variation was significantly higher than ITV for LMA (62.30 %) and for leaf C concentration on a mass (C) and area (C) basis (C 70.40 %; C 65.30 %). ITV was particularly low for LMA (<20 %). Species rankings were highly modified by phenology for a number of leaf traits (P, N:P ratio) but were relatively well conserved throughout the growing season for others (LMA, N). Patterns of ITV across the growing season differed significantly between deciduous and evergreen species for all traits except leaf P but did not vary between native and exotic species. Overall, our results show that intraspecific phenological variation in leaf traits may be similar to or greater than interspecific variation and that temporal patterns of ITV vary considerably among traits and species, especially for leaf nutrient concentrations, factors which can potentially affect quantitative interspecific relationships. [ABSTRACT FROM AUTHOR]

Background and Aims Disturbances, dispersal and biotic interactions are three major drivers of the spatial distribution of genotypes within populations, the last of which has been less studied than the other two. This study aimed to determine the role of competition and facilitation in the degree of conspecific genetic relatedness of nearby individuals of tree populations. It was expected that competition among conspecifics will lead to low relatedness, while facilitation will lead to high relatedness (selection for high relatedness within clusters). Methods The stand structure and spatial genetic structure (SGS) of trees were examined within old-growth and second-growth forests (including multi-stemmed trees at the edge of forests) of Nothofagus pumilio following large-scale fires in Patagonia, Chile. Genetic spatial autocorrelations were computed on a spatially explicit sampling of the forests using five microsatellite loci. As biotic plant interactions occur among immediate neighbours, mean nearest neighbour distance (MNND) among trees was computed as a threshold for distinguishing the effects of disturbances and biotic interactions. Key Results All forests exhibited a significant SGS for distances greater than the MNND. The old-growth forest genetic and stand structure indicated gap recolonization from nearby trees (significantly related trees at distances between 4 and 10 m). At distances smaller than the MNND, trees of the second-growth interior forest showed significantly lower relatedness, suggesting a fading of the recolonization structure by competition, whereas the second-growth edge forest showed a positive and highly significant relatedness among trees (higher among stems of a cluster than among stems of different clusters), resulting from facilitation. Conclusions Biotic interactions are shown to influence the genetic composition of a tree population. However, facilitation can only persist if individuals are related. Thus, the genetic composition in turn influences what type of biotic interactions will take place among immediate neighbours in post-disturbance forests. [ABSTRACT FROM AUTHOR]

* Premise of the study: Southern South American (SA) Proteaceae species growing in volcanic soils have been proposed as potential ecosystem engineers by tapping phosphorus (P) from soil through their cluster roots and shedding nutrient-rich litter to the soil, making it available for other species. We tested whether Embothrium coccineum (Proteaceae) has effectively lower P nutrient resorption efficiency and higher litter P concentrations than co-occurring, non-Proteaceae species. * Methods: In southern Chile, we assessed the P and nitrogen (N) resorption efficiency of senescent leaves and fresh litter of E. coccineum and co-occurring tree species in a soil fertility and moisture gradient (600-3000 mm of annual precipitation) in Patagonia, Chile. We determined P and N concentrations, leaf mass per area (LMA), and ratios of N/P and C/N in mature and senescent leaf cohorts and fresh litter. * Key results: Embothrium coccineum showed significantly higher P and N resorption efficiency than co-occurring species; in fact, E. coccineum fresh litter had the lowest P-content. While E. coccineum showed significantly lower fresh litter P concentrations than the rest of the species, it showed significantly higher N concentrations. Embothrium coccineum also had lower LMA and similar N/P and C/N ratios when compared with co-occurring tree species. * Conclusions: We found that E. coccineum efficiently mobilized P and, to a lesser extent, N before leaf shedding. We did not find support for the ecosystem engineering hypothesis via shedding P-rich litter. We suggest that southern South American Proteaceae may be taking up other nutrients besides P, probably N, from the young, volcanic soils of this region. [ABSTRACT FROM AUTHOR]

