Difference in tree growth responses to climate at the upper treeline: Qilian Juniper in the Anyemaqen Mountains

Three ring-width chronologies were developed from Qilian Juniper (Sabina przewalskii Kom.) at the upper treeline along a west-east gradient in the Anyemaqen Mountains.Most chronological statistics,except for mean sensitivity (MS),decreased from west to east.The first principal component (PC1) Ioadings indicated that stands in a similar climate condition were most important to the variability of radial growth.PC2 Ioadings decreased from west to east,suggesting the difference of tree-growth between eastern and western Anyemaqen Mountains.Correlations between standard chronologies and climatic factors revealed different climatic influences on radial growth along a west-east gradient in the study area.Temperature of warm season (July-August) was important to the radial growth at the upper treeline in the whole study area.Precipitation of current May was an important limiting factor of tree growth only in the western (drier) upper treeline,whereas precipitation of current September limited tree growth in the eastern (wetter) upper treeline.Response function analysis results showed that there were regional differences between tree growth and climatic factors in various sampling sites of the whole study area.Temperature and precipitation were the important factors influencing tree growth in western (drier) upper treeline.However,tree growth was greatly limited by temperature at the upper treeline in the middle area,and was more limited by precipitation than temperature in the eastern (wetter) upper treeline.     

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.     

Shrubline but not treeline advance matches climate velocity in montane ecosystems of south‐central Alaska

Tall shrubs and trees are advancing into many tundra and wetland ecosystems but at a rate that often falls short of that predicted due to climate change. For forest, tall shrub, and tundra ecosystems in two pristine mountain ranges of Alaska, we apply a Bayesian, error‐propagated calculation of expected elevational rise (climate velocity), observed rise (biotic velocity), and their difference (biotic inertia). We show a sensitive dependence of climate velocity on lapse rate and derive biotic velocity as a rigid elevational shift. Ecosystem presence identified from recent and historic orthophotos ~50 years apart was regressed on elevation. Biotic velocity was estimated as the difference between critical point elevations of recent and historic logistic fits divided by time between imagery. For both mountain ranges, the 95% highest posterior density of climate velocity enclosed the posterior distributions of all biotic velocities. In the Kenai Mountains, mean tall shrub and climate velocities were both 2.8 m y^?1. In the better sampled Chugach Mountains, mean tundra retreat was 1.2 m y^?1 and climate velocity 1.3 m y^?1. In each mountain range, the posterior mode of tall woody vegetation velocity (the complement of tundra) matched climate velocity better than either forest or tall shrub alone, suggesting competitive compensation can be important. Forest velocity was consistently low at 0.1–1.1 m y^?1, indicating treeline is advancing slowly. We hypothesize that the high biotic inertia of forest ecosystems in south‐central Alaska may be due to competition with tall shrubs and/or more complex climate controls on the elevational limits of trees than tall shrubs. Among tall shrubs, those that disperse farthest had lowest inertia. Finally, the rapid upward advance of woody vegetation may be contributing to regional declines in Dall's sheep (Ovis dalli), a poorly dispersing alpine specialist herbivore with substantial biotic inertia due to dispersal reluctance.     

Among‐tree variability and feedback effects result in different growth responses to climate change at the upper treeline in the Swiss Alps

Upper treeline ecotones are important life form boundaries and particularly sensitive to a warming climate. Changes in growth conditions at these ecotones have wide‐ranging implications for the provision of ecosystem services in densely populated mountain regions like the European Alps. We quantify climate effects on short‐ and long‐term tree growth responses, focusing on among‐tree variability and potential feedback effects. Although among‐tree variability is thought to be substantial, it has not been considered systematically yet in studies on growth–climate relationships. We compiled tree‐ring data including almost 600 trees of major treeline species (Larix decidua, Picea abies, Pinus cembra, and Pinus mugo) from three climate regions of the Swiss Alps. We further acquired tree size distribution data using unmanned aerial vehicles. To account for among‐tree variability, we employed information‐theoretic model selections based on linear mixed‐effects models (LMMs) with flexible choice of monthly temperature effects on growth. We isolated long‐term trends in ring‐width indices (RWI) in interaction with elevation. The LMMs revealed substantial amounts of previously unquantified among‐tree variability, indicating different strategies of single trees regarding when and to what extent to invest assimilates into growth. Furthermore, the LMMs indicated strongly positive temperature effects on growth during short summer periods across all species, and significant contributions of fall (L. decidua) and current year's spring (L. decidua, P. abies). In the longer term, all species showed consistently positive RWI trends at highest elevations, but different patterns with decreasing elevation. L. decidua exhibited even negative RWI trends compared to the highest treeline sites, whereas P. abies, P. cembra, and P. mugo showed steeper or flatter trends with decreasing elevation. This does not only reflect effects of ameliorated climate conditions on tree growth over time, but also reveals first signs of long‐suspected negative and positive feedback of climate change on stand dynamics at treeline.     

