Variation in carbohydrate source–sink relations of forest and treeline white spruce in southern, interior and northern Alaska

Two opposing hypotheses have been presented to explain reduced tree growth at the treeline, compared with growth in lower elevation or lower latitude forests: the carbon source and sink limitation hypotheses. The former states that treeline trees have an unfavorable carbon balance and cannot support growth of the magnitude observed at lower elevations or latitudes, while the latter argues that treeline trees have an adequate carbon supply, but that cold temperatures directly limit growth. In this study, we examined the relative importance of source and sink limitation in forest and treeline white spruce (Picea glauca) in three mountain ranges from southern to northern Alaska. We related seasonal changes in needle nonstructural carbohydrate (NSC) content with branch extension growth, an approach we argue is more powerful than using needle NSC concentration. Branch extension growth in the southernmost Chugach Mountains was much greater than in the White Mountains and the Brooks Range. Trees in the Chugach Mountains showed a greater seasonal decline in needle NSC content than trees in the other mountain ranges, and the seasonal change in NSC was correlated with site-level branch growth across mountain ranges. There was no evidence of a consistent difference in branch growth between the forest and treeline sites, which differ in elevation by approximately 100 m. Our results point to a continuum between source and sink limitation of growth, with high-elevation trees in northern and interior Alaska showing greater evidence of sink limitation, and those in southern Alaska showing greater potential for source limitation.     

Recent climate warming forces contrasting growth responses of white spruce at treeline in Alaska through temperature thresholds

Northern and high‐latitude alpine treelines are generally thought to be limited by available warmth. Most studies of tree‐growth–climate interaction at treeline as well as climate reconstructions using dendrochronology report positive growth response of treeline trees to warmer temperatures. However, population‐wide responses of treeline trees to climate remain largely unexamined. We systematically sampled 1558 white spruce at 13 treeline sites in the Brooks Range and Alaska Range. Our findings of both positive and negative growth responses to climate warming at treeline challenge the widespread assumption that arctic treeline trees grow better with warming climate. High mean temperatures in July decreased the growth of 40% of white spruce at treeline areas in Alaska, whereas warm springs enhance growth of additional 36% of trees and 24% show no significant correlation with climate. Even though these opposing growth responses are present in all sampled sites, their relative proportion varies between sites and there is no overall clear relationship between growth response and landscape position within a site. Growth increases and decreases appear in our sample above specific temperature index values (temperature thresholds), which occurred more frequently in the late 20th century. Contrary to previous findings, temperature explained more variability in radial growth after 1950. Without accounting for these opposite responses and temperature thresholds, climate reconstructions based on ring width will miscalibrate past climate, and biogeochemical and dynamic vegetation models will overestimate carbon uptake and treeline advance under future warming scenarios.     

Nitrogen and carbon source-sink relationships in trees at the Himalayan treelines compared with lower elevations

No single hypothesis or theory has been widely accepted for explaining the functional mechanism of global alpine/arctic treeline formation. The present study tested whether the alpine treeline is determined by (1) the needle nitrogen content associated with photosynthesis (carbon gain); (2) a sufficient source-sink ratio of carbon; or (3) a sufficient C-N ratio. Nitrogen does not limit the growth and development of trees studied at the Himalayan treelines. Levels of non-structural carbohydrates (NSC) in trees were species-specific and site-dependent; therefore, the treeline cases studied did not show consistent evidence of source/carbon limitation or sink/growth limitation in treeline trees. However, results of the combined three treelines showed that the treeline trees may suffer from a winter carbon shortage. The source capacity and the sink capacity of a tree influence its tissue NSC concentrations and the carbon balance; therefore, we suggest that the persistence and development of treeline trees in a harsh alpine environment may require a minimum level of the total NSC concentration, a sufficiently high sugar:starch ratio, and a balanced carbon source-sink relationship.     