An innovative hypothesis to explain the higher carbon (C) and nitrogen (N) storage in woody tissues of winter deciduous species as compared to evergreen species is that these storages reflect an adaptation to tolerate herbivory. Support for this hypothesis has been little when manipulative defoliations were partial and/or applied in a single season. Given that repeated defoliations throughout a single season are common in temperate forests and tend to be more severe in deciduous than in evergreen species, we tested this hypothesis considering complete and recurrent defoliation in two sympatric Nothofagus species with contrasting foliar habit., In the field, we applied three defoliation intensities for 3 years in naturally coexisting juvenile trees of Nothofagus betuloides (evergreen) and Nothofagus pumilio (deciduous). Defoliation intensities included complete defoliation (100%) twice during the growing season, partial defoliation (50%) twice during the whole experiment and no defoliation. We evaluated survival, regrowth and C- and N-storage in the leaves, stems and roots of each tree., Complete defoliation caused 100% mortality in N. betuloides after the first year and no mortality in N. pumilio after 3 years; it induced higher C reductions in N. pumilio roots, supported by greater C-storage. Partial defoliation caused no interspecific differences in survival, though it produced a stronger decrease in C-storage in N. betuloides than in N. pumilio. N concentrations in woody tissues were significantly higher in N. pumilio than in N. betuloides, and only in the former did they decrease with the defoliation intensity., Synthesis. We found a potential functional link between leaf habit, defoliation tolerance and C- and N-storage. The deciduous species tolerated complete and recurrent defoliations better than the evergreen species, which was associated with higher C- and N-storage in stems and roots of the former. This link was not detected under partial defoliation. We suggest that the higher C- and N-storage in the woody tissues of deciduous species as compared to evergreen species is an adaptation to tolerate complete and recurrent defoliations under which temperate winter deciduous species may have evolved. [ABSTRACT FROM AUTHOR]

* Premise of the study: The growth limitation hypothesis (GLH) is the most accepted mechanistic explanation for treeline formation, although it is still uncertain whether it applies across taxa. The successful establishment of Pinus contorta--an exotic conifer species in the southern hemisphere--above the Nothofagus treeline in New Zealand may suggest a different mechanism. We tested the GLH in Nothofagus pumilio and Pinus contorta by comparing seedling performance and carbon (C) balance in response to low temperatures. * Methods: At a southern Chilean treeline, we grew seedlings of both species 2 m above ground level, to simulate coupling between temperatures at the meristem and in the air (colder), and at ground level, i.e., decoupling air temperature (relatively milder). We recorded soil and air temperatures as well. After 3 yr, we measured seedling survival and biomass (as a surrogate of growth) and determined nonstructural carbohydrates (NSC). * Key results: Nothofagus and Pinus did not differ in survival, which, as a whole, was higher at ground level than at the 2-m height. The root-zone temperature for the growing season was 6.6°C. While biomass and NSC decreased significantly for Nothofagus at the 2-m height compared with ground level (C limitation), these trends were not significant for Pinus. * Conclusions: The treeline for Nothofagus pumilio is located at an isotherm that fully matches global patterns; however, its physiological responses to low temperatures differed from those of other treeline species. Support for C limitation in N. pumilio but not in P. contorta indicates that the physiological mechanism explaining their survival and growth at treeline may be taxon-dependent. [ABSTRACT FROM AUTHOR]

403I.403II.405III.410IV.411V.412VI.412VII.413VIII.413413References414 Summary: Models describing the biotic drivers that create and maintain biological diversity within trophic levels have focused primarily on negative interactions (i.e. competition), leaving marginal room for positive interactions (i.e. facilitation). We show facilitation to be a ubiquitous driver of biodiversity by first noting that all species use resources and thus change the local biotic or abiotic conditions, altering the available multidimensional niches. This can cause a shift in local species composition, which can cause an increase in beta, and sometimes alpha, diversity. We show that these increases are ubiquitous across ecosystems. These positive effects on diversity occur via a broad host of disparate direct and indirect mechanisms. We identify and unify several of these facilitative mechanisms and discuss why it has been easy to underappreciate the importance of facilitation. We show that net positive effects have a long history of being considered ecologically or evolutionarily unstable, and we present recent evidence of its potential stability. Facilitation goes well beyond the common case of stress amelioration and it probably gains importance as community complexity increases. While biodiversity is, in part, created by species exploiting many niches, many niches are available to exploit only because species create them. [ABSTRACT FROM AUTHOR]

