Growth delay by winter precipitation could hinder Juniperus sabina persistence under increasing summer drought

Plants in Mediterranean mountains are particularly vulnerable to climatic change. In these environments, low temperature is combined with water shortage during summer, and as a result, the positive effect of global warming theoretically expanding the growing season length may be counterbalanced by rising drought stress. These circumstances may be exacerbated in the rear edge of species distribution, where warmer conditions occur. Here, we examined the climate-growth relationships of Juniperus sabina, a major prostrate shrub above the treeline in Mediterranean mountains, to investigate climate sensitivity and long-term signals stability in four rear-edge populations from southern Spain. We demonstrate that, over recent decades, local climatic conditions have modulated the response of J. sabina secondary growth to the ongoing climate change. We observed a negative effect of winter-spring moisture on secondary growth that suggests a limitation for earlier growth activity at higher elevation, potentially hindering the ability of J. sabina to compensate forthcoming increases in summer drought. At the driest site, we also detected a positive effect of October precipitation, suggesting a second growth pulse by early autumn. Our results provide an example of how local climatic conditions may limit plastic responses of secondary growth to climatic variability. The implications of growth limitation in J. sabina exceed the species-scale level, since these prostrate shrubs play a critical role as nurse plants and local biodiversity foci in Mediterranean mountains.     

Recent climate hiatus revealed dual control by temperature and drought on the stem growth of Mediterranean Quercus ilex

A better understanding of stem growth phenology and its climate drivers would improve projections of the impact of climate change on forest productivity. Under a Mediterranean climate, tree growth is primarily limited by soil water availability during summer, but cold temperatures in winter also prevent tree growth in evergreen forests. In the widespread Mediterranean evergreen tree species Quercus ilex, the duration of stem growth has been shown to predict annual stem increment, and to be limited by winter temperatures on the one hand, and by the summer drought onset on the other hand. We tested how these climatic controls of Q. ilex growth varied with recent climate change by correlating a 40‐year tree ring record and a 30‐year annual diameter inventory against winter temperature, spring precipitation, and simulated growth duration. Our results showed that growth duration was the best predictor of annual tree growth. We predicted that recent climate changes have resulted in earlier growth onset (?10 days) due to winter warming and earlier growth cessation (?26 days) due to earlier drought onset. These climatic trends partly offset one another, as we observed no significant trend of change in tree growth between 1968 and 2008. A moving‐window correlation analysis revealed that in the past, Q. ilex growth was only correlated with water availability, but that since the 2000s, growth suddenly became correlated with winter temperature in addition to spring drought. This change in the climate–growth correlations matches the start of the recent atmospheric warming pause also known as the ‘climate hiatus’. The duration of growth of Q. ilex is thus shortened because winter warming has stopped compensating for increasing drought in the last decade. Decoupled trends in precipitation and temperature, a neglected aspect of climate change, might reduce forest productivity through phenological constraints and have more consequences than climate warming alone.     

Diverging shrub and tree growth from the Polar to the Mediterranean biomes across the European continent

Climate warming is expected to enhance productivity and growth of woody plants, particularly in temperature‐limited environments at the northernmost or uppermost limits of their distribution. However, this warming is spatially uneven and temporally variable, and the rise in temperatures differently affects biomes and growth forms. Here, applying a dendroecological approach with generalized additive mixed models, we analysed how the growth of shrubby junipers and coexisting trees (larch and pine species) responds to rising temperatures along a 5000‐km latitudinal range including sites from the Polar, Alpine to the Mediterranean biomes. We hypothesize that, being more coupled to ground microclimate, junipers will be less influenced by atmospheric conditions and will less respond to the post‐1950 climate warming than coexisting standing trees. Unexpectedly, shrub and tree growth forms revealed divergent growth trends in all the three biomes, with juniper performing better than trees at Mediterranean than at Polar and Alpine sites. The post‐1980s decline of tree growth in Mediterranean sites might be induced by drought stress amplified by climate warming and did not affect junipers. We conclude that different but coexisting long‐living growth forms can respond differently to the same climate factor and that, even in temperature‐limited area, other drivers like the duration of snow cover might locally play a fundamental role on woody plants growth across Europe.     