Back to the Future: The Responses of Alpine Treelines to Climate Warming are Constrained by the Current Ecotone Structure

Alpine treeline ecotones are considered early-warning monitors of the effects of climate change on terrestrial ecosystems, but it is still unclear how accurately treeline dynamics may track the expected temperature rises. Site-specific abiotic constraints, such as topography and demographic trends may make treelines less responsive to environmental fluctuations. A better understanding on how local processes modulate treelines’ response to warming is thus required. We developed a model of treeline dynamics based on individual data of growth, mortality and reproduction. Specifically, we modeled growth patterns, mortality rates and reproductive size thresholds as a function of temperature and stand structure to evaluate the influence of climate- and stand-related processes on treeline dynamics. In this study, we analyze the dynamics of four Pyrenean mountain pine treeline sites with contrasting stand structures, and subjected to differing rates of climate warming. Our models indicate that Pyrenean treelines could reach basal areas and reproductive potentials similar to those currently observed in high-elevation subalpine forest by the mid twenty-first century. The fastest paces of treeline densification are forecasted by the late twenty-first century and are associated with higher warming rates. We found a common densification response of Pyrenean treelines to climate warming, but contrasting paces arise due to current size structures. Treelines characterized by a multistratified stand structure and subjected to lower mean annual temperatures were the most responsive to climate warming. In monostratified stands, tree growth was less sensitive to temperature than in multistratified stands and trees reached their reproductive size threshold later. Therefore, our simulations highlight that stand structure is paramount in modulating treeline responsiveness to ongoing climate warming. Synthesis. Treeline densification over the twenty-first century is likely to occur at different rates contingent on current stand structure and its effects on individual-level tree growth responses to warming. Accurate projections of future treeline dynamics must thus incorporate site-specific factors other than climate, specifically those related to stand structure and its influence on tree growth.     

Limited prospects for future alpine treeline advance in the Canadian Rocky Mountains

Treeline advance has occurred throughout the twentieth century in mountainous regions around the world; however, local variation and temporal lags in responses to climate warming indicate that the upper limits of some treelines are not necessarily in climatic equilibrium. These observations suggest that factors other than climate are constraining tree establishment beyond existing treelines. Using a seed addition experiment, we tested the effects of seed availability, predation and microsite limitation on the establishment of two subalpine tree species (Picea engelmannii and Abies lasiocarpa) across four treelines in the Canadian Rocky Mountains. The effect of vegetation removal on seedling growth was also determined, and microclimate conditions were monitored. Establishment limitations observed in the field were placed in context with the effects of soil properties observed in a parallel experiment. The seed addition experiment revealed reduced establishment with increasing elevation, suggesting that although establishment within the treeline ecotone is at least partially seed limited, other constraints are more important beyond the current treeline. The effects of herbivory and microsite availability significantly reduced seedling establishment but were less influential beyond the treeline. Microclimate monitoring revealed that establishment was negatively related to growing season temperatures and positively related to the duration of winter snow cover, counter to the conventional expectation that establishment is limited by low temperatures. Overall, it appears that seedling establishment beyond treeline is predominantly constrained by a combination of high soil surface temperatures during the growing season, reduced winter snowpack and unfavourable soil properties. Our study supports the assertion that seedling establishment in alpine treeline ecotones is simultaneously limited by various climatic and nonclimatic drivers. Together, these factors may limit future treeline advance in the Canadian Rocky Mountains and should be considered when assessing the potential for treeline advance in alpine systems elsewhere     

Are gas exchange responses to resource limitation and defoliation linked to source:sink relationships?

Productivity of trees can be affected by limitations in resources such as water and nutrients, and herbivory. However, there is little understanding of their interactive effects on carbon uptake and growth. We hypothesized that: (1) in the absence of defoliation, photosynthetic rate and leaf respiration would be governed by limiting resource(s) and their impact on sink limitation; (2) photosynthetic responses to defoliation would be a consequence of changing source:sink relationships and increased availability of limiting resources; and (3) photosynthesis and leaf respiration would be adjusted in response to limiting resources and defoliation so that growth could be maintained. We tested these hypotheses by examining how leaf photosynthetic processes, respiration, carbohydrate concentrations and growth rates of Eucalyptus globulus were influenced by high or low water and nitrogen (N) availability, and/or defoliation. Photosynthesis of saplings grown with low water was primarily sink limited, whereas photosynthetic responses of saplings grown with low N were suggestive of source limitation. Defoliation resulted in source limitation. Net photosynthetic responses to defoliation were linked to the degree of resource availability, with the largest responses measured in treatments where saplings were ultimately source rather than sink limited. There was good evidence of acclimation to stress, enabling higher rates of C uptake than might otherwise have occurred.     

Temperature thresholds for the onset of xylogenesis in alpine shrubs on the Tibetan Plateau

Key message

The threshold minimum air temperature driving xylem growth of alpine  Rhododendron aganniphum is lower than that commonly observed at the treeline of conifers.