Recent functional trait studies have shown that trait differences may favour certain species (environmental filtering) while simultaneously preventing competitive exclusion (niche partitioning). However, phenomenological trait-dispersion analyses do not identify the mechanisms that generate niche partitioning, preventing trait-based prediction of future changes in biodiversity. We argue that such predictions require linking functional traits with recognised coexistence mechanisms involving spatial or temporal environmental heterogeneity, resource partitioning and natural enemies. We first demonstrate the limitations of phenomenological approaches using simulations, and then (1) propose trait-based tests of coexistence, (2) generate hypotheses about which plant functional traits are likely to interact with particular mechanisms and (3) review the literature for evidence for these hypotheses. Theory and data suggest that all four classes of coexistence mechanisms could act on functional trait variation, but some mechanisms will be stronger and more widespread than others. The highest priority for future research is studies of interactions between environmental heterogeneity and trait variation that measure environmental variables at within-community scales and quantify species' responses to the environment in the absence of competition. Evidence that similar trait-based coexistence mechanisms operate in many ecosystems would simplify biodiversity forecasting and represent a rare victory for generality over contingency in community ecology. [ABSTRACT FROM AUTHOR]

Background and Aims The most plausible explanation for treeline formation so far is provided by the growth limitation hypothesis (GLH), which proposes that carbon sinks are more restricted by low temperatures than by carbon sources. Evidence supporting the GLH has been strong in evergreen, but less and weaker in deciduous treeline species. Here a test is made of the GLH in deciduous–evergreen mixed species forests across elevational gradients, with the hypothesis that deciduous treeline species show a different carbon storage trend from that shown by evergreen species across elevations. Methods Tree growth and concentrations of non-structural carbohydrates (NSCs) in foliage, branch sapwood and stem sapwood tissues were measured at four elevations in six deciduous–evergreen treeline ecotones (including treeline) in the southern Andes of Chile (40°S, Nothofagus pumilio and Nothofagus betuloides; 46°S, Nothofagus pumilio and Pinus sylvestris) and in the Swiss Alps (46°N, Larix decidua and Pinus cembra). Key Results Tree growth (basal area increment) decreased with elevation for all species. Regardless of foliar habit, NSCs did not deplete across elevations, indicating no shortage of carbon storage in any of the investigated tissues. Rather, NSCs increased significantly with elevation in leaves (P < 0·001) and branch sapwood (P = 0·012) tissues. Deciduous species showed significantly higher NSCs than evergreens for all tissues; on average, the former had 11 % (leaves), 158 % (branch) and 103 % (sapwood) significantly (P < 0·001) higher NSCs than the latter. Finally, deciduous species had higher NSC (particularly starch) increases with elevation than evergreens for stem sapwood, but the opposite was true for leaves and branch sapwood. Conclusions Considering the observed decrease in tree growth and increase in NSCs with elevation, it is concluded that both deciduous and evergreen treeline species are sink limited when faced with decreasing temperatures. Despite the overall higher requirements of deciduous tree species for carbon storage, no indication was found of carbon limitation in deciduous species in the alpine treeline ecotone. [ABSTRACT FROM PUBLISHER]