Tree species from contrasting hydrological niches show divergent growth and water-use efficiency

Tree species occupy different hydrological niches and climate warming may affect tree performance in those niches through increased drought stress. However, the effects of climate warming on growth, carbon and water fluxes would differ depending on species’ hydrological niche. Species from wet sites should show a lower growth dependence on precipitation and also lower intrinsic water-use efficiency (WUEi), as compared with species from dry sites which should improve more the WUEi. We test these ideas by comparing radial-growth rates (basal-area increment), climate- and drought-growth associations and WUEi of hackberry (Celtis australis) vs. Aleppo pine (Pinus halepensis) and maritime pine (Pinus pinaster) in two Mediterranean sites located in Spain. Species are subjected to similar regional climate conditions in each site but occupy contrasting local hydrological niches (hackberry in wet sites and pines in dry sites). Climate is warming in both study sites and drought-avoiding pines are responding by showing higher growth rates and improved WUEi. We also found a similar growth dependency on winter-spring precipitation and summer drought of all species and sites and comparable WUEi values and trends, excepting in hackberry from southern Spain which grew more, and showed a higher growth resistance to drought and lower and more stable WUEi values. Variables inferred from tree rings as growth rates and WUEi allow characterizing the hydrological niche of tree species, which may be contingent on site conditions and climate warming.     

Seasonal precipitation and continentality drive bimodal growth in Mediterranean forests

Tree phenology is sensitive to climate warming and changes in seasonal precipitation. Long xylogenesis records are scarce, thus limiting our ability to analyse how radial growth responds to climate variability. Alternatively, process-based growth models can be used to simulate intra-annual growth dynamics and to better understand why growth bimodality varies along temperature and precipitation gradients. We used the Vaganov-Shashkin (VS) growth model to analyse the main climatic drivers of growth bimodality in eight trees and shrubs conifers (four pines and four junipers) across Spain. We selected eleven sites with different continentality degree and spring/autumn precipitation ratios since we expected to find pronounced bimodal growth in less continental sites with spring and autumn precipitation peaks. The VS model successfully simulated annual growth rates at all sites as a function of daily temperature and soil moisture data. Bimodal growth patterns clustered into less continental sites showing low spring/autumn precipitation ratios. This finding agrees with observed climate-growth associations showing that growth was enhanced by wet-cool winter-to-spring conditions, but also by wet autumn conditions in the most bimodal sites. We observed a stronger growth bimodality in pines compared to junipers. We discuss the spatial variability of climate drivers in bimodality growth pattern and how increasing continentality and shifts in seasonal precipitation could affect growth patterns. Bimodality could be an advantageous response to overcome summer drought in Mediterranean forests. The ability of some species to reactivate growth during autumn might determine their capacity to withstand increasing summer aridity.     

Reproductive output in Mediterranean shrubs under climate change experimentally induced by drought and warming

The effects of climate change on plant reproductive performance affects the sequence of different plant reproductive stages from flowering to seed production and viability, as well as the network of relationships between them. These effects are expected to respond to different components of climate change, such as temperature and water availability, and may be sensitive to differences in species phenology.We used long-term experimental drought and warming treatments to study the effect of climate change on flower production, fruit and seed-set, seed size and seed germination rate (proportion of germinating seeds) in three Mediterranean shrubs coexisting in a coastal shrubland.Larger plants produced significantly more flowers in all three species, and higher fruit-set in Dorycnium pentaphyllum. Flower production was reduced in drought and warming treatments in the spring-flowering species D. pentaphyllum and Helianthemum syriacum, but not in the autumn–winter species Erica multiflora, which increased flowering in the warming treatment. However, the drought treatment eventually resulted in a decreased seed-set in E. multiflora. Structural equation modelling revealed strong correlations between the sequential reproductive stages. Specifically, flower density in inflorescences determined seed-set in H. syriacum, and seed size and germination rate in E. multiflora. Nevertheless, the relevance of relationships between reproductive traits changed between climatic treatments: in D. pentaphyllum a direct relationship between plant size and seed size only arised in the drought treatment, while in H. syriacum climate treatments resulted in a stronger relationship between the number of flowers and seed-set.This experimental study shows the ability of changing climatic variables to determine the reproductive sequential process of woody species. We show that several parameters of the reproductive performance of some Mediterranean species are affected by drought and warming treatments simulating climate change, highlighting the importance of changes in both water availability and temperature, and the sequential relationship between reproductive stages. Phenological patterns also contribute to species’ differential responses to climatic change, due to the relationship of these patterns with resource availability, environmental conditions and plant–pollinator interactions.     