Recent invasion of the mountain birch Betula pubescens ssp. tortuosa above the treeline due to climate change: genetic and ecological study in northern Sweden

Mountain birch, Betula pubescens ssp. tortuosa, forms the treeline in northern Sweden. A recent shift in the range of the species associated with an elevation of the treeline is commonly attributed to climate warming. Using microsatellite markers, we explored the genetic structure of populations along an altitudinal gradient close to the treeline. Low genetic differentiation was found between populations, whereas high genetic diversity was maintained within populations. High level of gene flow compensated for possible losses of genetic diversity at higher elevations and dissipated the founding effect of newly established populations above the treeline. Spatial autocorrelation analysis showed low spatial genetic structure within populations because of extensive gene flow. At the treeline, significant genetic structure within the juvenile age class at small distances did not persist in the adult age class, indicating recent expansion of young recruits due to the warming of the climate. Finally, seedling performance above the treeline was positively correlated with parameters related to temperature. These data confirm the high migration potential of the species in response to fluctuating environmental conditions and indicate that it is now invading higher altitudes due to the recent warming of the climate.     

川西高山林线土壤活性碳、氮对短期增温的响应

随着温室效应的加剧,受低温限制的高山林线生态系统对全球气候变暖较为敏感,可能直接影响到植物的生长和土壤碳氮过程.本研究假设气候变暖会改变高山生态系统土壤活性碳氮含量,在四川省理县米亚罗高山生态系统定位站,采用开顶式模拟增温装置(OTC)模拟增温对土壤活性碳、氮的短期影响.分别于2017年4、7和10月,采集OTC以及对照样地(CK)内土壤有机层和矿质土壤层的原状土壤,测定土壤可溶性有机碳(DOC)、土壤微生物生物量碳(MBC)、土壤可溶性有机氮(DON)和土壤微生物生物量氮(MBN)含量.结果表明: 模拟增温使年均气温升高0.88 ℃,土壤有机层和矿质土壤层的年均温度分别提高0.48和0.23 ℃.模拟增温没有显著改变土壤有机质和含水量,但显著提高了矿质土壤层的pH值,同时显著降低了非生长季矿质土壤层的DOC、DON含量;季节变化对两个层次的DOC、DON和MBN含量有极显著影响,而MBC没有明显的季节动态;增温和季节交互作用对矿质土壤层的DOC和DON有显著影响.土壤有机层的MBC、MBN含量显著高于矿质土壤层.土壤活性碳、氮与土壤有机质和含水量呈极显著正相关,MBC、MBN与土壤pH呈极显著正相关,MBN与土壤温度呈显著负相关.     

Temperature‐growth divergence in white spruce forests of Old Crow Flats,Yukon Territory,and adjacent regions of northwestern North America

We present a new 23‐site network of white spruce ring‐width chronologies near boreal treeline in Old Crow Flats, Yukon Territory, Canada. Most chronologies span the last 300 years and some reach the mid‐16th century. The chronologies exhibit coherent growth patterns before the 1930s. However, since the 1930s, they diverge in trend and exhibit one of two contrasting, but well‐replicated patterns we call Group 1 and Group 2. Over the instrumental period (1930–2007) Group 1 sites were inversely correlated with previous‐year July temperatures while Group 2 sites were positively correlated with growth‐year June temperatures. At the broader northwestern North America (NWNA) scale, we find that the Group 1 and Group 2 patterns are common to a number of white spruce chronologies, which we call NWNA 1 and NWNA 2 chronologies. The NWNA 1 and NWNA 2 chronologies also share a single coherent growth pattern prior to their divergence (ca. 1950s). Comparison of the NWNA 1/NWNA 2 chronologies against gridded 20th‐century temperatures for NWNA and reconstructed northern hemisphere summer temperatures (ad 1300–2000) indicates that all sites responded positively to temperature prior to the mid‐20th century (at least back to ad 1300), but that some changed to a negative response (NWNA 1) while others maintained a positive response (NWNA 2). The spatial extent of divergence implies a large‐scale forcing. As the divergence appears to be restricted to the 20th century, we suggest that the temperature response shift represents a moisture stress caused by an anomalously warm, dry 20th‐century climate in NWNA, as indicated by paleoclimatic records. However, because some sites do not diverge and are located within a few kilometres of divergent sites, we speculate that site‐level factors have been important in determining the susceptibility of sites to the large‐scale drivers of divergence.     