Abstract: It has been predicted that subalpine forests will be negatively affected by global warming; however, direct responses to experimental warming have been scarcely examined in these systems. In this study we evaluated the effects of higher temperatures with and without water addition on the survival and growth of recently emerged (small) and large seedlings of the widely distributed species Nothofagus pumilio in subalpine forests of the southern Chilean Andes. We also examined the variations in seedling traits related to carbon balance in order to infer the causal mechanisms of survival and growth responses. Treatments of open top chambers (OTCs) were combined with watering in two locations with differing climates: Antillanca (40°S, humid) and Cerro Castillo (46°S, drier). OTCs increased mean and maximum air temperatures by 0.6°C and 2–3°C, respectively, and decreased soil humidity by 56% in Antillanca and 30% in Cerro Castillo, fulfilling methodological expectations and climate model predictions. After two complete growing seasons, the survival, relative growth rate (RGR), biomass, and a suite of seedling traits were measured and analyzed using mixed-effects models. Warming and warming in combination with watering significantly increased large seedling survival in Cerro Castillo. In Antillanca, warmer conditions increased the height, biomass, and leaf area of small seedlings, and the RGR of large seedlings. In this location, warming also caused lower leaf carbon isotopic composition in both age classes and higher specific leaf area in small seedlings, suggesting whole-plant carbon gain improvements; warming did not produce any drought effects. Our results indicate that warming produces positive effects on the seedling establishment of N. pumilio in the southern Andes, highlighting the importance of site-specific effects in response to climate change in widespread species. Site-specific effects can most likely explain the discrepancies between the results of this study and the predictions outlined by previous studies for these forests. [Copyright &y& Elsevier]

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]

Background and Aims Multi-stemmed trees (tree clusters) in Nothofagus pumilio, a dominant tree species in Patagonia, are very uncommon and are restricted to the edge of second-growth forests following human-provoked fires. No vegetative reproduction has been reported so far. The genetic structure of multi-stemmed trees of this species was investigated and it was hypothesized that genets within a cluster were more closely related than average in the population. Methods Fifteen clusters (composed of at least three purported stems) and 15 single trees were sampled at the edge of a second-growth forest and genotyped using two amplified fragment length polymorphism (AFLP) primer pairs. We obtained 119 polymorphic markers that allowed clonality to be determined, together with sibship structure and relatedness among samples. Key Results Clonality was detected in seven clusters but all clusters had at least two different genotypes. Full sibs were found exclusively within clusters and in all clusters. Within a cluster, stems that were not identified as full sibs were often half sibs. Relatedness values for the full sibs and half sibs were higher than the theoretical values of 0·5 and 0·25 but the relatedness between clusters was very low. Conclusions Tree clusters that are merged at the edge of the second-growth forest of N. pumilio are composed of stems of the same genotype and of other genotypes that are highly related (but not always). It is suggested that this peculiar genetic structure results from a combination of several causes, including selection for merging of related individuals. [ABSTRACT FROM AUTHOR]

Nonstructural carbohydrates (NSCs) mediate plant survival when the plant's carbon (C) balance is negative, suggesting that NSCs could predict plant survival under C stress. To examine this possibility, we exposed saplings of six temperate tree species to diverse levels of C stress created by the combination of two light conditions (full light availability and deep shade) and two defoliation levels (severe defoliation and nondefoliation). We then measured survival, biomass and total NSCs and soluble sugars (SSs) concentrations in different organs of both dead and live saplings.We estimated mean NSCs and SSs contents and concentrations per sapling and fitted logistic generalized mixed‐effects models to determine if NSCs and SSs predict survival. Using inverse prediction modelling, we also determined whether there is a common NSCs and SSs threshold across species at the time of sapling's death.Defoliation and shade reduced the mean sapling's NSCs and SSs contents, indicating C stress. Mean sapling NSCs and SSs contents and concentrations predicted survival and the robustness of the models improved with the inclusion of species. At death, saplings of the exotic deciduous tree species Acer pseudoplatanus exhibited significantly lower mean NSCs and SSs contents than saplings of the evergreen conifer species Podocarpus nubigenus and lower stem NSCs and SSs concentrations than the broadleaf evergreen species Drimys winteri.The energetic role that NSCs and SSs play in plants under C stress was evidenced by the capacity of these compounds to predict sapling survival under C stress. No common threshold of NSCs and SSs contents or concentrations for sapling survival amongst species was found, indicating that the level of these compounds may not be good proxies for interspecific comparisons of tolerance to C stress. Presumably, there are species‐specific limits for the mobilization and use of NSCs and SSs in metabolism.Our results anticipate that the inclusion of NSCs and SSs in modelling will improve predictions regarding tree responses to ongoing climate change. Nonetheless, a better understanding of the many roles that carbohydrates play in plant survival under C stress is required to scale predictions up to the community level. Read the free Plain Language Summary for this article on the Journal blog. [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]