Variation in woody plant mortality and dieback from severe drought among soils, plant groups, and species within a northern Arizona ecotone

Vegetation change from drought-induced mortality can alter ecosystem community structure, biodiversity, and services. Although drought-induced mortality of woody plants has increased globally with recent warming, influences of soil type, tree and shrub groups, and species are poorly understood. Following the severe 2002 drought in northern Arizona, we surveyed woody plant mortality and canopy dieback of live trees and shrubs at the forest–woodland ecotone on soils derived from three soil parent materials (cinder, flow basalt, sedimentary) that differed in texture and rockiness. Our first of three major findings was that soil parent material had little effect on mortality of both trees and shrubs, yet canopy dieback of trees was influenced by parent material; dieback was highest on the cinder for pinyon pine (Pinus edulis) and one-seed juniper (Juniperus monosperma). Ponderosa pine (Pinus ponderosa) dieback was not sensitive to parent material. Second, shrubs had similar mortality, but greater canopy dieback, than trees. Third, pinyon and ponderosa pines had greater mortality than juniper, yet juniper had greater dieback, reflecting different hydraulic characteristics among these tree species. Our results show that impacts of severe drought on woody plants differed among tree species and tree and shrub groups, and such impacts were widespread over different soils in the southwestern U.S. Increasing frequency of severe drought with climate warming will likely cause similar mortality to trees and shrubs over major soil types at the forest–woodland ecotone in this region, but due to greater mortality of other tree species, tree cover will shift from a mixture of species to dominance by junipers and shrubs. Surviving junipers and shrubs will also likely have diminished leaf area due to canopy dieback.     

Growth phenological variations in the narrow-leaved ash (Fraxinus angustifolia) over the Mediterranean region: A simulation study

Tree species inhabiting riparian forests under Mediterranean climate have evolved to face summer water shortage but may fail to cope with future increases in drought severity. Thus, understanding tree growth phenological variations in response to environmental conditions is necessary to assess the impact of seasonal drought in riparian forests. In this study, we investigated the response of stem radial growth to climate in the narrow-leaved ash (Fraxinus angustifolia) over its distribution in southern Europe. We simulated intra- and inter-annual growth patterns using the Vaganov-Shashkin (VS) model considering five sites subjected to summer drought but showing different climate conditions. The growth pattern in this species varied from unimodal in cool-wet sites to facultative bimodal in warm-dry sites. Bimodal patterns were characterized by two growth peaks coinciding with favorable climate conditions in spring and autumn. The spring growth peak occurs earlier (May) in warm-dry sites than in wet-cool sites (June–July). The variation in the season growth length and growth timing suggests different strategies adopted by this species to cope with summer drought. The VS model revealed different growth patterns across which would be relevant in predicting the response of this and other riparian tree species to climate warming and aridification. Differences in the length of the growing season, timings of growth peaks and the shift from unimodal to bimodal growth patterns should be considered when assessing growth adjustments to future climate scenarios.     

Growth response of Abies georgei to climate increases with elevation in the central Hengduan Mountains,southwestern China

Climatic harshness is expected to increase at higher elevations; however, elevational trends of tree radial growth response of high-elevation forests to climate change need to be investigated at different locations because of existing local variability in site-specific climatic conditions. We developed tree-ring width chronologies of Yunnan fir (Abies georgei) along elevation gradients at two sites in the central Hengduan Mountains (HM). High-elevation forests of A. georgei showed growth synchronicity and common growth signals along elevation gradients, indicating a common climatic forcing, although tree radial growth rates decreased with increasing elevation. Radial growth of Yunnan fir showed positive correlations with summer temperatures and February precipitation and moisture availability, but were negatively correlated with spring temperatures. The strongest positive relationship indicated summer (July) mean and minimum temperatures are the most important growth determining climatic factors for tree radial growth in the cold environment of HM, and this relationship revealed a clear elevational trend with stronger correlations at higher altitudes. In contrast, tree radial growth was negatively correlated with June precipitation and moisture availability. The whole study period 1954–2015 was split in two sub-periods of equal length. Comparing the early sub-period (1954–1984) to the later sub-period (1985–2015), tree growth response to the summer temperatures strongly increased, while it became weaker to June precipitation and moisture availability. High-elevation Yunnan fir forests in the HM currently benefit from elevated growing season temperatures under humid summer conditions. However, increasing temperatures may induce drought stress on tree radial growth if the observed decreasing trend in humidity and precipitation continues.     