Abstract

Understanding how alpine shrubs grow and which environmental factors drive their biomass gain could help to functionally differentiate trees and shrubs. The cambium is the main meristem responsible for wood formation in trees and shrubs. Thus, a better knowledge of cambium growth dynamics in alpine shrubs would allow explaining why shrubs displace trees above the treeline. Here, we aim to investigate the timings and dynamics of xylogenesis and to identify the thermal thresholds controlling the onset of xylem growth of Rhododendron aganniphum, a tall shrub growing above the alpine treeline on the Tibetan Plateau. Timings of xylogenesis and radial growth rates were assessed from anatomical observations of the developing xylem during three growing seasons (2011, 2012, and 2013). The threshold temperature at which xylogenesis had a 0.5 probability of being active was calculated with logistic regressions. The onset of xylogenesis was observed between mid and late June, whereas the end of xylogenesis lasted from mid to late September. Overall, the duration of xylem growth lasted 88–101 days, and 94 % of the ring was formed from June to August. The threshold for the onset of xylem growth was observed at 2.0 ± 0.6 °C for the minimum air temperature, lower than that commonly observed for treeline conifers (ca. 6 °C). This low thermal threshold allows alpine shrubs to have a growing season long enough to complete xylem production and maturation during the warmest summer months. Our results suggest that the time required to complete xylogenesis is critical to understand why shrubs displace trees above the treeline.

     

Temporal variations of mobile carbohydrates in Abies fargesii at the upper tree limits

Low temperatures are associated high‐altitude treelines, but the functional mechanism of treeline formation remains controversial. The relative contributions of carbon limitation (source activity) and growth limitation (sink activity) require more tests across taxa and regions. We examined temporal variations of mobile carbon supply in different tissues of Abies fargesii across treeline ecotones on north‐ and south‐facing slopes of the Qinling Mountains, China. Non‐structural carbohydrate (NSC) concentrations in tissues along the altitudinal gradient on both slopes changed significantly in the early and late growing season, but not in the mid‐growing season, indicating the season‐dependent carbon supply status. Late in the growing season on both slopes, trees at the upper limits had the highest NSC concentrations and total soluble sugars and lowest starch concentrations compared to trees at the lower elevations. NSC concentrations tended to increase in needles and branches throughout the growing season with increasing elevation on both slopes, but declined in roots and stems. NSC concentrations across sampling dates also indicated increases in needles and branches, and decreases in roots and stem with increasing elevation. Overall altitudinal trends of NSC in A. fargesii revealed no depletion of mobile carbon reserves at upper elevation limits, suggesting limitation of sink activity dominates tree life across treeline ecotones in both north‐ and south‐facing slopes. Carbon reserves in storage tissues (especially roots) in the late growing season might also play an important role in winter survival and early growth in spring at upper elevations on both slopes, which define the uppermost limit of A. fargesii.     

Climate change and the northern Russian treeline zone

The Russian treeline is a dynamic ecotone typified by steep gradients in summer temperature and regionally variable gradients in albedo and heat flux. The location of the treeline is largely controlled by summer temperatures and growing season length. Temperatures have responded strongly to twentieth-century global warming and will display a magnified response to future warming. Dendroecological studies indicate enhanced conifer recruitment during the twentieth century. However, conifers have not yet recolonized many areas where trees were present during the Medieval Warm period (ca AD 800-1,300) or the Holocene Thermal Maximum (HTM; ca 10,000-3,000 years ago). Reconstruction of tree distributions during the HTM suggests that the future position of the treeline due to global warming may approximate its former Holocene maximum position. An increased dominance of evergreen tree species in the northern Siberian forests may be an important difference between past and future conditions. Based on the slow rates of treeline expansion observed during the twentieth century, the presence of steep climatic gradients associated with the current Arctic coastline and the prevalence of organic soils, it is possible that rates of treeline expansion will be regionally variable and transient forest communities with species abundances different from today's may develop.     