Many studies at the regional scale have found either negative or hump-shaped relationships between productivity and diversity, and some theories propose that these occur because soil resource heterogeneity is either lower or less important in more productive environments. However, there have been few explicit tests of these theories in natural ecosystems. We evaluated the relationship between soil resource heterogeneity and plant richness within a well characterized system of 30 islands in northern Sweden across which soil fertility and productivity declines, and species richness increases, as a consequence of ecosystem retrogression. On each island we created a spatially explicit grid consisting of 49 sampling points in a 9.5 m quadrat, which we used to quantify spatial heterogeneity of five soil variables (NH4+-N, amino N, PO4−-P, microbial biomass, and decomposition), and plant community composition. Using a hierarchical Bayesian approach, we estimated mean semivariograms of each variable for each island size class to compare three components of spatial heterogeneity: total variability, spatial grain, and patchiness. This analysis showed that variability within islands was usually lowest on small islands, where species richness was highest and productivity lowest; however, NH4+-N and amino N had greater patchiness and spatial grain on small islands. We did not detect any significant across-island correlations between whole-plot plant species richness and either whole-plot standard deviation or coefficient of variation of any soil variable. Using partial Mantel tests, we found that mean correlation coefficients between within-plot plant community composition and the soil variables were never significant for any island size class, and did not differ between island size classes. Our findings do not provide any evidence that soil resource heterogeneity controls the productivity-diversity relationship in this system, and suggests other mechanisms are primarily responsible. [ABSTRACT FROM AUTHOR]

Background and Aims Trees universally decrease their growth with age. Most explanations for this trend so far support the hypothesis that carbon (C) gain becomes limited with age; though very few studies have directly assessed the relative reductions of C gain and C demand with tree age. It has also been suggested that drought enhances the effect of C gain limitation in trees. Here tests were carried out to determine whether C gain limitation is causing the growth decay with tree age, and whether drought accentuates its effect. Methods The balance between C gain and C demand across tree age and height ranges was estimated. For this, the concentration of non-structural carbohydrates (NSCs) in stems and roots of trees of different ages and heights was measured in the deciduous temperate species Nothofagus pumilio. An ontogenetic decrease in NSCs indicates support for C limitation. Furthermore, the importance of drought in altering the C balance with ontogeny was assessed by sampling the same species in Mediterranean and humid climate locations in the southern Andes of Chile. Wood density (WD) and stable carbon isotope ratios (δ13C) were also determined to examine drought constraints on C gain. Key Results At both locations, it was effectively found that tree growth ultimately decreased with tree age and height. It was found, however, that NSC concentrations did not decrease with tree age or height when WD was considered, suggesting that C limitation is not the ultimate mechanism causing the age/height-related declining tree growth. δ13C decreased with tree age/height at the Mediterranean site only; drought effect increased with tree age/height, but this pattern was not mirrored by the levels of NSCs. Conclusions The results indicate that concentrations of C storage in N. pumilio trees do not decrease with tree age or height, and that reduced C assimilation due to summer drought does not alter this pattern. [ABSTRACT FROM AUTHOR]