Age-dependency,climate, and environmental controls of recent tree growth trends at subarctic and alpine treelines

Spatial and temporal variability in growth and climate response of trees at and near treeline was investigated in the western Mackenzie Mountains, Northwest Territories, and the Hudson Bay Lowlands of northern Manitoba. Residual ring width chronologies were constructed using cores extracted from 108 trees in the mountains and 170 from the lowlands, and compared to historical climate data. Growth of most trees exhibited significant correlations with summer and autumn temperatures, and the growth–climate relationship did not differ noticeably between trees at and distal to treeline. Most mountain trees had significant positive growth trends from 1851 to 2006 that corresponded with warming over the same period, while growth trends varied among sites and species in the lowlands. Regionally, growth of all species responded positively to warming during the 20th century with the exception of lowland Picea mariana, which exhibited little response. Growth response for most trees was age-dependent, with trees established after 1920 demonstrating improved growth and sensitivity to temperature than older individuals, and growth of most species since the 1990s was greater than any time during the last 250 years, particularly for lowland Larix laricina. This study suggests that site factors and tree age can be more important drivers of local-scale growth trends than regional climate at arctic treelines where temperature is often assumed to be the main constraint on tree growth.     

Plastic bimodal growth in a Mediterranean mixed-forest of Quercus ilex and Pinus halepensis

Mediterranean tree species have evolved to face seasonal water shortages, but may fail to cope with future increases in drought frequency and intensity. We investigated stem radial increment dynamics in two typical Mediterranean tree species, Aleppo pine (Pinus halepensis), a drought-avoiding species, and holm oak (Quercus ilex), a drought-tolerant species, in a mixed forest and on contrasting slope aspects (south- and north-facing). Intra- and inter-annual growth patterns were modelled using the VS-Lite2 model for each tree species and slope-aspect. Both species showed a bimodal growth pattern, with peaks coinciding with favourable conditions in spring and autumn. A bimodal growth pattern is always observed in P. halepensis, while in Q. ilex is facultative, which suggests different strategies adopted by these species to cope with summer drought. More specifically, trees on south-facing slope showed a more evident bimodal pattern and more intra-annual density fluctuations. In recent decades, the intensity of both growth peaks has diminished and drifted away due to the increased summer drought. The VS-Lite2 model reveals a niche partitioning between both species. Differences in growing season’s length and timings of growth peaks in both species are relevant for their coexistence and should be considered for estimating mixed-forest responses under climate change scenarios.     

Differential impact of hotter drought on seedling performance of five ecologically distinct pine species

Increasing temperature and drought intensity is inducing the phenomenon of the so-called “hotter drought”, which is expected to increase in frequency over the coming decades across many areas of the globe, and is expected to have major implications for forest systems. Consequences of hotter drought could be especially relevant for closely related species overlapping their distributions, since differences in response can translate into range shifts. We assessed the effect of future climatic conditions on the performance of five ecologically distinct pine species common in Europe: Pinus halepensis, P. pinaster, P. nigra, P. sylvestris and P. uncinata. We hypothesised that Mediterranean species inhabiting dry, low-elevation sites will be less affected by the expected warming and drought increase than species inhabiting cold-wet sites. We performed a controlled conditions experiment simulating current and projected temperature and precipitation, and analysed seedling responses in terms of survival, growth, biomass allocation, maximum photochemical efficiency (F _v/F _m) and plant water potential (Ψ). Either an increase in temperature or a reduction in water input alone reduced seedling performance, but the highest impact occurred when these two factors acted in combination. Warming and water limitation reduced Ψ, whereas warming alone reduced biomass allocation to roots and F _v/F _m. However, species responded differentially to warmer and drier conditions, with lowland Mediterranean pines (P. halepensis and P. pinaster) showing higher survival and performance than mountain species. Interspecific differences in response to warmer, drier conditions could contribute to changes in the relative dominance of these pine species in Mediterranean regions where they co-occur and a hotter, drier climate is anticipated.     