Treeline dynamics in relation to climatic variability in the central Tianshan Mountains, northwestern China

Aim Climate variability may be an important mediating agent of ecosystem dynamics in cold, arid regions such as the central Tianshan Mountains, north‐western China. Tree‐ring chronologies and the age structure of a Schrenk spruce (Picea schrenkiana) forest were developed to examine treeline dynamics in recent decades in relation to climatic variability. Of particular interest was whether tree‐ring growth and population recruitment patterns responded similarly to climate warming. Location The study was conducted in eight stands that ranged from 2500 m to 2750 m a.s.l. near the treeline in the Tianchi Nature Reserve (43°45′?43°59′ N, 88°00′?88°20′ E) in the central Xinjiang Uygur Autonomous Region, northwestern China. Methods Tree‐ring cores were collected and used to develop tree‐ring chronologies. The age of sampled trees was determined from basal cores sampled as close as possible to the ground. Population age structure and recruitment information were obtained using an age–d.b.h. (diameter at breast height) regression from the sampled cores and the d.b.h. measured on all trees in the plots. Ring‐width chronologies and tree age structure were both used to investigate the relationship between treeline dynamics and climate change. Results Comparisons with the climatic records showed that both the radial growth of trees and tree recruitment were influenced positively by temperature and precipitation in the cold high treeline zone, but the patterns of their responses differed. The annual variation in tree rings could be explained largely by the average monthly minimum temperatures during February and August of the current year and by the monthly precipitation of the previous August and January, which had a significant and positive effect on tree radial growth. P. schrenkiana recruitment was influenced mainly by consecutive years of high minimum summer temperatures and high precipitation during spring. Over the last several decades, the treeline did not show an obvious upward shift and new recruitment was rare. Some trees had established at the treeline at least 200 years ago. Recruitment increased until the early 20th century (1910s) but then decreased with poor recruitment over the past several decades (1950–2000). Main conclusions There were strong associations between climatic change and ring‐width patterns, and with recruitments in Schrenk spruce. Average minimum temperatures in February and August, and total precipitation in the previous August and January, had a positive effect on tree‐ring width, and several consecutive years of high minimum summer temperature and spring precipitation was a main factor favouring the establishment of P. schrenkiana following germination within the treeline ecotone. Both dendroclimatology and recruitment analysis were useful and compatible to understand and reconstruct treeline dynamics in the central Tianshan Mountains.     

Warming‐induced upward migration of the alpine treeline in the Changbai Mountains,northeast China

Treeline responses to environmental changes describe an important phenomenon in global change research. Often conflicting results and generally too short observations are, however, still challenging our understanding of climate‐induced treeline dynamics. Here, we use a state‐of‐the‐art dendroecological approach to reconstruct long‐term changes in the position of the alpine treeline in relation to air temperature at two sides in the Changbai Mountains in northeast China. Over the past 160 years, the treeline increased by around 80 m, a process that can be divided into three phases of different rates and drives. The first phase was mainly influenced by vegetation recovery after an eruption of the Tianchi volcano in 1702. The slowly upward shift in the second phase was consistent with the slowly increasing temperature. The last phase coincided with rapid warming since 1985, and shows with 33 m per 1°C, the most intense upward shift. The spatial distribution and age structure of trees beyond the current treeline confirm the latest, warming‐induced upward shift. Our results suggest that the alpine treeline will continue to rise, and that the alpine tundra may disappear if temperatures will increase further. This study not only enhances mechanistic understanding of long‐term treeline dynamics, but also highlights the effects of rising temperatures on high‐elevation vegetation dynamics.     

Tree recruitment at the treeline across the Continental Divide in the Northern Rocky Mountains,USA: the role of spring snow and autumn climate

ABSTRACT

Background: Topoclimate can influence tree establishment within treeline ecotones. Yet much less is known about how regional topography, such as the Continental Divide, Rocky Mountains, mediates the role of climate in governing treeline dynamics.

Aims: To utilise the Continental Divide to test whether contrasts in growing-season moisture regimes to the west (summer-dry) and east (summer-wet) impact the spatio-temporal patterns of tree establishment and rates of treeline advance in the Northern Rocky Mountains.

Methods: We sampled trees at sites on north- and south-facing slopes, west and east of the Continental Divide. We used dendroecological techniques to reconstruct patterns of tree establishment. Age-structure data were quantitatively compared with climate to evaluate possible mechanistic linkages.

Results: Across all sites, 96% of trees established after 1950. There was a treeline advance (range = 39–140 m) accompanied by increases in tree density. Significantly more trees established during wet springs on both sides of the Divide.

Conclusions: Overall, snow duration in spring and autumn temperatures appear to influence patterns of tree recruitment at the treeline. Continued warming will likely amplify the role of autumn climate in regulating tree establishment throughout treeline ecotones in the Northern Rocky Mountains, particularly west of the Divide where summer-dry conditions persist.     