Facilitation (positive interactions) has emerged as a dominant ecological mechanism in many ecosystems. Its importance has recently been expanded to include intraspecific interactions, creating the potential for higher-level natural selection within species. Using multiple lines of evidence, we show that conspecific facilitation within the southern beech tree, Nothofagus pumilio, appears to overcome competition in two life phases. In a seedling experiment addressing stress and planting-density effects, we found that mortality was lowest (∼0%) where there was no stress and was indistinguishable across densities. Furthermore, in mature forests (45 years old), genetically variable, merged individuals had lower mortality (- 50%) than unmerged individuals in locations without identifiable stress. Thus, a full understanding of the occurrence of facilitation may require a more general model of resource improvements than the commonly cited stress gradient hypothesis. Additionally, the merged trees showed a density-dependent mortality pattern at the level of the group. These data demonstrate a potential mechanism (facilitation) driving natural selection at this higher level, via stem merging. These merged “superorganisms” would confirm theoretical predictions whereby facilitation acts as an ecological mechanism driving group selection. [ABSTRACT FROM AUTHOR]

Competition among conspecifics of the same cohort has been traditionally thought to be a main process driving population dynamics. In this classical view, however, the role of facilitation in stressful conditions has rarely been considered. Here, using a transplant experiment across a forest-prairie gradient, we test whether the stress gradient hypothesis (SGH) extends to individuals thought to be strongly competing. We transplanted 2-year-old seedlings of Nothofagus pumilio at two different densities (clusters of 10 and isolated) and at different distances from the forest edge (from 30 m inside the forest up to 50 m outside the forest in the prairie). We further stem-mapped all seedlings belonging to the clusters and computed a competition index (CI). After 3 years of growing, survival and increment growth in diameter and height were measured and analysed using mixed-effects models. We conducted a nearest-neighbour analysis using seedlings' CI and growth and computed model fit using the area under the curve (AUC) method. Seedlings planted in dense clusters had significantly higher survival than solitary seedlings at the stressful end of the gradient. This trend was reversed at the opposite end of the gradient, supporting the SGH at the intraspecific level. Pursuing this at the level of the individual, we found that higher CIs (more neighbours) in seedlings predicted higher probabilities of their survival (facilitation) in stressful conditions. Seedlings diameter and height increment growth were not affected by planting density and only diameter varied along the stress gradient; seedlings had higher diameter increments in growth outside the forest. Finally, when compared with conceptual models, our results mostly support predictions of a higher facilitation at intermediate position along the gradient. Synthesis. We showed that facilitation overrides competition among tree seedlings even at locations under moderate stress; the facilitation process occurs in resource-mediated interactions (niche overlapping). These results represent an important shift in our way to understand the density-dependent mortality process, and calls for a model reformulation including positive interactions even when competition is expected to be strongest (conspecifics of the same cohort). [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]

The focus of the trait-based approach to study community ecology has mostly been on trait comparisons at the interspecific level. Here we quantified intraspecific variation and covariation of leaf mass per area (LMA) and wood density (WD) in monospecific forests of the widespread tree species Nothofagus pumilio to determine its magnitude and whether it is related to environmental conditions and ontogeny. We also discuss probable mechanisms controlling the trait variation found. We collected leaf and stem woody tissues from 30-50 trees of different ages (ontogeny) from each of four populations at differing elevations (i.e. temperatures) and placed at each of three locations differing in soil moisture. The total variation in LMA (coefficient of variation (CV) = 21.14%) was twice that of WD (CV = 10.52%). The total variation in traits was never less than 23% when compared with interspecific studies. Differences in elevation (temperature) for the most part explained variation in LMA, while differences in soil moisture and ontogeny explained the variation in WD. Traits covaried similarly in the altitudinal gradient only. Functional traits of N. pumilio exhibited nonnegligible variation; LMA varied for the most part with temperature, while WD mostly varied with moisture and ontogeny. We demonstrate that environmental variation can cause important trait variation without species turnover. [ABSTRACT FROM AUTHOR]