Differential climate–growth relationships in Abies alba Mill. and Fagus sylvatica L. in Mediterranean mountain forests

Pointer year analysis, simple correlations, and response functions were combined in a dendroecological study to evaluate climate–growth relationships over the last century in two Abies alba Mill. and Fagus sylvatica L. mixed stands in Southern Italy mountainous areas. Analyses revealed species-specific attributes at the two study sites, i.e. Molise and Basilicata. Growth divergence between the two species emerged based on three primary climatic drivers, including drought stress and spring warmer temperatures during the current growing season for F. sylvatica, and water availability in the previous growing season for A. alba. However, despite the microclimatic differences between the two study sites, F. sylvatica showed similar climate–growth patterns, while differences were indicated for A. alba, due to its minor susceptibility to drought stress during the current growing season at the Basilicata site. Indeed, at the southernmost geographic limits of A. alba drought avoidance mechanisms were confirmed, consistent with traits considered diagnostic for the species in the Mediterranean region.     

Effects of vegetation canopy and climate on seedling establishment in Mediterranean shrubland

Abstract. Question: Does the influence of plant canopy on seedling establishment interact with climate conditions, and particularly, do intensified drought conditions, enhance a positive effect of the vegetation canopy on seedlings in Mediterranean‐type ecosystems. Location: Mediterranean shrubland near Barcelona, Spain at 210 m a.s.l. Methods: Over the course of four years we recorded seedling emergence and survival in open areas and below vegetation under control, drier and warmer experimental climatic conditions. Results: Seedling emergence is more sensitive to climate conditions than later stages of growth. When considering the whole set of species, the total number of established seedlings at the end of the experiment was lower in the drought and warming stands than in control ones, and vegetation canopy increased the number of these seedlings in the drought stands. Drought reduced seedling emergence but not warming, while the interaction between climate treatments and vegetation canopy was not significant. Seedling survival was lower in the warming treatment than in the control. Under drought conditions, vegetation canopy increased seedling emergence of the dominant Globularia alypum. In control stands, vegetation canopy reduced their survival. Vegetation canopy increased the survival of the dominant Erica multiflora in warming stands, and it reduced the survival of G alypum in drought stands. No significant effects of drought and warming were observed in the seed rain of these two species. Conclusions: The balance of the facilitation‐competition interactions between vegetation canopy and seedling establishment in Mediterranean‐type ecosystems determined by water availability, and drought conditions enhance the positive effect of vegetation canopy. This interaction is species‐specific and shows important between‐year variability.     

Growth of 19 conifer species is highly sensitive to winter warming,spring frost and summer drought

Background and AimsConifers are key components of many temperate and boreal forests and are important for forestry, but species differences in stem growth responses to climate are still poorly understood and may hinder effective management of these forests in a warmer and drier future.MethodsWe studied 19 Northern Hemisphere conifer species planted in a 50-year-old common garden experiment in the Netherlands to (1) assess the effect of temporal dynamics in climate on stem growth, (2) test for a possible positive relationship between the growth potential and climatic growth sensitivity across species, and (3) evaluate the extent to which stem growth is controlled by phylogeny.Key resultsEighty-nine per cent of the species showed a significant reduction in stem growth to summer drought, 37 % responded negatively to spring frost and 32 % responded positively to higher winter temperatures. Species differed largely in their growth sensitivity to climatic variation and showed, for example, a four-fold difference in growth reduction to summer drought. Remarkably, we did not find a positive relationship between productivity and climatic sensitivity, but instead observed that some species combined a low growth sensitivity to summer drought with high growth potential. Both growth sensitivity to climate and growth potential were partly phylogenetically controlled.ConclusionsA warmer and drier future climate is likely to reduce the productivity of most conifer species. We did not find a relationship between growth potential and growth sensitivity to climate; instead, some species combined high growth potential with low sensitivity to summer drought. This may help forest managers to select productive species that are able to cope with a warmer and drier future.     