Negative impacts of high temperatures on growth of black spruce forests intensify with the anticipated climate warming

An increasing number of studies conclude that water limitations and heat stress may hinder the capacity of black spruce (Picea mariana (Mill.) B.S.P.) trees, a dominant species of Canada's boreal forests, to grow and assimilate atmospheric carbon. However, there is currently no scientific consensus on the future of these forests over the next century in the context of widespread climate warming. The large spatial extent of black spruce forests across the Canadian boreal forest and associated variability in climate, demography, and site conditions pose challenges for projecting future climate change responses. Here we provide an evaluation of the impacts of climate warming and drying, as well as increasing [CO₂], on the aboveground productivity of black spruce forests across Canada south of 60°N for the period 1971 to 2100. We use a new extensive network of tree‐ring data obtained from Canada's National Forest Inventory, spatially explicit simulations of net primary productivity (NPP) and its drivers, and multivariate statistical modeling. We found that soil water availability is a significant driver of black spruce interannual variability in productivity across broad areas of the western to eastern Canadian boreal forest. Interannual variability in productivity was also found to be driven by autotrophic respiration in the warmest regions. In most regions, the impacts of soil water availability and respiration on interannual variability in productivity occurred during the phase of carbohydrate accumulation the year preceding tree‐ring formation. Results from projections suggest an increase in the importance of soil water availability and respiration as limiting factors on NPP over the next century due to warming, but this response may vary to the extent that other factors such as carbon dioxide fertilization, and respiration acclimation to high temperature, contribute to dampening these limitations.     

Stand basal area and solar radiation amplify white spruce climate sensitivity in interior Alaska: Evidence from carbon isotopes and tree rings

The negative growth response of North American boreal forest trees to warm summers is well documented and the constraint of competition on tree growth widely reported, but the potential interaction between climate and competition in the boreal forest is not well studied. Because competition may amplify or mute tree climate‐growth responses, understanding the role current forest structure plays in tree growth responses to climate is critical in assessing and managing future forest productivity in a warming climate. Using white spruce tree ring and carbon isotope data from a long‐term vegetation monitoring program in Denali National Park and Preserve, we investigated the hypotheses that (a) competition and site moisture characteristics mediate white spruce radial growth response to climate and (b) moisture limitation is the mechanism for reduced growth. We further examined the impact of large reproductive events (mast years) on white spruce radial growth and stomatal regulation. We found that competition and site moisture characteristics mediated white spruce climate‐growth response. The negative radial growth response to warm and dry early‐ to mid‐summer and dry late summer conditions intensified in high competition stands and in areas receiving high potential solar radiation. Discrimination against ¹³C was reduced in warm, dry summers and further diminished on south‐facing hillslopes and in high competition stands, but was unaffected by climate in open floodplain stands, supporting the hypothesis that competition for moisture limits growth. Finally, during mast years, we found a shift in current year's carbon resources from radial growth to reproduction, reduced ¹³C discrimination, and increased intrinsic water‐use efficiency. Our findings highlight the importance of temporally variable and confounded factors, such as forest structure and climate, on the observed climate‐growth response of white spruce. Thus, white spruce growth trends and productivity in a warming climate will likely depend on landscape position and current forest structure.     