Carbon sink limitation and frost tolerance control performance of the tree <Emphasis Type="Italic">Kageneckia angustifolia</Emphasis> D. Don (Rosaceae) at the treeline in central Chile

The decrease in temperature with increasing elevation may determine the altitudinal tree distribution in different ways: affecting survival through freezing temperatures, by a negative carbon balance produced by lower photosynthetic rates, or by limiting growth activity. Here we assessed the relative importance of these direct and indirect effects of altitudinal decrease in temperature in determining the treeline in central Chile (33°S) dominated by Kageneckia angustifolia. We selected two altitudes (2000 and 2200 m a.s.l.) along the treeline ecotone. At each elevation, leaf non-structural carbohydrates (NSC) and gas exchange parameters were measured on five individuals during the growing season. We also determined the cold resistance of K.␣angustifolia, by measuring temperatures that cause 50% seedling mortality (LT₅₀) and ice nucleation (IN). No differences in net photosynthesis were found between altitudes. Although no differences were detected on NSC concentration on a dry matter basis between 2000 and 2200 m, when NSC concentration was expressed on a leaf area basis, higher contents were found at the higher elevation. Thus, carbon sink limitations may occur at the K. angustifolia’s upper altitudinal limit. For seedlings derived from seeds collected at the 2200 m, LT₅₀ of cold-acclimated and non-acclimated plants were −9.5 and −7 °C, respectively. However, temperatures as low as −10 °C can frequently occur at this altitude during the end of winter. Therefore, low temperature injury of seedlings seems also be involved in the treeline formation in this species. Hence, a confluence of global (carbon sink limitation) and regional (freezing tolerance) mechanisms explains the treeline formation in the Mediterranean-type climate zone of central Chile.     

Variation of mobile carbon reserves in trees at the alpine treeline ecotone is under environmental control

? 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.     

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.     

Dynamic relationships between Picea crassifolia growth and climate at upper treeline in the Qilian Mts., Northeast Tibetan Plateau, China

Knowledge of the spatial pattern and temporal relationships between tree-growth and climatic factors are important not only for the projection of forest growth under varying climate but for dendroclimatology in general. Here, we systematically investigated tree-growth climate relationships of Picea crassifolia at upper treeline in the Qilian Mts., northwestern China. 297 trees from eleven sites, covering a large part of the natural range of this species, show increasing and partly divergent correlations with temperature in the most recent decades. The dominant signal at all sites was a strengthening of negative correlations of annual radial growth with summer temperature. In a subset of trees at six sites, a strengthening positive correlation with summer temperatures existed as well. Wetter and high altitude sites tended to show a higher percentage of trees that are positively correlated with warming temperatures, indicating that some individuals there may take advantage of warmer conditions. Divergent responses between the two sub-populations clustered by their response to climate factor were significantly stronger in the last 30 years compared to earlier time slices. In the same time frame, hydrothermal conditions of the investigation area changed to a drier and warmer combination. Drought conditions, most likely affecting the radial growth of most P. crassifolia, have been intensifying over time and expanding spatially from the middle Qilian Mts. to most of our study area during the last half century. While explanations such as methodological effects due to trend removal or human disturbance at the sampling sites might be able to explain the result at single sites, the spatial and temporal co-occurrence of large scale changes in climate and tree growth suggests a causal link between them.     