Forest composition in Mediterranean mountains is projected to shift along the entire elevational gradient under climate change

Aim Species distribution models have been used frequently to assess the effects of climate change on mountain biodiversity. However, the value and accuracy of these assessments have been hampered by the use of low‐resolution data for species distributions and climatic conditions. Herein we assess potential changes in the distribution and community composition of tree species in two mountainous regions of Spain under specific scenarios of climate change using data with a high spatial resolution. We also describe potential changes in species distributions and tree communities along the entire elevational gradient. Location Two mountain ranges in southern Europe: the Central Mountain Range (central west of the Iberian Peninsula), and the Iberian Mountain Range (central east). Methods We modelled current and future distributions of 15 tree species (Eurosiberian, sub‐Mediterranean and Mediterranean species) as functions of climate, lithology and availability of soil water using generalized linear models (logistic regression) and machine learning models (gradient boosting). Using multivariate ordination of a matrix of presence/absence of tree species obtained under two Intergovernmental Panel on Climate Change (IPCC) scenarios (A2 and B2) for two different periods in the future (2041–70 and 2071–2100), we assessed the predicted changes in the composition of tree communities. Results The models predicted an upward migration of communities of Mediterranean trees to higher elevations and an associated decline in communities of temperate or cold‐adapted trees during the 21st century. It was predicted that 80–99% of the area that shows a climate suitable for cold–wet‐optimum Eurosiberian coniferous and broad‐leaved species will be lost. The largest overall changes were predicted for Mediterranean species found currently at low elevations, such as Pinus halepensis, Pinus pinaster, Quercus ilex ssp. ballota and Juniperus oxycedrus, with sharp increases in their range of 350%. Main conclusions It is likely that areas with climatic conditions suitable for cold‐adapted species will decrease significantly under climate warming. Large changes in species ranges and forest communities might occur, not only at high elevations within Mediterranean mountains but also along the entire elevational gradient throughout this region, particularly at low and mid‐elevations. Mediterranean mountains might lose their key role as refugia for cold‐adapted species and thus an important part of their genetic heritage.     

A multi-scale dendroclimatological analysis of four common species in the southern Canadian boreal forest

A comprehensive assessment of the tree growth/climate relationship was undertaken to better understand the potential impacts of climate change on the growth dynamics of four widespread and common boreal tree species, namely jack pine (Pinus banksiana), black spruce (Picea mariana), eastern larch (Larix laricina), and trembling aspen (Populus tremuloides), located at the southern limits of the Canadian boreal forest. Over intra-annual time scales, results show that precipitation is likely the main driver of stem radius change (∆R), with jack pine radius exhibiting the most consistent positive relationship. Precipitation had a stronger relationship with stem radius variation in black spruce and eastern larch during periods when volumetric water content (VWC) in the root zone was below average, pointing to the likelihood that certain species rely more heavily on available moisture in the uppermost layers of the soil column to replenish stem water, especially during extended dry periods. Warm air temperatures had an immediate negative impact on stem water content due to transpiration. This was most marked during periods of reduced moisture availability in the root zone, when trees are more susceptible to net water volume loss. During periods when moisture was not limiting, a positive relationship between lagged air temperature and ∆R was detected. Warm air temperatures may therefore play an important role in stimulating radial growth when moisture requirements are met. At annual temporal resolution, the growth/climate relationship changed over the lifetime of our study species. Over the last several decades, the relationship between precipitation and annual radial tree growth has weakened, while positive relationships between spring and summer air temperature and annual radial tree growth have emerged, likely signaling a decrease in moisture limitations, and a positive response to spring warming. Our findings reveal that boreal forest tree species may benefit from spring and summer warming over the near term, providing there is sufficient moisture to support growth. Over the long term, rates of evapotranspiration are expected to overshadow gains in moisture related to an increase in precipitation. Under these circumstances, we are likely to see reduced growth rates and an increasingly negative response of boreal tree species growth to warm air temperatures.     

Species-specific growth-climate responses of Dahurian larch (Larix gmelinii) and Mongolian pine (Pinus sylvestris var. mongolica) in the Greater Khingan Range,northeast China