• Premise of the Study: Negative density-dependent processes have been thought to be the primary cause of shifting spatial patterns of tree populations through time. The existence of adult tree clusters might challenge this classical prediction. Here, we document the prevalence of merged stems (clustering of mature trees leading to stem fusion) in second-growth forests of Nothofagus pumilio and hypothesize that it is nonrandom but predictable in space. • Methods: We stem-mapped nine sites in second-growth edge and interior forests of fire origin and in mature forests of N. pumilio (>3500 trees) in central Patagonia, Chile. The spatial structure of stand-level and individual-level features was estimated with spatial analyses (pair-correlation function and nearest-neighbor distances). • Key Results: Multistemmed trees were merged clusters of separate individuals. Merged trees were predominantly found at the edge of the second-growth forests. We found strong clustering (≤5 m) at forest edge sites and none at interior sites. Nearest-neighbor distance distributions were unimodal for unmerged trees and monotonically decreasing for merged trees; interstem distances were much smaller at the edge sites than at the interior sites. • Conclusions: The occurrence of merged trees at the forest edge, and the resulting high spatial aggregation of stems, is consistent with the hypothesis that establishment was probably aggregated. The spatial pattern found at the forest edge changes the standard spatial pattern sequence through time in temperate forests, altering traditional forest-stand-dynamics models. [ABSTRACT FROM AUTHOR]

Questions: Fire appears to affect both replacement patterns and coexistence of Araucaria araucana- Nothofagus pumilio forests in the Andean Araucarian region of south-central Chile. A quantitative assessment of coexistence in the absence of recent fires, however, is lacking. In this study, we considered the life-history attributes, time of recruitment and spatial pattern of individuals of both tree species to address the following questions. How regular has recruitment of both species been in time? Is there any temporal niche differentiation? Are the two species positively or negatively associated in space and, if so, at what scale and for what age and size classes? Is there any spatial niche differentiation? Location: Andean Araucarian region of Chile, Villarrica National Park (39°35′S, 71°31′W; 1300 m a.s.l.). Methods: We stem-mapped and cored a total of 1073 trees in a 1 -ha plot in a late-successional post-fire stand to examine spatiotemporal patterns of establishment. We used semivariogram modelling and the pair-correlation function to distinguish between regeneration modes and describe species interactions. Results: The two species differ in their regeneration mode: whereas A. araucana appeared to recruit more continuously in time and space, episodic pulses of establishment were dominant for N. pumilio. At small scales, younger age -class stems of A. araucana were randomly distributed, while older age -class stems were aggregated. This was in contrast to common patterns for temperate tree species, including N. pumilio, following processes of self-thinning. Younger age classes of A. araucana were distributed independently of older trees of both species, but younger age classes of N. pumilio had a negative association with older conspecifics at scales similar to crown diameter. Conclusions: In the absence of recent fires, it is likely that A. araucana would dominate the stand alone, given its greater shade tolerance, greater longevity and continuous recruitment. However, while canopy closure is still incomplete, the shade-intolerant N. pumilio will be able to recruit in those open areas after seed masting and will coexist with A. araucana. [ABSTRACT FROM AUTHOR]

We developed multiple a priori hypotheses to link the observed spatial patterns with colonisation processes in the high alpine cushion plant, Azorella madreporica. We conducted this study in the Molina River basin (33°20′ S, 70°16′ W, 3600 m a.s.l.), in the Andes of central Chile, approximately 50 km east of Santiago. We mapped and measured size (as a surrogate for age) of individual cushions in two populations and used a standard spatial analytical tool (semivariograms) to test our alternative a priori hypotheses related to colonisation mode of the cushion species. In both populations, the size distribution of A. madreporica reflected a negative exponential or inverse-J pattern, typical of uneven-aged populations, where most of the cushions belonged to relatively smaller size classes, in effect, a regular success in the establishment of seedlings, where all size classes of cushions were represented in the population. The results were site-specific, where best-fit semivariograms for spatial cushion's size distribution suggested a gradual colonisation in one population and an episodic colonisation in the other population. Microsite distribution proved to be homogeneous at both sites. Thus, the study of the spatial explicit size-age population distribution of an alpine species provides valuable information about the frequency, magnitude and site variation of the reproductive pulses in these harsh environments. [ABSTRACT FROM AUTHOR]