青藏高原东缘两处高山树线交错带时空动态及其建群种的生态学特征

热量匮乏是高山树线的主要成因, 在全球变暖趋势下对高山树线及其建群种的生态学过程及特征的研究具有重要意义。该文以青藏高原东缘的折多山和剪子弯山两处高山树线(海拔分别为4 265 m和4 425 m)作为研究对象, 通过设置垂直样带, 同时结合区域温度、降水的长时间序列分析, 探究两处树线的时空动态过程, 并明确了建群种冷杉(Abies spp.)的生态学特征。结果表明: 1)折多山和剪子弯山区域的气温在过去58年均存在显著的上升趋势(分别上升了0.72和0.91 ℃), 而折多山和剪子弯山区域降水均存在微弱的降低趋势。2)折多山的峨眉冷杉(A. fabri)龄级结构呈反J形, 剪子弯山的鳞皮冷杉(A. squamata)龄级结构呈双峰形, 二者种群结构均相对稳定。3)在小尺度上, 种子扩散限制使得两处树线的冷杉聚集分布。在大尺度上, 折多山峨眉冷杉亦呈聚集分布, 而剪子弯山鳞皮冷杉受生长环境以及种内或种间关系的影响呈随机分布。4)两处样地建群树种的树高和基径均随海拔升高而降低, 位于树线交错带上部的冷杉均呈现树高生长大于径向生长的异速生长关系, 而位于样地中、下部位的冷杉大部分呈等速生长关系。5)相比10年前, 折多山和剪子弯山的树线及树种线位置均无明显变化, 剪子弯山鳞皮冷杉种群的树木密度亦无明显变化, 而折多山的树木个体数提高了约25%; 相比20年前, 折多山和剪子弯山的树种线分别上移了50和30 m, 树线位置分别升高了75和40 m, 树木个体数亦明显增加, 分别提高了约220%和100%。树线及其建群种在较大时空尺度上主要受热量的控制, 而在较小时空尺度上受温度及生长环境共同作用的影响。     

Responses of white spruce (Picea glauca) to experimental warming at a subarctic alpine treeline

From 2001 to 2004 we experimentally warmed 40 large, naturally established, white spruce [Picea glauca (Moench) Voss] seedlings at alpine treeline in southwest Yukon, Canada, using passive open‐top chambers (OTCs) distributed equally between opposing north and south‐facing slopes. Our goal was to test the hypothesis that an increase in temperature consistent with global climate warming would elicit a positive growth response. OTCs increased growing season air temperatures by 1.8°C and annual growing degree‐days by one‐third. In response, warmed seedlings grew significantly taller and had higher photosynthetic rates compared with control seedlings. On the south aspect, soil temperatures averaged 1.0°C warmer and the snow‐free period was nearly 1 month longer. These seedlings grew longer branches and wider annual rings than seedlings on the north aspect, but had reduced Photosystem‐II efficiency and experienced higher winter needle mortality. The presence of OTCs tended to reduce winter dieback over the course of the experiment. These results indicate that climate warming will enhance vertical growth rates of young conifers, with implications for future changes to the structure and elevation of treeline contingent upon exposure‐related differences. Our results suggest that the growth of seedlings on north‐facing slopes is limited by low soil temperature in the presence of permafrost, while growth on south‐facing slopes appears limited by winter desiccation and cold‐induced photoinhibition.     

Spatio-temporal dynamics of two alpine treeline ecotones and ecological characteristics of their dominate species at the eastern margin of Qinghai-Xizang Plateau

热量匮乏是高山树线的主要成因, 在全球变暖趋势下对高山树线及其建群种的生态学过程及特征的研究具有重要意义。该文以青藏高原东缘的折多山和剪子弯山两处高山树线(海拔分别为4 265 m和4 425 m)作为研究对象, 通过设置垂直样带, 同时结合区域温度、降水的长时间序列分析, 探究两处树线的时空动态过程, 并明确了建群种冷杉(Abies spp.)的生态学特征。结果表明: 1)折多山和剪子弯山区域的气温在过去58年均存在显著的上升趋势(分别上升了0.72和0.91 ℃), 而折多山和剪子弯山区域降水均存在微弱的降低趋势。2)折多山的峨眉冷杉(A. fabri)龄级结构呈反J形, 剪子弯山的鳞皮冷杉(A. squamata)龄级结构呈双峰形, 二者种群结构均相对稳定。3)在小尺度上, 种子扩散限制使得两处树线的冷杉聚集分布。在大尺度上, 折多山峨眉冷杉亦呈聚集分布, 而剪子弯山鳞皮冷杉受生长环境以及种内或种间关系的影响呈随机分布。4)两处样地建群树种的树高和基径均随海拔升高而降低, 位于树线交错带上部的冷杉均呈现树高生长大于径向生长的异速生长关系, 而位于样地中、下部位的冷杉大部分呈等速生长关系。5)相比10年前, 折多山和剪子弯山的树线及树种线位置均无明显变化, 剪子弯山鳞皮冷杉种群的树木密度亦无明显变化, 而折多山的树木个体数提高了约25%; 相比20年前, 折多山和剪子弯山的树种线分别上移了50和30 m, 树线位置分别升高了75和40 m, 树木个体数亦明显增加, 分别提高了约220%和100%。树线及其建群种在较大时空尺度上主要受热量的控制, 而在较小时空尺度上受温度及生长环境共同作用的影响。