Responses of tree growth to climate are usually spatially heterogeneous. Besides regionally varying external environments, species specificity is a crucial factor in determining said spatial heterogeneity. A better understanding of this species specificity would improve our estimations of the warming effects on forests. In this study, we selected two widely-distributed boreal conifers, Dahurian larch (Larix gmelinii) and Mongolian pine (Pinus sylvestris var. mongolica), to compare their growth-climate responses, including long-term growth-climate correlations and short-term growth resilience to drought. We sampled 160 trees and 481 tree-ring cores from the two species in two pure and two mixed forests, located in the Greater Khingan Range, northeast China. We found that Dahurian larch was generally positively correlated with spring temperature and negatively correlated with summer temperature. In contrast, Mongolian pine was more sensitive to summer moisture. Our results suggest that the main climatic limitations were low spring temperatures for Dahurian larch and summer moisture deficits for Mongolian pine. Dahurian larch represented higher growth resistance to drought, while Mongolia pine represented higher recovery. Based on this, we inferred that Dahurian larch was more vulnerable to extreme droughts, while Mongolian pine was more vulnerable to frequent droughts. We also demonstrated the effects of forest type on growth-climate responses. The negative effects of summer temperatures on Mongolian pine seemed to be more significant in mixed forests. As warming continued, Mongolian pine in this area would suffer severer moisture deficits, especially when coexisting with Dahurian larch. Our results suggest that Dahurian larch gained an advantage in the competition with Mongolian pine during high moisture stress. Driven by the warming trends, the species specificity in growth response would ultimately promote the separation of the two species in distribution. This study will help improve our estimations of the warming effects on forests and develop more species-targeted forest management practices.     

Abies spectabilis shows stable growth relations to temperature,but changing response to moisture conditions along an elevation gradient in the central Himalaya

A better understanding of growth-climate responses of high-elevation tree species across their distribution range is essential to devise an appropriate forest management and conservation strategies against adverse impacts of climate change. The present study evaluates how radial growth of Himalayan fir (Abies spectabilis D. Don) and its relation to climate varies with elevation in the Manaslu Mountain range in the central Himalaya. We developed tree-ring width chronologies of Himalayan fir from three elevational belts at the species’upper distribution limit (3750−3900 m), in the middle range (3500−3600 m), and at the lower distribution limit (3200−3300 m), and analyzed their associations with climatic factors. Tree growth of Himalayan fir varied synchronously across elevational belts, with recent growth increases observed at all elevations. Across the elevation gradient, radial growth correlated positively (negatively) with temperature (precipitation and standardized precipitation-evapotranspiration index, SPEI-03) during the summer (July to September) season. However, the importance of summer (July to September) temperatures on radial growth decreased with elevation, whereas correlations with winter (previous November to current January) temperatures increased. Correlations with spring precipitation and SPEI-03 changed from positive to negative from low to high elevations. Moving correlation analysis revealed a persistent response of tree growth to May and August temperatures. However, growth response to spring moisture availability has strongly increased in recent decades, indicating that intensified spring drought may reduce growth rates of Himalayan fir at lower elevations. Under sufficient moisture conditions, increasing summer temperature might be beneficial for fir trees growing at all elevations, while trees growing at the upper treeline will take additional benefit from winter warming.     

Assessing forest vulnerability to climate warming using a process‐based model of tree growth: bad prospects for rear‐edges

Growth models can be used to assess forest vulnerability to climate warming. If global warming amplifies water deficit in drought‐prone areas, tree populations located at the driest and southernmost distribution limits (rear‐edges) should be particularly threatened. Here, we address these statements by analyzing and projecting growth responses to climate of three major tree species (silver fir, Abies alba; Scots pine, Pinus sylvestris; and mountain pine, Pinus uncinata) in mountainous areas of NE Spain. This region is subjected to Mediterranean continental conditions, it encompasses wide climatic, topographic and environmental gradients, and, more importantly, it includes rear‐edges of the continuous distributions of these tree species. We used tree‐ring width data from a network of 110 forests in combination with the process‐based Vaganov–Shashkin‐Lite growth model and climate–growth analyses to forecast changes in tree growth during the 21st century. Climatic projections were based on four ensembles CO₂ emission scenarios. Warm and dry conditions during the growing season constrain silver fir and Scots pine growth, particularly at the species rear‐edge. By contrast, growth of high‐elevation mountain pine forests is enhanced by climate warming. The emission scenario (RCP 8.5) corresponding to the most pronounced warming (+1.4 to 4.8 °C) forecasted mean growth reductions of ?10.7% and ?16.4% in silver fir and Scots pine, respectively, after 2050. This indicates that rising temperatures could amplify drought stress and thus constrain the growth of silver fir and Scots pine rear‐edge populations growing at xeric sites. Contrastingly, mountain pine growth is expected to increase by +12.5% due to a longer and warmer growing season. The projections of growth reduction in silver fir and Scots pine portend dieback and a contraction of their species distribution areas through potential local extinctions of the most vulnerable driest rear‐edge stands. Our modeling approach provides accessible tools to evaluate forest vulnerability to warmer conditions.