Expanding forests and changing growth forms of Siberian larch at the Polar Urals treeline during the 20th century

The ongoing climatic changes potentially affect plant growth and the functioning of temperature‐limited high‐altitude and high‐latitude ecosystems; the rate and magnitude of these biotic changes are, however, uncertain. The aim of this study was to reconstruct stand structure and growth forms of Larix sibirica (Ledeb.) in undisturbed forest–tundra ecotones of the remote Polar Urals on a centennial time scale. Comparisons of the current ecotone with historic photographs from the 1960s clearly document that forests have significantly expanded since then. Similarly, the analysis of forest age structure based on more than 300 trees sampled along three altitudinal gradients reaching from forests in the valleys to the tundra indicate that more than 70% of the currently upright‐growing trees are <80 years old. Because thousands of more than 500‐year‐old subfossil trees occur in the same area but tree remnants of the 15–19th century are lacking almost entirely, we conclude that the forest has been expanding upwards into the formerly tree‐free tundra during the last century by about 20–60 m in altitude. This upward shift of forests was accompanied by significant changes in tree growth forms: while 36% of the few trees that are more than 100 years old were multi‐stem tree clusters, 90% of the trees emerging after 1950 were single‐stemmed. Tree‐ring analysis of horizontal and vertical stems of multi‐stemmed larch trees showed that these trees had been growing in a creeping form since the 15th century. In the early 20th century, they started to grow upright with 5–20 stems per tree individual. The incipient vertical growth led to an abrupt tripling in radial growth and thus, in biomass production. Based on above‐ and belowground biomass measurements of 33 trees that were dug out and the mapping of tree height and diameter, we estimated that forest expansion led to a biomass increase by 40–75 t ha^?1 and a carbon accumulation of approximately 20–40 g C m^?2 yr^?1 during the last century. The forest expansion and change in growth forms coincided with significant summer warming by 0.9 °C and a doubling of winter precipitation during the 20th century. In summary, our results indicate that the ongoing climatic changes are already leaving a fingerprint on the appearance, structure, and productivity of the treeline ecotone in the Polar Urals.     

Autumn photosynthetic decline and growth cessation in seedlings of white spruce are decoupled under warming and photoperiod manipulations

Climate warming is expected to increase the seasonal duration of photosynthetic carbon fixation and tree growth in high‐latitude forests. However, photoperiod, a crucial cue for seasonality, will remain constant, which may constrain tree responses to warming. We investigated the effects of temperature and photoperiod on weekly changes in photosynthetic capacity, leaf biochemistry and growth in seedlings of a boreal evergreen conifer, white spruce [Picea glauca (Moench) Voss]. Warming delayed autumn declines in photosynthetic capacity, extending the period when seedlings had high carbon uptake. While photoperiod was correlated with photosynthetic capacity, short photoperiods did not constrain the maintenance of high photosynthetic capacity under warming. Rubisco concentration dynamics were affected by temperature but not photoperiod, while leaf pigment concentrations were unaffected by treatments. Respiration rates at 25 °C were stimulated by photoperiod, although respiration at the growth temperatures was increased in warming treatments. Seedling growth was stimulated by increased photoperiod and suppressed by warming. We demonstrate that temperature is a stronger control on the seasonal timing of photosynthetic down‐regulation than is photoperiod. Thus, while warming can stimulate carbon uptake in boreal conifers, the extra carbon may be directed towards respiration rather than biomass, potentially limiting carbon sequestration under climate change.     

Cytochrome and alternative pathway respiration in white spruce (Picea glauca) roots. Effects of growth and measurement temperature

Interactions between growth temperature and measurement temperature were examined for their effects on white spruce [ Picea glauca (Moench) Voss] root respiration. Total dark respiration rates increased with measurement temperature and were unaffected by growth temperature. Partitioning of respiratory electron flow between the cytochrome and alternative pathways was also unaffected by growth temperature. The proportion of respiration mediated by the alternative pathway was constant at measurement temperatures between 4°C and 18°C, but was increased at higher temperatures. Changes in alternative pathway activity were paralleled by changes in capacity, and the alternative pathway was almost fully engaged at all temperatures. Roots grown at low temperature displayed higher carbohydrate levels than roots grown at higher temperatures, but respiration rate was unaffected. Spruce root respiration did not appear to acclimate to growth temperature, and the alternative pathway was not preferentially engaged at low temperature.     

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.     

Recent climatic drying leads to age‐independent growth reductions of white spruce stands in western Canada

Since 2001, climatic conditions have been notably drier than normal across large areas of the western Canadian interior, leading to widespread impacts on the forests of this region. This poses a major concern for the future, given climate change projections for continued warming and drying. We conducted tree‐ring analysis in 75 pure stands of white spruce (Picea glauca) across Alberta and west‐central Saskatchewan to examine the effects of recent climatic drying on the growth of this important boreal tree species. Allometric equations were used to calculate annual growth in aboveground tree biomass (G_BM) from ring width measurements. Results showed an increasing trend in G_BM from the 1960s to the 1990s, followed by a sharp decline during the severe drought of 2001–2002. Of the 75 stands, only 18 recovered sufficiently to cause an increase in mean G_BM from the predrought decade of 1991–2000 to the subsequent decade of 2001–2010. The remaining 57 stands exhibited a decline in mean G_BM between these decades. Climatic drying was a major cause of the growth decline, as shown by the significant stand‐level relationship between percentage change in decadal mean G_BM and the change in decadal mean values of a climate moisture index from 1991–2000 to 2001–2010. Subsequent analyses of boreal stands sampled across Alberta during 2015 revealed that white spruce growth had declined even further as drought conditions intensified during 2014–2015. Overall, there was a 38% decrease in mean G_BM between 1997 and 2015, but surprisingly, the percentage decrease was not significantly different for young, productive stands compared with older, less productive stands. Thus, stand ageing cannot explain the observed decline in white spruce growth during the past quarter century, suggesting that these forests are at risk if the trend towards more frequent, severe drought continues in the region.     

玉龙雪山不同海拔丽江云杉径向生长对气候变异的响应

为研究滇西北高原树木径向生长与气候关系随海拔的变化规律, 分别在玉龙雪山低、中、高海拔采集丽江云杉(Picea likiangensis)年轮样本, 建立了不同海拔丽江云杉树轮宽度残差年表, 将年轮指数与气候因子进行响应分析、冗余分析以及滑动响应分析。结果表明: 玉龙雪山丽江云杉径向生长受气温和降水共同影响, 但不同海拔径向生长响应模式存在差异。其中当年1-3月降水与不同海拔丽江云杉径向生长均呈显著正相关关系; 当年生长季后期降水与中、低海拔树木生长呈显著负相关关系, 与高海拔树木生长呈显著正相关关系; 中、低海拔树木生长还受当年春季干旱胁迫; 而当年7月气温升高促进高海拔丽江云杉生长。冗余分析与响应分析结果基本一致, 说明冗余分析能够有效量化树轮宽度指数与气候因子的关系。滑动响应分析显示气温和降水在小时间尺度上的变化也会影响树木生长。结合不同海拔丽江云杉生长对气候因子的响应模式及未来气候预测, 玉龙雪山高海拔丽江云杉生长将得到加强, 而中、低海拔丽江云杉生长则表现出不确定性。     

Plant phenological responses to a long‐term experimental extension of growing season and soil warming in the tussock tundra of Alaska

Climate warming is strongly altering the timing of season initiation and season length in the Arctic. Phenological activities are among the most sensitive plant responses to climate change and have important effects at all levels within the ecosystem. We tested the effects of two experimental treatments, extended growing season via snow removal and extended growing season combined with soil warming, on plant phenology in tussock tundra in Alaska from 1995 through 2003. We specifically monitored the responses of eight species, representing four growth forms: (i) graminoids (Carex bigellowii and Eriophorum vaginatum); (ii) evergreen shrubs (Ledum palustre, Cassiope tetragona, and Vaccinium vitis‐idaea); (iii) deciduous shrubs (Betula nana and Salix pulchra); and (iv) forbs (Polygonum bistorta). Our study answered three questions: (i) Do experimental treatments affect the timing of leaf bud break, flowering, and leaf senescence? (ii) Are responses to treatments species‐specific and growth form‐specific? and (iii) Which environmental factors best predict timing of phenophases? Treatment significantly affected the timing of all three phenophases, although the two experimental treatments did not differ from each other. While phenological events began earlier in the experimental plots relative to the controls, duration of phenophases did not increase. The evergreen shrub, Cassiope tetragona, did not respond to either experimental treatment. While the other species did respond to experimental treatments, the total active period for these species did not increase relative to the control. Air temperature was consistently the best predictor of phenology. Our results imply that some evergreen shrubs (i.e., C. tetragona) will not capitalize on earlier favorable growing conditions, putting them at a competitive disadvantage relative to phenotypically plastic deciduous shrubs. Our findings also suggest that an early onset of the growing season as a result of decreased snow cover will not necessarily result in greater tundra productivity.